Ancient Tethyan Vicariance and Long-Distance Dispersal Drive Global Diversification and Cryptic Speciation in the Red Seaweed Pterocladiella.

We investigated the globally distributed red algal genus Pterocladiella, comprising 24 described species, many of which are economically important sources of agar and agarose. We used DNA-based species delimitation approaches, phylogenetic, and historical biogeographical analyses to uncover cryptic diversity and infer the drivers of biogeographic patterns. We delimited 43 species in Pterocladiella, of which 19 are undescribed. Our multigene time-calibrated phylogeny and ancestral area reconstruction indicated that Pterocladiella most likely originated during the Early Cretaceous in the Tethys Sea. Ancient Tethyan vicariance and long-distance dispersal have shaped current distribution patterns. The ancestor of Eastern Pacific species likely arose before the formation of the formidable Eastern Pacific Barrier-a first confirmation using molecular data in red algae. Divergences of Northeast and Southeast Pacific species have been driven by the Central American Seaway barrier, which, paradoxically, served as a dispersal pathway for Atlantic species. Both long- and short-distance dispersal scenarios are supported by genetic relationships within cosmopolitan species based on haplotype analysis. Asymmetrical distributions and the predominance of peripatry and sympatry between sister species suggest the importance of budding speciation in Pterocladiella. Our study highlights the underestimation of global diversity in these crucial components of coastal ecosystems and provides evidence for the complex evolution of current species distributions.

Molecular phylogeny of the Siphonocladales (Chlorophyta: Cladophorophyceae).

The Siphonocladales are tropical to warm-temperate, marine green macro-algae characterized by a wide variety of thallus morphologies, ranging from branched filaments to pseudo-parenchymatous plants. Phylogenetic analyses of partial large subunit (LSU) rDNA sequences sampled from 166 isolates revealed nine well-supported siphonocladalean clades. Analyses of a concatenated dataset of small subunit (SSU) and partial LSU rDNA sequences greatly clarified the phylogeny of the Siphonocladales. However, the position of the root of the Siphonocladales could not be determined unambiguously, as outgroup rooting and molecular clock rooting resulted in a different root placement. Different phylogenetic methods (likelihood, parsimony and distance) yielded similar tree topologies with comparable internal node resolution. Likewise, analyses under more realistic models of sequence evolution, taking into account differences in evolution between stem and loop regions of rRNA, did not differ markedly from analyses using standard four-state models. The molecular phylogeny revealed that all siphonocladalean architectures may be derived from a single Cladophora-like ancestor. Parallel and convergent evolution of various morphological characters (including those traditionally employed to circumscribe the families and genera) have occurred in the Siphonocladales. Consequently, incongruence with traditional classifications, including non-monophyly in all families and most genera, was shown.

Species boundaries and phylogenetic relationships within the green algal genus Codium (Bryopsidales) based on plastid DNA sequences.

Despite the potential model role of the green algal genus Codium for studies of marine speciation and evolution, there have been difficulties with species delimitation and a molecular phylogenetic framework was lacking. In the present study, 74 evolutionarily significant units (ESUs) are delimited using 227 rbcL exon 1 sequences obtained from specimens collected throughout the genus' range. Several morpho-species were shown to be poorly defined, with some clearly in need of lumping and others containing pseudo-cryptic diversity. A phylogenetic hypothesis of 72 Codium ESUs is inferred from rbcL exon 1 and rps3-rpl16 sequence data using a conventional nucleotide substitution model (GTR+Gamma+I), a codon position model and a covariotide (covarion) model, and the fit of a multitude of substitution models and alignment partitioning strategies to the sequence data is reported. Molecular clock tree rooting was carried out because outgroup rooting was probably affected by phylogenetic bias. Several aspects of the evolution of morphological features of Codium are discussed and the inferred phylogenetic hypothesis is used as a framework to study the biogeography of the genus, both at a global scale and within the Indian Ocean.

Evolution and phylogeography of Halimeda section Halimeda (Bryopsidales, Chlorophyta).

Nuclear ribosomal and plastid DNA sequences of specimens belonging to section Halimeda of the pantropical green seaweed genus Halimeda show that the group under scrutiny contains many more genetically delineable species than those recognized by classical taxonomy. Discordances between phylograms inferred from nuclear and plastid DNA sequences suggest that reticulate evolution has been involved in speciation within the clade. Nonetheless, our data do not allow ruling out certain alternative explanations for the discordances. Several pseudo-cryptic species are restricted to the margins of the generic distribution range. In a clade of H. cuneata sibling species from widely separated subtropical localities in the Indian Ocean, the South African sibling branches off first, leaving the Arabian and West Australian species as closest relatives. We hypothesize that geographic isolation of the siblings may have taken place following Pleistocene or Pliocene periods of climatic cooling during which subtropical species occupied larger distribution ranges. A more basal separation of Atlantic, Indo-Pacific, and Mediterranean species indicates vicariance. The alternative events that could have caused this vicariance are discussed.

Molecular systematics, historical ecology, and phylogeography of Halimeda (Bryopsidales).

Halimeda (Bryopsidales), a genus of calcified, segmented green seaweeds, abounds in reefs and lagoons throughout the tropics. To investigate phylogenetic, phylogeographic, and historic ecological relationships of the genus, the nuclear rDNA including the SSU and both ITS regions were sequenced. A maximum likelihood tree revealed the following: (1) there were anatomical and morphological synapomorphies for five well-supported lineages; (2) the last common ancestor of one lineage invaded sandy substrata; those of two other lineages established in wave-affected habitats, whereas the cenancestor of the remaining two lineages occupied sheltered rocky slopes. Yet, several species adapted to new habitats subsequently, resulting in several cases of convergence; (3) all lineages separated into Atlantic and Indo-Pacific daughters, likely resulting from the rise of the Panamanian Isthmus. Each daughter pair gave rise to additional convergent species in similar habitats in different oceans; (4) Halimeda opuntia, the only monophyletic pantropical species detected so far, dispersed from the Indo-Pacific into the Atlantic well after the closure event; (5) minor SSU-sequence differences across species and phylogeographic patterns of vicariance indicated a relatively recent diversification of the extant diversity. Cretaceous and Early Tertiary fossil look-alikes of modern species must then have resulted from iterative convergence.