William J. Woelkerling

CHOREONEMA (CORALLINALES, RHODOPHYTA): 18S rDNA PHYLOGENY AND RESURRECTION OF THE HAPALIDIACEAE FOR THE SUBFAMILIES CHOREONEMATOIDEAE, AUSTROLITHOIDEAE, AND MELOBESIOIDEAE

Phylogenetic analyses of 18S rDNA gene data for Choreonema thuretii (Corallinales, Rhodophyta) and available data for other coralline red algae indicated that Choreonema belongs to the same lineage as other taxa of Corallinales possessing tetra/bisporangial conceptacles with multiporate plates. These results, when integrated with extant morphological/anatomical data, ultrastructural data, and taxonomic data led to the conclusion that all taxa of Corallinales possessing multiporate conceptacles belong to a distinct family, the Hapalidiaceae. Recognition of the Hapalidiaceae as a distinct family was supported both phylogenetically and phenetically. The Hapalidiaceae includes those taxa of Corallinales whose tetrasporangia produce zonately arranged spores and whose tetra/bisporangia are borne in conceptacles, produce apical plugs, and develop beneath multiporate plates. The Hapalidiaceae includes the subfamilies Choreonematoideae, Melobesioideae, and Austrolithoideae, formerly placed in the Corallinaceae sensu lato. The Choreonematoideae lack cell connections between adjacent vegetative filaments and have a multiporate plate that is acellular at maturity, consisting only of a calcium carbonate matrix. The Austrolithoideae and Melobesioideae both have cellular pore plates; taxa of Melobesioideae have cell fusions between cells of adjacent vegetative filaments, whereas taxa of Austrolithoideae lack cellular connections between adjacent vegetative filaments. Inclusion of the Austrolithoideae in the Hapalidiaceae was based entirely on morphological/anatomical evidence; molecular evidence currently is lacking. Relevant historical and nomenclatural data are included.

Utility of psbA and nSSU for phylogenetic reconstruction in the Corallinales based on New Zealand taxa

A number of molecular studies of the Corallinales, a calcified order of the red algae, have used the conservative nSSU gene to investigate relationships within the order. However interspecific variation at this locus is low for closely related species, limiting resolution of recently diverged groups. In this study, we obtained psbA sequence data from specimens of the order from New Zealand that had been identified according to current taxonomic criteria. We compared phylogenetic analyses based on psbA with those based on nSSU for the same dataset, and also analysed nSSU sequences of the New Zealand material with nSSU sequences of Corallinales taxa from other parts of the world. Our study shows that psbA has considerable potential as a marker for this group, being easily amplified and considerably more variable than nSSU. Combined analyses using both markers provide significant support for relationships at both distal and terminal nodes of the analysis. Our analysis supports the monophyly of all three families currently defined in Corallinales: the Sporolithaceae, Hapalidiaceae and Corallinaceae, and indicates cryptic speciation in Mesophyllum and Spongites.

Rhodolith beds (Corallinales, Rhodophyta) and their physical and biological environment at 80°31′N in Nordkappbukta (Nordaustlandet, Svalbard Archipelago, Norway)

Teichert S., Woelkerling W., Rüggeberg A., Wisshak M., Piepenburg D., Meyerhöfer M., Form A., Büdenbender J. and Freiwald A. 2012. Rhodolith beds (Corallinales, Rhodophyta) and their physical and biological environment at 80°31′N in Nordkappbukta (Nordaustlandet, Svalbard Archipelago, Norway). Phycologia 51: 371–390. DOI: 10.2216/11-76.1 Polar coralline red algae (Corallinales, Rhodophyta) that form rhodoliths have received little attention concerning their potential as ecosystem engineers and carbonate factories; although, recent findings revealed that they are much more widespread in polar waters than previously thought. The present study deals with the northernmost rhodolith communities currently known, discovered in 2006 at 80°31′N in Nordkappbukta (North Cape Bay) at Nordaustlandet, Svalbard. These perennial coralline algae must be adapted to extreme seasonality in terms of light regime (c. 4 months winter darkness), sea ice coverage, nutrient supply, turbidity of the water column, temperature and salinity. The rhodolith communities and their environment were investigated using multibeam swath bathymetry, CTD measurements, recordings of the photosynthetic active radiation (PAR) and determination of the water chemistry, seabed imaging and targeted sampling by means of the manned submersible JAGO as well as benthic collections with a dredge. The coralline flora was composed mainly of Lithothamnion glaciale, with a lesser amount of Phymatolithon tenue. Based on their distribution and development at different depth levels, a facies model was developed. Rhodoliths occurred between 30 and 51 m, while coralline algae attached to cobbles were present as deep as 78 m. Measurements of the PAR indicated their adaptation to extreme low light levels. Ambient waters were always saturated with reference to calcite and aragonite for the whole area. The rhodolith-associated macrobenthic fauna samples yielded 59 species, only one of which was typically Arctic, and the concomitant appearance of corallines and grazers kept the corallines free from epiphytes and coequally provided feeding grounds for the grazers. Overall, L. glaciale and P. tenue appeared to be well adapted to the extreme environment of the Arctic.

Paulsilvella huveorum gen. & sp. nov. (Corallinaceae, Rhodophyta) from the Holocene of Somalia and Kenya, with a reassessment of Lithothrix antiqua from the Late Pleistocene of Mauritius‡

Abstract A morphological, anatomical and taxonomic account of Paulsilvella Woelkeriing, Sartoni & Boddi gen. nov. (Corallinales, Rhodophyta) is provided. Paulsilvella belongs to the Corallinaceae, subfamily Lithophylloideae sensu lato (encompassing the Amphiroideae) and differs from other lithophylloid genera (Amphiroa, Ezo, the Lithophyllum-Titanoderma complex, Lithothrix, Tenarea) in possessing nongeniculate protuberant branches that resemble a series of fused beads. The branches are entirely monomerous, with a core region composed of single arching tiers of elongate, columnar cells alternating with two tiers of short, noncolumnar cells. Paulsilvella includes one known extant species, P. huveorum Woelkerling, Sartoni & Boddi sp. nov. from Somalia and Kenya, and one known fossil species, P. antiqua (G.F. Elliott) Woelkerling, Sartoni & Boddi comb. nov. from the Late Pleistocene of Mauritius. The latter was originally described as Lithothrix antiqua (‘antiquum’). In P. huveorum, the uniporate carposporangial and tetrasporangial conceptacle chambers are largely buried within the thallus, whereas in P. antiqua they protrude above the surrounding thallus surface. A key to species is provided, and generic relationships within the Lithophylloideae are discussed.

Recognition of Pachyarthron and Bossiella as Distinct Genera in the Corallinaceae, Subfamily Corallinoideae (Corallinales, Rhodophyta)

W.J. Woelkerling, A.J.K. Millar, A. Harvey and M. Baba. 2008. Recognition of Pachyarthron and Bossiella as distinct genera in the Corallinaceae, subfamily Corallinoideae (Corallinales, Rhodophyta). Phycologia 47: 265–293. DOI: 10.2216/07-85.1 New evidence supports recognition of both Pachyarthron and Bossiella as distinct genera within the Corallinaceae, subfamily Corallinoideae (Corallinales, Rhodophyta). Pachyarthron cretaceum, the type species of Pachyarthron, is confirmed to occur in Australia, and a new study of the designated lectotype of P. cretacea and of plants from Australia, Japan, Canada (British Columbia) and the United States (Alaska, Washington State) has shown that male conceptacles are produced both axially at branch tips and laterally on intergenicula. By contrast, only lateral conceptacles occur in Bossiella. Differences between Pachyarythron and Serraticardia & Marginisporum, the only genera of Corallinoideae known to produce both axial and lateral conceptacles, are outlined, and a comparison of and key to all genera of Corallinoideae are included. An emended generic description of Pachyarthron and an account of P. cretaceum are provided. Brief accounts of four infraspecific taxa never transferred into Pachyarthron but attributed to the homotypic synonym Amphiroa cretaceum also are included. These are Amphiroa cretacea f. breviarticulata (Areschoug) Yendo from the Caribbean; A. cretacea f. capensis (Areschoug) Yendo from South Africa; A. cretacea f. rosariformis Yendo from Japan; and A cretacea f. tasmanica (Sonder) Yendo from Australia. The biogeographic implications for Pachyarthron and Bossiella are reviewed; Bossiella should be removed from the floras of Japan, Korea and China, whereas Pachyarthron should be added.

Typification and reassessment of seventeen species of coralline red algae (Corallinales and Sporolithales, Rhodophyta) described by W. Ishijima during 1954–1978

Type specimens of 17 species of non-geniculate coralline red algae belonging to the Corallinales and Sporolithales (Corallinophycidae, Rhodophyta) and established between 1954 and 1978 by Wataru Ishijima have been re-examined, focusing on characters relevant to modern coralline taxonomy. The species are from the Early Cretaceous, Eocene, Oligocene, Miocene and Pleistocene rocks of southern and central Japan, Indonesia, Malaysia, Indian Ocean, Philippines, Pakistan and the Galapagos Archipelago, and were attributed by Ishijima to Lithothamnion (as Lithothamnium), Mesophyllum, Lithophyllum and Archaeolithothamnion (as Archaeolithothamnium). With two exceptions, all species are reassigned to different genera or are of uncertain generic placement within particular subfamilies, families or orders. The type specimen of Lithophyllum oboraensis is retained in Lithophyllum and the type specimen of Mesophyllum izuensis is retained in Mesophyllum. The type specimens of Lithophyllum izuensis (Ishijima) comb nov. (= Lithothamnion izuensis), Phymatolithon tokumanensis (Ishijima) comb. nov. (= Mesophyllum tokumanensis), Sporolithon borneoensis (Ishijima) comb. nov. (= Archaeolithothamnium borneoense) and Sporolithon galapagosensis (Ishijima) comb. nov. (= Archaeolithothamnion galapagosensis) possess features justifying their assignment to various genera within a particular family and subfamily. A specimen ascribed by Ishijima to Archaeolithothamnion galapagosensis (as Archaeolithothamnium) in the protologue of that species is a misidentified specimen of Sporolithon ptychoides Heydrich. The type specimens of Lithophyllum minoensis and Mesophyllum indicum belong to the Corallinaceae, subfamily Mastophoroideae but are of uncertain generic placement. The type specimens of Lithothamnion makinogoensis, L. nodai, L. tenuicrustatum, Mesophyllum niinoi and M. yuyashimaensis belong to the Hapalidiaceae (Melobesioideae) but are of uncertain generic placement. The type specimens of Lithophyllum hashimotoi, Lithophyllum minae, Lithothamnion huseinii and Lithothamnion tokiensis lacked sufficient data to allow for order, family, subfamily or genus placement within the Corallinophycidae. Lectotype specimens have been designated as necessary, and relevant historical, repository, geological, morphological and other data are presented.

The Genus Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the Temperate Coasts of the Australian Continent, Including the Newly Described A. klochkovana

Harvey A.S., Woelkerling W.J. and Millar A.J.K. 2009. The genus Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the temperate coasts of the Australian continent, including the newly described A. klochkovana. Phycologia 48: 258–290. DOI: 10.2216/08-84.1 Studies of Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the temperate coasts of Australia provide new evidence that differences in tetrasporangial conceptacle pore canal anatomy are diagnostically significant in delimiting species within the genus. Differences in overall morphology and genicular anatomy are also reliable for delimiting species. These data are supported by examination of relevant type specimens. Four species occur in temperate Australian waters. Three (Amphiroa anceps, Amphira beauvoisii, and the newly described Amphiroa klochkovana) occur in southeastern Australia, and three (A. anceps, A. beauvoisii, and Amphiroa gracilis) occur in southern and southwestern Australia. Comparisons of A. beauvoisii and A. anceps have shown that they cannot be separated at species level morphologically but clearly differ in tetrasporangial conceptacle pore canal anatomy. This has important flow-on implications concerning specimen identification, reported geogeographic distribution and putative heterotypic synonymy of the two species. Relevant historical data, a species key and a synoptic description of Amphiroa also are included.

ANALYSIS OF ANCIENT DNA FROM FOSSIL CORALLINES (CORALLINALES, RHODOPHYTA)1

The field of molecular paleontology has recently made significant contributions to anthropology and biology. Hundreds of ancient DNA studies have been published, but none has targeted fossil coralline algae. Using regions of the SSU gene, we analyzed rDNA from fossil coralline algae of varying ages and states of preservation from Spain, Papua New Guinea (PNG), and the Great Barrier Reef (GBR). Specimens from PNG, GBR, and some localities from Spain did not contain endogenous ancient DNA. Reproducible sequence data were obtained from specimens ∼550 years old from near Cadiz, Spain, and from rocky‐shore deposits in Carboneras, Almeria Province of Spain (∼78,000 years before present [YBP]). Based on BLAST searches and a phylogenetic analysis of sequences, an undescribed coralline alga belonging to the Melobesioideae was discovered in the Carboneras material as well as the following coralline genera: Jania, Lithophyllum, Lithothamnion, Mesophyllum, and Phymatolithon. DNA from fleshy brown and red macroalgae was also discovered in the specimens from Carboneras. The coralline algae identified using molecular techniques were in agreement with those based on morphological methods. The identified taxa are common in the present‐day southeastern Spain littoral zone. Amino acid racemization, concentration ratios, and specific concentrations failed to show a correlation between biomolecular preservation and PCR amplification success. Results suggest that molecular investigations on fossil algae, although limited by technical difficulties, are feasible. Validity of our results was established using authentication criteria and a self‐critical approach to compliance.

Hydrolithon braganum sp. nov. (Corallinaceae, Rhodophyta), the first known exclusively fossil semi-endophytic coralline red alga

Woelkerling W.J., Bassi D. and Iryu Y. 2012. Hydrolithon braganum sp. nov. (Corallinaceae, Rhodophyta), the first known exclusively fossil semi-endophytic coralline red alga. Phycologia 51: 604–611. DOI: 10.2216/11-119.1 Hydrolithon braganum sp. nov., the first known exclusively fossil semi-endophytic species of Corallinaceae (Corallinales, Rhodophyta), is based on material from Integrated Ocean Drilling Program Expedition 310 cores of last deglacial age [c. 20,000–10,000 years before present] from reef terraces around Tahiti, French Polynesia. A detailed morphological–anatomical account is provided and morphological–anatomical similarities to other species are discussed. The taxonomic position of H. braganum is problematic in relation to recent proposals for changes to the circumscription of Hydrolithon. Evidence is presented to show that the morphological–anatomical criteria proposed in recent molecular-based studies are untenable for circumscribing genera called Hydrolithon and Porolithon. Thus, pending further research, the two genera are best treated as the Hydrolithon–Porolithon complex with Hydrolithon having nomenclatural priority, being polyphyletic, and encompassing one or several cryptic genera distinguishable on presently known molecular criteria but not distinguishable using presently known morphological–anatomical criteria.

The genus Mesophyllum (Hapalidiaceae, Corallinales, Rhodophyta) : typification update

W.J. Woelkerling and L.M. Irvine. 2007. The genus Mesophyllum (Hapalidiaceae, Corallinales, Rhodophyta): typification update. Phycologia 46: 230–231. DOI: 10.2216/06-71.1 The 1986 neotypification of Mesophyllum lichenoides (J. Ellis) Me. Lemoine, type species of Mesophyllum (Hapalidiaceae, Corallinales, Rhodophyta), has to be superseded because the International Code of Botanical Nomenclature rules governing the selection of types have changed, and the new rules are retroactive. In accordance with the present rules, an illustration in the original account of the species is designated here as lectotype, and the superseded neotype is designated here as epitype. Relevant historical information and nomenclatural details associated with these changes are presented.