MOLECULAR SYSTEMATICS OF THE XANTHOPHYCEAE

The Corallinales includes ca. 40 genera of calcified red seaweeds. Species are of two distinct morphotypes; those that possess genicula (uncalcified nodes) and those that lack genicula. Most nongeniculate species take the form of crusts. The presence (or absence) of genicula, secondary pit connections, and tetrasporangial conceptacle features have traditionally been used as key characters for delimiting coralline subfamilies. In this study, nuclear encoded 18S and 26S r RNA gene sequences were determined and used to reexamine relationships among coralline taxa. Separate and combined phylogenetic analyses of these data yielded similar trees in which four major lineages are resolved. Heydrichia and Sporolithon (Sporolithaceae) are positioned at the base of the tree and appear to be distantly related to other species examined. Within the Corallinaceae, the nongeniculate Melobesioideae is resolved as a monophyletic group. All members of this subfamily produce mutiporate tetraspoangial conceptacles. The Corallinoideae, which are characterized by unizonate genicula, are resolved as sister to a clade containing species placed in the Lithophylloideae, Mastophoroideae and Metagoniolithoideae. The molecular data indicate that geniculate and nongeniculate species characterized by the presence of secondary pit connections are closely related. For example, both data sets robustly support a sister taxon relationship between Amphiroa and Titanoderma. Our results indicate that: 1) all taxa in which secondary pit connections are present should be referred to the Lithophylloideae and, 2) genicula are nonhomologous structures that are independently derived in Amphiroa, Lithothrix, Metagoniolithon and the last common ancestor of the Corallinoideae.     

A PHYLOGENETIC STUDY OF THE CORALLINALES (RHODOPHYTA) BASED ON NUCLEAR SMALL-SUBUNIT rRNA GENE SEQUENCES

Conflicting classifications for the Corallinales were tested by analyzing partial sequences for the nuclear small-subunit ribosomal RNA (SSU) gene of 35 species of coralline algae. Parsimony and likelihood analyses of these data yielded congruent hypotheses that are inconsistent with classifications for the group that include as many as eight subfamilies. Four major clades are resolved within the order, including the early-diverging Sporolithaceae as well as the Melobesioideae and Corallinoideae. The fourth clade, which is supported robustly, includes both nongeniculate and geniculate species classified in the subfamilies Mastophoroideae, Metagoniolithoideae, Lithophylloideae, and Amphiroideae. Molecular and morphological data support the proposal that the latter two subfamilies are sister taxa. Although relationships among some genera are not resolved clearly, the order of branching of taxa among and within the four principal lineages is concordant with paleontological evidence for the group. Relationships inferred among genera within each of the clades is discussed. Seven morphological characters delimiting higher taxonomic groups within the order were combined with the sequence data, analyzed, and optimized onto the resulting tree(s). Except for the presence or absence of genicula, all other characters were found to be phylogenetically informative. Genicula are nonhomologous structures that evolved independently in the Amphiroideae, Corallinoideae, and Metagoniolithoideae. The phenetic practice of separating coralline algae into two categories solely on the basis of the presence or absence of genicula does not accurately reflect the evolutionary history of the group.     

Crusticorallina gen. nov., a nongeniculate genus in the subfamily Corallinoideae (Corallinales,Rhodophyta)

Molecular phylogenetic analyses of 18S rDNA (SSU) gene sequences confirm the placement of Crusticorallina gen. nov. in Corallinoideae, the first nongeniculate genus in an otherwise geniculate subfamily. Crusticorallina is distinguished from all other coralline genera by the following suite of morpho‐anatomical characters: (i) sunken, uniporate gametangial and bi/tetrasporangial conceptacles, (ii) cells linked by cell fusions, not secondary pit connections, (iii) an epithallus of 1 or 2 cell layers, (iv) a hypothallus that occupies 50% or more of the total thallus thickness, (v) elongate meristematic cells, and (vi) trichocytes absent. Four species are recognized based on rbcL, psbA and COI‐5P sequences, C. painei sp. nov., the generitype, C. adhaerens sp. nov., C. nootkana sp. nov. and C. muricata comb. nov., previously known as Pseudolithophyllum muricatum. Type material of Lithophyllum muricatum, basionym of C. muricata, in TRH comprises at least two taxa, and therefore we accept the previously designated lectotype specimen in UC that we sequenced to confirm its identity. Crusticorallina species are very difficult to distinguish using morpho‐anatomical and/or habitat characters, although at specific sites, some species may be distinguished by a combination of morpho‐anatomy, habitat and biogeography. The Northeast Pacific now boasts six coralline endemic genera, far more than any other region of the world.     

The occurrence of Ezo epiyessoense Adey,Masaki & Akioka (Rhodophyta,Corallinaceae) in England with a summary of parasitism and endophytism in nongeniculate Corallinaceae

The parasitic, nongeniculate, coralline red alga Ezo epiyessoense (Rhodophyta, Corallinaceae) was described in 1974 by Adey, Masaki & Akioka on the basis of specimens growing on Lithophyllum yessoense in Japan. The authors considered Ezo to be an adelphoparasite because it resembled its host taxonomically in being a member of the coralline subfamily Lithophylloideae. The species had not been recorded outside Japan until the present observation in England where it was found growing on another lithophylloid species, Titanoderma pustulatum. The structure of the English material of E. epiyessoense is described and shown to closely resemble that of the type material despite its occurrence on a different host species. Tetrasporangia and trisporangia are recorded for the first time in Ezo. A summary is given of known nongeniculate coralline parasites, semi-endophytes and outgrowths.     

Lithophyllum cuneatum sp. nov. (Corallinaceae, Rhodophyta), a new species of non-geniculate coralline alga semi-endophytic in Hydrolithon onkodes and Neogoniolithon sp. from Fiji, South Pacific

A new species of semi-endophytic coralline alga, Lithophyllum cuneatum (Corallinaceae: Lithophylloideae), is described from Fiji. The species is characterized by a wedge-like thallus that is partially buried in the thallus of the host coralline, Hydrolithon onkodes (Heydrich) Penrose et Woelkerling or occasionally Neogoniolithon sp., and that appears at the surface of the host as a small pustule that is usually paler in color than the host. The thallus consists of erect filaments that are derived from a single cell. The basal cell, when visible, is non-palisade, and areas of bistratose margin are absent. Cells of contiguous erect filaments are joined by secondary pit connections. Epithallial cells are present in 2–3 layers, and individual trichocytes are common. Gametangial plants are dioecious. Male conceptacles have simple spermatangial systems that are confined to the floors of their elliptical chambers. Carposporangial conceptacles contain 5–8 celled gonimoblast filaments that are borne at the margin of a more-or-less discoid fusion cell, and so occupy the periphery of the elliptical conceptacle chambers. Tetrasporangial conceptacles are uniporate, with roofs formed from peripheral filaments, and chambers lack a central columella of sterile filaments. Despite its semi-endophytic nature, haustorial cells are absent, and plastids and pigmentation are present.     

PARASITIC INTERACTIONS AND VEGETATIVE ULTRASTRUCTURE OF CHOREOAEMA THURETII (CORALLINALES,RHODOPHYTA)1

The monotypic coralline red alga, Choreonema thuretii (Bornet) Schmitz (Choreonematoideae), grows endophytically within three geniculate genera of the Corallinoideae. Although the thallus of Choreonema is reduced, lacks differentiated plastids, and is endophytic except for its conceptacles, its status as a parasite has been questioned because cellular connections to the host had not been ob served. Transmission electron microscopy, however, disclosed a previously undescribed type of parasitic interaction in which Choreonema interacts with its host through specialized cells known as lenticular cells. These small, lens-shaped cells are produced from the single file of host-penetrating vegetative cells. Pit plug morphology between vegetative and lenticular cells is polarized. Plug caps facing the vegetative cell have normal coralline morphology, while those facing the lenticular cell are composed of three layers. Regions of lenticular cells near host cells protrude toward the host cell; upon encountering the host cell wall, the prolrusion produces numerous finger-like fimbriate processes that make cellular connections with the host cell. Lenticular cells may extend several protrusions toward a host cell or penetrate more than one host cell; two or more lenticular cells may also penetrate the same host cell. The lack of secondary pit connections, cell fusions, and passage of parasitic nuclei suggest that this parasitic relationship may be evolutionarily older than previously reported cases of parasitism in red algae.     

CHIHARAEA AND YAMADAIA (CORALLINALES,RHODOPHYTA) REPRESENT REDUCED AND RECENTLY DERIVED ARTICULATED CORALLINE MORPHOLOGIES1

Phycologists have hypothesized that the diminutive fronds produced by species in the genera Chiharaea and Yamadaia, which are composed of comparatively few genicula and intergenicula, represent morphological intermediates in the evolution of articulated corallines from crustose ancestors. We test this “intermediate frond hypothesis” by comparing rbcL sequences from the generitype species Chiharaea bodegensis and Yamadaia melobesioides to sequences from other coralline genera. We demonstrate that Chiharaea includes two other NE Pacific species, Arthrocardia silvae and Yamadaia americana. Chiharaea species are characterized morphologically by inflated intergenicula and axial conceptacles with apical or acentric pores. Although relationships among the three species are unresolved, Chiharaea bodegensis, C. americana comb. nov., and C. silvae comb. nov. are distinguished from one another by DNA sequences, morphology, habitat, and biogeography. Chiharaea occurs together with Alatocladia, Bossiella, Calliarthron, and Serraticardia macmillanii in a strongly supported clade of nearly endemic north Pacific articulated coralline genera and species that have evolved relatively recently compared to other articulated corallines. In contrast, NW Pacific Yamadaia melobesioides belongs in a clade with Corallina officinalis, the generitype species of Corallina, and therefore we reduce Yamadia to a synonym of Corallina and propose Corallina melobesioides comb. nov. We reject the ‘intermediate frond hypothesis’ and conclude that Chiharaea and Yamadaia are recently derived taxa that evolved from articulated coralline ancestors and represent a reduction in the number of genicula and intergenicula.     

Timing of the evolutionary history of Corallinaceae (Corallinales,Rhodophyta)

The temporal dimension of the most recent Corallinaceae (order Corallinales) phylogeny was presented here, based on first occurrence time estimates from the fossil record. Calibration of the molecular clock of the genetic marker SSU entailed a separation of Corallinales from Hapalidiales in the Albian (Early Cretaceous ~105 mya). Neither the calibration nor the fossil record resolved the succession of appearance of the first three emerging subfamilies: Mastophoroideae, Corallinoideae, and Neogoniolithoideae. The development of the tetra/bisporangial conceptacle roofs by filaments surrounding and interspersed among the sporangial initials was an evolutionary novelty emerging at the Cretaceous–Paleogene boundary (~66 mya). This novelty was shared by the subfamilies Hydrolithoideae, Metagoniolithoideae, and Lithophylloideae, which diverged in the early Paleogene. Subclades within the Metagoniolithoideae and Lithophylloideae diversified in the late Oligocene–middle Miocene (~28–12 mya). The most common reef corallinaceans (Hydrolithon, Porolithon, Harveylithon, “Pneophyllum” conicum, and subclades within Lithophylloideae) appeared in this interval in the Indo‐Australian Archipelago.     

Community structure of rhodolith-forming beds on the central Brazilian continental shelf

The community structure of rhodoliths beds in the central Brazilian continental shelf was studied under the hypothesis that nongeniculate coralline algae are the major contributors of the individual rhodoliths. Samples were collected from five localities within a single area at 17–18 m depth. At each locality, rhodoliths were collected in 10 random quadrat samples along a 20-m transect. Our results show that dead cores of rhodoliths were significantly composed by nongeniculate coralline red algae rather than bryozoans, corals, or inorganic material. The live outer layers of the rhodoliths are composed mainly of 7 species of nongeniculate red coralline algae (Lithophyllum coralline, L. johansenii, L. depressum, L. stictaeformis, Neogoniolithon brassica-florida, Spongites fruticosus, and Lithothamnion muellerii) associated with other encrusting organisms such as bryozoans, sponges, corals, barnacles, and Peyssonnelia red algae. Significant differences were found in the proportion of Lithophyllum species in relation to other red coralline algae found in this study. Our results show that on the Brazilian continental shelf, the rhodolith-forming species are quite higher in size than in any other studied areas in the world. There was no difference in the proportion of live-to-dead rhodolith materials, suggesting an old bed deposit. Also, the amount of calcium carbonate material in the specimens is relevant to take in account in terms of the CO₂ balance worldwide.     

Bending strategies of convergently evolved,articulated coralline algae

The evolution of uncalcified genicula in upright calcified corallines has occurred at least three times independently, resulting in articulated corallines within Corallinoideae, Lithophylloideae, and Metagoniolithoideae. Genicula confer flexibility to otherwise rigid thalli, and the localization of bending at discrete intervals amplifies bending stress in genicular tissue. Genicular morphology must, therefore, be balanced between maintaining flexibility while mitigating or resisting stress. Genicula in the three articulated lineages differ in both cellular construction and development, which may result in different constraints on morphology. By studying the interaction between flexibility and morphological variation in multiple species, we investigate whether representatives of convergently evolving clades follow similar strategies to generate mechanically successful articulated fronds. By using computational models to explore different bending strategies, we show that there are multiple ways to generate flexibility in upright corallines but not all morphological strategies are mechanically equivalent. Corallinoids have many joints, lithophylloids have pliant joints, and metagoniolithoids have longer joints—while these strategies can lead to comparable thallus flexibility, they also lead to different levels of stress amplification in bending. Moreover, genicula at greatest risk of stress amplification are typically the strongest, universally mitigating the trade‐off between flexibility and stress reduction.     

Taxonomic and biostratigraphical re-assessments of Subterraniphyllum Elliott (Corallinales, Rhodophyta)

A taxonomic and biostratigraphical re‐assessment of Subterraniphyllum Elliott (Corallinales, Rhodophyta) is presented. Results from studies of the type collection and of newly collected material from north‐eastern Italy and northern Slovenia have shown that this taxon is not a geniculate coralline red alga as originally suggested by Elliott and most subsequent authors. Vegetatively, Subterraniphyllum most closely resembles certain living members of the Corallinales; however, the phenetic and phylogenetic relationships of Subterraniphyllum to other Corallinales cannot be determined with greater certainty. The exclusion of Subterraniphyllum from any group of Corallinaceae with genicula is based on unequivocal evidence that branch formation does not involve the occurrence of genicula. Subterraniphyllum seems to be restricted to the Oligocene. Reports of occurrences in Upper Eocene and Lower Miocene sediments cannot be substantiated. Subterraniphyllum, however, cannot be considered a useful stratigraphical marker until further data on its occurrence in well‐dated carbonate sequences are acquired. This study illustrates the problems associated with placing fossil coralline algal specimens into geniculate genera without the preservation of relevant morphological characters. This is especially true in the absence of the careful assessment of fossil material with respect to current taxonomic concepts of geniculate coralline genera, all of which are based on studies of living species. According to the current concepts for geniculate coralline genera, the placing of fossil specimens into geniculate genera without appropriate evidence must be avoided by grouping all potentially geniculate fragments under the informal group ‘Geniculate sensu lato’. Furthermore, for all those many fossil specimens where unequivocal evidence is not present, it is possible to utilize ‘form genera’ based on characters that are normally preserved. This leads to creating a consistent, workable system of applying names to most fossil corallines so that they can be reliably used in relation to stratigraphical and palaeoecological studies.     

Kelp versus coralline: cellular basis for mechanical strength in the wave‐swept seaweed Calliarthron (Corallinaceae,Rhodophyta)1

Previous biomechanical studies of wave‐swept macroalgae have revealed a trade‐off in growth strategies to resist breakage in the intertidal zone: growing in girth versus growing strong tissues. Brown macroalgae, such as kelps, grow thick stipes but have weak tissues, while red macroalgae grow slender thalli but have much stronger tissues. For example, genicular tissue in the articulated coralline Calliarthron cheilosporioides Manza is more than an order of magnitude stronger than some kelp tissues, but genicula rarely exceed 1 mm in diameter. The great tissue strength of Calliarthron genicula results, at least in part, from a lifelong strengthening process. Here, a histological analysis is presented to explore the cellular basis for mechanical strengthening in Calliarthron genicula. Genicula are composed of thousands of fiber‐like cells, whose cell walls thicken over time. Thickening of constitutive cell walls likely explains why older genicula have stronger tissues: a mature geniculum may be >50% cell wall. However, the material strength of genicular cell wall is similar to the strength of cell wall from a freshwater green alga, suggesting that it may be the quantity—not the quality—of cell wall material that gives genicular tissue its strength. Apparent differences in tissue strength across algal taxa may be a consequence of tissue construction rather than material composition.     

Unveiling commonalities in understudied habitats of boulder-reefs: life-history traits of the widespread invertebrate and algal inhabitants

Compared to stable reef habitats, dynamic boulder-reefs (commonly called boulder-fields when intertidal) host many habitat specialist species. Most occur underneath boulders where they are largely hidden from view; only limited research has assessed their life-histories despite their widespread importance for biological diversity. But some abundant under-boulder species likely structuring this system are habitat generalists widely researched elsewhere. Here we review this research, focusing on three widespread under-boulder sessile taxa: spirorbids, serpulids (tubeworms) and nongeniculate coralline algae, and three mobile taxa: sea urchins, chitons and crabs. Spirorbids have extensive reproductive/colonization capabilities but are readily out-competed. We thus characterize spirorbids as mostly early-successional, while serpulids often have greater competitiveness. Nongeniculate corallines occur underneath boulders where light reaches, although they can withstand low levels of that and most other resources. Such traits characterize nongeniculate corallines as late-successional. Thus, succession underneath boulders may shift deterministically from early tubeworms to late nongeniculate corallines. Habitat generalist sea urchin and chiton species often have strong inter-specific interactions in exposed habitats. Future experiments may find that under-boulder aggregations of these taxa, and also crabs, are impacting algal and invertebrate assemblages. These experiments will be required if dynamic boulder-reefs are to be as thoroughly understood as other benthic systems.     

DIFFERENCES IN POLYSACCHARIDE STRUCTURE BETWEEN CALCIFIED AND UNCALCIFIED SEGMENTS IN THE CORALLINE CALLIARTHRON CHEILOSPORIOIDES (CORALLINALES,RHODOPHYTA)1

The articulated coralline Calliarthron cheilosporioides Manza produces segmented fronds composed of calcified segments (intergenicula) separated by uncalcified joints (genicula), which allow fronds to bend and reorient under breaking waves in the wave‐swept intertidal zone. Genicula are formed when calcified cells decalcify and restructure to create flexible tissue. The present study has identified important differences in the main agaran disaccharidic repeating units [→3)‐β‐d ‐Galp (1→ 4)‐α‐l ‐Galp(1→] synthesized by genicular and intergenicular segments. Based on chemical and spectroscopical analyses, we report that genicular cells from C. cheilosporioides biosynthesize a highly methoxylated galactan at C‐6 position with low levels of branching with xylose side stubs on C‐6 of the [→3)‐β‐d ‐Galp (1→] units, whereas intergenicular segments produce xylogalactans with high levels of xylose and low levels of 6‐O‐methyl β‐d ‐Gal units. These data suggest that, during genicular development, xylosyl branched, 3‐linked β‐d ‐Galp units present in the xylogalactan backbones from intergenicular walls are mostly replaced by 6‐O‐methyl‐d‐ galactose units. We speculate that this structural shift is a consequence of a putative and specific methoxyl transferase that blocks the xylosylation on C‐6 of the 3‐linked β‐d ‐Galp units. Changes in galactan substitutions may contribute to the distinct mechanical properties of genicula and may lend insight into the calcification process in coralline algae.     

Cover

ON THE COVER: Articulated coralline Johansenia macmillanii (Corallinoideae), growing epilithically in the intertidal, Botanical Beach, Vancouver Island, BC, Canada. [Vol. 54, No. 3, pp. 305–316 ]     

CONCEPTACLE STRUCTURE OF THE PARASITIC CORALLINE RED ALGA CHOREONEMA THURETII (CORALLINALES) AND ITS TAXONOMIC IMPLICATIONS1

The major diagnostic features for erecting the red algal subfamily Choreonematoideae (Corallinales) were a combination of 1) absence of both cell fusions and secondary pit connections, 2) conceptacle roof and wall comprised of a single cell layer, and 3) presence of tetrasporangial pore plugs within a uniporate conceptacle in the monotypic taxon Choreonema thuretii (Bornet) Schmitz. Because this alga is a parasite, the absence of secondary cell connections is most likely an adaptation to a reduced thallus. This study shows that all conceptacles are not composed of a file of cells but rather a single layer of epithallial cells that are underlain by a thick layer of calcified acellular material; both epithallial cells and the calcified layer are produced by peripheral sterile cells. Although the outermost tetrasporangial pore canal is uniporate, there is a calcified acellular multiporate plate recessed just below the rim. The plate is produced by interspersed sterile cells and is continuous with the calcified layer supporting the conceptacle. These unique structures are likely due to parasitism rather than to the ancestral state. Based on these results and a reexamination of published micrographs depicting lenticular cells in Austrolithon intumescens Harvey et Woelkerling, we propose that both subfamily Choreonematoideae and Austrolithoideae are closely allied with subfamily Melobesioideae. This distant relationship to its host (Corallinoideae) plus a combination of unique conceptacle and unusual type of parasitism indicates that C. thuretii is an alloparasite and that it is likely the most ancient red algal parasite studied to date.     

COMPOSITION,DISTRIBUTION, AND ABUNDANCE OF DEEP‐WATER (>30 m) MACROALGAE IN CENTRAL CALIFORNIA1

The deep‐water macroalgal assemblage was described at 14 sites off the central California coast during 1999 and 2000 from SCUBA and remotely operated vehicle sampling. The stipitate kelp Pleurophycus gardneri Setchell & Gardner, previously thought to be rare in the region, was abundant from 30 to 45 m, forming kelp beds below the well‐known giant kelp forests. Macroalgae typically formed three broadly overlapping zones usually characterized by one or a few visually dominant taxa: 1) the upper “Pleurophycus zone” (30–45 m) of stipitate kelps and Desmarestia spp. with a high percent cover of corallines, low cover of uncalcified red algae, and rare green algae; 2) a middle “Maripelta zone” (40–55 m) with other uncalcified red algae and infrequent corallines and green algae; and 3) a zone (55–75 m) of infrequent patches of nongeniculate coralline algae. The green alga Palmophyllum umbracola Nelson & Ryan, not previously reported from the Northeast Pacific, was found over the entire geographical range sampled from 35 to 54 m. Year‐round profiles of water column irradiance revealed unexpectedly clear water with an average K₀ of 0.106·m ^{? 1} Received 18 January 2002. Accepted 16 December 2002. . The low percent surface irradiance found at the average lower macroalgal depth limits in this study (0.56% for brown algae, 0.12% for uncalcified red algae, and 0.01% for nongeniculate coralline algae) and lack of large grazers suggest that light controls the lower distributional limits. The ubiquitous distribution, perennial nature, and similar lower depth limits of deep‐water macroalgal assemblages at all sites suggest that these assemblages are a common persistent part of the benthic biota in this region.     

Phylogeny of Cyperaceae Based on DNA Sequence Data: Current Progress and Future Prospects

In the last decade, efforts to reconstruct suprageneric phylogeny of the Cyperaceae have intensified. We present an analysis of 262 taxa representing 93 genera in 15 tribes, sequenced for the plastid rbcL and trnL-F (intron and intergenic spacer). Cyperaceae are monophyletic and resolved into two clades, here recognised as Mapanioideae and Cyperoideae, and the overall topology is similar to results from previous studies. Within Cyperoideae, Trilepideae are sister to rest of taxa whereas Cryptangieae, Bisboeckelerieae and Sclerieae are resolved within Schoeneae. Cladium and Rhynchospora (and Pleurostachys) are resolved into clades sister to the rest of Schoeneae, lending support to the recognition of these taxa in separate tribes. However, we retain these taxa in Schoeneae pending broader sampling of the group. The phylogenetic position of 40 species in 21 genera is presented in this study for the first time, elucidating their position in Abildgaardieae (Trachystylis), Cryptangieae (Didymiandrum, Exochogyne), Cypereae (Androtrichum, Volkiella), Eleocharideae (Chillania), and Schoeneae (Calyptrocarya, Morelotia). More sampling effort (more taxa and the use of more rapidly evolving markers) is needed to resolve relationships in Fuireneae and Schoeneae.     

Evolutionary history of the Corallinales (Corallinophycidae, Rhodophyta) inferred from nuclear, plastidial and mitochondrial genomes

Systematics of the red algal order Corallinales has a long and convoluted history. In the present study, molecular approaches were used to assess the phylogenetic relationships based on the analyses of two datasets: a large dataset of SSU sequences including mainly sequences from GenBank; and a combined dataset including four molecular markers (two nuclear: SSU, LSU; one plastidial: psbA; and one mitochondrial: COI). Phylogenetic analyses of both datasets re-affirmed the monophyly of the Corallinales as well as the two families (Corallinaceae and Hapalidiaceae) currently recognized within the order. Three of the four subfamilies of the Corallinaceae (Corallinoideae, Lithophylloideae, Metagoniolithoideae) were also resolved as a monophyletic lineage whereas members of the Mastophoroideae were resolved as four distinct lineages. We therefore propose to restrict the Mastophoroideae to the genera Mastophora, Metamastophora, and possibly Lithoporella in the aim of rendering this subfamily monophyletic. In addition, our phylogenies resolved the genus Hydrolithon in two unrelated lineages, one containing the generitype Hydrolithon reinboldii and the second containing Hydrolithon onkodes, which used to be the generitype of the now defunct genus Porolithon. We therefore propose to resurrect the genus Porolithon for the second lineage encompassing those species with primarily monomerous thalli, and trichocyte arrangements in large pustulate horizontal rows. Moreover, our phylogenetic analyses revealed the presence of cryptic diversity in several taxa, shedding light on the need for further studies to better circumscribe species frontiers within the diverse order Corallinales, especially in the genera Mesophyllum and Neogoniolithon.     

Descriptions and phylogenetic relationships of a new genus and two new species of Oligo‐Miocene cormorants (Aves: Phalacrocoracidae) from Australia

Tertiary cormorant fossils (Aves: Phalacrocoracidae) from Late Oligocene deposits in Australia are described. They derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations in the Lake Eyre and Lake Frome Basins, South Australia, respectively. A new genus, Nambashag gen. nov. , with two new species ( Nambashag billerooensis sp. nov. , 30 specimens; Nambashag microglaucus sp. nov. , 14 specimens), has been established. Phylogenetic analyses based on 113 morphological and two integumentary characters indicated that Nambashag is the sister taxon to the Early Miocene Nectornis miocaenus of Europe and all extant phalacrocoracids. As Nambashag, Nectornis, and extant phalacrocoracids constitute a strongly supported clade sister to Anhinga species, the fossil taxa have been referred to Phalacrocoracidae. Sulids and Fregata were successive sister taxa to the Phalacrocoracoidea, i.e. phalacrocoracids + Anhinga. As phalacrocoracids lived in both Europe and Australia during the Late Oligocene and no older phalacrocoracid taxa are known, the biogeographical origin of cormorants remains unanswered. The phylogenetic relationships of extant taxa were not wholly resolved, but contrary to previous morphological analyses, considerable concordance was found with relationships recovered by recent molecular analyses. Microcarbo is sister to all other extant phalacrocoracids, and all Leucocarbo species form a well‐supported clade. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 277–314.