PHYLOGENY AND CLASSIFICATION OF REEF‐BUILDING CORALLINE ALGAE (CORALLINALES,RHODOPHYTA)

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 rRNA 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.     

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.     

Coralline red algae from the Silurian of Gotland indicate that the order Corallinales (Corallinophycidae,Rhodophyta) is much older than previously thought

Aguirrea fluegelii gen. et sp. nov. (Corallinales, Corallinophycidae, Rhodophyta) is described from the mid‐Silurian of Gotland Island, Sweden (Högklint Formation, lower Wenlock). The holotype is of dimerous construction and includes a uniporate conceptacle with a sporangium, thus providing evidence that taxa of the Corallinales/Corallinaceae existed at least 300 million years earlier than previously documented. Aguirrea fluegelii cannot be unequivocally placed in any of seven currently recognized lineages/subfamilies/groups of the Corallinaceae as not all diagnostic characters are preserved, and thus is accorded incertae sedis status within the family Corallinaceae and order Corallinales. Extant evolutionary history studies of Corallinophycidae involving molecular clocks now require updating using new calibration points to take account of the much earlier unequivocal mid‐Silurian record of uniporate conceptacle‐bearing taxa of Corallinales/Corallinaceae as well as the parallel record of Graticula, a genus attributed to the Sporolithales.     

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.     

植物分类学在化石珊瑚藻(珊瑚藻目,红藻门)中的应用

最近有人认为将化石藻类的分类归入现生藻类分类单元有利于珊瑚藻作为古环境的标志,便于理解该类群的演化。然而,这样分类可能很难,因为并不是所有现生藻类分类特征都能在化石种中保存下来。Sporolithacea科的钙化部分(独立或者聚集的孢子囊群)的出现,可以把它们与这个类群的另一个现生科Corallinaceae区别开,这个科在生殖窠中产生孢子囊。节片的有无,丝间细胞的联系类型,生殖窠中孢子囊释放的数目都是用来划分Coral1inaceae科的亚科的标准,在化石样品中也可以用合适的条件进行观察。在大多数情况下,对现生珊瑚藻类属的划分特征可以在化石藻类中鉴别出来,但在几种现生珊瑚藻没有钙化的生殖结构或发育特征。因此,它们生殖结构无法与相应的化石藻类进行对比,也不能进行化石藻类的分类。近年来的趋势认为生殖结构和发育特征是对现生珊瑚藻进行分类的优先鉴定标准,然而,某些特征的稳定性在属的划分上仍然存在争论。在许多情况下,现生藻类的分类标准特征都不能在化石中保存,对古生物化石的分类标准的最佳选择是在化石藻类中选择辅助的,并且可以识别的其它鉴定特征,或者应用非正式的比现生藻类代表定义更宽的属名。     

Phylomineralogy of the Coralline red algae: Correlation of skeletal mineralogy with molecular phylogeny

The coralline algae in the orders Corallinales and Sporolithales (subclass Corallinophycidae), with their high degree of mineralogical variability, pose a challenge to projections regarding mineralogy and response to ocean acidification. Here we relate skeletal carbonate mineralogy to a well-established phylogenetic framework and draw inferences about the effects of future changes in sea-water chemistry on these calcified red algae. A collection of 191 coralline algal specimens from New Zealand, representing 13 genera and 28 species, included members of three families: Corallinaceae, Hapalidiaceae, and Sporolithaceae. While most skeletal specimens were entirely calcitic (range: 73–100 wt.% calcite, mean 97 wt.% calcite, std dev = 5, n = 172), a considerable number contained at least some aragonite. Mg in calcite ranged from 10.5 to 16.4 wt.% MgCO₃, with a mean of 13.1 wt.% MgCO₃ (std dev = 1.1, n = 172). The genera Mesophyllum and Lithophyllum were especially variable. Growth habit, too, was related to mineralogy: geniculate coralline algae do not generally contain any aragonite. Mg content varied among coralline families: the Corallinaceae had the highest Mg content, followed by the Sporolithaceae and the Hapalidiaceae. Despite the significant differences among families, variation and overlap prevent the use of carbonate mineralogy as a taxonomic character in the coralline algae. Latitude (as a proxy for water temperature) had only a slight relationship to Mg content in coralline algae, contrary to trends observed in other biomineralising taxa. Temperate magnesium calcites, like those produced by coralline algae, are particularly vulnerable to ocean acidification. Changes in biomineralisation or species distribution may occur over the next few decades, particularly to species producing high-Mg calcite, as pH and CO₂ dynamics change in coastal temperate oceans.     

Inventario delle Corallinales del Mar Mediterraneo: considerazioni tassonomiche

Abstract

Inventory of Corallinales of the Mediterranean Sea: taxonomic aspects. - The aim of the present work is the drawing up of a critical inventory of Corallinales of the Mediterranean Sea. This research has been based upon a census of 320 works published since 175 years, upon direct collections and sample exchanges. A census of 88 species of Corallinales (with forms and varieties), made up on the basis of about 5000 records, has been taken. 34 of the 88 species are considered “species dubiae aut rariter inventae”, it has been reported also “species inquirendae” and “species addendae”. From this census it results that there are some taxonomic problems about genera attributions (Lithophyllum, Pseudolithophyllum, Titanoderma, Neogoniolithon, Spongites, Goniolithion, Leptophytum), species attributions of articulated Corallinales (Corallina elongata, C. officinalis e Haliptilon squamatum) and non-articulated Corallinales (Fosliella ischitana, Lithophyllum (Titanoderma) trochanter, Spongites fruticulosus). Definition of these taxonomic attributions might condition not only studies of phytogeographical distribution of these algae but also every successive interpretation.     

Plio-Pleistocene cercopithecids from Kanam East, western Kenya

This paper describes cercopithecid craniodental and postcranial fossils recovered by L. S. B. Leakey at Kanam East, Kenya during the early 1930s. These fossil monkeys have been generally assumed to have been derived from early Pliocene horizons, but their exact geographical and stratigraphical provenience is unknown. Although the question of the evolutionary significance of these specimens must await the recovery of more securely dated material from Kanam East, some general conclusions can be drawn concerning their taxonomic affinities and paleobiology. Based on comparative studies of the craniodental material, at least three extant genera are represented—Colobus,Lophocebus, andCercopithecus. The postcranial fossils include a number of hindlimb specimens, as well as the manubrium of a sternum and a caudal vertebra. Identification of the postcranial remains to particular genera is not possible, but they are similar in morphology to modern arboreal and semiterrestrial cercopithecid monkeys of small to medium size. It is evident that Kanam East had a diverse cercopithecid community, similar to those found today in forested and woodland habitats, and this may be of some significance in reconstructing the paleoecology of the site. Because the fossil record of most extant cercopithecid genera is rather sparse at Plio-Pleistocene sites in Africa, Kanam East represents one of only a few sites that has yielded material that can be assigned toColobus,Lophocebus, orCercopithecus. The fossil monkeys from the site, therefore, provide additional evidence to help reconstruct the paleobiology, as well as the patterns of species diversity and community structure that characterized the cercopithecid radiation during the Plio-Pleistocene.     

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.     

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.     

The taxonomic status ofGraecopithecus

In 1944 a single mandibular specimen was found in Pyrgos, southern Greece.von Koenigswald (1972) described this specimen and allocated it to a new taxon,Graecopithecus freybergi.de Bonis and colleagues later described new hominid specimens from northern Greece and allocated them to the new taxon,Ouranopithecus macedoniensis.Martin andAndrews (1984), however, argued that there was no reason for maintaining a generic or specific distinction between the northern and southern Greek fossil populations as suggested byde Bonis andMelentis. This paper reexamines the taxonomic status of the southern and northern Greek hominids in light of recent fossil discoveries and concludes that there is no evidence for recognizing two hominine genera from the Miocene of Greece. There is evidence, however, to suggest that the Macedonian hominid is specifically distinct from the southern Greek hominid. Thus, becauseGraecopithecus has taxonomic priority overOuranopithecus, it is concluded that the Macedonian specimens should be allocated to a separate species ofGraecopithecus.     

AN SEM‐BASED ANALYSIS OF THE MORPHOLOGY,ANATOMY, AND REPRODUCTION OF LITHOTHAMNION TOPHIFORME (ESPER) UNGER (CORALLINALES,RHODOPHYTA), WITH A COMPARATIVE STUDY OF ASSOCIATED NORTH ATLANTIC ARCTIC/SUBARCTIC MELOBESIOIDEAE1

Lithothamnion tophiforme (Esper) Unger is a dominant, arctic, saxicolous species that extends southward, albeit with reduced cover, into the deeper colder waters of the North Atlantic subarctic, where it also occurs in significant rhodolith deposits with L. glaciale. The external appearance of L. tophiforme is distinctive, but typification, anatomy, reproduction, ecology, and biogeography have not been previously analyzed. These topics are now addressed, with extensive use of SEM, in comparison with other North Atlantic arctic and subarctic melobesioid genera and species. The species considered in this article comprise 95% of the coralline biomass of the colder North Atlantic and adjacent arctic (i.e. less than 12° C summer and less than 0° C winter). In the outer thallus region of coralline algae, crust extension proceeds, calcification develops, surface sloughing and grazing occur, and reproductive structures are initiated. Analysis of the ultrastructure of the outer thallus region (epithallium, meristem, and perithallium) of L. tophiforme shows distinctive generic similarities and specific differences from the other Lithothamnion species discussed here. Considerable generic differences from the Clathromorpum and Leptophytum species also encountered in the region considered are highlighted as well. We discuss the functional and taxonomic implications of these distinguishing features and recommend that they be more widely considered in future research on coralline algae to understand more fully the ecology and evolution of the Corallinales.     

Taxonomic biodiversity of geniculate coralline red algae (Corallinales,Rhodophyta) from the Macaronesian region: summary and analysis

A catalog and critical review of species and infraspecific taxa of non-fossil geniculate coralline red algae (Corallinales, Rhodophyta) previously reported from the Macaronesian region are presented along with an assessment of species diversity in the region. Published records of geniculate coralline algae are included along with comments relating to type material. Within the catalog, taxa are organized alphabetically by genus and within this by final epithet. From the 31 taxa recorded, 4 are based on type collections from Macaronesian localities. The types of most species and infraspecific taxa reported from the region have yet to be re-examined in a modern context, and most Macaronesian records require verification. The biodiversity of Macaronesian geniculate coralline algae may be lower than current information indicates.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: RHAMNUS MARGINATUS (RHAMNACEAE) REEXAMINED

Leaf compressions, previously assigned to Rhamnus marginatus Lesquereux, were collected from the Middle Eocene Claiborne Formation of western Kentucky and Tennessee. The leaf architecture and cuticular features of over 40 compressions were carefully examined and compared to those of many extant species of Rhamnaceae and related families as well as fossil specimens previously assigned to this taxon. This leaf type appears to belong to the Rhamnaceae, however, it conforms more closely to species of several genera in the tribe Zizypheae than to those of Rhamnus or other genera in the tribe Rhamneae. Confident assignment to any specific genus within this complex of genera cannot be made on the basis of leaf characteristics alone and would require discovery and analysis of additional vegetative and reproductive organs. Because this fossil leaf form cannot be confidently assigned to any modern genus and earlier classifications appear to be improper, this leaf type has been reassigned to the taxon Berhamniphyllum claibornense gen. et sp. nov. The transfer of this leaf form at the tribe level reaffirms the need for close examination of taxonomic determinations made by early workers.     

An appraisal of the fossil record for the Cirolanidae (Malacostraca: Peracarida: Isopoda: Cymothoida), with a description of a new cirolanid isopod crustacean from the early Miocene of the Vienna Basin (Western Carpathians)

Abstract:  Isopod crustaceans are rarely preserved in the fossil record. Herein, an appraisal of the fossil record for the cirolanid isopods is presented. Five genera are briefly discussed, including Bathynomus, Brunnaega, Palaega, Pseudopalaega and Cirolana. A key for the cirolanid genera known to date from the fossil record is provided based mostly on pleotelson characters. From the early Miocene of the Slovak part of the Vienna Basin, Cirolana feldmanni sp. nov. is described being only the fifth fossil Cirolana species known to date and one of the few with preserved appendages. The material exhibits preservation suggesting biphasic moulting; the mode of preservation suggests a rather short time between shedding the posterior and anterior parts of the exoskeleton instead of hours or even days known in extant taxa. As no subsequent transport or physical disturbance was inferred, the specimens can be stated as in situ preservation. From the palaeoecological point of view, it is concluded that Cirolana feldmanni sp. nov. is the first unequivocal fossil deep‐water Cirolana as suggested by the accompanied fauna.     

Palaeobiology of tanaidaceans (Crustacea: Peracarida) from Cretaceous ambers: extending the scarce fossil record of a diverse peracarid group

Diverse assemblages of tanaidacean peracarid crustaceans from western Tethyan continental deposits suggest that the group was relatively common in or around ancient resin‐producing forests. Here we report the results of an examination of 13 tanaidacean specimens from three Cretaceous (Albian to Turonian) French amber deposits. Two new species of the fossil family Alavatanaidae are placed in the previously described Early Cretaceous genus Eurotanais: Eurotanais pyrenaensis sp. nov. from Cenomanian Pyrenean amber (Fourtou, Aude) and Eurotanais seilacheri sp. nov. from Turonian Vendean amber (La Garnache, Vendée). The remaining specimens are placed in three newly erected genera and species (but family incertae sedis): Arcantitanais turpis gen. et sp. nov. from Albian–Cenomanian Charentese amber (Archingeay, Charente‐Maritime), and Tytthotanais tenvis gen. et sp. nov. and Armadillopsis rara gen. et sp. nov. from Pyrenean amber. These are the first formally described fossils that might be related to the paratanaoidean families Nototanaidae and Paratanaidae, sharing with these some putatively derived features and providing possible evidence for the antiquity and morphological stability of these families and the suborder Tanaidomorpha. The distinctive features and character combinations of these fossil taxa are discussed in connection with possible relationships to the living lineages of tanaidaceans. Propagation phase‐contrast X‐ray synchrotron microtomography was used to obtain high‐quality 3D images for some fossils. A discussion is provided on the putative palaeobiology of tanaidaceans and the French resiniferous forest ecosystem. The discovery of these new tanaidaceans extends the palaeogeographical distribution and stratigraphical range of the family Alavatanaidae and sheds new light on the palaeoecology and diversity of tanaidaceans in pre‐angiospermous woodlands.     

Establishing temperate crustose early Holocene coralline algae as archives for palaeoenvironmental reconstructions of the shallow water habitats of the Mediterranean Sea

Over the past decades, coralline algae have increasingly been used as archives of palaeoclimate information due to their seasonal growth bands and their vast distribution from high latitudes to the tropics. Traditionally, these reconstructions have been performed mainly on high latitude species, limiting the geographical area of their potential use. Here we assess the use of temperate crustose fossil coralline algae from shallow water habitats for palaeoenvironmental reconstruction to generate records of past climate change. We determine the potential of three different species of coralline algae, Lithothamnion minervae, Lithophyllum stictaeforme and Mesophyllum philippii, with different growth patterns, as archives for pH (δ¹¹B) and temperature (Mg/Ca) reconstruction in the Mediterranean Sea. Mg concentration is driven by temperature but modulated by growth rate, which is controlled by species-specific and intraspecific growth patterns. L. minervae is a good temperature recorder, showing a moderate warming trend in specimens from 11.37 cal ka BP (from 14.2 ± 0.4°C to 14.9 ± 0.15°C) to today. In contrast to Mg, all genera showed consistent values of boron isotopes (δ¹¹B) suggesting a common control on boron incorporation. The recorded δ¹¹B in modern and fossil coralline specimens is in agreement with literature data about early Holocene pH, opening new perspectives of coralline-based, high-resolution pH reconstructions in deep time.     

Phylogenetic analysis of rhodolith formation in the Corallinales (Rhodophyta)

Although the ecological importance of rhodolith (maerl, free-living coralline algae) beds is well-known, rhodolith-forming species have been neglected in molecular phylogenetic studies. This is the first molecular systematic study aimed at understanding whether the rhodolith habit is a fixed feature in lineages and determining the relationship (phylogenetic vs. environmental) between rhodolith and crustose habits. Phylogenetic relationships of rhodolith-forming species and encrusting coralline algae at generic and species levels were analysed using SSU rDNA and psbA sequences. Extensive sampling in the European North Atlantic, Pacific and Caribbean Mexico of Phymatolithon, Lithothamnion, Lithophyllum and Neogoniolithon taxa forming rhodoliths and crusts was accompanied by examination of type or topotype material. Phylogenetic reconstruction showed that Neogoniolithon contained a monophyletic group of rhodolith-forming species whereas other rhodolith-formers were closely related to encrusting forms in the genera Phymatolithon, Lithothamnion, Mesophyllum, Hydrolithon, Spongites and Sporolithon. DNA analysis showed that the crust-forming Lithophyllum cf. incrustans/dentatum also forms rhodoliths with a stone nucleus that occur on rocky shores. In contrast, species that form beds of non-nucleate rhodoliths (e.g. Neogoniolithon spectabile, N. strictum, Lithophyllum cf. incrustans/dentatum or sp. 1 and Phymatolithon calcareum) rarely form crusts. The rhodolith habit cannot be used to delimit species for taxonomic or identification purposes. Extensive taxonomic revision will be required to deal with problems such as the position of specimens identified as Lithophyllum margaritae in two unrelated lineages.