A morphological study and hybridization analysis of Caloglossa leprieurii (Ceramiales, Rhodophyta) from Japan, Singapore and Australia

Morphological and hybridization experiments were performed on Caloglossa leprieurii (Montagne) J. Agardh collected from Japan, Singapore and Australia in order to evaluate taxonomic characters of this species. Within C. leprieurii at least four mating groups were recognized from the Indo-Pacific region. These mating groups could be characterized by the blade width at the inter-node and the cell-row numbers on the opposite side derived from the first axial cell at the main axis, though these properties showed a certain variability even in the same plant under both field and culture conditions. The phylogenetic relationship and geographic distribution of the four mating groups are discussed.     

EVOLUTIONARY DIVERGENCE IN THE RED ALGAE CALOGLOSSA LEPRIEURII AND C. APOMEIOTICA

Morphological comparisons, hybridization experiments, and molecular phylogenetic analyses using the RUBISCO spacer region were undertaken on 12 populations of Caloglossa leprieurii (Montagne) J. Agardh in order to clarify their relationships. In addition, data from one population of the morphologically similar but asexual species, C. apomeiotica (West et Zuccarello), were included in the assessment. Three morphological types were recognized on the basis of the number of cell rows at nodes of the main axis opposite to the lateral branch and blade width: single/ broad (with three mating groups), multiple/broad (three mating groups) and multiple/slender (one mating group). In the molecular analyses, C. leprieurii was resolved as two clusters that correspond phenetically to the single and multiple cell row types. Both the morphological and molecular data indicate that the asexual species was derived from sexual plants of the multiple cell row type. The reproductive compatibility correlates with genetic distance rather than geographical distance. Sympatric mating groups are completely incompatible and have 10–21 nucleotide changes in the examined region, whereas mating groups that produce abnormal progeny or pseudocystocarps are allopatrically distributed with 5–7 nucleotide changes. The present data suggest that the two populations, one with single and the other with multiple cell rows, which are sympatrically distributed in southeastern Japan, have probably evolved by allopatric speciation. The single/broad type that is restricted to the western Pacific, may have diverged genetically between eastern and western Australia, with subsequent dispersal from the western population as far as Japan.     

DICROGLOSSUM CRISPATULUM GEN. ET COMB. NOV. FROM WESTERN AUSTRALIA,REPRESENTING A NEW TRIBE WITHIN THE DELESSERIACEAE (RHODOPHYTA)1

Dicroglossum gen. nov. (Delesseriaceae, Ceramiales) is a monotypic genus based on Delesseria crispatula, a species originally described by Harvey for plants collected from southwestern Western Australia. Distinctive features of the new genus include exogenous indeterminate branches; growth by means of a single transversely dividing, apical cell; absence of intercalary divisions in the primary, secondary, and tertiary cell rows; lateral pericentral cells not transversely divided; not all cells of the secondary cell rows producing tertiary cells rows; all tertiary initials reaching the thallus margin; midrib present but lateral nerves absent; determinate lateral bladelets arising endogenously; blades monostromatic, except, at the midrib; carpogonial branches restricted to primary cell rows, on both surfaces of unmodified blades; procarps produced on both blade surfaces, each procarp consisting of a supporting cell that bears two four-celled carpogonial branches and one sterile-cell group of three to four cells; and tetrasporangia borne in two layers, separated by a central row of sterile cells. The combination of exogenous indeterminate branching and bicarpogonial procarps is considered to warrant the recognition of a new tribe, the Dicroglosseae, within the subfamily Delesserioideae.     

THE GENUS HYPOGLOSSUM KÜTZING (DELESSERIACEAE,RHODOPHYTA) IN THE TROPICAL WESTERN ATLANTIC,INCLUDING H. ANOMALUM SP. NOV.1

Observations are made on the occurrence and distribution of the red algal genus Hypoglossum Kützing (Delesseriaceae, Ceramiales) in the tropical western Atlantic. In addition to the type of the genus, H. hypoglossoides (Stackh.) Coll. & Herv., three other species are reported: H. anomalum sp. nov., H. involvens (Harv.) J. Ag., and H. tenuifolium (Harv.) J. Ag. A key is presented to distinguish these four species. The newly described species, H. anomalum, is like other species in the genus in that its branches arise endogenously from the primary axial row but it is unique in that the branches emerge from the parent blade at some point between the midline and the margin of the blade. The new species is reported from Puerto Rico and Florida.     

AUGOPHYLLUM,A NEW GENUS OF THE DELESSERIACEAE (RHODOPHYTA) BASED ON rbcL SEQUENCE ANALYSIS AND CYSTOCARP DEVELOPMENT1

A new genus, Augophyllum Lin, Fredericq et Hommersand gen. nov. related to Nitophyllum, tribe Nitophylleae, subfam. Nitophylloideae of the Delesseriaceae, is established to contain the type species Augophyllum wysorii Lin, Fredericq et Hommersand sp. nov. from Caribbean Panama; Augophyllum kentingii Lin, Fredericq et Hommersand sp. nov. from Taiwan; Augophyllum marginifructum (R. E. Norris et Wynne) Lin, Fredericq et Hommersand comb. nov. (Myriogramme marginifructa R. E. Norris et Wynne 1987) from South Africa, Tanzania, and the Sultanate of Oman; and Augophyllum delicatum (Millar) Lin, Fredericq et Hommersand comb. nov. (Nitophyllum delicatum Millar 1990 ) from southeastern Australia. Like Nitophyllum, Augophyllum is characterized by a diffuse meristematic region, the absence of macro‐ and microscopic veins, procarps consisting of a supporting cell bearing a slightly curved four‐celled carpogonial branch flanked laterally by a cover cell and a sterile cell, a branched multicellular sterile group after fertilization, absence of cell fusions between gonimoblast cells, and tetrasporangia transformed from multinucleate surface cells. Augophyllum differs from Nitophyllum by the blades becoming polystromatic inside the margins, often with a stipitate cylindrical base, the possession of aggregated discoid plastids neither linked by fine strands nor forming bead‐like branched chains, spermatangia and procarps initiated at the margins of blades, not diffuse, and a cystocarp composed of densely branched gonimoblast filaments borne on a conspicuous persistent auxiliary cell with an enlarged nucleus. Analyses of the rbcL gene support the separation of Augophyllum from Nitophyllum. An investigation of species attributed to Nitophyllum around the world is expected to reveal other taxa referable to Augophyllum.     

Caloglossa intermedia, sp. nov. (Rhodophyta) from the western atlantic coast: molecular and morphological analyses with special reference to C. leprieurii and C. monosticha

Caloglossa intermedia , sp. nov. is described from estuaries and salt marshes of Atlantic North America. This species is related to C. leprieurii (Montagne) G. Martens based on the position of rhizoids, whereas it is more similar to C. monosticha Kamiya in the number of cell rows at the nodes. Rhizoidal development of this alga also shows an intermediate characteristic between C. leprieurii and C. monosticha. Crosses were successful between the strains of C. intermedia from Georgia and South Carolina, but neither strain was interfertile with morphologically related species. C. intermedia from Georgia, New Jersey, and South Carolina had identical DNA sequences in the Rubisco spacer and flanking regions of rbc L and rbc S. This alga was more closely related to C. leprieurii in a molecular phylogenetic analysis. The evolution of each morphological character is discussed.     

SYSTEMATICS OF THE DELESSERIACEAE (CERAMIALES, RHODOPHYTA) BASED ON LARGE SUBUNIT rDNA AND rbcL SEQUENCES, INCLUDING THE PHYCODRYOIDEAE, SUBFAM. NOV.

The present classification of the Delesseriaceae retains the essential features of Kylin's system, which recognizes two subfamilies Delesserioideae and Nitophylloideae and a series of “groups” or tribes. In this study we test the Kylin system based on phylogenetic parsimony and distance analyses inferred from two molecular data sets and morphological evidence. A set of 72 delesseriacean and 7 additional taxa in the order Ceramiales was sequenced in the large subunit rDNA and rbcL analyses. Three large clades were identified in both the separate and combined data sets, one of which corresponds to the Delesserioideae, one to a narrowly circumscribed Nitophylloideae, and one to the Phycodryoideae, subfam. nov., comprising the remainder of the Nitophylloideae sensu Kylin. Two additional trees inferred from rbcL sequences are included to provide broader coverage of relationships among some Delesserioideae and Phycodryoideae. Belonging to the Delesserioideae are the Caloglosseae with Caloglossa; an expanded Hemineureae that includes Hemineura, Patulophycus, Marionella, Laingia, Botryocarpa, and Pseudophycodrys; the Delesserieae with Delesseria and Membranoptera; the Apoglosseae with Apoglossum and a group of southern hemisphere species presently placed in Delesseria that belong in Paraglossum; the Hypoglosseae with Hypoglossum, Branchioglossum, Zellera, and Bartoniella; and the Grinnellieae with Grinnellia. The revised Nitophylloideae contains the Nitophylleae with Nitophyllum, Valeriemaya, Polyneuropsis, and Calonitophyllum and the Martensieae with Opephyllum and Martensia. A new subfamily, Phycodryoideae, is proposed to include the Phycodryeae with Phycodrys, Polyneura, Nienburgia, Cladodonta, Heterodoxia, and Womersleya; the Cryptopleureae with Cryptopleura, Hymenena, Acrosorium, and Botryoglossum; the Myriogrammeae with Myriogramme and Haraldiophyllum; and the Schizoserideae with Schizoseris, Neuroglossum, Drachiella, Abroteia, and species from South America placed in Platyclinia. This research promotes the correlation of molecular and morphological phylogenies.     

CHARACTERIZATION OF SCHIZOSERIS CONDENSATA,SCHIZOSERIDEAE TRIB. NOV. (DELESSERIACEAE,RHODOPHYTA)1

The Myriogramme group of Kylin contains two distinct clusters of genera that merit recognition at the tribal level. We previously established the tribe Myriogrammeae, and in this paper we erect the Schizoserideae based on a study of the type species of Schizoseris, S. laciniata (=S. condensata), from the southern hemisphere. The Schizoserideae is characterized by 1) marginal and diffuse intercalary meristems; 2) nuclei initially arranged in a plate in the median plane in meristematic and mature cells; 3) chloroplasts one to few, lobed or dissected; 4) microscopic veins absent; 5) procarps scattered, formed singly on either side of the blade with cover cells absent and consisting of a one- to two-celled lateral sterile group, a one- to two-celled basal sterile group, and a four-celled carpogonial branch in which the trichogyne passes beneath the lateral sterile group and emerges anterior to it; 6) auxiliary cell diploidized by a connecting cell cut off posteriolaterally from the fertilized carpogonium; 7) gonimoblast initial cut off laterally from one side of the auxiliary cell and giving rise to unilaterally branched gonimoblast filaments bearing carposporangia in branched chains; 8) gonimoblast fusion cell highly branched, candelabra-like, incorporating all but the basalmost cells of the carposporangial chains and radiating through the central cells in the floor of the cystocarp; 9) spermatangial and tetrasporangial sori formed from surface cells in both monostromatic and polystromatic portions on both sides of the blade; and 10) tetrasporangia formed primarily from cortical rather than from central cells. The Schizoserideae presently includes Schizoseris Kylin, Neuroglossum Kützing, Abroteia J. Agardh, and Polycoryne Skottsberg in Kylin and Skottsberg.     

THE GENUS BALLIELLA ITONO & TANAKA (RHODOPHYTA: CERAMIACEAE) FROM EASTERN AUSTRALIA1

Three new species of the genus Balliella Itono & Tanaka are described from eastern Australia. Balliella amphiglanda from Lord Howe I. and Port Hacking, N.S.W., is distinctive in producing either abaxial or adaxial gland cells from the basal cells of lateral branches and in having tetrasporangia restricted to short branches. Balliella repens, from Tryon I., Heron I and Wistari Reef Qld., and Lord Howe I., N.S.W., is distinguished from the other species of the genus by its regularly developed prostrate systems and clustered tetrasporangia. Balliella grandis, from Wistari Reef, North West I. and One Tree I., Qld., is the largest recorded species of Balliella. It has correspondingly large gland cells and abaxial as well as adaxial tetrasporangia. Our work supports the placement of Balliella in the tribe Antithamnieae, a move which necessitates emending Wollaston's definition of the tribe to include species with procarps that form at successive levels along main axes rather than at only one or two points behind the apices.     

EFFECTS OF SALINITY ON THE DISTRIBUTION OF CALOGLOSSA LEPRIEURII (RHODOPHYTA) IN THE BRISBANE RIVER,AUSTRALIA1

The role of salinity as a factor determining the distribution of two red algal taxa, Caloglossa leprieurii (Mont.) J. Ag. var. leprieurii and Caloglossa leprieurii var. angusta Jao, along the course of the Brisbane River, Queensland, Australia, was investigated. In the field, C. leprieurii var. angusta tolerated a narrower salinity range (mean salinity = 0.0–18.9) than C. leprieurii var. leprieurii (mean salinity = 2.0–33.8) and occupied areas of lower salinity (salinity expressed according to the Practical Salinity Scale of 1978). Both taxa coexisted for a distance of 23 km along the middle reaches of the river. Cell measurements of specimens of both taxa collected along the river showed an increase in cell sizes upstream from the mouth. Results of a reciprocal transplant experiment and growth responses in a series of laboratory culture studies of the two taxa in a range of salinities are presented. These could be correlated with the field distribution of the algae, demonstrating their euryhalinity and the presence of distinct salinity ecotypes.     

MORPHOLOGY AND TAXONOMY OF MERISTIELLA GEN. NOV. (SOLIERIACEAE,RHODOPHYTA) 1

Eucheuma acanthocladum (Harvey) J. Agardh, E. gelidium (J Agardh) J. Agardh, E. echinocarpum Areschoug and E. schrammii(P. et H. Crouan) J. Agardh from the tropical and warm temperate waters of the western Atlantic Ocean and Caribbean Sea are transferred to a new genus, Meristiella. Meristiella exhibits the following Unique combination of characters among genera in the Solieriaceae: (1) rotated periaxial cells, (2) a loosely filamentous medulla. (3) an auxiliary cell complex, (4) Single and twin connecting filaments and (5) spinose cystocarps composed of a central, small-celled placentum, based on its reproductive features, Meristiella. is assigned to the tribe Agardhielleae. Culture experiments and herbarium studies provide evidence that E, gelidium and E. acantghocladum are conspecific. Lectotypes are designated for the included species.     

MORPHOLOGY AND SYSTEMATICS OF SCIUROTHAMNION STEGENGAE GEN. ET SP. NOV. (CERAMIACEAE,RHODOPHYTA) FROM THE INDO‐PACIFIC1

A new ceramiaceous alga, Sciurothamnion stegengae De Clerck et Kraft, gen. et sp. nov., is described from the western Indian Ocean and the Philippines. Sciurothamnion appears related to the tribe Callithamnieae on the basis of the position and composition of its procarps and by the majority of post‐fertilization events. It differs, however, from all current members of the tribe by the presence of two periaxial cells bearing determinate laterals per axial cell. Additionally, unlike any present representative of the subfamily Callithamnioideae, no intercalary foot cell is formed after diploidization of the paired auxiliary cells. The genus is characterized by a terminal foot cell (“disposal cell”), which segregates the haploid nuclei of the diploidized auxiliary cell from the diploid zygote nucleus. The nature of three types of foot cells reported in the Ceramiaceae (intercalary foot cells containing only haploid nuclei, intercalary foot cells containing haploid nuclei and a diploid nucleus, and terminal foot cells containing only haploid nuclei) is discussed.     

CHARACTERIZATION OF MARTENSIA (DELESSERIACEAE,RHODOPHYTA) BASED ON A MORPHOLOGICAL AND MOLECULAR STUDY OF THE TYPE SPECIES,M. ELEGANS,AND M. NATALENSIS SP. NOV. FROM SOUTH AFRICA1

An examination of a series of collections from the coast of Natal, South Africa, has revealed the presence of two species of Martensia C. Hering nom. cons: M. elegans C. Hering 1841, the type species, and an undescribed species, M. natalensis sp. nov. The two are similar in gross morphology, with both having the network arranged in a single band, and with reproductive thalli of M. elegans usually larger and more robust than those of M. natalensis. Molecular studies based on rbcL sequence analyses place the two in separate, strongly supported clades. The first assemblage occurs primarily in the Indo‐West Pacific Ocean, and the second is widely distributed in tropical and warm‐temperate waters. Criteria that have been used in the past for separating the two, namely, the number and shape of the blades, the presence of a single‐ versus a multiple‐banded network, and blade margins entire or toothed, were determined to be unreliable. Although the examination of additional species is required, the morphology and position of procarps and cystocarps, whether at or near the corners of the longitudinal lamellae and the cross‐connecting strands or along the lobed, membranous edges of the longitudinal lamellae or on the thallus margins, may prove to be diagnostic at the subgenus level. We recognize subg. Martensia, including the type of Martensia: M. elegans and subg. Mesotrema (J. Agardh) De Toni based on Martensia pavonia (J. Agardh) J. Agardh.     

NEW GENERA AND SPECIES OF CERAMIACEAE (RHODOPHYTA) FROM THE SOUTHEASTERN UNITED STATES1

Five Ceramiaceae (Rhodophyta) are reported from the offshore waters of the southeastern, warm temperate coast of the United States. These include two new monotypic genera, Calliclavula trifurcate Schneider in the Griffithsieae and Nwynea grandispora Searles in the Sphondylothamnieae, and three new species, Callithamniella silvae Searles, Ptilothamnion occidentale Searles, and Lejolisia exposita Schneider et Searles.     

MORPHOLOGY,MOLECULAR PHYLOGENY AND TAXONOMY OF TWO NEW SPECIES OF PLEODORINA (VOLVOCEAE,CHLOROPHYCEAE)1

The volvocacean genus Pleodorina has been morphologically characterized as having small somatic cells in spheroidal colonies and anisogamous sexual reproduction with sperm packets. In this study we examined two new species that can be assigned to the genus Pleodorina based on morphology: P. starrii H. Nozaki et al. sp. nov. and P. thompsonii F. D. Ott et al. sp. nov. P. starrii was collected from Japan and had 32‐ or 64‐celled colonies with anterior somatic cells and spheroidal individual cellular sheaths that were weakly attached to each other within the colonial envelope. P. thompsonii from Texas (USA) exhibited four or 12 somatic cells in the anterior pole of 16‐ or 32‐celled colonies, respectively, and had a single large pyrenoid in the chloroplast of mature reproductive cells. The chloroplast multigene phylogeny placed P. starrii and P. indica (Iyenger) H. Nozaki in a clade that was robustly separated from the type species P. californica Shaw and P. japonica H. Nozaki. Pleodorina thompsonii was resolved as a basal branch within a large monophyletic group (Eudorina group) composed of Eudorina, Pleodorina and Volvox (excluding section Volvox). Thus, Pleodorina was found among three separate lineages within the Eudorina group in which Eudorina and Volvox were also resolved as nonmonophyletic. The DNA sequences from additional species/strains as well as recognition of morphological attributes that characterize the monophyletic groups within the Eudorina group are needed to construct a natural generic classification within these members of the Volvocaceae.     

THE GENUS PIKEA (DUMONTIACEAE,RHODOPHYTA) IN ENGLAND AND THE NORTH PACIFIC: COMPARATIVE MORPHOLOGICAL,LIFE HISTORY,AND MOLECULAR STUDIES1

A Pikea species attributed to Pikea californica Harvey has been established in England since at least 1967. Previously, this species was believed to occur only in Japan and Pacific North America. Comparative morphological studies on field-collected material and cultured isolates from England, California, and Japan and analysis of organellar DNA restriction fragment length polymorphisms, detected using labeled organellar DNA as a non-radioactive probe, showed that English Pikea is conspecific with P. californica from California. Both populations consist of dioecious gametophytes with heteromorphic life histories involving crustose tetrasporophytes; 96% of organellar DNA bands were shared between interoceanic samples. A second dioecious species of Pikea, P. pinnata Setchell in Collins, Holden et Setchell, grows sympatrically with P. californica near San Francisco but can be distinguished by softer texture, more regular branching pattern, and elongate cystocarpic axes. Pikea pinnata and P. californica samples shared 49–50% of organellar DNA bands, consistent with their being distinct species. Herbarium specimens of P. robusta Abbott resemble P. pinnata in some morphological features but axes are much wider; P. robusta may represent a further, strictly sub-tidal species but fertile material is unknown. Pikea thalli from Japan, previously attributed to P. californica and described here as Pikea yoshizakii sp. nov., are monoecious and show a strikingly different type of life history. After fertilization, gonimoblast filaments grow outward through the cortex and form tetrasporangial nemathecia; released tetraspores develop directly into erect thalli. Tetrasporoblastic life histories are characteristic of certain members of the Phyllophoraceae but were previously unknown in the Dumontiaceae. Japanese P. yoshizakii shared 55 and 56% of organellar DNA bands with P. californica and P. pinnata, respectively; phylogenetic analysis indicated equally distant relationships to both species. Pikea yoshizakii or a closely similar species with the same life history occurs in southern California and Mexico.     

TWO NEW SPECIES OF TAYLORIELLA (RHODOMELACEAE,RHODOPHYTA) FROM THE NORTHEASTERN NORTH PACIFIC1

Two new species are recognized in the rhodomelacean genus Tayloriella Kylin: T. divaricata sp. nov. and T. abyssalis sp. nov. These two taxa are distributed in the northeastern North Pacific, the former ranging from Amchitka Island in the Aleutians through southcentral Alaska to northern British Columbia, and the latter ranging also from Amchitka Island through southcentral Alaska and British Columbia into northern Washington. A characteristic of these two species shared with the type of Tayloriella is that the abaxial lateral always overtops the monopodially developed axes in every order of branching. The laterals have little congenital fusion with the parent axes. A common feature in these two species is that the laterals are terminated in a relatively long monosiphonous portion (usually 6 or 7 cells). The relationship of Tayloriella to Pterosiphonia and Pterosiphoniella is discussed.     

MORPHOLOGY, REPRODUCTION, AND THE 18S rRNA GENE SEQUENCE OF PIHIELLA LIAGORACIPHILA GEN. ET SP. NOV. (RHODOPHYTA), THE SO-CALLED 'MONOSPORANGIAL DISCS' ASSOCIATED WITH MEMBERS OF THE LIAGORACEAE (RHODOPHYTA), AND PROPOSAL OF THE PIHIELLALES ORD. NOV.

Pihiella liagoraciphila gen. et sp. nov. (Rhodophyta) is described for a minute endo/epiphyte that is commonly associated with members of the Liagoraceae ( Nemaliales, Rhodophyta). Algae are discoid or subspherical and grow to a maximum diameter of 400 μm. Attachment is via isolated elongate rhizoids that penetrate into the loosely filamentous structure of the host or by a pad of several coalesced rhizoids where the host has a more cohesive cortex. Elongate surface hairs are common. Gametophytes are dioecious, the spermatangia arising on surface cells, and carpogonia with elongate trichogynes borne directly on undifferentiated surface supporting cells. Large sporangia form on stalk cells across the upper surface of the plants, these appearing to be either monosporangial or the result of fertilization of the carpogonia and equivalent to undivided zygotosporangia. Carposporophytes and tetrasporangia are unknown. 18S rRNA gene sequence analyses indicate that Pihiella constitutes a clade of long branch length most closely related to the Ahnfeltiales. The unique morphology and reproduction of Pihiella , combined with a substantial genetic divergence from the Ahnfeltiales, suggest that it is sufficiently distinct to warrant placement in a new family and order. We therefore describe the family Pihiellaceae and the order Pihiellales to accommodate the new genus.     

THE NEW GENUS AND SPECIES OSSIELLA PACIFICA (GRIFFITHSIEAE, RHODOPHYTA) FROM HAWAII AND NORFOLK ISLAND, PACIFIC OCEAN

Ossiella pacifica gen. et sp. nov. is described from sub tidal habitats (5–19 m) in the central (Hawaiian Islands) and southwestern (Norfolk Island) Pacific Ocean. Plants consist of limited prostrate axes that rapidly become erect and form ecorticate, subdichotomously branched axes bearing up to seven (mostly six) di-/to trichotomously branched, pigmented, determinate, quickly caducous whorled laterals from subapical axial cells. Two to five tetrasporangza are borne directly on the inflated basal cells of pigmented determinate laterals. One to two spermatangial fascicles are similarly placed in male plants. Subapical procarps are borne on a three-celled fertile axis, which occupies the same position as an indeterminate branch, being itself displaced laterally by the continued growth of the indeterminate axis. Procarps are bicarpogonial, the sterile cell normally associated with the supporting cell in procarps of the Griffithsieae being converted into a functional four-celled carpogonial branch . Ossiella is compared with the other membus of the tribe, and a dichotomous key to the genera is offered. On morphological grounds , Ossiella seemingly forms a perfect intermediary link along the phylogenetic lineage between the eastern Australian genus Baldockia A. Millar, the eastern American genus Calliclavula G. W. Schneid, and the widespread Anotrichium Nägeli .