New and old problems in the taxonomy of the Gelidiales (Rhodophyta)

The order Gelidiales includes over 140 agar-producing taxa. Many species are taxonomically confused; the boundaries of one family and all but four genera recently have been contested, and the controversy over ordinal status has lasted for over 25 years. This study reviews recent developments in the taxonomy of the group and suggests future areas for studies. The order cannot be defined by a few exclusive characters, as intended in the past, but it shows a unique combination of characters. Additional studies on Gelidiella and Acanthopeltis seem advisable to clarify family limits. The segregation of Onikusa and Pterocladiastrum is doubtful. Only three of six characters discriminate Gelidium from Pterocladia. None allows complete generic segregation and all need variability studies. Analysis of nomenclatural types in the light of morphological variation would permit an understanding of species limits in Gelidium and Pterocladia. Similar studies are needed in Gelidiella and Ptilophora.     

Advances in cultivation of Gelidiales

Currently, Gelidium and Pterocladia (Gelidiales) are collected or harvested only from the sea. Despite several attempts to develop a cultivation technology for Gelidium, no successful methodology has yet been developed. Initial steps towards developmental efforts in Portugal, Spain, South Africa and Israel have been published. More developments have probably been performed but have not been published. Two different technological concepts have been tested for Gelidium cultivation: (1) the attachment of Gelidium fragments to concrete cylinders floating in the sea, and (2) free-floating pond cultivation technology. These vegetative cultivation technologies might be partially optimized by controlling physical, chemical and biological growth factors. The pond cultivation technology is the much more controllable option. The effects of all factors are discussed in detail in this review. It seems that the main difficulty with cultivation of Gelidium is its low growth rate. The claimed yields of the two technologies are far from being economically attractive at this stage of their development. It seems that in order to introduce Gelidium into commercial cultivation, major efforts in genetic improvement through selection or genetic engineering will be required. Only high yield strains will have the potential to compete economically with the present harvesting tradition. However, accumulated experience with genetic improvement of other useful seaweed species suggests that this is possible.     

Gelidiaceae (Rhodophyta) in Bahía de Banderas,Western Pacific,México

We present the results of a floristic study of intertidal species of Gelidiaceae collected in ten localities in the Bahía de Banderas region. The study area is within an extensive overlapping transitional zone between the tropical and subtropical regions of the Mexican Pacific coast. Species of the genera Gelidium and Pterocladia were found, G. microdentatum and G. pusillum being the most abundant and widely distributed in the Bay. The local and Mexican Pacific distribution for each species is given. The number of species found in this study is compared with the numbers reported in the literature for the Mexican Pacific.     

Architectural models for apical patterns in Gelidium (Gelidiales, Rhodophyta): hypothesis of growth

The great morphological variation in the apical region of main axes and branches in species of Gelidium and Pterocladia suggests different patterns of growth. Those patterns have been reconstructed from the ontogenetic sequence of morphological stages and regeneration of the apex. Using the architectural approach, models for four species of Gelidium from the Pacific coast of central Chile and the Pacific coast of Mexico's tropical region, were constructed based on the following criteria: growth pattern, specialization of laterals, symmetry, apical dominance and rhythm of apical and lateral growth. Those models allow for a formal evaluation of apical differences between species.     

Marine plant harvest in Portugal

This work brings together the scattered information on marine plant harvests and the colloid extraction industry in Portugal, as an initial contribution to the improvement of resource management. The first phase of exploitation of marine plant resources started prior to the 14th century, with the gathering and sale of storm-tossed seaweeds for fertilizer. The harvest of seagrasses and algae at Ria de Aveiro was of great economic importance. The second phase of resource exploitation began with the wider scale harvest of agarophyte species for colloid extraction. Portugal is at present the third largest harvester of the agarophytes Gelidium and Pterocladia (2500 t annually), and it is the fifth largest agar producer (350 t annually). Other colloid-producing species, including Chondrus crispus and Mastocarpus stellatus, are also harvested for export. The total agarophyte landings, agar production and income from agar exports is far below the maximum levels attained in the early 1970s. The status of stocks in each different harvest zone on the continental coast and the Azores is examined. Although there is an effective management structure for the Portuguese marine plant resource, research is needed to provide a sound biological basis for management. author for correspondence     

The Structure and Development of the Thallus in the British Species of Gelidium and Pterocladia

The apical structure and the development of the thalli of allthe British species of Gelidium and Pterocladia have been investigated;the development of G. pulchellum is described in detail. Eachaxis is terminated by one or more apical cells, which by theirsegmentation form the tissues of the thallus. An axial filamentis distinct for a short distance behind each apical cell, butsecondary pit-connexions develop rapidly so that in sectionthe mature axis has the appearance of a multi-axial structure. Lateral branches of the frond develop by the segmentation ofthe lateral branch apical cells, which are formed by the transformationof superficial cortical cells, either in the meristematic ormature parts of the axes. The extreme variability of externalappearance is due principally to the indeterminate origin ofall lateral branches. The thallus in the British species of Gelidium and Pterocladiaconsists of erect fronds borne on creeping axes. The relativeproportions of the frond and creeping axes in various speciesand their survival through adverse conditions are discussed.     

Artificial sporeling and field cultivation of Gelidium in China

The supply of Gelidium resources is dependent on collection from natural habitats. As these resources are very limited, one possible means of augmenting production is by implementation of artificial cultivation as has been done with Laminaria, Porphyra and Eucheuma. Chinese phycologists have conducted research on sporeling and field cultivation of Gelidium for many years. Trials have been made using: (1) raft cultivation based on vegetative propogation of thallus fragments, (2) cultivation based on spore-collecting and land-based tank culture to provide seedstock, (3) cultivation based on regeneration from small thallus fragments to provide seedstock. Some of these results have been promising, but development remains at the experimental stage and methods of cultivation need to be improved. This paper is an updated review of the research and development on artificial sporeling production and field cultivation of Gelidium in China.     

Medullary structure differences in Gelidium and Pterocladia (Gelidiales, Rhodophyceae): Taxonomic meaning

Study of the apical portions of vegetative thalli of 21 species of the genera Gelidium and Pterocladia (Gelidiales-Rhodophyta) revealed the existence of six patterns of medullary structure. Each pattern is characterized by the organization of the medullary tissue and the changes in cellular dimensions from apex to subapex. These characters can be used to separate groups of species within each genus.     

Phylogenetic Diversity of New Zealand Gelidiales as Revealed by rbcL Sequence Data

Diversity and phylogenetic relationships of New Zealand representatives of the red algal order Gelidiales have been examined using rbcL sequence data. Extensive field collections have been made from throughout the New Zealand region. Six genera have been reported previously from New Zealand (Capreolia, Gelidium, Pterocladia, Pterocladiella, Pterocladiastrum, Ptilophora). This research has revealed species with very restricted local distributions, as well as the discovery of several undescribed, cryptic taxa. The common and widespread Gelidium caulacantheum is confirmed to be more closely related to Capreolia than to other species of Gelidium. The generic concept of Capreolia, based on life history characters, will need to be modified to accommodate additional species possessing “Gelidium” life histories. A species endemic to New Zealand, Gelidium ceramoides, has been found to differ significantly from all other members of the Gelidiales and requires reclassification in another genus and order. Examination of field collections and herbarium specimens in addition to molecular sequence data have led us to conclude that specimens previously placed in the genera Ptilophora and Pterocladiastrum belong within Pterocladia lucida.     

Chemical structure and quality of agars from Gracilaria

Agar polymers synthesized by species of the genus Gracilaria constitute a complex mixture of molecules, containing several extremes in structure. Sulphate hemi-esters, methyl ethers and pyruvic ketals can alter in a number of ways the structural regularity of agar based on strictly 3-O-linked β-l-galactopyranose and 4-O-linked α-l-galactopyranose residues. In comparison with agars from Gelidium and Pterocladia, agars from Gracilaria can have higher degrees of sulphation, methoxylation and pyruvylation. The gelling ability of agars from most of Gracilaria species is considerably improved by adopting, before extraction, an alkali pretreatment which converts α-l-galactose 6 sulphate into 3,6-anhydro-α-l-galactose. Native agars obtained from Gracilaria cannot be classified, with few exceptions, as bacteriological grade agar as they have a high content of methoxyls and consequently high gelling temperatures. On the contrary, the genus Gracilaria is considered the most important source of food and sugar-reactive grade agars. Among techniques which can be used to study algal polysaccharides, combined ¹H and ¹³C nuclear magnetic resonance spectroscopy represent the most effective and powerful method for the investigation of the chemical structure of agarocolloids.     

The harvesting of macroalgae in New Zealand

Several species of algae have been commercially harvested in New Zealand, mainly for extraction of agar and alginates. In the past, the harvest was comprised mostly of shore-cast plants. There has been more recent interest, however, in harvesting attached plants of Pterocladia spp., Porphyra spp., Gracilaria sordida, Durvillaea spp., Macrocystis pyrifera, and Ecklonia radiata. The ecological effects of harvesting attached algae depend largely on the sizes of plants, the season of removal, the patch size of clearances, and the proximity and identity of mature plants. These have not been well-studied for seaweeds in New Zealand, but population and life history studies indicate that harvesting methods affect the continuity of algal resources, at least on a local scale, and are crucial factors in their management.     

Three new <Emphasis Type="Italic">Ophiostoma</Emphasis> species isolated from Japanese red pine

Three new species of Ophiostoma found on Japanese red pine are described as Ophiostoma pusillum sp. nov., O. botuliforme sp. nov., and O. nigrogranum sp. nov. Ophiostoma pusillum is characterized by oblong ascospores and a Hyalorhinocladiella anamorph. Ophiostoma botuliforme has ostioles covered with a hyaline gelatinous cap, allantoid ascospores, and a Pesotum anamorph with hyaline to pale brown stipes. Ophiostoma nigrogranum has hyaline ostiolar hyphae with rounded tips, allantoid ascospores, and sclerotium-like structures.Contribution no. 172, Laboratory of Plant Pathology and Mycology, Institute of Agriculture and Forestry, University of Tsukuba     

Observations on Flattened Species of Gracilaria (Gracilariaceae, Rhodophyta) from Taiwan

Four flattened Gracilaria species have been reported from Taiwan: G. spinulosa, G. vieillardii, G. textorii and G. punctata, identified based on branching pattern, the presence or absence of spines, and characters that often vary seasonally. Gracilaria spinulosa was originally described from the type locality, Tainan. Species with toothed margins are usually referred to G. “vieillardii”; those with smooth margins to G. “textorii”, and those with smooth margins and dark spots scattered over the blade to G. “punctata”. Molecular analyses show that specimens with marginal teeth cluster in three different groups: a G. “vieillardii” clade, a G. spinulosa clade, and a clade sister to G. spinulosa. An undescribed species comprises the third clade, which is distinguished by its relatively large gonimoblast cells and weakly developed tubular nutritive cells. The three clades can be separated by the character of the tubular nutritive cells, the size of gonimoblast cells and certain vegetative features. Plants with entire margins form a single clade characterized by cystocarps with basal tubular nutritive cells and their absence in the cystocarp cavity. They are nested in the Hydropuntia complex and are referred to as Gracilaria “punctata” here. The records of G. textorii and G. punctata from Taiwan require reinvestigation in comparison with the Japanese species.     

Production ecology of Gelidium

Available data on determinants of production in species of Gelidium suggest several general patterns. Species diversity is higher in tropical latitudes, whereas in temperate latitudes the size of the fronds is larger, the species are ecologically dominant and commercially viable. Typically, the species occur on rocky substrate, often on coralline crusts, associated with rapid water movement and arranged in successive belts that can extend down to 25 m depth. Yields vary among species, to a maximum of 2.0 kg m^–2 y^–1. Growth and production in many species can best be explained by complex interactions between irradiance and nutrients. Temperature can interact synergistically with irradiance, while water movement compensates for nutrient limitations. Increased water movement or the addition of nutrients can prevent, to an extent, bleaching by high light and high temperature. Available data suggest the existence of at least eight biological factors affecting predicted productivity of Gelidium crops: morphology, age of the frond, thallus part, reproductive state, seasonality, crop density, life history phase and geographic and ecological origin of the species. At least four events can remove or destroy Gelidium crops: extreme low tides, storms, grazing and careless harvesting. Only the last-named factor has been analyzed over more extensive experimental periods.     

中国四川报春花属一新种

报道了中国四川报春花属(Primula L.)一新种——小繸瓣报春(P.hydrocotylifolia G.Hao,C.M.Hu&Y.Xu)。小繸瓣报春植株无毛、无粉,叶片多少呈圆形,具纤细的叶柄,开花期叶丛基部无鳞片,球形蒴果藏于宿存花萼,成熟时裂成不规则碎片,显然属于脆蒴报春组(sect.Petiolares Pax)革叶报春亚组(subsect.Chartacea W.W.Sm.&Forrest),并与该亚组的川西繸瓣报春(P.veitchiana Petitm.)最为接近,但后者植物体的各部分均远大于此新种,花序多花,花亦较大,二者之间无过渡类型。     

Biological criteria for the exploitation of the commercially important species of Gelidium in Spain

Exploitation of the commercially-important species of Gelidium in Spain (G. sesquipedale and G. latifolium) and development of the industry occurred after World War II, as a consequence of their use as resources for the extraction of agar. This resulted in the implementation of several harvesting methods, the most important of which is the gathering of cast seaweeds, both from the shore and the sea. From the very beginning, direct exploitation of these species (i.e. plucking) was controversial because of possible adverse ecological effects. Consequently, several biological and ecological studies of both species of Gelidium were begun. This included such aspects as growth, biomass production, productivity, reproduction, regeneration capacity and agar yield. Recently, the growing interest in using and conserving this resource has led to increased knowledge of the biology and ecology of these species. In this paper, we provide an overview of the research carried out by different Spanish groups and suggest guidelines for rational management of these resources.     

Current state of seaweed resources in Spain

Commercial seaweeds in Spain are harvested on the north and northwest coasts. They are mainly agarophytes and carrageenophytes (Gelidium spp. and some Irish moss-like species, respectively), although some Phaeophyceae species (Fucus spp. and Laminaria spp.) are also exploited for alginates. No industrial seaweed cultivation is carried out in Spain at present. Spain's total commercial seaweed harvest can be estimated at 6,528 ± 2,076 t dry wt year^–1. Gelidium spp. are by far the most harvested, attaining 5,135 ± 1,761 t dry wt y^–1. To date, Spain's commercial seaweed harvest is low compared with that of other countries, but data on field standing crops and productivities of commercial taxa suggest that harvesting could be increased greatly.     

Distribution and Recent Reduction of Gelidium Beds in Toyama Bay, Japan

The distribution and recent reduction of Gelidium beds, i.e. mat-like beds dominated by the agarophyte G. elegans Kützing in Toyama Bay (Sea of Japan), in which 95% of the coastline is protected artificially, are reported. Gelidium beds were common in shallow waters (usually < 10 m deep); most of the large beds (> 1 ha) were restricted to the inner coasts of the bay. In calm and eutrophic areas, however, G. elegans was heavily colonized by epiphytes. In the last decade, two beds were buried in situ and beds in their vicinity were damaged by the stagnation of coastal water and/or sedimentation by silts which accompanied land reclamation. At the other two beds monitored since 1988, Gelidium declined a few times but most prominently in 1998, when episodic long summer rain was recorded. This is the first report, not only on the current status of Gelidium beds other than for the central Pacific Coast of Honshu in Japan, but also concerning reduction of the beds caused by both anthropogenic and natural events.     

Mating System and Clonal Architecture: A Comparative Study in Two Marine Angiosperms

In the present study, we compared the clonal architecture between two seagrass species, the dioecious Cymodocea nodosa and the hermaphroditic, self-compatible Zostera noltii, in order to verify the hypothesis that clonal growth strategies and resulting genet architecture are associated with mating system in clonal plants. It is expected that self-incompatible species should be associated to a guerrilla growth form, because of pollen limitation due to obligate outbreeding, while the ecologically advantageous phalanx strategy can be adopted in self-compatible species. Genotypic diversity and heterozygosity were also assessed in the two species. Sampling has been conducted in mixed stands, collecting shoots of the two species at the same points of the sampling grid, in order to even out any effects of environmental heterogeneity. Species-specific microsatellite loci have been used as molecular markers to identify clones and assess their spatial distribution in both species. As expected, we found an intermingled configuration of genets in the dioecious C. nodosa while Z. noltii was characterized by a clumped, `phalanx-type' distribution of clones. C. nodosa was characterized by higher genotypic diversity with regard to Z. noltii, while heterozygosity levels were comparable in the two species. Coordinating Editor: Dr J. Tuomi     

Gelidium pristoides in South Africa

Gelidium pristoides has been harvested commercially from the eastern Cape, South Africa, since 1951, with 40–80 t y^–1 (dry wt) collected in recent years. This species has been intensively studied since 1983, and we briefly review knowledge of its biology in relation to harvesting. We describe a new study of intertidal epiphytic animals, showing that none is specific to G. pristoides, and that only 2.8% of these animals (numbers) inhabit this agarophyte, while the rest are found in other intertidal algal communities: harvesting is considered to have negligible effects on epifauna. Over the past 3 y, we have monitored, at two sites, the effects of the harvesting of G. pristoides on other benthic algae and animals. In only two of the seven main components analysed, did we find any difference between harvested and control plots. At one site only, the number of limpets and percentage cover of Gelidium was higher in harvested plots. These results show that harvesting has no significant biological effect. Regulations governing seaweed exploitation in South Africa were amended in 1988, to encourage local processing of products, and these changes are discussed in relation to the local Gelidium industry. Despite experimental results predicting a higher yield per unit effort if harvesting is limited to summer, harvesting continues throughout the year for practical reasons.