CHEMOTAXONOMY IN MARINE ALGAE: CHEMICAL SEPARATION OF SOME LAURENCIA SPECIES (RHODOPHYTA) FROM THE GULF OF CALIFORNIA1

Species separation in the genus Laurencia (Rhodomelaceae, Rhodophyta) is complicated by the high degree of morphological variation within the species. Chemical investigations on a worldwide basis of over 15 species indicate that 1 or more of the halogenated natural products synthesized by Laurencia are unique to each species. Our chemical investigations of Laurencia pacifica, as presently understood from the Gulf of California, indicate that more than 1 species had been included under this name. Thin layer chromatographic (TLC) comparisons of the halogenated components of 3 recognizable forms of “L. pacifica” were completed. The results revealed 3 distinct forms, with halogenated products unique to each form. In each form the observed chemical characters had been previously isolated and identified and could now be positively assigned to their algal source. Comparisons were also conducted with L. pacifica Kylin (1941) from the type locality of the species, La Jolla, California, and revealed that it contained another halogenated product different from those isolated from the Gulf species. We conclude that 3 species of Laurencia have been elucidated in the Gulf of California and these are separate from L. pacifica Kylin. Each species can be distinguished by its characteristic array of halogenated compounds. Comparative thin layer chromatography of the lipid components of morphologically similar Laurencia species should prove to be a useful new taxonomic aid.     

On the absence of previously reported Japanese and Peruvian species of Prionitis (Halymeniaceae, Rhodophyta) in the northeast Pacific

Partial rbc L sequences of type specimens, historically significant specimens and recently collected comparative specimens are used to demonstrate that Prionitis decipiens (Montagne) J. Agardh (type locality: Paita, Peru), Prionitis angusta (Harvey) Okamura (type locality: Shimoda, Japan, currently known as Grateloupia angusta (Harvey) Kawaguchi et Wang) and Prionitis cornea (Okamura) E. Y. Dawson (type locality: Shizuoka, Japan, currently known as Grateloupia cornea Okamura), all previously reported from the northeast (NE) Pacific (California, USA and Baja California, Mexico), were not, and currently are not, present. Historically significant specimens from the NE Pacific specifically annotated by E. Y. Dawson as belonging to P. cornea or specifically annotated by I. A. Abbott as belonging to P. angusta were sequenced and shown to belong to either Prionitis linearis Kylin (type locality: La Jolla, California, USA) or to Prionitis filiformis Kylin (type locality: San Francisco, California, USA). The lectotype of Gelidium decipiens Montagne (basionym of Prionitis decipiens ) is narrowed to a single specimen illustrated by Montagne, and a lectotype is designated for P. linearis . The utility of partial rbc L sequences (less than 200 base pairs) for determining the identity of specimens is discussed, as is the importance of sequencing type and historically significant specimens.     

Chemical variation in a new bromochamigrene derivative from the red seaweed Laurencia pacifica

The secondary metabolite chemistry of the red seaweed Laurencia pacifica Kylin varies geographically. The La Jolla, Calif. population (type locality) contains the sesquiterpene prepacifenol, while L. pacifica collected south of Ensenada, Mexico contains a new bromochamigrene derivative, the structure of which is described herein. Since chamigrene synthesis in Laurencia species has been demonstrated to be reasonably species-specific, L. pacifica may be a mixture of morphologically similar forms.     

Characterization of the edible red alga Meristotheca papulosa (Solieriaceae, Gigartinales) from Japan

The vegetative and reproductive morphology of the edible red alga Meristotheca papulosa (Montagne) J. Agardh (Solieriaceae) was reexamined based on material collected from various localities in Japan. Although the habit of the blades is variable according to the length and width of the axes, the frequency of branching and the abundance of proliferations, rbcL sequence analyses indicate their conspecificity. M. papulosa displays four distinctive reproductive features (presence of an auxiliary cell complex, occurrence of cystocarps on marginal proliferations and the blade surface (although very rare) in addition to the margins of axes, frequent production of spinose outgrowths on the pericarp and tetrasporangial initials typically basally attached to their parental cells) that have not been reported for M. papulosa from other areas. Although these features might warrant recognition of the Japanese entity as a separate species, a better understanding of their possible taxonomic value requires comparisons with M. papulosa from other geographic regions, including the type locality.     

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.     

LIFE HISTORY AND HYBRIDIZATION STUDIES ON GIGARTINA STELLATA AND PETROCELIS CRUENTA (RHODOPHYTA) IN THE NORTH ATLANTIC1

Fourteen isolates of the crustose marine red alga Petrocelis cruenta J. Agardh from various localities in the British Isles, France (including the type locality), Spain and Portugal gave rise in culture to dioecious foliose plants identifiable as Gigartina stellata (Stackhouse) Batters although two isolates formed only sterile foliose blades. A total of 145 isolates of Gigartina stellata were also grown in culture from various localities in the U.S.A. (Maine), the British Isles, Iceland, Denmark, France, Spain and Portugal using both carpospores and vegetative blade apices. Two basic types of life history were found among these isolates: a direct-type life history involving the formation of further foliose plants from carpospores, some isolates of which also form spermatangia on the same papillae as the cystocarps; and a heteromorphic-type in which only crustose plants resembling Petrocelis cruenta are formed from carpospores. Only heteromorphic-type life histories were found from Spain and Portugal. Both life history types were found in plants from the U.S.A., the British Isles and northern France. Only direct-type life histories were found in plants from Iceland and Denmark. Some Petrocelis-like crusts derived from field collected G. stellata carpospores and Petrocelis crusts of hybrid progeny formed tetrasporangia in 8:16 h LD, 10° C but not in 8:16 h LD, 15° C; 16:8 h LD 10° C or 15° C; and 10:6.5:1: 6.5 h LDLD, 10° C. The spores thus formed were viable and produced normal dioecious male and female gametophytes. Short day and low temperature conditions appear necessary for tetrasporogenesis. The results from crossing experiments with 32 male and 27 female isolates of the heteromorphic-type derived from both G. stellata and P. cruenta showed that two virtually non-interbreeding populations with a high degree of geographical separation exist in the north-eastern Atlantic. Morphological differences between plants from each population are described. On the basis of culture and crossing results, Petrocelis cruenta J. Agardh is placed in synonymy with Gigartina stellata (Stackhouse in Withering) Batters.     

The distribution of benthic marine algae

Hitherto only gametangial and carposporangial phases of Thuretellopsis peggiana Kylin have been reported. A study of this rare member of the Dumontiaceae (Cryptonemiales) in culture has disclosed the existence of an encrusting tetrasporangial phase, previously unknown, whilst the life history has been shown to be of the ‘Bonnemaisonia’ type.     

THERMAL ECOPHYSIOLOGY OF LAURENCIA PACIFICA AND LAURENCIA NIDIFICA (CERAMIALES,RHODOPHYTA) FROM TROPICAL AND WARM‐TEMPERATE REGIONS1

Laurencia is a globally distributed genus with about 80 species (order Ceramiales) that inhabit tropical, subtropical, and warm‐temperate regions of both sides of the Atlantic and Indo‐Pacific oceans. This study investigated how two species of Laurencia distributed in different thermal environments (California and Hawaii) varied in their photosynthetic responses to temperature. The thermal ecophysiology of both species was investigated at different temporal scales (short‐term responses and seasonal acclimatization) using oxygen evolution and pulse‐amplitude‐modulated (PAM) fluorometry. Our results indicated that seasonal acclimatization of both species of Laurencia influenced the short‐term photosynthetic response at both locations. Greater seasonal differences in the photosynthetic performance were observed for L. pacifica Kylin, which reflects the ability of this species to acclimatize to local environmental conditions characterized by short‐term fluctuations and a broader annual temperature range. Photosynthetic performance of L. nidifica J. Agardh was consistent with the less variable local environment (no short‐term fluctuations and a narrower temperature range). These results suggest that acclimatization to temperature variability in the environment can influence the degree of flexibility of physiological responses of species in this genus.     

TAXONOMY OF THE FLATTENED SOLIERIACEAE (RHODOPHYTA) IN BRAZIL: AGARDHIELLA AND MERISTIELLA1

Taxonomic studies of an extensive collection of flattened members of the Solieriaceae considered by many authors to be species of Agardhiella, Eucheuma, Meristotheca, and Meristiella lead us to conclude that they represent only two genera in Brazil: Agardhiella, including A. ramosissima (Harvey) Kylin and A. floridana (Kylin) Gabrielson ex Guimarães et Oliveira, and Meristiella, with only one species, M. gelidium (J. Agardh) Cheney et Gabrielson. Accordingly, we propose that 1) Meristotheca gigartinoides Joly et Ugadim be synonymized with Agardhiella floridana, and 2) Meristiella echinocarpa (Areschoug) Cheney et Gabrielson and M. schrammii (P. et H. Crouan) Cheney et Gabrielson be synonymized with Meristiella gelidium.     

STUDIES IN SOME FOLIOSE RED ALGAE OF THE PACIFIC COAST. I. CRYPTONEMIACEXE1

Pacific coast species of the red algae Halymenia and Cryptonemia are described Of the 8 previously reported species of Halymenia 3 (H. abyssicola Dawson, H. megaspore Dawson, H refugiensis Dawson) hive been placed in synonymy with other species. Halymenia hollenbergii from southern California is described as new; H cocinea is transferred from Schizymenia (and includes Aeodes gardneri Kylin) and H. templetonii is a transfer from Weeksia. Five species of Cryptonemia are described; one of them, C. taylorii from the Revillagigedo Archipelago, as new. Wider distribution ranges are given for the remaining species, C. ovalifolia, C. obovata, C. bore alis and C. angustata. Keys to genera of the west coast Cryptonemiaceae and to the species of Halymenia and Crypronemia are provided.     

CULTURE AND HYBRIDIZATION STUDIES ON PETROCELIS (RHODOPHYTA) FROM ALASKA AND CALIFORNIA1,2

Cultured tetraspores of Petrocelis middendorffii (Ruprecht) Kjellman from Amchitka Island, Alaska, gave rise to foliose, dioecious gametophytes similar to cultured gametophytes of P. franciscana Setchell & Gardner. A 1:1 ratio male:female gametophytes was obtained. Fertilized female plants produced cystocarps and carpospores that gave rise to crustose plants anatomically similar to field-collected Petro-celis sporophytes. Cultured male gametophytes of P. middendorffii were interfertile with cultured female blades of field-collected Gigartina pacifica Kjellman. Cultured P. middendorffii gametophytes from Amchitka were interfertile with cultured gametophytes of P. franciscana from 2 localities in California. Hybrid carpospores gave rise to crustose sporophytes that have not reproduced. Anatomical comparisons of P. middendorffii from Amchitka with P. franciscana from California showed no important differences in the characters originally used to separate these species. The interfertility of cultured Petrocelis gametophytes from california and Amchitka as well as the similarities of the history and anatomy suggests that a single species is involved. P. franciscana is reduced to a synonym of P. middendorfii.     

OBSERVATIONS ON THE GENUS BLIDINGIA (CHLOROPHYTA) IN CALIFORNIA1

The green algal genus Blidingia (Ulvales), reported for the first time in California only 4 years ago, is now found to be fairly widely distributed there as B. minima (Nägeli ex Kützing) Kylin var. minima. B. minima var. subsalsa (Kjellman) Seagal is now recorded for California. Ulva vexata Setchell and Gardner [Enteromorpha vexata (Setch. & Gardn.) Doty] is reduced to varietal status as B. minima var. vexata, and is reported, to be found in 2 ecological forms. Forms with expanded, ulvoid blades are common to exposed localities and is the form described by Setchell & Gardner. Smaller, narrower, enteromorphoid forms are found near river mouths and are the kind referred to by Doty. There are many intermediate forms of this variety. Characters which are used to separate Blidingia from Enteromorpha are critically evaluated.     

The life history of Callophyllis firma (Kylin) Norris in laboratory culture

Callophyllis firma (Kylin) Norris has been shown to have a life history in laboratory culture in which the tetrasporangial and gametangial phases are morphologically identical. This is the first published report of such a Polysiphonia-type of life history in a member of the Cryptos nemiales.     

PHYLOGENETIC RELATIONSHIPS WITHIN THE GENUS HYPNEA (GIGARTINALES,RHODOPHYTA), WITH A DESCRIPTION OF H. CAESPITOSA SP. NOV.1

Species discrimination within the gigartinalean red algal genus Hypnea has been controversial. To help resolve the controversy and explore phylogeny within the genus, we determined rbcL sequences from 30 specimens of 23 species within the genus, cox1 from 22 specimens of 10 species, and psaA from 16 species. We describe H. caespitosa as a new species characterized by a relatively slender main axis; a pulvinate growth habit with entangled, anastomosing, and subulate uppermost branches; and unilaterally borne tetrasporangial sori. The new species occurs in the warm waters of Malaysia, the Philippines, and Singapore. The phylogenetic trees of rbcL, psaA, and cox1 sequences showed a distant relationship of H. caespitosa to H. pannosa J. Agardh from Baja California and the marked differentiation from other similar species. The rbcL + psaA tree supported monophyly of the genus with high bootstrap values and posterior probabilities. The analysis revealed three clades within the genus, corresponding to three sections, namely, Virgatae, Spinuligerae, and Pulvinatae first recognized by J. G. Agardh. Exceptions were H. japonica T. Tanaka in Pulvinatae and H. spinella (C. Agardh) Kütz. in Spinuligerae.     

Taxonomic features of the red alga Meristotheca coacta (Solieriaceae, Gigartinales)

Vegetative and reproductive structures of the little‐known red alga, Meristotheca coacta Okamura (Solieriaceae, Gigartinales) were described on the basis of specimens collected from three localities in southern Japan, including an island close to the lectotype locality. Meristotheca coacta was shown to be a strongly supported monophyletic clade in our rbcL analyses. The taxonomic features of this species were reassessed. This species is characterized as follows: (i) the thalli consist of relatively thick (300–1400 μm from the uppermost to basal regions), elastic and rose‐red blades; (ii) the blades are irregularly lobed and branched into variously shaped segments with undulate and crispate margins; (iii) tetrasporangial initials are laterally attached to their parental cells and the lateral pit‐connections remain in mature tetrasporangia; (iv) gametophytes are dioecious; (v) carpogonial branches are three‐celled (occasionally two‐ or four‐celled); (vi) a darkly staining auxiliary cell complex is present and recognizable prior to diploidization; and (vii) cystocarps are produced along (or near) the margins of the blades and on the marginal proliferations and lack spinous outgrowths.     

LATITUDINAL TRENDS IN THE GROWTH AND REPRODUCTIVE SEASONALITY OF DELESSERIA SANGUINEA,MEMBRANOPTERA ALATA,AND PHYCODRYS RUBENS (RHODOPHYTA)1

The seasonality of Delesseria sanguinea, Membranoptera alata, and Phycodrys rubens (Rhodophyta) was studied at Helgoland (North Sea, Germany) and Roscoff (Brittany, France). Plants were collected at bimonthly intervals, and growth and reproduction were monitored. Growth of blades was observed mainly in spring, although small blades were found on plants of M. alata and P. rubens all year round. In summer, plants started to degenerate and in autumn they became fertile. The reproductive season of D. sanguinea lasted from October to February/April at both locations, whereas reproductive plants of M. alata and P. rubens were found until April at Helgoland and until June and August, respectively, in Roscoff_: Lower winter temperatures at Helgoland than at Rascoff may have caused these differences in the duration of the reproductive season. Using published data, the seasonal patterns at Helgoland and Roscoff were compared to those found at other locations (e.g. Barents Sea; Maine, USA; Isle of Man, UK) and local temperature/daylength conditions. Blade growth was synchronized across all populations and occurred in spring, when temperatures were usually still suboptimal for growth. Maximum reproduction was generally found in the colder half of the year but started earlier in autumn in the Barents Sea. Adaptive strategies in the seasonal control of growth and reproduction are discussed. Adequate timing of the history events (e.g. appearance of juveniles in spring) appears more important than maximal growth and reproduction of adults during the season with the most favorable temperatures.     

Tylotus laqueatus,a new species of Dicranemataceae (Gigartinales,Rhodophyta) from the Hawaiian Islands

A rarely collected shallow‐subtidal Hawaiian macroalga has been determined anatomically and molecularly to belong to an undescribed species of Tylotus J. Agardh, the most widely distributed genus of the small, mostly Australian‐endemic family Dicranemataceae. Thalli are repent and imbricate on calcareous boulders at the type locality on O‘ahu, and are anchored both basally and by haptera arising marginally and ventrally on the (sub‐)dichotomous, linear axes. Simple or forked terete haptera can be a means of perennation by the occasional direct issuing of adventitious blades. Fronds are multiaxial and consist of a broad pseudoparenchymatous medulla of thick‐walled cells surrounded on both sides by a two‐ or three‐layered small‐celled pigmented cortex in which numbers of ‘glandular’ hairs are embedded. Tetrasporangia are zonate, and gametophytes are monoecious. Carpogonial branches are three‐celled, directed to the thallus surface, and borne laterally on inner‐cortical supporting cells; cystocarps are prominently protuberant and scattered sparingly on dorsal frond surfaces, the carposporophytes directed outwardly beneath an ostiolate pericarp and connected to the parent gametophyte across a broad placental base in which the remnant auxiliary cell persists centrally. The inner surface of the pericarp is unusual in producing extensive patches or isolated islands of short gonimoblast filaments with terminal carposporangia as an apparent result of the implantation of gonimoblasts into the tissue of the lining. Anatomy indicates that the new species is more closely related to the East‐Asian Tylotus lichenoides Okamura than to the only other described member of the genus, the type species T. obtusatus (Sonder) J. Agardh from southern Australia. An rbcL phylogeny supports placement of sequences for Hawaiian specimens within the genus Tylotus but distinct from all previously recorded sequences of Tylotus. As is widely reported in other molecular‐phylogenetic analyses of the Gigartinales, we find that support for generic and familial relationships within the order is strong whereas that for between‐family relationships is low.     

CENTROCERAS INTERNITENS N. SP. (RHODOPHYCEAE,CERAMIACEAE) FROM THE WESTERN TROPICAL NORTH ATLANTIC OCEAN1

Centroceras internitens n. sp. (Rhodophyceae, Ceramiaceae) was found in the Florida Keys and Bermuda only as an epiphyte on Ceramium nitens (C. Agardh) J. Agardh. It is described on the basis of many collections, including an abundance of reproductive material representing all stages of the life history. Laboratory culture has not been successful. The type locality is a well-known limestone platform on the Atlantic Ocean side of West Summerland Key, Florida Keys. In addition to living material, C. internitens was found on eight herbarium specimens of Ceramium nitens at the United States National Museum, thereby establishing records of its occurrence throughout the Caribbean Sea and dating back to 1887.