The life history of Ahnfeltiopsis paradoxa (Gigartinales, Rhodophyta) in laboratory culture

The life history of the red alga Ahnfeltiopsis paradoxa (Suringar) Masuda (Phyllophoraceae, Gigartinales) from Japan was completed in laboratory culture. Carpospores isolated from field-collected plants germinated to form circular crusts that were composed of a monostromatic hypothallium consisting of radiating filaments, a polystromatic perithallium consisting of tightly coalescent erect filaments, and hypobasal tissue derived from the hypothallium. The crusts were induced to sporulate by transferring them from short-day to long-day regimes at 15° and 2°C. Each crust produced several nemathecia along 1-4 concentric rings. Intercalary, cruciately or decussately divided tetrasporangia were formed in 4-6 (1-2 at the margin of the nemathecium) successive cells of a single filament of the nemathecia. Tetraspore germlings gave rise to basal discs from which upright axes developed. The upright axes first grew without branches or were sparsely branched and later bore many marginal reproductive proliferations. Procarps and spermatangia were formed in the proliferations on different individuals. Carposporophytes developed on female plants that were co-cultured with male plants. Gonimoblast filaments were formed from an auxiliary cell that fused with a carpogonium. Carposporangia developed from gonimoblast filaments and medullary cells contacted by the gonimoblast filaments. Carpospores were discharged through carpostomes formed in the thickened cortex. Tetraspores were cultured from field-collected crusts of a morphology similar to that of cultured tetrasporophytes. They gave rise to upright gametophytic axes similar in morphology to this species as seen in the field.     

CULTURE AND HYBRIDIZATION STUDIES ON GIGARTINA PAPILLATA (RHODOPHYTA)1

The foliose red alga Gigartina papillata (C. Ag.) J. Ag. was studied in culture to determine its life history and possible relationship to the life history of Petrocelis middendorffii (Ruprecht) Kjellman. Carpospores cultured from individual female plants gave rise to either crustose Petrocelis-like plants that reproduced by tetraspores, or to another generation of foliose female (cystocarpic) plants that reproduced by carpospores. Apices cultured from blades of individual field-collected female plants produced either papillae with many procarps that developed cystocarps only when crossed with male plants, or papillae with few procarps that produced cystocarps in the absence of male plants. The results are interpreted to demonstrate that two types of life history occur in G. papillata: one, a sexual life history involving a crustose tetrasporophyte; the other, a possibly apomictic life history involving only cystocarpic plants. Hybridization experiments demonstrated, that G. papillata is interfertile with Gigartina-phase gametophytes cultured from tetraspores of P. middendorffii. Sexual plants of G. papillata are postulated to represent the naturally-occurring gametophyte of P. middendorffii in California. The possible relationships of the sexual and apomictic plants of G. papillata are discussed.     

On the life history ofTurnerella mertensiana (Rhodophyta,Gigartinales)

Turnerella (Gigartinales) withT. mertensiana known from northern Japan was studied to determine its life history. Carpospores cultured from foliose female plants gave rise on germination to crustose plants containing tetrasporangia, as noted previously inT. pennyi from the Atlantic. The crusts were slow to develop and required 3–5 years to achieve reproductive maturity. Tetraspores liberated in culturedT. mertensiana gave rise on germination eventually to thalli similar to gametophyticT. mertensiana. Thus this species may be said to adhere to the pattern of life history in which a large, foliose gametophyte alternates with a small crustose tetrasporophyte. The erect filaments of the small sporophytes branch laterally and tetrasporangial mother cells are found there, exemplifying an unusual method for crustose species to produce tetrasporangia.     

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.     

A MOLECULAR AND MORPHOLOGICAL ANALYSIS OF THE GYMNOGONGRUS DEVONIENSIS (RHODOPHYTA) COMPLEX IN THE NORTH ATLANTIC1

The Gymnogongrus devoniensis (Greville) Schotter complex in the North Atlantic Ocean was elucidated by comparative molecular, morphological, and culture studies. Restriction fragment length patterns and hybridization data on organellar DNA revealed two distinct taxa in samples from Europe and eastern Canada. Nucleotide sequences for the intergenic spacer between the large and small subunit genes of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and the adjoining regions of both genes, differed by 12.5–13.4% between the two taxa. One of the taxa, which included material from the type locality of G. devoniensis at Torbay, Devon, England, was taken to represent authentic G. devoniensis. Within this taxon, samples from Ireland, England, northern France, northern Spain, and southern Portugal showed great morphological variation, particularly in habit, but their Rubisco spacer sequences were identical or differed by only a single nucleotide. Constant morphological features included the development, from a single auxiliary cell, of the spherical cystocarp with a thick mucilage sheath that appears to be typical of Gymnogongrus species with internal cystocarps. Two life-history types were found. Northern isolates underwent a direct-type life history, recycling apomictic females by carpospores, whereas the Portuguese isolate followed a heteromorphic life history in which carpospores gave rise to a crustose tetrasporophyte. The second group of samples, from Nova Scotia and Northern Ireland, provisionally referred to as Gymnogongrus sp., showed little morphological variation. The life history in both areas consists of apomictically reproducing diploid female gametophytes and diploid crustose bisporophytes and tetrasporophytes. Rubisco spacer sequences of the samples were identical, and the plasmid previously described in the Nova Scotian samples was also present in the Northern Ireland population. This species is widely distributed in the western Atlantic, from Newfoundland to Massachusetts. In Europe, gametophytes are known only at one site, but crusts are distributed from Denmark, Scotland (and probably Norway) to France. It is very likely that this species was introduced from one side of the North Atlantic to the other by shipping during the early nineteenth century. Several morphological features are unusual within the genus but are shared with G. leptophyllus J. Agardh from the eastern Pacific Ocean, and further work is necessary to determine whether Gymnogongrus sp. and G. leptophyllus are conspecific.     

The life history of Petrocelis franciscana

Tetraspores from Petrocelis franciscana Setch. et Gardn. collected at Rockaway Beach, San Mateo County, California, 11 January 1971 were isolated into unialgal culture with Provasoli's enriched seawater medium at 10° and 15° C, 16:8 daily photoregime, 2000–2500 lx cool white fluorescent lighting. The germlings developed as crustose discs with marginal meristems until about 300–500 µm in diameter when erect multiaxial blades were established. The erect plants grew to reproductive maturity in 15° C but not in 10° C. By the fifth month of growth the erect axes flattened and branched dichotomously. At the end of 7 months the plants were reproductively mature. These cultured plants were morphologically similar to Gigartina agardhii Setch. et Gardn. (subgenus Mastocarpus) which occurs abundantly at Rockaway Beach. Male plants were non-papillate with spermatangia forming on the entire blade surface except at the growing tip and near the stipe. The female plants developed papillae which bore many procarps. Many of these procarps were penetrated and seemingly destroyed by intrusive filaments from adjacent vegetative branches but apparently a number were functional since a number of cystocarps developed on plants in both stationary and shake culture after 5 months. Some viable spores were released and germinated.     

CULTURE STUDIES ON THE RELATIONSHIP BETWEEN SCHIZYMENIA AND HAEMATOCELIS (GIGARTINALES,RHODOPHYCEAE) FROM THE PACIFIC COAST OF NORTH AMERICA1

Carpospores from Schizymenia pacifica (Kylin) Kylin (Gymnophlaeaceae) from California formed crusts anatomically identical to Haematocelis rubens J. Agardh (Cruoriaceae). Tetraspores of H. rubens from Monterey, California, and Baja California, Mexico, germinated to form basal discs from which arose upright multiaxial blades with a filamentous medulla and cortical gland cells. Pro-carps and spermatangia were present on the same blades; subsequently, cystocarps characteristic of Schizymenia pacifica developed. Re-examination of herbarium specimens suggests that the foliose tetrasporangial phases previously reported as S. pacifica are referable to Halymenia, Dilsea, Cryptonemia, or Turnerella. Schizymenia pacifica (type locality: San Juan Islands, Washington) thus is considered to be the gametophyte in the life history of Haematocelis rubens (type locality: Brest, France), which has also been reported to be the tetrasporophyte of S. dubyi (Chauvin ex Duby) J. Agardh (type locality: Cherbourg, France). Atlantic and Pacific gametophytes and tetrasporophytes are anatomically very similar, suggesting that only one species is involved, but critical studies must be made before a decision on this taxonomic question can be reached. Haematocelis zonalis Dawson et Neushul (type locality: Anacapa Island, California) is considered to be a growth form of the tetrasporangial phase of S. pacifica.     

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.     

FIELD AND CULTURE STUDIES ON GIGARTINA AGARDHII (RHODOPHYTA)1,2

Cultured carpospores from field-collected plants of Gigartina agardhii Selfchell & Gardner gave rise to either Petrocelis-like crustose plants, or basal discs with erect Gigartina-like blades. Made tips excised from field-collected female plants and cultured in the laboratory also showed two patterns of reproduction: procarpic papillae that only produced cystocarps in the presence of male plants with spermatid, or cystocarpic papillae that formed and released carpospores in the absence of male plants. The first pattern in each case above is evidence for a sexual life history and the second for an apomictic life history similar to that for G. papillata (C. Agardh) J. Agardh. The resorcinol lest for k-carrageenan was negative for Petrocelis-like plants and positive for foliose plants providing addition evidence that the former are haploid and the latter diploid as is characteristic for the Gigartinaceae. Crossing experiments between sexual male and female isolates of G. agardhii from several locations indicated that free interbreeding occurs. All crosses between G. agardhii and Gigartina-phase gametophytes cultured from tetraspores of Petrocelis middendorffii (Ruprecht) Kjellman were negative demonstrating that G. agardhii is reproductively isolated from G. papillata/P. middendorffii and represents a distinct species. The tetrasporophytic phase of G. agardhii in nature is still not known. The significance of reproductive isolation in Gigartina subgenus Mastocarpus is discussed. On the bosis of nomenclatural priority G. agardhii Setchell & Gardner 1933 is placed in synonymy under G. jardinii J . Agardh 1876.     

THE LIFE HISTORIES OF RHODOCHORTON PURPUREUM AND R. TENUE IN CULTURE

The life histories of the red algae Rhodochorton purpureum and R. tenue were studied in unialgal culture. Telrasporophytes produced sporangia in short day regimes (8–12 hr) with 50–200 ft-c cool white light at 10 or 15 C. Mature gametophytes were not observed in the Washington and Alaska clones of R. purpureum. Tetraspores from R. tenue and the California clones of R. purpureum give rise to unisexual gametophytes that are reproductive when smaller than the tetrasporophytes. The tetrasporophytes develop directly from the gonimoblast cells. Because of similarities in morphology and life histories, it is proposed that R. tenue be placed in synonomy with R. purpureum.     

GEOGRAPHIC VARIATION IN THE LIFE HISTORY OF MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Variation in the geographic distribution of the life histories of Mastocarpus papillatus was investigated. Carpospores were isolated from 377 female gametophytes collected from eight localities on the Pacific coast of Baja California, Mexico, and California, U.S.A., and grown in laboratory culture. All carpospores from a single female gave rise either to basal discs with gametophyte-like uprights or crustose plants formerly referred to the genus Petrocelis. Early stages in the development of each type of germling were observed, and environmental factors affecting development were suggested. Based on carpospore germlings, females from each location were scored as having either the 1) sexual life history (crustose germlings) or 2) direct-development life history (discoid germlings with uprights). All females from the two southernmost locations in Baja California exhibited the sexual life history. In the three locations from the central-southern California coast, 70-95% of the females exhibited the sexual life history and the remainder exhibited the direct-development life history. In two of the three populations from the central-northern California coast, 70-90% of the females exhibited the direct-development life history and the remainder the sexual life history. In the third location from the central-northern California coast, the northernmost location sampled in the current study, 60% of the females exhibited the sexual life history and 40% the direct-development life history. The relative ecological advantages and disadvantages of the life histories are unknown as are the environmental factors that produced the ratios of sexual to direct-development females observed at each location.     

WAERNIA MIRABILIS GEN. NOV., SP. NOV. (DUMONTIACEAE,GIGARTINALES): A NEW NONCORALLINE CRUSTOSE RED ALGA FROM THE NORTHWESTERN ATLANTIC OCEAN AND ITS RELATIONSHIP TO GAINIA AND BLINKSIA1

Waernia mirabilis Wilce, Maggs et Sears, gen. et sp. nov., is described for an encrusting red alga endemic to the northeastern North Atlantic Ocean. Its perennial thalli form major understory populations that may outcompete Corallinaceae and Hildenbrandiaceae in some habitats. Crustose red algae with isomorphic gametophytes and tetrasporophytes other than Corallinaceae are poorly known; Waernia differs vegetatively and reproductively from any other such algae. Gametangia are similar to those of the monotypic uncalcified crustose genera Blinksia and Gainia endemic to the Antarctic and central coastal California, respectively. Comparison of Waernia with type material of Blinksia californica Abbott & Hollenberg and Gainia mollis Moe shows that it differs from Blinksia in lacking cohesive crusts in which adjacent cells cohere by direct fusions and by its strongly recurved, rather than straight, carpogonial branches. Gainia and Waernia both have strongly recurved carpogonial branches and laxly constructed crusts that lack secondary pit connections. Gainia differs from Waernia in its considerably larger and thicker crusts, non‐nemathecial tetrasporangia, shorter carpogonial branches, and its consistently single gonimoblast initial. We continue to recognize the family Blinksiaceae as a probable member of the Gigartinales with uncertain affinities. Waernia is assigned to the Dumontiaceae (Gigartinales), as its first described isomorphic crustose member. It is also the only genus and one of few algal species endemic to northeastern North America. Waernia either arrived at its present distribution via migration from a Tertiary polar ocean or evolved from an unknown ancestor in a newly formed boreal northwest Atlantic Ocean during post‐Oligocene millennia.     

MORPHOLOGY AND DEVELOPMENT OF AHNFELTIA PLICATA (RHODOPHYTA): PROPOSAL OF AHNFELTIALES ORD. NOV.1

Ahnfeltia plicata (Hudson) Fries, the type species of Ahnfeltia Fries, is currently assigned to the Phyllophoraceae (Gigartinales). Several morphological and biochemical characters distance A. plicata from the Phyllophoraceae but, because sexual reproduction has never been demonstrated, an alternative placement has not been possible. A. plicata now is shown to have a heteromorphic sexual life history. Erect branched gametophytes are dioecious. In male sori, spermatangia are cut off transversely from spermatangial mother cells. Female sori form numerous terminal sessile carpogonia. Following fertilization, several zygotes in each sorus fuse facultatively with undifferentiated intercalary cells of the female sorus and cut off gonimoblast initials obliquely outwards. These initials give rise to branching gonimoblast filaments that fuse with apical and intercalary female sorus cells and with each other, then grow radially outward in the compound external carposporophyte and terminate in carposporangia. Carpospores develop in culture into crustose tetrasporophytes identical to Porphyrodiscus simulans Batters. Field-collected P. simulans tetraspores grew into erect A. plicata axes. Tetrasporangia are formed by division and enlargement of crust apical cells followed by sequential enlargement and maturation of tetrasporocytes in an erosive process. Monosporangia are formed in sori on male gametophytes. Pit plugs of both gametophyte and tetrasporophyte phases consist of naked plug cores without cap layers of membranes. Gametophytes exhibit both cell fusions and secondary pit connections whereas tetrasporophytes form cell fusions but lack secondary pit connections. On the basis of the unique female and postfertilization reproductive development and in conjunction with the pit plug structure which is unique among florideophytes, the order Ahnfeltiales, containing the family Ahnfeltiaceae, is proposed.     

UNUSUAL PHENOMENA IN THE LIFE HISTORIES OF FLORIDEAE IN CULTURE1

Several Florideae grown in natural seawater media under defined laboratory conditions have interesting and unusual life histories. Antithamnion occidentale males of one generation produced tetraspores that gave rise to nonsporangiate males. The functional females of A. pygmaeum developed spermatangia and tetrasporangia; the tetraspores formed new females. Antithamnion defectum tetrasporophytes of one generation bore spermatangia in addition to tetrasporangia; the tetraspores gave rise to typical gametophytes. Tetraspores from successive generations of Callitham-nion sp. developed into tetrasporophytes and males but no females were produced. Functional female gametophytes of Platythamnion sp. bore abortive tetrasporangia. Field-collected plants of two species of Fauchea produced tetraspores that yielded additional sporangiate plants: those of F. pygmaea being bispo-rangiate and tetrasporangiate, and those of F. lacini-ata being strictly tetrasporangiate. Male plants of Pleonosporium vancouverianum from a running seawater table bore spermatangia and polysporangia when collected. The same plants in unialgal culture produced only spermatangia.     

The occurrence of a Porphyrodiscus simulans Batt. phase in the life history of Ahnfeltia plicata (Huds.) Fries

Plants of Ahnfeltia plicata (Huds.) Fries collected from various localities in the North Atlantic were found to have arisen from basal tetrasporangial crusts identifiable as Porphyrodiscus simulans Batt. Tetraspores from the Porphyrodiscus crusts were cultured for the first time and were found to give rise to discs morphologically similar to those produced by the monospores of Ahnfeltia plicata.     

THE LIFE HISTORY OF POLYSIPHONIA DENUDATA (DILLWYN) KÜTZING IN CULTURE1

The sequence of somatic phases in the life history of the red alga Polysiphonia denudata has been determined in unialgal culture, using supplemented seawater media. Tetraspores gave rise to sexual plants bearing either antheridia or carpogonia in 50 days. Fertilization occurred within 1 week and the fertilized carpogonium developed to produce the mature carposporophytic phase in approximately another week. Carpospores formed by the carposporophyte germinated to produce the tetrasporangium-bearing phase, which released tetraspores in 44 days, thus completing the life history in 3^1/2% months. No deviations from this pattern have so far been observed.     

Comparative reproductive patterns in culture of differentGigartina subgenusMastocarpus andPetrocelis populations from northern Japan

Samples of theGigartina pacifica-ochotensis complex were collected at 21 localities around Hokkaido and northern Honshu. The carpospore and blade tip cultures showed 3 reproductive patterns. (1) 237 (86.8%) of the 273 cultured isolates derived from single plants have a direct type of life history. (2) 29 (10.6%) isolates exhibited a heteromorphic type with the alternation of foliose gametophytes and crustose tetrasporophytes. (3) 7 (2.6%) isolates showed a mixed pattern in which carposporelings developed intoPetrocelis-like crusts, basal discs with uprightGigartina blades, or chimera-like discs with compositePetrocelis-Gigartina anatomy. CulturedGigartina blades derived from bothG. pacifica andG. ochotensis were similar in morphology. In 18 cultures from 5 localitiesPetrocelis tetraspores developed into dioeciousGigartina gametophytes. A single tetrasporeling grew into aGigartina plant that reproduced directly. In hybridization experiments with 8 male and 14 female isolates from 4 localities on Hokkaido 85 (78.0%) of 109 were positive. On the basis of these and earlier studies it is concluded that a single species is present in northern Japan:G. pacifica Kjellm. has priority overG. ochotensis (Rupr.) Rupr. ex Yendo.     

LIFE HISTORY OF BONNEMAISONIA CENICULATA (RHODOPHYTA): A LABORATORY AND FLELD STUDY1, 2

Cystocarpic and spermatangial plants of rarely reported red alga Bonnemaisonia geniculata Gardner, epiphytic on Odonthalia Aoccosa (Esp.) Falk, were collected from june to September 1975 at shell Beach, california. Carpospores inoculated into unialgal culture divided, upon germination, in to two daughter cells, both of which formed erect and rhizoidal axes, Erect axes were uniseriale and alternately branched with a distictive zigzag pattern of axial cells. No tetrasporangia developed in culture. The presumptive tetrasporangia developed in culture to a described genus. Plants morphologically similar to those cultured from carpospores were found at the collection site; they bore tetrasporangia from February to june. Cullured letrasporews gave rise to male and female plants similar to those of field-collected B. geniculate in ca. a I:I ratio. Fertile female plants in the presence of male plants formed cystocarps. Carpospores gave rise to the alternately branched tetrasporophyte phase. Bonnemaisonia geniculate has a heteromorphic life history involving a previously undescribed tetrasporophyte.     

PHOTOSYNTHETIC RESPONSES TO LIGHT AND TEMPERATURE OF THE HETEROMORPHIC MARINE ALGA MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Physiological differentiation of the heteromorphic life-history phases of the red alga Mastocarpus papillatus (C. Agardh) Kützing was assessed. Photosynthetic responses to light and temperature of the erect, foliose gametophyte were compared to those of the crustose tetrasporophyte. Plants of both phases were collected from four locations on the Pacific coast of Baja California, Mexico, and California, USA, between 32–4l° N latitude. Within each location, the chlorophyll-specific, light-saturated photosynthetic rates of gametophytes were generally five times greater than those of tetrasporophytes. Initial slopes of photosynthesis: irradiance curves were greater for gametophytes than for tetrasporophytes. The crust and the blade from each location were similar with respect to dark respiration rates. For tetrasporophytes from all locations, the photosynthetic temperature optima were between 12–15° C. The photosynthetic temperature optima for gametophytes ranged from 15–17° C for plants from Trinidad Head (41° N) to 22–25° C for plants from Punta Descanso (32° N). Both gametophytes and tetrasporophytes from the northernmost location had significantly higher photosynthetic rates than the same phase from the other three locations. The photosynthetic responses to light support models for the life history in which life history phases have different ecological roles. The gametophyte, thought to be specialized for rapid growth and competition, may allocate more resources to photosynthetic machinery, hence the higher maximum photosynthetic rates. The tetrasporophyte, thought to be specialized for resistance to herbivores, may allocate more resources to structural or chemical defenses in preference to photosynthetic machinery. Consequently, the tetrasporophyte has lower photosynthetic capacity.     

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.