LIFE HISTORY AND SYSTEMATIC POSITION OF PSEUDOCHORDA GRACILIS SP. NOV. (LAMINARIALES,PHAEOPHYCEAE)1

Pseudochorda gracilis sp. Nov. (Pseudochordacease, Laminariales) is described from the Japan Sea coast of Hokkaido/ the species is subtidal, epilithic and annual, appearing in spring and maturing in winter. Erect thalli grow solitary or in tufts on a small discoid holdfast. They are simple, cord-shaped and hollow, with inner hyphal filaments, cylindrical medullary cells and paraphyses consisting of 3–6 cells. Hair tufts are observed only in young thalli. Unilocular sporangia are sessile and narrowly ovate. In culture, P. gracilis shows a heteromorphic life history with oogamy, characteristic of the order Laminariales. Gametophytes are dioecious and dimorphic. Gametophytes mature under lower temperature conditions (usually below 10°C), and sporophytes mature under low temperature and short-day conditions (5°C, SD). The seasonal growth pattern of the species results from the photoperiod-temperature conditions controlling saprophyte maturation.     

LIFE HISTORY OF COLPOMENIA SINUOSA (SYCTOSIPHONACEAE,PHAEOPHYCEAE) IN THE AZORES1

Colpomenia sinuosa (Mertens ex Roth) Derbès and Solier (Scytosiphonaceae, Phaeophyceae) is a common species on the rocky intertidal shores of the Azores, where reproductive gametophytes occur throughout the year. Life‐history studies of this species were carried out in culture, and both sexual and asexual reproduction were observed. Anisogamous gametes fused to form zygotes. The zygotes gave rise to a filamentous prostrate sporophyte generation bearing unilocular sporangia, under both short‐day and long‐day conditions at 15 and 22°C, and to both unilocular and plurilocular sporangia, under the lower temperature condition. Unispores developed into gametophytes, and plurispores gave rise to filamentous sporophytes. Asexual reproduction was carried out by unfused female gametes and asexual plurispores produced from the same gametophyte. Unfused gametes developed into filamentous prostrate sporophytes producing unilocular sporangia in both culture conditions, and unispores released from the sporangia gave rise to gametophytes. Asexual plurispores from field gametophytes, under both culture conditions, developed directly into new gametophytes. The species exhibited three types of life history: a heteromorphic, diplohaplontic; a heteromorphic, monophasic (both with alternation between the erect and filamentous prostrate thalli); and a monomorphic, monophasic.     

MORPHOLOGY AND LIFE HISTORY OF SCYTOSIPHON LOMENTARIA (SCYTOSIPHONACEAE,PHAEOPHYCEAE) FROM THE AZORES1

Morphological and culture studies of Scytosiphon lomentaria (Lyngbye) Link and Microspongium gelatinosum Reinke (Scytosiphonaceae, Phaeophyceae) were undertaken on material collected on the Island of São Miguel, Azores, where both species were commonly found. Erect thalli of S. lomentaria collected in the field were up to 33 cm long and 2.3 mm wide, tubular, hollow, and commonly constricted at intervals. The plurilocular sporangia were positioned in continuous sori on the thallus surface. Ascocysts were present. In the field, M. gelatinosum formed crustose to slightly pulvinate plants, were spongy in texture, and dark brown to black in color, which were circular or irregularly spreading over several centimeters and firmly attached to the substratum. Sessile unilocular sporangia were located in sori on the crust surface. In culture S. lomentaria plurispores developed into Microspongium‐like crustose prostrate thalli that formed unilocular sporangia. Unispores developed into new erect thalli that formed plurilocular sporangia. Sexual reproduction was not observed. In culture, M. gelatinosum unispores developed into erect thalli identical with S. lomentaria. These results are similar to those reported for other areas and suggest the occurrence in the Azorean plants of a monophasic and heteromorphic life history, involving both entities studied.     

OBSERVATIONS ON THE LIFE HISTORY OF PAPENFUSSIELLA CALLITRICHA (PHAEOPHYCEAE,CHORDARIALES) IN CULTURE1

Papenfussiella callitricha (Rosenv.) Kylin from eastern Canada was studied in culture. Zoids from unilocular sporangia develop into microscopic, filamentous, dioecious gametophytes which produce isogametes in filament cells and few-chambered plurilocular gametangia. Unfused gametes germinate to reproduce the gametophytes. Fusion takes place between a settled (“female”) and a motile (“male”) gamete. The zygote gives rise to a filamentous plethysmothallus that reproduces asexually by zoids formed in thallus cells and in few-chambered plurilocular zoidangia. Erect macrothalli are produced on the plethysmothallus, beginning with the formation of upright filaments. Later on, these filaments become the terminal assimilators of the macrothalli. Further assimilatory filaments, rhizoids, and unilocular sporangia are produced in a branching region at the base of the terminal assimilator. Zoids from unilocular sporangia formed in culture germinate to reestablish the gametophyte phase. Chromosome counts yielded n = 19 ± 3 for the gametophytes, and 32 ± 6 for the sporophyte, both plethysmothallus and macrothallus.     

CULTURE STUDIES ON THE LIFE HISTORY OF CHORDARIA LINEARIS (PHAEOPHYCEAE) FROM TIERRA DEL FUEGO,SOUTH AMERICA1

Studies of laboratory cultures of Chordaria linearis (Hooker et Harvey) Cotton from southernmost South America revealed that this species has an obligate sexual life history in which a macroscopic sporophyte alternates with a monoecious microscopic gametophyte. Sexual reproduction is isogamous and under photoperiodic control. Gametes are produced only in short days, whereas in long days, asexual zoospores are formed that recycle the gametophyte generation. Unfused gametes develop into gametophytes, and sporophytes originate only from zygotes. Unlike other sexual members of the Chordariales, gametes of C. linearis have a reduced stigma and do not show phototaxis. They are released at the beginning of the night, not in the morning. In nature, C. linearis seems to be regularly infected by a dictyosiphonalean epiphyte resembling the rare arctic species Trachynema groenlandicum (Lund) Pedersen. The epiphyte is responsible for previous contradictory results obtained in laboratory cultures of C. linearis. This is the first record of Trachynema in the southern hemisphere.     

MORPHOLOGY,LIFE HISTORY,AND MOLECULAR PHYLOGENY OF STSCHAPOVIA FLAGELLARIS (TILOPTERIDALES,PHAEOPHYCEAE) AND THE ERECTION OF THE STSCHAPOVIACEAE FAM. NOV.1

The phenology, life history, ultrastructure of reproductive structures, and molecular phylogeny using rbcL and rDNA (5.8S, internal transcribed spacer 2, and partial 26S) gene sequences of Stschapovia flagellaris, endemic to the northwestern Pacific Ocean, were studied. This species was first classified in the order Delamareales together with Delamarea, Coelocladia, and Cladothele. Those three genera, however, were later transferred to Dictyosiphonales, whereas the systematic position of Stschapovia remained unclear. At Abashiri, Hokkaido, Japan, the species regenerated by forming a new erect thallus from a perennial crustose holdfast or by presumably parthenogenetic development of eggs released from the erect thallus. There was no alternation of generations. In winter, the monoecious erect thallus formed reproductive structures (i.e. plurilocular antheridia and oogonia) in the thickened part of the thallus. Sperm had a chloroplast with an eyespot and a long anterior and short posterior flagellum. Eggs contained numerous disc‐shaped chloroplasts, physodes, and vacuoles. Neither sexual attraction of the presumptive sperm by eggs nor their sexual fusion was observed. Molecular phylogenetic analyses revealed the closest phylogenetic relationship between Stschapovia and Halosiphonaceae, and they grouped with Phyllariaceae and Tilopteridaceae (Tilopteridales s. s.). Stschapovia and Tilopteridaceae have several important morphological similarities: chloroplasts lacking pyrenoids, lack of sexual reproduction despite the release of obvious sperm, occurrence of monoecious gametophytes, and similarity in the early developmental pattern of the erect thallus. In conclusion, we propose the establishment of the new family Stschapoviaceae to accommodate Stschapovia and the placement of the family in the order Tilopteridales together with Tilopteridaceae, Halosiphonaceae, and Phyllariaceae.     

DIFFERENTIAL LIFE HISTORY PHASE EXPRESSION IN TWO COEXISTING SPECIES OF SCYTOSIPHON (PHAEOPHYCEAE) IN NORTHERN CHILE1

The identity of two phaeophycean taxa that monopolized the middle‐lower rocky intertidal zone of a coastal area chronically exposed to copper mine wastes in northern Chile was unraveled. One of them was preliminarily identified as the gametophytic stage of Scytosiphon lomentaria (Lyngbye) Link. The other, a dark crust, resembled the alternate stage of some Scytosiphon species. Comparative analysis of morphology, life history, and DNA sequences strongly suggests that crusts corresponded to sporophytic S. tenellus Kogame and confirm that erect thalli belonged to S. lomentaria. A clear segregation of erect and crustose thalli was found using internal transcribed spacer region 1 and RUBISCO spacer sequences. Furthermore, whereas crusts always grouped with S. tenellus, erect thalli always grouped with S. lomentaria. Life history studies failed to connect the two entities. First, field‐collected S. tenellus produced progeny that either recycled the crust, which reproduced by unilocular zoidangia, or developed into erect thalli. The latter, unlike typical gametophytic S. lomentaria, developed patchy sori of plurilocular zoidangia. Second, S. lomentaria displayed a direct‐type life cycle, in which progeny from erect individuals only developed into erect thalli and produced only plurilocular zoidangia. This constitutes the first experimental study on Scytosiphon from the Pacific coast of South America and the first report of S. tenellus on this coast. It is also the first report of the crustose stage of Scytosiphon appearing as a perennial and dominant algal species in a temperate rocky intertidal system.     

CULTURE STUDIES ON REPRODUCTION OF SPERMATOCHNUS PARADOXUS (PHAEOPHYCEAE,CHORDARIALES)1

The life history of Spermatochnus paradoxus (Roth) Kütz. isolated from the Mediterranean Sea was studied in culture. Meiospores develop to a microscopic stage (microthallus) which at 20°C perpetuates asexually by plurilocular sporangia and formation of new microthalli. At 9°C microthalli act as homothallic gametophytes. Fusion of isogametes results in a diploid microthallus which, after differentiation of an apical cell, leads back to Spermatochnus plants. In addition, gametes develop without fusion to form haploid macrothalli, the further fate of which has not been determined. Chromosome numbers alternate between n = 20 ± 2 in the microthalli and 2n = 41 ± 4 in macrothalli.     

LIFE HISTORY AND GROWTH OF LAURENCIA BRACHYCLADOS (RHODOPHYTA,CERAMIALES)1

Laurencia brachyclados Pilger from Hawai'i completed a “Polysiphonia-type” tri-phasic life history in 21 weeks in laboratory culture. Tetraspores developed into gametophytes in a nearly 1:1:1 ratio of females: males: non-reproductive. Carpospores were released as early as 21 days after mixing virgin female and male gametophytes. Cultured thalli showed a “guerilla type” growth form. Other Hawaiian Laurencia species in culture had longer maturation times or remained non-reproductive. Variation in life history schedules may influence Laurencia species coexistence and algal community structure.     

MORPHOLOGY, LIFE HISTORY, AND MOLECULAR PHYLOGENY OF CHORDA RIGIDA, SP. NOV. (LAMINARIALES, PHAEOPHYCEAE) FROM THE SEA OF JAPAN AND THE GENETIC DIVERSITY OF CHORDA FILUM

Chorda rigida Kawai et Arai, sp. nov. (Chordaceae, Laminariales) is described from the Sea of Japan, NW Pacific. This species resembles Chorda filum (Linnaeus) Stackhouse but is distinguished by the following characteristics: 1) the sporophytes grow on more or less exposed rocks at 2–7 m depth and do not form dense tufts; 2) compared with C. filum , sporophytes of C. rigida are much more rigid and are composed of denser cortical layers (6–18 cells thick); 3) C. filum becomes fertile and disappears in late spring to summer, whereas C. rigida appears in early summer, oversummers, and becomes fertile only in late autumn at the same localities; 4) in culture, C. rigida sporophytes tolerate higher temperature conditions (20 and 25° C) than C. filum; and 5) C. rigida has considerably longer sequences of the rDNA ITS region than does C. filum . The independence of the species is further supported by molecular phylogenetic analyses using sequence of the ITS + 5.8S ribosomal DNA. Interestingly, C. filum is shown to be genetically diverse and possibly paraphyletic, and it may require subdivision into several species or subspecies. The rbc L and associated spacer sequence data established monophyly of the genus Chorda among Laminariales, but the resolution was limited for discussing the phylogenetic relationships within the genus.     

LIFE HISTORY AND THE EVOLUTION OF PARENTAL CARE

Patterns of parental care are strikingly diverse in nature, and parental care is thought to have evolved repeatedly multiple times. Surprisingly, relatively little is known about the most general conditions that lead to the origin of parental care. Here, we use a theoretical approach to explore the basic life‐history conditions (i.e., stage‐specific mortality and maturation rates, reproductive rates) that are most likely to favor the evolution of some form of parental care from a state of no care. We focus on parental care of eggs and eggs and juveniles and consider varying magnitudes of the benefits of care. Our results suggest that parental care can evolve under a range of life‐history conditions, but in general will be most strongly favored when egg death rate in the absence of care is high, juvenile survival in the absence of care is low (for the scenario in which care extends into the juvenile stage), adult death rate is relatively high, egg maturation rate is low, and the duration of the juvenile stage is relatively short. Additionally, parental care has the potential to be favored at a broad range of adult reproductive rates. The relative importance of these life‐history conditions in favoring or limiting the evolution of care depends on the magnitude of the benefits of care, the relationship between initial egg allocation and subsequent offspring survival, and whether care extends into the juvenile stage. The results of our model provide a general set of predictions regarding when we would expect parental care to evolve from a state of no care, and in conjunction with other work on the topic, will enhance our understanding of the evolutionary dynamics of parental care and facilitate comparative analyses.     

ULTRASTRUCTURE OF SWARMERS IN THE LAMINARIALES (PHAEOPHYCEAE). II. SPERM1

The ultrastructure of sperm from 13 species in 11 genera of Laminariales collected in the northeast Pacific Ocean is unique in the brown algae. The sperm are elongate, and possess a nucleus, several mitochondria and two or three chloroplasts, but no eyespot. The anterior flagellum bears mastigonemes on the proximal half of its length; a distal “whiplash” portion lacks mastigonemes and is an extension of only the two central singlet microtubules of the axoneme. A peculiar feature of these sperm is the posterior flagellum, which is longer than the anterior flagellum and tapers distally as the doublet microtubules become singlets and decrease in number. This feature contrasts with the laminarialean zoospore, which possesses a short posterior flagellum with the usual “9 + 2” axoneme. The structure of these sperm differs from that reported for Chorda, the sperm of which resembles a primitive brown algal zoospore. The facts support the concept that Chorda is the most primitive member of the Laminariales.     

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.     

ARE LIFE HISTORY CHARACTERISTICS GOOD PREDICTORS OF GENETIC DIVERSITY AND STRUCTURE? A CASE STUDY OF THE INTERTIDAL ALGA FUCUS SPIRALIS (HETEROKONTOPHYTA; PHAEOPHYCEAE)1

Characteristics of an organism's life history are often good predictors of genetic diversity and genetic structure. We tested hypotheses about genetic structure and diversity in an intertidal alga based on life history and life form. Fucus spiralis L. is a perennial monoecious alga that is abundant on the shores of Maine. Reproduction in fucoids is highly sensitive to water motion, resulting in fertilization success close to 100%. Given these life history characteristics, we predicted genetic structure among populations to be high and genetic diversity within populations to be low. We used five microsatellite loci to analyze genetic structure and diversity in F. spiralis from four sites on each of two coastal points, Maine, USA. Observed heterozygosities were relatively low (0.23 to 0.56), and F_IS estimates were usually significantly large, ranging from 0.021 to 0.476. This suggests that selfing and/or inbreeding may occur. Contrary to predictions, genetic differentiation between the two coastal points was insignificant. Moreover, few sites were genetically different from one another. Pairwise tests revealed complex patterns among sites. Genetic differentiation was not correlated with distance (P>0.05). Life history characteristics are good predictors of genetic diversity but not of population genetic structure in F. spiralis. We suggest that long distance dispersal of F. spiralis via drifting algal rafts increases gene flow. In addition, low levels of genetic structure may arise due to episodic recruitment or recent recolonization events. We discuss the implications of our results in terms of using life history characteristics as predictors of genetic diversity and structure in algae.     

REPRODUCTION AND LIFE HISTORY OF THE RED ALGA GALAXAURA OBLONGATA (NEMALIALES,GALAXAURACEAE)1

Culture and morphological studies showed that Galaxaura oblongata (Ellis et Solander) Lamouroux has a triphasic life history with conspicuous gametophytes and small filamentous tetrasporophytes. Development of male and female reproductive structures is very similar and both begin with the enlargement of a terminal cell of a filament branch occupying a normal vegetative position within the apical pit of a thallus branch. In male thalli this modified branch forms a conceptacle in which spermatangia are produced. In female thalli, this modified branch forms a three-celled carpogonial branch consisting of a carpogonium, hypogynous cell and basal cell. Filament branches from the basal cell form a pericarp and the gonimoblast develops directly from the carpogonium. Carposporangia are produced in conceptacles which resemble the male conceptacles. About the time the first carposporangia are produced, the carpogonium, hypogynous cell and basal cell form a large fusion cell. Released carpospores germinate in a unipolar or bipolar manner and form small filamentous thalli. Under short day conditions, cruciate tetrasporangia are produced in small clusters. Tetraspores germinate similarly to carpospores and also form small filamentous thalli. Under low nutrient conditions, small cylindrical thalli develop on the filaments and these appear similar to gametophytes collected in nature.     

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.     

THE RECLASSIFICATION OF LESSONIA LAMINARIOIDES (LAMINARIALES,PHAEOPHYCEAE): PSEUDOLESSONIA GEN. NOV.1

A new genus, Pseudolessonia, is proposed for the kelp Lessonia laminarioides Postels et Ruprecht (Laminariales, Phaeophyceae), which occurs on the northwest side of the Sea of Okhotsk, in the northwest Pacific Ocean. Pseudolessonia is monotypic and differs from Lessonia in its short primary stipes and its corrugated, unilaterally arranged blades with entire margins. This species is transferred on the basis of morphology and plastid gene sequence comparisons. We determined psaA and rbcL gene sequences from 17 taxa of Pseudolessonia, Lessonia, and putative relatives. Analyses of individual and combined data sets resulted in congruent trees showing a clear separation of Pseudolessonia laminarioides from Lessonia, but suggesting its sister relationships with the clade of Nereocystis, Macrocystis, Pelagophycus, and Postelsia in the North Pacific Ocean. On the other hand, Lessonia species from the South Pacific Ocean formed a strongly supported clade. The results indicate that the basal splitting of the blade, which has been considered a diagnostic character for the family Lessoniaceae, is a result of convergent evolution.     

NORTHERN AND SOUTHERN HEMISPHERE HYBRIDS OF MACROCYSTIS (PHAEOPHYCEAE)1

All combinations of individuals of Macrocystis pyrifera (L.) C. Agardh, M. integrifolia Bory, and M. angustifolia Bory hybridized. Gametophyte isolates obtained from 18 individuals were used, including M. pyrifera and M. integrifolia from the extremes of their Northern Hemisphere ranges along the Pacific Coast of North America and M. pyrifera and M. angustifolia from Tasmania, Australia. All combinations of gametophytes produced sporophytes of normal morphology, with the exception of crosses involving three gametophyte isolates. One female (M. integrifolia) and two male (M. pyrifera and M. angustifolia)gametophyte isolates were unable to produce normal sporophytes in combination with gametophytes of the opposite sex. Some cultures of female gametophytes produced abnormally shaped parthenogenetic sporophytes. Gametophytes and sporangia of M. pyrifera had n= 16 chromosomes. The M. integrifolia female gametophyte that was unable to produce normal sporophytes had n = ca. 32 chromosomes. These results show that these species of Macrocystis have not become reproductively isolated. Although these species may be considered conspecific according to the biological species concept, we recommend that they continue to be recognized as separate species based on morphological differences.     

卵形异绒螨的形态和生活史研究(真螨目：绒螨科)

卵形异绒螨Allothrombium ovatum Zhang et Xin,1992是我国北方地区棉蚜Aphis gossypii Giover,1877的一种外寄生性天敌。通过室内饲养和田间观察结果表明：该螨一年发生一代,以卵在土壤内越冬。在年平均温度13.7℃条件下,其卵、前幼螨、幼螨、若蛹、若螨、成蛹和成螨的发育历期平均为220.8d、19.5d、22.0 d、12.0 d、13.0 d、13.0 d和59.O d。另外,本文还对卵形异绒螨主要虫态的形态和习性进行了描述和研究。     

ULTRASTRUCTURE OF SWARMERS IN THE LAMINARIALES (PHAEOPHYCEAE). I. ZOOSPORES1

Zoospores of 17 species in 14 genera of Laminariales, collected in the northeast Pacific Ocean, were studied by electron microscopy. These zoospores are unique in the brown algae in lacking both an eyespot in the single chloroplast and any associated swelling at the base of the shorter, posterior flagellum. Spores of all species examined possess a distal whiplash portion on the longer, mastigoneme-bearing anterior flagellum. This appendage may sometimes be as long as the mastigoneme-bearing portion of the flagellum, but it is only seldom preserved in the preparations for electron microscopy. A microtubular cytoskeleton is probably responsible for maintaining the shape of the spore. It consists of a short band of about 10 microtubules between the two basal bodies, scattered tubules converging at the anterior of the spore, a band of 7–9 tubules directed anteriorly from the anterior basal body, and a band directed posteriorly from the posterior basal body. These anterior and posterior bands may form one continuous band looping around the periphery of the spore. Variation with possible taxonomic significance was found in the ultrastructure of vesicles which apparently contain adhesive material, and which are extruded through the plasmalemma when the zoospores settle.