PHENOTYPIC PLASTICITY RECONCILES INCONGRUOUS MOLECULAR AND MORPHOLOGICAL TAXONOMIES: THE GIANT KELP,MACROCYSTIS (LAMINARIALES,PHAEOPHYCEAE), IS A MONOSPECIFIC GENUS1

The giant kelp genus Macrocystis C. Agardh (Laminariales, Phaeophyceae) is one of the world’s most ecologically and economically important seaweed taxa, yet its taxonomy remains uncertain. Although the genus currently contains four accepted species based on variable holdfast and blade morphology [M. pyrifera (L.) C. Agardh, M. integrifolia Bory, M. angustifolia Bory, and M. laevis C. H. Hay], numerous recent studies on Macrocystis interfertility, genetic relatedness, and morphological plasticity all suggest that the genus is monospecific. We reviewed this evidence and present an explanation for the extreme phenotypic plasticity that results in morphological variability within Macrocystis, driven by the effects of environmental factors on early development of macroscopic sporophytes. We propose that the genus be collapsed back to a single species, with nomenclatural priority given to M. pyrifera.     

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.     

THE EFFECT OF SEAWEED DIETS ON GROWTH OF GREEN ABALONE,HALIOTIS FULGENS,FROM BAJA CALIFORNIA,MEXICO

Two abalone species: green Haliotis fulgens and yellow Halioti corrugata represent nearly 97% of the total production in the Mexican abalone fishery. It has been assumed that abalone feed on the kelp algae Macrocystis pyrifera. Regional hatcheries use this species as a main source of natural food. M. pyrifera does not occur at the southern limit of the distribution of abalone species along the Baja California Peninsula. In this study, growth rates of juveniles H. fulgens, 17.3 ± 2.2 mm shell length and 0.4 ± 0.2 g body weight, were evaluated. Juveniles were fed with common species in the benthic environments inhabited by abalone along the western coast of Baja California during 191 days. Three diets were based on algae: palm kelp, Eisenia arborea, giant kelp, M. pyrifera and Gelidium robustum, and one on seagrass, Phyllospadix torreyi. Shell length and body growth rates varied between 21.5 μm day^?1 and 2.2 mg day^?1 for E. arborea and between 45.9 μm day^?1 and 6.7 mg day^?1 for M. pyrifera. Higher specific growth rates (SGR) in length and weight were determined for M. pyrifera: 0.2% and 0.7% day^?1. Significant differences between values of juveniles fed M. pyrifera with the rest of the diets were found. The highest mortality (21%) was in juveniles fed the red algae G. robustum.     

PHOTOSYNTHETIC PHYSIOLOGY AND CHEMICAL COMPOSITION OF SPORES OF THE KELPS MACROCYSTIS PYRIFERA,NEREOCYSTIS LUETKEANA,LAMINARIA FARLOWII,AND PTERYGOPHORA CALIFORNICA (PHAEOPHYCEAE)1

Recently released spores of the kelps Macrocystis pyrifera (L.) C. Ag., Nereocystis luetkeana (Mert.) Post. and Rupr., Laminaria farlowii Setch., and Pterygophora californica Rupr. had different levels of net photosynthesis. Spore-specific photosynthesis–irradiance relationships were similar in many respects for M. pyrifera, N. luetkeana, and L. farlowii spores. All three species had low rates of net light-saturated photosynthesis. In contrast, spores of P. californica had higher photosynthetic potential and overall net photosynthesis than the other three species. On a cell carbon basis, however, photosynthetic rates in N. luetkeana spores were similar to those of P. californica spores and higher than those of M. pyrifera spores. Chlorophyll a content of spores varied 10-fold among species. The rank order of significant differences in chlorophyll a content was P. californica > L. farlowii > N. luetkeana > M. pyrifera. As a result, chlorophyll-specific measurements suggest M. pyrifera and N. luetkeana spores had much higher quantum efficiency and photosynthetic potential than either P. californica or L. farlowii spores. Maternal carbon and nitrogen investment significantly differed in spores of M. pyrifera, N. luetkeana, and P. californica with P. californica > M. pyrifera > N. luetkeana. Carbon content in spores of each of these three species increased by about 30% during 12 h of saturating irradiance. We suggest that the photosynthetic capabilities of and maternal investment in spores may be related to the spore as a unit of dispersal, to the reproductive ecology of the parental sporophytic stages, and to the growth and physiology of the germling gametophyte stages.     

On an unknown lamprey holotype (Petromyzon macrostomus Burmeister 1868) and the genus Exomegas Gill 1883 from South America

The taxonomy of the South American genus Exomegas Gill 1883 has been unstable owing to the unknown status of the type specimen of its type species, Petromyzon macrostomus Burmeister 1868. Here the authors announce the finding of the holotype in the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” ichthyology collection. Whereas Exomegas type species is recognized based on its holotype, Exomegas gallegensis (Smitt 1901) new status is proposed. These preliminary results resolve a century-long taxonomic uncertainty and foster a nomenclatural arrangement for a Southern Hemisphere lamprey genus.     

PURIFICATION OF SPECIES SPECIFIC ANTIBODIES TO CARBOHYDRATE COMPONENTS OF MACROCYSTIS PYRIFERA (PHAEOPHYTA)1

Polyclonal rabbit antibodies to cell wall components were produced against gametophytes of the giant kelp Macrocystis pyrifera (Linnaeus) C. Agardh. These antibodies were found to react with carbohydrates extracted from M. pyrifera and Pterygophora californica Ruprecht by carbohydrate based enzyme immunoassay (EIA). The antibodies reacted with carbohydrates from both species. After affinity purification on a column with M. pyrifera carbohydrate coupled to AH-Sepharose, the eluted antibody was specific for M. pyrifera carbohydrate with little cross reactivity to P. californica carbohydrate in the EIA test. In experiments carried out to characterize the antigenic specificity of unfractionated antibody using commercially prepared carbohydrates in the EIA, the antibodies were shown to react primarily with fucoidan and to a lesser degree, alginate. The unfractionated antibody was also shown to bind to proteins from both M. pyrifera and P. californica. These results indicate that species specific carbohydrate determinants may be present in the kelp cell wall.     

PHOTOSYNTHETIC PERFORMANCE,LIGHT ABSORPTION,AND PIGMENT COMPOSITION OF MACROCYSTIS PYRIFERA (LAMINARIALES,PHAEOPHYCEAE) BLADES FROM DIFFERENT DEPTHS1

Macrocystis pyrifera (L.) C. Agardh is a canopy‐forming species that occupies the entire water column. The photosynthetic tissue of this alga is exposed to a broad range of environmental factors, particularly related to light quantity and quality. In the present work, photosynthetic performance, light absorption, pigment composition, and thermal dissipation were measured in blades collected from different depths to characterize the photoacclimation and photoprotection responses of M. pyrifera according to the position of its photosynthetic tissue in the water column. The most important response of M. pyrifera was the enhancement of photoprotection in surface and near‐surface blades. The size of the xanthophyll cycle pigment pool (XC) was correlated to the nonphotochemical quenching (NPQ) of chl a fluorescence capacity of the blades. In surface blades, we detected the highest accumulation of UV‐absorbing compounds, photoprotective carotenoids, ΣXC, and NPQ. These characteristics were important responses that allowed surface blades to present the highest maximum photosynthetic rate and the highest PSII electron transport rate. Therefore, surface blades made the highest contribution to algae production. In contrast, basal blades presented the opposite trend. These blades do not to contribute significantly to photosynthetate production of the whole organism, but they might be important for other functions, like nutrient uptake.     

Nuclear DNA level and life cycle of kelps: Evidence for sex‐specific polyteny in Macrocystis (Laminariales,Phaeophyceae)

Giant kelp, Macrocystis pyrifera (Linnaeus) C. Agardh, is the subject of intense breeding studies for marine biomass production and conservation of natural resources. In this context, six gametophyte pairs and a sporophyte offspring of Macrocystis from South America were analyzed by flow cytometry. Minimum relative DNA content per cell (1C) was found in five males. Unexpectedly, nuclei of all female gametophytes contained approximately double the DNA content (2C) of males; the male gametophyte from one locality also contained 2C, likely a spontaneous natural diploid variant. The results illustrate a sex‐specific difference in nuclear DNA content among Macrocystis gametophytes, with the chromosomes of the females in a polytenic condition. This correlates with significantly larger cell sizes in female gametophytes compared to males and resource allocation in oogamous reproduction. The results provide key information for the interpretation of DNA measurements in kelp life cycle stages and prompt further research on the regulation of the cell cycle, metabolic activity, sex determination, and sporophyte development.     

Effects of chelated iron on oogenesis and vegetative growth of kelp gametophytes (Phaeophyceae)

The supply of iron has been reported to affect gametogenesis in the gametophytes of some species of kelps (order Laminariales). Spores of the kelps Alaria marginata Postels & Ruprecht, Dictyoneurum californicum Ruprecht, Egregia menziesii (Turner) Areschoug, Laminaria setchellii Silva, and Macrocystis pyrifera (L.) C. Agardh were cultured in enriched seawater with and without added chelated iron (Fe‐ethylenediaminetetraacetate) to determine the effects of iron on oogenesis. All species showed a decrease in oogenesis without added Fe‐ethylenediaminetetraacetic acid (EDTA). Gametophytes of E. menziesii showed predominant gametogenesis with or without supplied iron, resulting in all cells being converted to gametes so that vegetative growth did not continue. Vegetative gametophytes were obtained in the other species used. Gametophytes of M. pyrifera did not show any oogenesis without added Fe‐EDTA, while those of L. setchellii, A. marginata and D. californicum were intermediate in their response, showing some gametogenesis without added Fe‐EDTA. When Fe‐EDTA supply was delayed by 6, 13 and 20 days with spores of M. pyrifera, the gametophytes produced fewer eggs, with a greater decrease as the delay grew longer. A range of Fe‐EDTA concentrations was investigated using isolated female gametophytes of two strains of M. pyrifera and one of Macrocystis integrifolia Bory. None of these three strains produced gametes without the addition of Fe‐EDTA. Gametophytes of M. integrifolia required the least amount of added Fe‐EDTA to achieve gametogenesis while gametophytes of M. pyrifera required higher amounts, with the two strains showing somewhat different responses. Iron nutrition appears to be an essential factor for gametogenesis in several species of kelps.     

The giant cryptic amphipod species of the subterranean genus Niphargus (Crustacea,Amphipoda)

Amphipods from the genus Niphargus represent an important part of the Western Palearctic subterranean fauna. The genus is morphologically diverse, comprising several distinct ecomorphs bound to microhabitats in the subterranean environment. The most impressive among them are “lake giants,” a series of massive, large‐bodied species. These range from morphologically distinct to morphologically cryptic taxa. We analysed the taxonomic structure of the Niphargus arbiter–Niphargus salonitanus species complex, belonging to “lake giants” from the Dinaric Karst (West Balkans), and assessed their phylogenetic, morphological and ecological diversity. Multilocus phylogeny suggested that the complex is monophyletic and nested within other cave lake ecomorphs. Unilocus and multilocus coalescence species delimitations indicated that the complex totals nine species. These species substantially overlap in morphology and cannot be unambiguously told apart without the use of molecular markers. An analysis of splitting events within a palaeogeological context, and modelling of environmental characteristics on the phylogeny unveiled a complex history of diversification. Part of this diversification might have been influenced by ecological divergence along the altitudinal gradient reaching from the Adriatic coast to inland Dinaric mountain chains and Poljes. Other splits coincide with the marine regression–transgression cycles during Pliocene. We describe Niphargus alpheus sp. n., Niphargus anchialinus sp. n., Niphargus antipodes sp. n., Niphargus arethusa sp. n., Niphargus doli sp. n., Niphargus fjakae sp. n. and Niphargus pincinovae sp. n., and by doing so hope to prompt their further research.     

EFFECT OF HIGH IRRADIANCE ON RECRUITMENT OF THE GIANT KELP MACROCYSTIS (PHAEOPHYTA) IN SHALLOW WATER1

Laboratory and field experiments were done hi Still-water Cove, Carmel Bay, California, and Monterey Harbor, California, to determine the effect of photosynthetically active radiation (PAR) on the shallow (upper) limit of giant kelp, Macrocystis pyrifera (L.) C. Agardh. At shallow depths, M. pyrifera did not recruit or grow to macroscopic size from gametophytes or embryonic sporophytes transplanted to vertical buoy lines; sharp decreases in PAR with depth coincided with observed recruitment and sporophyte distributions. Shade manipulations indicated that settlement of M. pyrifera zoospores was decreased, but not prohibited, by high PAR. Postsettlement stages (gametophytes and embryonic sporophytes), however, survived only under shade. These results suggest that high PAR can inhibit the recruitment of M. pyrifera to shallow water by killing its postsettlement stages; whether or not ultraviolet (UV) radiation also inhibits recruitment was not tested. In either case, however, it appears that high irradiance (PAR and/or UV) regulates the shallow limit of M. pyrifera prior to temperature and desiccation stresses inherent to intertidal regions. In an additional experiment, recruitment or growth of transplanted gametophytes or embryonic sporophytes of Macrocystis integrifolia Bory also did not occur at shallow depths, suggesting that this shallow water species accesses high irradiance regions via a method other than sexual reproduction.     

Mitochondrial gene rearrangements suggest a new genus in the subfamily Cantharinae (Coleoptera)

The Cephalomalthinus semifumatus species group, referred to as the “semifumatus” group henceforth, is interesting because of its heterogeneous morphology resembling either Cephalomalthinus Pic, 1921 or Rhagonycha Eschscholtz, 1830. To elucidate its phylogenetic status, mitochondrial genomes of four species of the “semifumatus” group, 11 Cephalomalthinus species, and 11 Rhagonycha species were sequenced and examined. All analysed mitogenomes were similar with respect to genome size, nucleotide composition, and AT content. Surprisingly, a rearrangement of the trnW-trnC and trnY genes was detected in the “semifumatus” group, presumably caused by tandem duplication and random loss events. Furthermore, genetic distance analyses showed that the proximity of the “semifumatus” group to Cephalomalthinus and to Rhagonycha was comparable to that between the latter two. Moreover, the produced phylogeny strongly supported the monophyly of the “semifumatus” group, and molecular clock analyses dated its divergence from Cephalomalthinus to 32.52 Ma. Thus, the new genus Amphimorphus gen. nov. is suggested to comprise the “semifumatus” group, in which the observed gene rearrangement was a synapomorphy. Moreover, morphological evidence regarding the unique structure of the aedeagus supported this separation. These results indicate that mitochondrial gene rearrangement provide important phylogenetic implications for revising Cephalomalthinus, a speciose genus that is puzzling in the morphology-based taxonomy.     

Leucothoe kawesqari,a new amphipod from Bernardo O’Higgins National Park (Chile), with remarks on the genus in the Magellan Region (Crustacea,Peracarida)

Although the genus Leucothoe has been reported repeatedly in the Magellan Region, the citations in the Channels and Fjords Ecoregion were either unidentified or attributed to the previously considered cosmopolitan Leucothoe spinicarpa. In this work, Leucothoe kawesqari sp. n. is described, which can be distinguished from other species of the genus in the Southern Ocean by having eyes present, epimeral plates with no setae, anterior coxae not acutely produced or excavate, coxa 5 slightly bilobed, accessory flagellum present, mandibular palp article 3 shorter than ½ article 2, pereopods 5–7 basis expanded, ovoid, posterior margin weakly crenulate and telson apex irregularly truncated. The new species was found in hard substrates, both unvegetated and with macroalgae, mainly in kelp forest of Macrocystis pyrifera.     

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.     

REPRODUCTIVE LONGEVITY OF DRIFTING KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) IN MONTEREY BAY,USA1

Drifting Macrocystis pyrifera (L.) C. Agardh sporophytes have long been viewed as the primary long‐distance dispersal vector; yet, few data exist that support the ability of reproductive viable sporophytes to actually travel the presumed hundreds to thousands of kilometers. This study addressed the reproductive longevity of experimental and naturally occurring M. pyrifera drifters. Temporal variability in sporophyte size and reproduction was estimated for experimental drifting sporophytes that were tethered to surface buoys and compared with attached plants (controls). Reproductive viability was also studied for beach‐cast drifters (BCD), and naturally drifting sporophytes observed during field surveys in Monterey Bay. Detached drifting sporophytes were tracked with radio transmitters to follow drifter trajectories and to measure drifting speed. Experimental drifters (ED) experienced a 74% reduction in frond length after 35 days, a 76% reduction in average frond number after 70 days, and a reduction in average sorus area by 83% after 28 days. Although zoospore production was reduced following detachment, sporophytes remained fertile with high zoospore germination success as long as sori were present (125 days). Zoospore production and germination success for natural and BCD was similar to ED. The average displacement of radio‐tagged drifters was 7.12 km·day^?1, suggesting that a sporophyte adrift for 125 days disperses viable propagules (zoospores) over 890 km (±363). Dispersal of propagules is important for population restoration, distribution, and genetic diversity. Such dispersal distances are long enough to connect potentially all Northern Hemisphere Macrocystis populations across a generational timescale and may facilitate inter‐hemispheric gene flow.     

Seascape genetics of the stalked kelp Pterygophora californica and comparative population genetics in the Santa Barbara Channel

We conducted a population genetic analysis of the stalked kelp, Pterygophora californica, in the Santa Barbara Channel, California, USA. The results were compared with previous work on the genetic differentiation of giant kelp, Macrocystis pyrifera, in the same region. These two sympatric kelps not only share many life history and dispersal characteristics but also differ in that dislodged P. californica does not produce floating rafts with buoyant fertile sporophytes, commonly observed for M. pyrifera. We used a comparative population genetic approach with these two species to test the hypothesis that the ability to produce floating rafts increases the genetic connectivity among kelp patches in the Santa Barbara Channel. We quantified the association of habitat continuity and oceanographic distance with the genetic differentiation observed in stalked kelp, like previously conducted for giant kelp. We compared both overall (across all patches) and pairwise (between patches) genetic differentiation. We found that oceanographic transit time, habitat continuity, and geographic distance were all associated with genetic connectivity in P. californica, supporting similar previous findings for M. pyrifera. Controlling for differences in heterozygosity between kelp species using Jost's D_EST, we showed that global differentiation and pairwise differentiation were similar among patches between the two kelp species, indicating that they have similar dispersal capabilities despite their differences in rafting ability. These results suggest that rafting sporophytes do not play a significant role in effective dispersal of M. pyrifera at ecologically relevant spatial and temporal scales.     

ROLE OF NUTRIENT FLUCTUATIONS AND DELAYED DEVELOPMENT IN GAMETOPHYTE REPRODUCTION BY MACROCYSTIS PYRIFERA (PHAEOPHYCEAE) IN SOUTHERN CALIFORNIA1

Organisms occurring in environments subject to severe disturbance and/or periods of poor environmental quality that result in severe adult mortality can survive these periods by relying on alternate life stages that delay their development in a resistant state until conditions improve. In the northeast Pacific, the forest‐forming giant kelp Macrocystis pyrifera (L.) C. Agardh periodically experiences widespread adult mortality during extended periods of extremely low nutrients and high temperatures, such as those associated with El Niño. Recovery following these periods is hypothesized to occur from microscopic life stages that delay their development until the return of favorable conditions. In the laboratory, we experimentally examined the environmental conditions responsible for regulating delayed development of the microscopic stages of M. pyrifera from Southern California, USA. Nutrients controlled the delay and resumption of gametophyte growth and reproduction, perhaps linked to the large fluctuations in nutrients occurring seasonally and interannually in this region. Although growth of gametophytes proceeded in the virtual absence of nitrate, both nitrate and other trace nutrients were necessary for gametogenesis. Upon exposure to elevated nutrients, delayed gametophytes produced sporophytes more quickly (5–20 d) and at smaller sizes (10–200 μm) than gametophytes that had never been delayed (18–80 d, 80–400 μm, respectively), reducing negative density‐dependent effects. This finding demonstrates that delayed gametophytes of M. pyrifera rapidly utilize increased resources to consistently produce sporophytes. Further work is needed to assess their potential role in population recovery following periods of poor environmental quality.     

PHYLOGENETIC PATTERNS AMONG NOSTOC CYANOBIONTS WITHIN BI‐ AND TRIPARTITE LICHENS OF THE GENUS PANNARIA1

Phylogenetic relationships between Nostoc cyanobionts in the lichen genus Pannaria were studied to evaluate their correlation to geography, habitat ecology, and other patterns previously reported. The 16S rRNA gene sequences of a total of 37 samples of 21 Pannaria species from seven countries from the Northern and Southern hemispheres were analyzed and compared with 69 free‐living and symbiotic cyanobacterial strains. The sequences from Pannaria were distributed throughout a branch of Nostoc sequences previously called “the Nephroma guild,” and within two subgroups from another branch, referred to as the “Peltigera guild,” although there was a gradual transition between the two major groups. There is a more diverse pattern of relationships between Nostoc sequences from bipartite versus tripartite lichen species in Pannaria, compared with other well‐studied genera, such as Nephroma and Peltigera. Cyanobionts from several tripartite Pannaria species from the Southern Hemisphere and corticolous bipartite species from both hemispheres were grouped together. Four sequences of Pannaria and Pseudocyphellaria cyanobionts from rocks in the Chilean Juan Fernández Islands were nested within corticolous cyanobionts, whereas the terricolous “Pannaria sphinctrina clade” was placed with other terricolous strains. The cluster patterns derived from phylogenetic analysis were partly reflecting lichen taxonomy, in two groups of lichen species, possibly indicating coevolution. The phylogram partly also reflected lichen ecology. Three Pannaria species have very different cyanobiont strains when they grow in different habitats.     

A MULTISPECIES LABORATORY ASSESSMENT OF RAPID SPOROPHYTE RECRUITMENT FROM DELAYED KELP GAMETOPHYTES1

Recent work suggests that the ability to delay reproduction as resistant haploid gametophytes may be important for seaweeds that experience unpredictable disturbances or seasonal periods of poor conditions that result in adult sporophyte absence. Further, delayed gametophytes of some kelp species (order Laminariales) may produce sporophytes more rapidly than if they had never experienced a delay, conferring a competitive advantage when conditions improve or after disturbance events. Here, it was determined that the gametophytes of the canopy‐forming kelp Macrocystis pyrifera (L.) C. Agardh could delay reproduction in a one‐ to two‐cell state (<50 μm) for at least 7 months when grown under nutrient‐limiting conditions. These stages retained reproductive viability and produced sporophytes within 5 d once nutrients were increased. This finding suggests that gametophytes could potentially promote recovery of M. pyrifera populations after extended periods of sporophyte absence. In addition, the time required for sporophyte production between gametophytes of the four most conspicuous kelp species in Southern California that had delayed reproduction and gametophytes that had not was compared. For these four kelp species, a delay of at least 30 d conferred a 40%–76% reduction in the time required for sporophyte production once nutrients were received. Fecundity did not decrease with delay duration, suggesting there is no apparent cost of delayed development for kelps as has been observed in other organisms. Thus, delayed development may be a viable strategy for surviving and initially dominating in environments with variable quality.     

<Emphasis Type="Italic">Macrocystis</Emphasis> mariculture in Chile: growth performance of heterosis genotype constructs under field conditions

Recent progress in Macrocystis mariculture is based on clonal stock cultures of gametophyte parents. Batches of up to 10⁵ genetically identical sporophyte seedlings can be produced at any time in the laboratory and explanted in the field for production of biomass. Sexual crosses of selected Macrocystis pyrifera gametophyte parents of different geographic origin along the coast of Chile showed heterosis and produced sporophyte batches with superior growth performance. Starting from zygotes, after 10 weeks in the laboratory and 5 months in the sea, our best hybrid genotypes grew up to 11 kg fresh weight per frond, which corresponds to 66 kg m⁻¹ of line in a commercial mariculture installation. In contrast, average yields of 14.4 and 22 kg m⁻¹ are reported in the literature for traditional methods. Additional experiments, including inter-specific crosses M. pyrifera × M. integrifolia and their performance in different climate zones of Chile, confirm that heterosis is a powerful tool for crop improvement in Macrocystis. It opens the possibility to construct tailor-made heterosis genotypes with maximum productivity and/or other desired properties for any given locality.