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1.
Rapid ocean warming is affecting kelp forests globally. While the sporophyte life stage has been well studied for many species, the microscopic life stages of laminarian kelps have been understudied, particularly regarding spatial and temporal variations in thermal tolerance and their interaction. We investigated the thermal tolerance of growth, survival, development, and fertilization of Ecklonia radiata gametophytes, derived from zoospores sampled from two sites in Tasmania, Australia, throughout a year, over a temperature gradient (3–30°C). For growth we found a relatively stable thermal optimum at ~20.5°C and stable thermal maxima (25.3–27.7°C). The magnitude of growth was highly variable and depended on season and site, with no consistent spatial pattern for growth and gametophyte size. Survival also had a relatively stable thermal optimum of ~17°C, 3°C below the optimum for growth. Gametophytes grew to single cells between 5 and 25°C, but sporophytes were only observed between 10 and 20°C, indicating reproductive failure outside this range. The results reveal complex effects of source population and season of collection on gametophyte performance in E. radiata, with implications when comparing results from material collected at different localities and times. In Tasmania, gametophytes grow considerably below the estimated thermal maxima and thermal optima that are currently only reached during summer heatwaves, whereas optima for survival (~17°C) are frequently reached and surpassed during heatwaves, which may affect the persistence and recruitment of E. radiata in a warmer climate.  相似文献   

2.
Warming ocean temperatures have been linked to kelp forest declines worldwide, and elevated temperatures can act synergistically with other local stressors to exacerbate kelp loss. The bull kelp Nereocystis luetkeana is the primary canopy-forming kelp species in the Salish Sea, where it is declining in areas with elevated summer water temperatures and low nutrient concentrations. To determine the interactive effects of these two stressors on microscopic stages of N. luetkeana, we cultured gametophytes and microscopic sporophytes from seven different Salish Sea populations across seven different temperatures (10–22°C) and two nitrogen concentrations. The thermal tolerance of microscopic gametophytes and sporophytes was similar across populations, and high temperatures were more stressful than low nitrogen levels. Additional nitrogen did not improve gametophyte or sporophyte survival at high temperatures. Gametophyte densities were highest between 10 and 16°C and declined sharply at 18°C, and temperatures of 20 and 22°C were lethal. The window for successful sporophyte production was narrower, peaking at 10–14°C. Across all populations, the warmest temperature at which sporophytes were produced was 16 or 18°C, but sporophyte densities were 78% lower at 16°C and 95% lower at 18°C compared to cooler temperatures. In the field, bottom temperatures revealed that the thermal limits of gametophyte growth (18°C) and sporophyte production (16–18°C) were reached during the summer at multiple sites. Prolonged exposure of bull kelp gametophytes to temperatures of 16°C and above could limit reproduction, and therefore recruitment, of adult kelp sporophytes.  相似文献   

3.
The annual kelp Eckloniopsis radicosa is distributed along Japanese coasts and occurs within the area with a February isotherm ranging 15–18°C and August isotherm ranging 25–28°C. In this study, the effects of temperature on the gametophyte growth and maturation, and the young sporophyte growth of E. radicosa were examined and the results are discussed in relation to the distribution of other warm‐adapted kelp species and the potential effects of climate change. The optimal temperature ranges for growth of male and female gametophytes were 23–27°C and 20–26°C, respectively. The upper survival temperature for gametophyte growth was 31°C for males and 30°C for females, respectively. The optimal temperature range for maturation of female gametophytes was ≤23°C. The optimal temperature range for growth of young sporophytes was 14–22°C. It was clarified that E. radicosa has the most warm‐adapted characteristics for growth and maturation of gametophytes among members of the Laminariales studied so far. The natural seawater temperature ranges during the growth and maturation seasons for gametophytes of E. radicosa, as well as the growth season for young sporophytes near to the northern and southern distribution limits (Izu‐Oshima: 14.9–24.5°C, Ichiki‐kushikino: 17.1–29.6°C), agreed with the experimentally determined temperature requirements. The warm‐adapted gametophyte stage and annual lifecycle are major factors enabling survival of E. radicosa in warm waters near tropical regions along the Japanese coast.  相似文献   

4.

The kelp Ecklonia radiata has become a target for controlled cultivation. However, to date there are no standardised protocols for the hatchery stage of this species that result in high rates of germination, gametophyte development and transition to sporophytes. Therefore, the objective of this study was to quantify the effect of photoperiod, light intensity, temperature, nutrient media and use of GeO2 on the key hatchery processes of germination, gametophyte development and transition to sporophytes in controlled laboratory experiments. Germination of E. radiata was high (≥?85%) throughout the study, regardless of treatments. Temperature had a major effect on the length of gametophytes, which increased with increasing temperature. The formation of sporophytes was favoured when individuals were maintained under 17 °C continuously, while reduced by approximately 30% when using F/2 compared to PES nutrient media. Overall, the recommended conditions for the hatchery stage of E. radiata are to maintain cultures under a 12 h L:12 h D photoperiod at 17 °C as this resulted in higher germination rates, good gametophyte development and higher transition to sporophytes compared to other treatments. Moreover, the use of GeO2 has to be limited to no more than 2 days as extended use has detrimental effects on the development of sporophytes. Finally, storage of sorus-bearing fronds of sporophytes up to 4 days after the collection from the field generally increased the number of released zoospores and is a simple mechanism to increase the fertility of brood stock.

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5.
Temperate kelp forests (Laminarians) are threatened by temperature stress due to ocean warming and photoinhibition due to increased light associated with canopy loss. However, the potential for evolutionary adaptation in kelp to rapid climate change is not well known. This study examined family‐level variation in physiological and photosynthetic traits in the early life‐cycle stages of the ecologically important Australasian kelp Ecklonia radiata and the response of E. radiata families to different temperature and light environments using a family × environment design. There was strong family‐level variation in traits relating to morphology (surface area measures, branch length, branch count) and photosynthetic performance (Fv/Fm) in both haploid (gametophyte) and diploid (sporophyte) stages of the life‐cycle. Additionally, the presence of family × environment interactions showed that offspring from different families respond differently to temperature and light in the branch length of male gametophytes and oogonia surface area of female gametophytes. Negative responses to high temperatures were stronger for females vs. males. Our findings suggest E. radiata may be able to respond adaptively to climate change but studies partitioning the narrow vs. broad sense components of heritable variation are needed to establish the evolutionary potential of E. radiata to adapt under climate change.  相似文献   

6.
We analysed the effects of temperature and photon fluence rate on meiospore germination, growth and fertility of gametophytes, and growth of young sporophytes of Laminaria ochroleuca. Maximum percentages of germination (91–98%) were obtained at 15°C and 18°C, independent of photon fluence rate. Optimal development of female gametophyte and maximum fecundity and reproductive success of gametophytes occurred at 15°C and 18°C and at 20 and 40 μmol m–2 s–1. Maximum relative growth rate of young sporophytes after 2 weeks of culture was achieved under the same conditions. L. ochroleuca gametophytes cannot reproduce and growth of its sporophytes is not competitive at temperatures close to 10°C. Received in revised form: 31 August 2001 Electronic Publication  相似文献   

7.
Climate change is affecting marine ecosystems in many ways, including raising temperatures and leading to ocean acidification. From 2014 to 2016, an extensive marine heat wave extended along the west coast of North America and had devastating effects on numerous species, including bull kelp (Nereocystis luetkeana). Bull kelp is an important foundation species in coastal ecosystems and can be affected by marine heat waves and ocean acidification; however, the impacts have not been investigated on sensitive early life stages. To determine the effects of changing temperatures and carbonate levels on Northern California's bull kelp populations, we collected sporophylls from mature bull kelp individuals in Point Arena, CA. At the Bodega Marine Laboratory, we released spores from field-collected bull kelp, and cultured microscopic gametophytes in a common garden experiment with a fully factorial design crossing modern conditions (11.63 ± 0.54°C and pH 7.93 ± 0.26) with observed extreme climate conditions (15.56 ± 0.83°C and 7.64 ± 0.32 pH). Our results indicated that both increased temperature and decreased pH influenced growth and egg production of bull kelp microscopic stages. Increased temperature resulted in decreased gametophyte survival and offspring production. In contrast, decreased pH had less of an effect but resulted in increased gametophyte survival and offspring production. Additionally, increased temperature significantly impacted reproductive timing by causing female gametophytes to produce offspring earlier than under ambient temperature conditions. Our findings can inform better predictions of the impacts of climate change on coastal ecosystems and provide key insights into environmental dynamics regulating the bull kelp lifecycle.  相似文献   

8.
The rates of net photosynthesis as a function of irradiance and temperature were determined for gametophytes and embryonic sporophytes of the kelp, Macrocystis pyrifera (L.) C. Ag. Gametophytes exhibited higher net photosynthetic rates based on oxygen and pH measurements than their derived embryonic sporophytes, but reached light saturation at comparable irradiance levels. The net photosynthesis of gametophytes reached a maximum of 66.4 mg O2 g dry wt?1 h?1 (86.5 mg CO2 g dry wt?1 h?1), a value approximately seven times the rate reported previously for the adult sporophyte blades. Gametophytes were light saturated at 70 μE m?2 s?1 and exhibited a significant decline in photosynthetic performance at irradiances 140 μE m?1 s?1. Embryonic sporophytes revealed a maximum photosynthetic capacity of 20.6 mg O2 g dry wt?1 h?1 (25.3 mg CO2 g dry wt?1 h?1), a rate about twice that reported for adult sporophyte blades. Embryonic sporophytes also became light saturated at 70 μE m?2 s?1, but unlike their parental gametophytes, failed to exhibit lesser photosynthetic rates at the highest irradiance levels studied; light compensation occurred at 2.8 μE m?2 s?1. Light-saturated net photosynthetic rates of gametophytes and embryonic sporophytes varied significantly with temperature. Gametophytes exhibited maximal photosynthesis at 15° to 20° C, whereas embryonic sporophytes maintained comparable rates between 10° and 20° C. Both gametophytes and embryonic sporophytes declined in photosynthetic capacity at 30° C. Dark respiration of gametophytes was uniform from 10° to 25° C, but increased six-fold at 30° C; the rates for embryonic sporophytes were comparable over the entire range of temperatures examined. The broader light and temperature tolerances of the embryonic sporophytes suggest that this stage in the life history of M. pyrifera is well suited for the subtidal benthic environment and for the conditions in the upper levels of the water column.  相似文献   

9.
A kelp/red algal symbiosis is described from nature based on extensive collections from the San Juan Islands, Washington. Kelp gametophytes were found as endophytes in the cell walls of seventeen species of red algae in three different kelp communities. Host red algae were mostly filamentous (e.g., Pleonosporium vancouverianum) or polysiphonous (e.g. Polysiphonia paniculata). The kelp gametophytes completed vegetative and reproductive development in the hosts with gametangia formed at the host surface and with sporophytes up to several mm in height being produced while still attached to the host. To date, none of the kelp gametophytes from nature have been identified to genus or species, although the gametophyte of Nereocystis luetkeana is a potential candidate for the symbiosis. Preliminary observations from Nova Scotia and the Isle of Man have not found the association in the Atlantic Ocean. Laboratory studies in Korea successfully reconstructed the symbiosis in the red alga Aglaothamnion oosumiense using zoospores of Undaria pinnatifida but not Laminaria religiosa. Here we outline the development of the symbiosis and discuss the potential adaptive significance of the kelp/red algal interaction.  相似文献   

10.
We tested the ability of sporophytes of a small kelp, Ecklonia radiata (C. Agardh) J. Agardh, to adjust their photosynthesis, respiration, and cellular processes to increasingly warm ocean climates along a latitudinal gradient in ocean temperature (~4°C). Tissue concentrations of pigment and nutrients decreased with increasing ocean temperature. Concurrently, a number of gradual changes in the metabolic balance of E. radiata took place along the latitudinal gradient. Warm‐acclimatized kelps had 50% lower photosynthetic rates and 90% lower respiration rates at the optimum temperature than did cool‐acclimatized kelps. A reduction in temperature sensitivity was also observed as a reduction in Q10‐values from cool‐ to warm‐acclimatized kelps for gross photosynthesis (Q10: 3.35 to 1.45) and respiration (Q10: 3.82 to 1.65). Respiration rates were more sensitive to increasing experimental temperatures (10% higher Q10‐values) than photosynthesis and had a higher optimum temperature, irrespective of sampling location. To maintain a positive carbon balance, E. radiata increased the critical light demand (Ec) exponentially with increasing experimental temperature. The temperature dependency of Ec was, however, weakened with increasing ocean temperature, such that the critical light demand was relaxed in kelp acclimated to higher ocean temperatures. Nevertheless, calculations of critical depth limits suggested that direct effects of future temperature increases are unlikely to be as strong as effects of reduced water clarity, another globally increasing problem in coastal areas.  相似文献   

11.
Delayed recruitment of microscopic stages in response to cyclical cues is critical to the population dynamics of many annual and seasonally reproducing perennial seaweeds. Microscopic stages may play a similar role in continuously reproducing perennials in which adult sporophytes are subject to episodic mortality, if they can respond directly to the unpredictable onset and relaxation of unfavorable conditions. We experimentally evaluated the potential for temporary reduction in limiting resources (light, nutrients) to directly delay recruitment of giant kelp (Macrocystis pyrifera (L.) C.A. Agardh) gametophytes and embryonic sporophytes. Laboratory cultures were subjected to limiting conditions of light and nutrients for 1 month and then exposed to nonlimiting conditions for 10 days. Gametophytes in all treatments failed to recruit to sporophytes after 2 weeks, suggesting they are not a source of delayed recruitment in giant kelp. Sporophytes in light‐limited treatments, however, survived and grew significantly slower than non–light‐limited controls. When stimulated with light, light‐limited sporophytes grew from 2 to>10 times faster than unstimulated controls depending on nutrient availability. These results suggest that limiting resources can delay recruitment of embryonic giant kelp sporophytes for at least 1 month. Flexible timing of recruitment from embryonic sporophytes may enhance persistence of continuously reproducing perennial species when mac‐ roscopic adults are subject to episodic large‐scale removals.  相似文献   

12.
The giant kelp Macrocystis pyrifera (L.) C. Agardh is widely distributed in the Northern Hemisphere and Southern Hemisphere, yet it exhibits distinct population dynamics at local to regional spatial scales. Giant kelp populations are typically perennial with the potential for year‐round reproduction and recruitment. In southern Chile, however, annual giant kelp populations exist and often persist entirely on secondary substrata (e.g., shells of the slipper limpet Crepipatella fecunda [Gastropoda, Calyptraeidae]) that can cover up to 90% of the rocky bottom. In these populations, the macroscopic sporophyte phase disappears annually during winter and early spring, leaving a 3–4 month period in which a persistent microscopic phase remains to support the subsequent year’s recruitment. We tested the effects of a suite of grazers on the recruitment success of this critical microscopic phase at two sites in southern Chile. Field experiments indicated that the snail Tegula atra negatively impacted M. pyrifera sporophyte recruitment, but that recruitment was highest in the presence of sessile female limpets, C. fecunda. Conversely, small male C. fecunda (biofilm grazers) did not regulate kelp recruitment. Laboratory observations showed that C. fecunda males only grazed on microscopic kelp gametophytes and small (<250 μm) sporophytes, rejecting larger sporophytes, whereas T. atra grazed on all the kelp stages. Recruitment to the C. fecunda treatments far exceeded that to bare rock in the absence of grazers but was not due to the physical presence of C. fecunda shells. We concluded that the key to M. pyrifera recruitment success in southern Chile is its capacity to colonize secondary substrates provided by the slipper limpet C. fecunda.  相似文献   

13.
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.  相似文献   

14.
At an early stage of growth gametophytes support the sporophytes of ferns. Young sporophytes become independent of gametophytes when the first leaves develop. Although large fern gametophytes produce multiple archegonia simultaneously, only one sporophyte is typically established on one gametophyte. The number of sporophytes is believed to be controlled in two possible directions, from gametophyte to sporophyte or from preceding sporophyte to another sporophyte. To investigate the effects of gametophytes on their sporophytes, we studied the relationship between organic matter production by gametophytes and the growth of young sporophytes of Thelypteris palustris. We cut gametophytes in half (CGs) to reduce the gametophytes’ production of matter. There was no significant difference between the growth of sporophytes on intact gametophytes (IGs) and that on CGs. According to our estimates, based on the rate of organic matter production, the large gametophyte was able to produce two or more sporophytes. The resources required for CGs to make similar-sized sporophytes was twice that for IGs. In polyembryony each of the multiple sporophytes was similar in size to the single sporophytes. Resource limitation does not seem to explain why fern gametophytes establish single sporophytes.  相似文献   

15.
Kelps are in global decline due to climate change, which includes ocean warming. To identify vulnerable species, we need to identify their tolerances to increasing temperatures and determine whether tolerances are altered by co-occurring drivers such as inorganic nutrient levels. This is particularly important for those species with restricted distributions, which may already be experiencing thermal stress. To identify thermal tolerance of the range-restricted kelp Lessonia corrugata, we conducted a laboratory experiment on juvenile sporophytes to measure performance (growth, photosynthesis) across its thermal range (4–22°C). We determined the upper thermal limit for growth and photosynthesis to be ~22–23°C, with a thermal optimum of ~16°C. To determine if elevated inorganic nitrogen availability could enhance thermal tolerance, we compared the performance of juveniles under low (4.5 μmol · d−1) and high (90 μmol · d−1) nitrate conditions at and above the thermal optimum (16–23.5°C). Nitrate enrichment did not enhance thermal performance at temperatures above the optimum but did lead to elevated growth rates at the thermal optimum. Our results indicate L. corrugata is likely to be extremely susceptible to moderate ocean warming and marine heatwaves. Peak sea surface temperatures during summer in eastern and northeastern Tasmania can reach up to 20–21°C, and climate projections suggest that L. corrugata's thermal limit will be regularly exceeded by 2050 as southeastern Australia is a global ocean-warming hotspot. By identifying the upper thermal limit of L. corrugata, we have taken a critical step in predicting the future of the species in a warming climate.  相似文献   

16.
Abstract The sea urchin Centrostephanus rodgersii grazes areas of reef free of large brown algae (the barrens habitat). Survival of the kelp Ecklonia radiata adjacent to patches of barrens habitat was compared to that in the centre of kelp forests (centre) and edges of forests not adjacent to patches of barrens habitat (ungrazed). Estimates of rates of instantaneous mortality for tagged plants, as described by the slope of a negative exponential model (± 95% CI) were: barrens, 0.078 ± 0.004; centre, 0.051 ± 0.004 and ungrazed edge, 0.065 ± 0.007. Survival of plants was greatest in the middle of forests and least on the margins of patches of barrens habitat. A significant proportion of mortality in the barrens and centre positions was caused by herbivorous fish. When these plants were excluded from analysis there were no significant differences in survival between the barrens and ungrazed positions. It is concluded that C. rodgersii has little impact on the abundance of Ecklonia outside sharply defined boundaries.  相似文献   

17.
Levenbach S 《Oecologia》2009,159(1):181-190
Recent studies have emphasized the role of positive interactions in ecological communities, but few have addressed how positive interactions are mediated by abiotic stress and biotic interactions. Here, I investigate the effect of a facilitator species on the abundance of macroalgae over a gradient of herbivory. Grazing by sea urchins can be intense on temperate reefs along the California coast, with benthic macroalgae growing exclusively in physical refuges and interspersed within colonies of the strawberry anemone, Corynactis californica. Field experiments indicated that the net effect of C. californica on turf algae was strongly nonlinear over a gradient in density of sea urchins. At low intensities of urchin grazing, the anemone and macroalgae competed for space, with algae capable of overgrowing C. californica. At intermediate grazing intensities, C. californica provided a refuge for turf algae but not for juvenile kelp. Neither turf algae nor kelp benefited from the presence of C. californica at the highest levels of grazing intensity, as sea urchins consumed nearly all macroalgae. The hump-shaped effect observed for C. californica contrasts with the prevailing view in ecological theory that positive interactions are more common in harsh environmental conditions. The results reported here qualify this view and underscore the need to evaluate positive interactions over a range of abiotic stress and consumer pressure.  相似文献   

18.
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.  相似文献   

19.

The kelp Lessonia corrugata (Ochrophyta, Laminariales) is being developed for integrated multi-trophic aquaculture (IMTA) trials in the vicinity of salmon cages in Tasmania, Australia. Gametophytes are vegetally maintained before seeding on hatchery twine; however, the optimal temperature and light conditions for growth and sexual development are unknown. We measured vegetative size of female and male gametophytes and sexual development of females over a range of temperatures and irradiances using a temperature gradient table and neutral density light filters. Over a 4-week experiment, gametophytes were exposed to a combination of thermal (5.7–24.9 °C) and irradiance (10–100 μmol photons m?2 s?1) gradients, to assess biological performance. At the temperature extremes (hottest = 24.9 °C, coldest = 5.7 °C), we observed the critical thermal limits for this species and the results reveal a narrow optimal temperature range for growth and sexual development between 15.7 and 17.9 °C, with irradiances between 40 and 100 μmol photons m?2 s?1 resulting in fertile female gametophytes. Lessonia corrugata inhabits a small geographic range, found only around Tasmania, south of the Australian mainland, hence oceanic changes such as ongoing increases in sea surface temperatures (SSTs), and altered irradiance regimes may limit recruitment of the early microscopic life stages in the future. Our findings provide optimised culture conditions for aquaculture and information to predict the future geographic range of L. corrugata under ocean global change.

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20.
This paper describes the formation of gametophytes and the early stages of the development of sporophytes in the kelp seaweed Alaria angusta from Kamchatka. To establish laboratory cultures we used zoospores obtained from A. angusta sporophytes collected on October 29, 2014. The gametophytes were grown under different conditions: at 6–7°С and natural light and at 10°С and illumination with cool white fluorescent bulbs, 30 μmol photon m–2 s–1, 12: 12 h L: D cycle. In the first case (natural light, 6–7°С), the vegetative growth of male and female gametophytes lasted for more than 4 months; maturation of sexual products occurred 144 days after germination of the embryospores. In the second case (artificial light, 10°С), rapid development of the gametophytes occurred; the first juvenile sporophytes appeared on the 10th day after the zoospores settled onto the substrate. Our data contribute to understanding of the regulatory effect of temperature and light on the peculiarities of species vegetation in the natural environment and formation of the age structure of the species population, one particular feature of this process is the continuous appearance of juvenile sporophytes in the warm period of the year.  相似文献   

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