FIELD MEASUREMENTS OF INORGANIC NITROGEN UPTAKE BY EPIFLORA COMPONENTS OF THE SEAGRASS POSIDONIA OCEANICA (MONOCOTYLEDONS,POSIDONIACEAE)1

Crustose corallines, crustose and erect brown algae, and sessile animals are major components of the epiphytic community of the Mediterranean seagrass Posidonia oceanica (L.) Delile. Production, biomass, and specific composition of this epiphyte–seagrass association are impacted by anthropogenic increase of nutrient load in this oligotrophic area. In this context, nitrogen uptake by P. oceanica and its epiflora was measured using the isotope ¹⁵N at a 10 m depth in the Revellata Bay (Corsica, Mediterranean Sea). Epiflora components showed various seasonal patterns of biomass and abundance. The epiphytic brown algae appeared at the end of spring, later than the crustose corallines, and after the nitrate peak in the bay. Because of their later development in the season, epiphytic brown algae mostly rely on ammonium for their N needs. We hypothesize that the temporal succession of epiphytic organisms plays a crucial role in the N dynamics of this community under natural conditions. The epiphytic brown algae, which have a growth rate one order of magnitude greater than that of crustose corallines, showed lower N‐uptake rates. The greater N‐uptake rates of crustose corallines probably reflect the greater N requirements (i.e., lower C/N ratios) of red algae. We determined that the epiflora incorporated ammonium and nitrate more rapidly than their host. Nevertheless, when biomass was taken into account, P. oceanica was the most important contributor to N uptake from the water column by benthic macrophytes in this seagrass bed.     

Limited flexibility in resource use in a coral reef grazer foraging on seasonally changing algal communities

Feeding ecology of three life phases of the parrotfish Scarus ferrugineus was studied on a southern Red Sea fringing reef by comparing availability and consumption of benthic algae during the monsoon hot and cool seasons. Dominant biota covering dead carbonate substrates were in decreasing order of importance: turfs on endoliths, turfs on crustose corallines, and crustose corallines. On the reef crest and shallow fore reef, composition of the biota changed seasonally. Cover of turfs on endoliths and turfs on crustose corallines was higher during the hot season, while crustose corallines and macroalgae (only on reef crest) increased during the cool season. Biota in the deep fore reef did not show seasonal variation. All life phases used similar resources and showed selective feeding in all zones. Turfs on endoliths, followed by turfs on crustose corallines, was the primary feeding substrate. These two sources represented over 92% of bites during both seasons. Crustose corallines, macroalgae, and living corals were negligible components being strongly avoided at all zones and seasons. Resource use varied seasonally on the reef crest and shallow fore reef, while it remained unchanged on the deep fore reef. Turfs on endoliths were consistently preferred in both seasons but their contribution increased from 45% in the cool to 70% of bites in the hot season. Electivity for turfs on crustose corallines shifted from random feeding in the hot (27% of bites) to selection in the cool season (47% of bites). Feeding pattern changed diurnally with more bites taken from crustose corallines and turfs on crustose corallines during morning. During the rest of the day, bites from turfs on endoliths predominate. S. ferrugineus shows limited capacity to exploit seasonal increases in the biomass of foliose and canopy forming macroalgae, despite indications of energetic limitation during the cool season.     

Temperature × light interaction and tolerance of high water temperature in the planktonic freshwater flagellates Cryptomonas (Cryptophyceae) and Dinobryon (Chrysophyceae)

Using microcosm experiments, we investigated the interactive effects of temperature and light on specific growth rates of three species each of the phytoplanktonic genera Cryptomonas and Dinobryon. Several species of these genera play important roles in the food web of lakes and seem to be sensitive to high water temperature. We measured growth rates at three to four photon flux densities ranging from 10 to 240 μmol photon · m^?2 · s^?1 and at 4–5 temperatures ranging from 10°C to 28°C. The temperature × light interaction was generally strong, species specific, and also genus specific. Five of the six species studied tolerated 25°C when light availability was high; however, low light reduced tolerance of high temperatures. Growth rates of all six species were unaffected by temperature in the 10°C–15°C range at light levels ≤50 μmol photon · m^?2 · s^?1. At high light, growth rates of Cryptomonas spp. increased with temperature until the temperature optimum was reached and then declined. The Dinobryon species were less sensitive than Cryptomonas spp. to photon flux densities of 40 μmol photon · m^?2 · s^?1 and 200 μmol photon · m^?2 · s^?1 over the entire temperature range but did not grow under a combination of very low light (10 μmol photon · m^?2 · s^?1) and high temperature (≥20°C). Among the three Cryptomonas species, cell volume declined with temperature and the maximum temperature tolerated was negatively related to cell size. Since Cryptomonas is important food for microzooplankton, these trends may affect the pelagic carbon flow if lake warming continues.     

Light and temperature dependence of the emission of cyclic and acyclic monoterpenes from holm oak (Quercus ilex L.) leaves

In a laboratory study, we investigated the monoterpene emissions from Quercus ilex, an evergreen sclerophyllous Mediterranean oak species whose emissions are light dependent. We examined the light and temperature responses of individual monoterpenes emitted from leaves under various conditions, the effect of heat stress on emissions, and the emission-onset during leaf development. Emission rate increased 10-fold during leaf growth, with slight changes in the composition. At 30 °C and saturating light, the monoterpene emission rate from mature leaves averaged 4·1 nmol m^–2 s^–1, of which α-pinene, sabinene and β-pinene accounted for 85%. The light dependence of emission was similar for all monoterpenes: it resembled the light saturation curve of CO₂ assimilation, although monoterpene emission continued in the dark. Temperature dependence differed among emitted compounds: most of them exhibited an exponential increase up to 35 °C, a maximum at 42 °C, and a slight decline at higher temperatures. However, the two acyclic isomers cis-β-ocimene and trans-β-ocimene were hardly detected below 35 °C, but their emission rates increased above this temperature as the emission rates of other compounds fell, so that total emission of monoterpenes exponentially increased from 5 to 45 °C. The ratio between ocimene isomers and other compounds increased with both absolute temperature and time of heat exposure. The light dependence of emission was insensitive to the temperature at which it was measured, and vice versa the temperature dependence was insensitive to the light regime. The results demonstrated that none of the models currently applied to simulate isoprene or monoterpene emissions correctly predicts the short-term effects of light and temperature on Q. ilex emissions. The percentage of fixed carbon lost immediately as monoterpenes ranged between 0·1 and 6·0% depending on temperature, but rose up to 20% when leaves were continuously exposed to temperatures between 40 and 45 °C.     

Temperature requirements for growth and survival of macroalgae from Disko Island (Greenland)

The temperature requirements for growth and upper temperature tolerance were determined in 16 macroalgal species collected on Disko Island (Greenland). The upper survival temperatures were examined in 1°C steps, and growth measured at 5°C intervals between 0 and 20°C using a refined method, where the fresh weight was determined weekly or fortnightly over a period of 5 or 6 weeks. To express temperature-growth responses, growth rates of temperature-acclimated plants were taken. Two groups with different temperature requirements were identified: (1) A stenothermal group includingAcrosiphonia arcta, Acrosiphonia sonderi, Urospora penicilliformis, Devaleraea ramentacea, Desmarestia aculeata, Pilayella littoralis, growing between 0 and (10 to) 15 (or 20)°C with optima between 0 and 10°C. The upper survival temperatures in these species and inChromastrum secundatum, Chromastrum virgatulum, Chordaria flagelliformis were between 17 and 23°C (duration of experiment: 2 weeks). (2) A eurythermal group includingEnteromorpha clathrata, Enteromorpha intestinalis andPolysiphonia urceolata growing between 0 and 20°C with growth optima at 10 or 15°C. The upper survival temperatures in these species and inChaetomorpha tortuosa, Bangia atropurpurea andEudesme virescens were between 24 and 31°C. These algal species showed little adaptation to the Arctic temperatures. In contrast, algae from the first group exhibited a relatively high adaptation to low temperatures — approaching the low temperature requirements of Antarctic algae. The results are discussed in relation to the geographic distribution of individual species.     

The genus phymatolithon in the Gulf of Maine

Summary New information on anatomy, cytology and the development of reproductive structures is presented to show that Phymatolithon is a genus distinct from both Clathromorphum and the branching members of Lithothamnium. Also, a new species of Phymatolithon, Ph. rugulosum, is described. The reproductive cycles and geographic and bathymetric distributions of Ph. laevigatum and Ph. rugulosum in the Gulf of Maine are presented and discussed. There is strong indication that the geographic distribution of crustose corallines in the region is controlled primarily by maximum summer temperatures. The depth distributions are apparently controlled primarily by decrease of light with depth, though temperatures and substrate are also factors.Now at Colorado School of Mines, Golden, Colorado, U.S.A. The necessary field work for this study was made possible through the generous support of: The American Petroleum Institute, The Geological Society of America, the ONR (through the Department of Geology and Geophysics, Massachusetts Institute of Technology) and the Woods Hole Oceanographie Institution (Contr. No. 1500).     

Growth temperature affects O2 consumption rates and plasticity of respiratory flux to support shoot growth at various growth temperatures

The temperature dependence of respiration rates and their acclimation to growth temperature vary among species/ecotypes, but the details remain unclear. Here, we compared the temperature dependence of shoot O₂ consumption rates among Arabidopsis thaliana ecotypes to clarify how the temperature dependence and their acclimation to temperature differ among ecotypes, and how these differences relate to shoot growth. We examined growth analysis, temperature dependence of O₂ consumption rates, and protein amounts of the respiratory chain components in shoots of twelve ecotypes of A. thaliana grown at three different temperatures. The temperature dependence of the O₂ consumption rates were fitted to the modified Arrhenius model. The dynamic response of activation energy to measurement temperature was different among growth temperatures, suggesting that the plasticity of respiratory flux to temperatures differs among growth temperatures. The similar values of activation energy at growth temperature among ecotypes suggest that a similar process may determine the O₂ consumption rates at the growth temperature in any ecotype. These results suggest that the growth temperature affects not only the absolute rate of O₂ consumption but also the plasticity of respiratory flux in response to temperature, supporting the acclimation of shoot growth to various temperatures.     

Aragonite infill in overgrown conceptacles of coralline Lithothamnion spp. (Hapalidiaceae,Hapalidiales, Rhodophyta): new insights in biomineralization and phylomineralogy

New empirical and quantitative data in the study of calcium carbonate biomineralization and an expanded coralline psbA framework for phylomineralogy are provided for crustose coralline red algae. Scanning electron microscopy (SEM) and energy dispersive spectrometry (SEM‐EDS) pinpointed the exact location of calcium carbonate crystals within overgrown reproductive conceptacles in rhodolith‐forming Lithothamnion species from the Gulf of Mexico and Pacific Panama. SEM‐EDS and X‐ray diffraction (XRD) analysis confirmed the elemental composition of these calcium carbonate crystals to be aragonite. After spore release, reproductive conceptacles apparently became overgrown by new vegetative growth, a strategy that may aid in sealing the empty conceptacle chamber, hence influencing the chemistry of the microenvironment and in turn promoting aragonite crystal growth. The possible relevance of various types of calcium carbonate polymorphs present in the complex internal structure and skeleton of crustose corallines is discussed. This is the first study to link SEM, SEM‐EDS, XRD, Microtomography and X‐ray microscopy data of aragonite infill in coralline algae with phylomineralogy. The study contributes to the growing body of literature characterizing and speculating about how the relative abundances of carbonate biominerals in corallines may vary in response to changes in atmospheric pCO₂, ocean acidification, and global warming.     

THE CRUSTOSE CORALLINES OF THE NORTHWESTERN NORTH ATLANTIC,INCLUDING LITHOTHAMNIUM LEMOINEAE N. SP.1

Reproductive, vegetative, anatomical, and depth distribution keys are presented for the saxicolous crustose corallines of the northwestern North Atlantic. The 3 species of Lithothamnium in the region are named, while Lithothamnium lemoineae is described as a new species.     

The cost of mutualism: interactions between Trollius europaeus and its pollinating parasites

Summary Photosynthetic acclimation to 5 light environments ranging from 2 to 60% full sun was determined in Alocasia macrorrhiza, a shade tolerant species from tropical forest understories, and Colocasia esculenta, a cultivated species which occurs naturally in open marshy areas. Photosynthetic capacities of both species increased nearly 3 fold with increased photon flux density (PFD). In a given environment, however, photosynthetic capacities of C. esculenta were double those of A. macrorrhiza. Stomatal limitations explained only a small part of this difference. Respiration rates and estimated biochemical capacities increased in parallel to photosynthetic capacity. No differences were observed either between species or environments in the ratio of RuBP regeneration capacity to carboxylation capacity as determined from the CO₂ dependence response of photosynthesis. Quantum yields of both species decreased only slightly with increasing growth PFD, providing little evidence for photoinhibition at high PFD. The results are discussed in terms of the mechanisms of and limitations on acclimation in these two species.     

THE EFFECT OF DAYLENGTH AND LIGHT INTENSITY ON THE GROWTH RATE OF THE MARINE DIATOM DETONULA CONFERVACEA (CLEVE) GRAN1

The influence of 40 combinations of temperature (2, 7 C), light intensity (50, 200, 600, 1200, 1800 ft-c), and photoperiod (24, 15:9, 12:12, and 8:16 LD) at 30% salinity on the rate of cell division of the Narragansett Bay clone of Detonula confervacea (Det-1) was examined following appropriate preconditioning. At 2 C Detonula is a long day species (24 L) and prefers low light intensities (200–600 ft-c); poorest growth occurred at 12:12 and 8:16 LD, and the compensation intensity was about 10 ft-c. Increasing temperature to 7 C increased the mean growth rate, reduced the optimal daylength (15:9 LD), even though Detonula remained a long day species and increased the optimal light intensity (600–800 ft-c). The compensation intensity varied with daylength and ranged from about 10–50 ft-c. Photoperiods of 12:12 and 8:16 LD were least favorable for growth at both temperatures; light limitation and inhibition were observed at 50 and 1800 ft-c. respectively; inhibition was less pronounced at 7 C. There is some indication that the conditions of growth that the stock cultures were exposed to prior to preconditioning for use in the experiments may have sometimes influenced response. Detonula produced resting spores without nutrient depletion at 2 and 7 C at all light intensities when the photoperiod was lengthened. Auxospore formation was also observed. Although short daylengths (9:15 LD) limit Detonula's growth during the early stages of the winter bloom, it competes successfully against Skeletonema costatum initially. This results from its higher rates of growth and of photosynthesis at the prevailing temperature and light conditions and a lower compensation intensity than reported for Skeletonema. The main causes of Detonula's growth inception and termination in Narragansett Bay differ.     

High thermal acclimation potential of both photosynthesis and respiration in two lowland Plantago species in contrast to an alpine congeneric

Thermal acclimation of photosynthesis and respiration can enable plants to maintain near constant rates of net CO₂ exchange, despite experiencing sustained changes in daily average temperature. In this study, we investigated whether the degree of acclimation of photosynthesis and respiration of mature leaves differs among three congeneric Plantago species from contrasting habitats [two fast‐growing lowland species (Plantago major and P. lanceolata), and one slow‐growing alpine species (P. euryphylla)]. In addition to investigating some mechanisms underpinning variability in photosynthetic acclimation, we also determined whether leaf respiration in the light acclimates to the same extent as leaf respiration in darkness, and whether acclimation reestablishes the balance between leaf respiration and photosynthesis. Three growth temperatures were provided: constant 13, 20, or 27°C. Measurements were made at five temperatures (6–34°C). Little acclimation of photosynthesis and leaf respiration to growth temperature was exhibited by P. euryphylla. Moreover, leaf masses per area (LMA) were similar in 13°C‐grown and 20°C‐grown plants of the alpine species. In contrast, growth at 13°C increased LMA in the two lowland species; this was associated with increased photosynthetic capacity and rates of leaf respiration (both in darkness and in the light). Alleviation of triose phosphate limitation and increased capacity of electron transport capacity relative to carboxylation were also observed. Such changes demonstrate that the lowland species cold‐acclimated. Light reduced the short‐term temperature dependence (i.e. Q₁₀) of leaf respiration in all three species, irrespective of growth temperature. Collectively, our results highlight the tight coupling that exists between thermal acclimation of photosynthetic and leaf respiratory metabolism (both in darkness and in the light) in Plantago. If widespread among contrasting species, such coupling may enable modellers to assume levels of acclimation in one parameter (e.g. leaf respiration) where details are only known for the other (e.g. photosynthesis).     

The influence of temperature on growth rate and length of larval life of the gastropod, Crepidula plana Say

The relationship between rate of larval development and the potential to prolong larval life was examined for larvae of the marine prosobranch gastropod Crepidula plana Say. Larvae were maintained in clean glass dishes at constant temperatures ranging from 12–29°C and fed either Isochrysis galbana Parke (ISO) or a Tahitian strain of Isochrysis species (T-ISO). Under all conditions, larvae grew at constant rates, as determined by measurements of shell length and tissue biomass. Most larvae eventually underwent spontaneous metamorphosis. Regardless of temperature, faster growth was associated with a shorter planktonic stage prior to spontaneous metamorphosis. Within an experiment, higher temperatures generally accelerated growth rates and reduced the number of days from hatching to spontaneous metamorphosis. However, growth rates were independent of temperature for larvae fed ISO at 25 and 29°C and for larvae fed T-ISO at 20 and 25°C. Where growth rates were unaffected by temperature, time to spontaneous metamorphosis was similarly unaffected. Maximum durations of larval life at a given temperature were shorter for larvae of Crepidula plana than for those of the congener C. fornicata (L.), although both species grew at comparable rates. Interpretations of the ecological significance of these interspecific differences in delay capabilities will require additional data on adult distributions and larval dispersal patterns in the field.     

Effects of temperature on growth rates of fungi from subantarctic Macquarie Island and Casey,Antarctica

Summary The linear growth rates of fungal isolates were measured on agar plates at temperatures ranging from 4° to 35°C. Fungi tested included the major fungal colonizers of leaves and litter of the three dominant plant species on subantarctic Macquarie Island, and major fungal species associated with plant and soil communities near Australia's Casey Station on the Antarctic Continent. All fungi grew at 4°C and were classified as psychrotrophs. Maximum growth rates were recorded at temperatures of 10° to 20°C for 13 of the 15 isolates from Macquarie Island and for all six isolates from Casey. Most of the leaf colonizing fungi from Macquarie Island had optimum growth temperatures of 15°C whereas all litter fungi from Macquarie Island and Casey fungi except Thelebolus microsporus had optimum growth temperatures of 20°C or above. Maximum growth of all species was at temperatures above those normally prevailing in their natural environments, with most species growing at 4°C at between 10% and 30% of their maximum rates. However, microclimatic effects may have resulted at times in temperatures near their growth optima. The highest growth rates at 4°C were recorded for Phoma spp. 1 and 2, Phoma exigua and Mortierella gamsii from Macquarie Island and Mortierella sp. 1 from Casey. Thelebolus microsporus and sterile sp. G from Casey also grew relatively fast at 4°C, and these species, and Phoma sp. 3 and Phoma exigua from Macquarie Island had the lowest Q-₁₀ values for the temperature range 4° to 15°C.     

Temperature dependence of nitrate reductase activity in marine phytoplankton: biochemical analysis and ecological implications

The temperature dependence of NADH:NR activity was examined in several marine phytoplankton species and vascular plants. These species inhabit divergent thermal environments, including the chromophytes Skeletonema costatum (12–15° C), Skeletonema tropicum (18–25° C), Thalassiosira antarctica (?2 to 4° C), and Phaeocystis antarctica (?2 to 4° C), the green alga Dunaliella tertiolecta (14–28° C), and the vascular plants Cucurbita maxima (20–35° C) and Zea mays (20–25° C). Despite the difference in growth habitats, similar temperature response curves were observed among the chromophytic phytoplankton, with temperatures optimal for NR activity being between 10–20° C. In contrast, the chlorophyll b‐containing alga and vascular plants exhibited optimal temperatures for NR activity above 30° C. Such dramatic differences in NR thermal characteristics from the two taxonomic groups reflect a divergence in NR structure that may be associated with the evolutionary diversification of chromophytes and chlorophytes. Further, it suggests a potential contribution of the thermal performance of NR to the geographic distributions, seasonal abundance patterns, and species composition of phytoplankton communities. NR partial activities, which assess the individual functions of Mo‐pterin and FAD domains, were evaluated on NR purified from S. costatum to determine the possible causes for high temperature (>20° C) inactivation of NR from chromophytes. It was found that the FAD domain and electron transport among redox centers were sensitive to elevated temperatures. S. costatum cells grown at 5, 15, and 25° C exhibited an identical optimal temperature (15° C) for NADH:NR activity, whereas the maximal NR activity and NR protein levels differed and were positively correlated with growth temperature and growth rate. These findings demonstrate that thermal acclimation of NO₃^? reduction capacity is largely at the level of NR protein expression. The consequences of these features on NO₃^? utilization are discussed.     

AUTECOLOGICAL STUDIES ON THE MARINE DIATOM RHIZOSOLENIA FRAGILISSIMA BERGON. I. THE INFLUENCE OF LIGHT,TEMPERATURE, AND SALINITY1

The influence of 113 combinations of temperature (9, 12, 18, 25, 30 C), salinity (5–35 ‰ at 5 ‰ intervals), and light (4 levels) on the mean daily cell division rate (K) of the Narragansett Bay clone of Rhizosolenia fragilissima was examined following appropriate preconditioning. Growth did not occur below 9 C, but was excellent (K =～1.2) under certain combinations of light and salinity at 12, 18, and 25 C. The optimal salinity of 20–25 ‰ was temperature independent. Growth was not measurable at 5 ‰, although survival occurred. At 20 ‰ and 1200 ft-c, K increased approximately 1.8-fold from 0.65 to ～1.2 between 9 and 18–25 C. The optimal light intensity was generally 600 ft-c, although several light-temperature-salinity trends were found. At 10 ‰ at all temperatures, the mean daily division rate decreased with increasing light above 600 ft-c, a response found at all salinities at 12 C, but not at other temperatures. Between 15 and 25 ‰, at 18 and 25 C, mean growth was independent of light intensity; at 30–35 ‰ a direct relationship with light may be present with maximum growth occurring at 1200–1800 ft-c. The in situ and in vitro responses of Rhiz. fragilissima to salinity and the optimum and upper temperature levels are in general agreement. However, growth failure below 9 C in vitro is at odds with reports that natural populations occur even at ?1.11 C. The questions of to what extent this discrepancy reflects the occurrence of thermal clones, different taxa, and/or experimental artifacts are briefly discussed. It is suggested that naturally occurring populations found below 9 C might be designated as Rhiz. fragilissima f. faeröensis, and that Rhiz. fragilissima f. bergonii be used for populations growing at higher temperatures, until this matter is resolved. Observations on auxospore formation are presented.     

Day and night drift-feeding by juvenile salmonids at low water temperatures

Drift-feeding salmonids in boreal streams face temperatures below physical optima for extensive periods of the year. Because juvenile salmonids react to low water temperatures by becoming nocturnal, knowledge about their foraging ability at low light intensities in cold water is needed to accurately estimate energy intake during non-summer conditions. In a laboratory stream channel, we studied temperature effects on the drift-feeding behaviour of juvenile Atlantic salmon, brown trout, and European grayling in simulated daylight and moonlight at temperatures ranging from 2 °C to 11 °C. Prey capture probability was positively related to temperature, but the temperature dependence did not agree with predictions of the Metabolic Theory of Ecology. Furthermore, reaction distance was positively related to temperature for the three species, which may be one of the underlying mechanisms responsible for the temperature effects on prey capture probability. Overall, the three species had similar capture rates at the different temperature and light levels, although there were species differences. European grayling had a slightly higher prey capture probability than brown trout, and brown trout had a shorter reaction distance than Atlantic salmon and European grayling. These results have implications for both energetics-based drift-foraging theory and for studies of winter ecology.     

The effect of light and temperature on the growth and photosynthesis of Gracilariopsis chorda (Gracilariales, Rhodophtya) from geographically separated locations of Japan

The effect of light and temperature on the growth and photosynthesis of the Japanese agarophyte, Gracilariopsis chorda (Gracilariaceae, Rhodophyta), was determined to better understand its physiology so that we could identify candidates for mass cultivation. Above the photosynthetic active radiation of 66 μmol photons m^?2 s^?1, photosynthetic rates saturated for all strains that were collected from six different locations (Hokkaido, Chiba, Tokushima, Saga, Kagoshima, and Okinawa); furthermore, either photosynthesis or growth was observed at all temperature treatments examined in our study (4–32 °C for photosynthesis, 16–32 °C for growth experiments). We identified a temperature range for optimal photosynthesis and growth, which occurred within 20.1–29.1 °C and roughly correlated with the water temperatures of the collection locations and strongly suggests that this species tolerates a wide variety of water temperature. In particular, the Kagoshima strain had the widest range of optimal temperatures (20.8–29.1 °C), whereas the Saga strain had the narrowest range (23.1–27.3 °C). It is important to note that all the optimal temperature ranges overlapped among the strains; therefore, no definitive distinction can be determined. The broad tolerance to temperatures commonly observed from northern to southern Japan suggests that the cultivation of this species should succeed during spring to summer in the majority of the coastal regions in Japan.