Thallus Size Research Articles

RETRACTED ARTICLE: Derbesia maxima sp. nov. (Bryopsidales, Chlorophyta) from Korea based on morphological and molecular data

A green alga was collected from Daejin, located on the eastern coast of Korea. It shows a typical siphonous thallus structure composed of erect and prostrate filaments like other Derbesia sporophytes and is characterized by a relatively large sporophytic thallus (6–8 cm) without calcified attachment disk, irregularly or subdichotomously branched filaments with a basal septum and 1(2) pyrenoid per ovoid to lenticular, occasionally irregular chloroplast. Sporangia have an obovoid or pyriform shape with a short pedicel and septum at the base. Vesicular gametophytes are spherical or ovoid, with short anchoring stalk and rhizoids. A heteromorphic life history alternating between sporophytic filaments and gametophytic Halicystis stage was observed. In a phylogenetic tree based on rbcL sequence, the Korean entity nests in a sister clade to several species groups including the type Derbesia marina, and of these species, D. fastigiata is closest to the Korean alga with a genetic divergence of 3.1%. However, it is distinguished from the Korean species by the size of sporophytic thalli, branching pattern, basal constriction of laterals, and shape of sporangia. The other species forming the sister clade are also distinguished from the Korean entity by general morphological features of sporophytes and gametophytes. The specific distinction of the Korean alga from those species is supported by its genetic divergence, which is considered to be involved in the inter-specific range for the genus based on the previous data. Accordingly, Derbesia maxima sp. nov. is proposed for this siphonous green alga from Korea.

Geographic variation in fitness-related traits of the bladderwrack Fucus vesiculosus along the Baltic Sea-North Sea salinity gradient.

In the course of the ongoing global intensification and diversification of human pressures, the study of variation patterns of biological traits along environmental gradients can provide relevant information on the performance of species under shifting conditions. The pronounced salinity gradient, co‐occurrence of multiple stressors, and accelerated rates of change make the Baltic Sea and its transition to North Sea a suitable region for this type of study. Focusing on the bladderwrack Fucus vesiculosus, one of the main foundation species on hard‐bottoms of the Baltic Sea, we analyzed the phenotypic variation among populations occurring along 2,000 km of coasts subjected to salinities from 4 to >30 and a variety of other stressors. Morphological and biochemical traits, including palatability for grazers, were recorded at 20 stations along the Baltic Sea and four stations in the North Sea. We evaluated in a common modeling framework the relative contribution of multiple environmental drivers to the observed trait patterns. Salinity was the main and, in some cases, the only environmental driver of the geographic trait variation in F. vesiculosus. The decrease in salinity from North Sea to Baltic Sea stations was accompanied by a decline in thallus size, photosynthetic pigments, and energy storage compounds, and affected the interaction of the alga with herbivores and epibiota. For some traits, drivers that vary locally such as wave exposure, light availability or nutrient enrichment were also important. The strong genetic population structure in this macroalgae might play a role in the generation and maintenance of phenotypic patterns across geographic scales. In light of our results, the desalination process projected for the Baltic Sea could have detrimental impacts on F. vesiculosus in areas close to its tolerance limit, affecting ecosystem functions such as habitat formation, primary production, and food supply.

Updating Ceramium (Rhodophyta, Ceramiales) biodiversity in the North Adriatic Sea (Mediterranean): Ceramium rothianum sp. nov. and rediscovery of three forgotten species

ABSTRACTThe cosmopolitan genus Ceramium Roth (Ceramiaceae), with over 200 currently accepted species, is one of the largest in the Rhodophyta. The genus is characterized by cylindrical or slightly compressed thalli, with axial cells incompletely to completely covered by cortical cells, alternate to pseudo-dichotomous branching and straight to inrolled apices. Despite recent studies, the nomenclature and taxonomy of this genus are still very problematic, linked to the high degree of variation in morphological characters, small thallus sizes, epiphytic habit and the existence of cryptic species. The Mediterranean Sea is reported to have a particularly high diversity of Ceramium species: the North Adriatic Sea, in particular Venice and nearby coasts, was a favourite classical collecting area from where several species were described. In this study we characterize Ceramium sampled from transitional waters of the Venice Lagoon (Italy) by molecular and morphological approaches. Through phylogenetic analyses, using the plastid ribulose-1,5-bisphosphate carboxylase/oxygenase gene (rbcL) as molecular marker, we identify six distinct taxonomic entities, of which four represent taxa not currently recognized in the Venice Lagoon or the wider Mediterranean Sea. One of these is the poorly known species Ceramium connivens Zanardini, which is non-spiny and fully corticated when mature with short naked internodes developing in older axes. Two entities with erect partially corticated thalli without spines, probably misidentified up to now as Ceramium diaphanum (Lightfoot) Roth, correspond to the new species Ceramium rothianum Wolf et al. and to the forgotten species Ceramium nodosum (Kützing) A.W. Griffiths & Harvey. The last entity, characterized by prostrate axes giving rise to narrow erect incompletely corticated, non-spiny axes, is identified as the rare, poorly known species Ceramium incospicuum Zanardini.

Gracilaria phuquocensis sp. nov., a new flattened Gracilaria species (Gracilariales, Rhodophyta), previously recognized as G. mammillaris, from the southern coast of Vietnam

SUMMARYFlattened Gracilaria species are widely distributed along the coasts of the South China Sea with more than 20 species recorded. Within the South China Sea, Gracilaria mammillaris has only been reported from Vietnam, but this species is likely restricted to the western Atlantic. This study aimed to reevaluate the taxonomic status of Vietnamese specimens of ‘G. mammillaris’ using combined morphological and molecular data. Our data clearly indicated that Vietnamese specimens were morphologically and genetically distinct from authentic G. mammillaris from the western Atlantic, and also other described flat Gracilaria species. We, therefore, propose that specimens from Vietnam originally identified as G. mammillaris be designated as a new species, Gracilaria phuquocensis sp. nov. Morphologically, G. phuquocensis can be distinguished from other flat Gracilaria species by its small thallus size, narrower blades, many medullary layers, abundant basal nutritive filaments within mature cystocarps, and tetrasporangial nemathecium. Our rbcL sequence analyses showed that the new species was sister to Gracilaria rhodymenioides from Thailand, and these two species formed a clade with cylindrical Gracilaria species. Our study contributes to clarification of the taxonomic status of misidentified specimens attributed to the flattened Gracilaria species in the South China Sea.

Effects of Exogenous Cytokinins on Spore Germination and Gametophyte Morphogenesis of Dryopteris filix-mas (L.) Schott in vitro Culture

The impact of exogenous cytokinins (kinetin, zeatin, 6-benzylaminopurine, and N6-2-isopentenyladenine) on the pattern of Dryopteris filix-mas (L.) Schott spore germination, gametophyte growth and morphology in vitro have been studied. It was found that all studied cytokinins significantly retarded spore germination, inhibited gametophyte growth, caused deformations and decrease in the thallus size, and suppressed the development of reproductive structures and sporophyte growth at the concentration of 10–5 М. The reduction of the hormone concentration to 10–8 М stimulated the gametophyte development, induced cell divisions, particularly in the apical zone, due to which some of thalli were deformed, promoted the production of rhizoids, affected the formation of antheridia and archegonia, and slowed the sporophyte development.

Wiesnerella denudata (Mitt.) Steph. (Marchantiales; Marchantiophyta) a rare species in the Western Ghats of India

Wiesnerella denudata (Mitt.) Steph., a complex thalloid liverwort in the family Wiesnerellaceae (Marchantiales), is reported as a new species to Peninsular India from the Anamudi Shola National park in the Western Ghats of Kerala. This species is easily confused with another thalloid liverwort, Dumortiera hirsuta (Sw.) Nees, due to similar morphological characters, especially size of thallus, colour and habit. Wiesnerella denudata is characterised by the presence of elevated air pores which can be seen with the help of a hand lens or even with naked eye, subsessile rotund male receptacle, raised female receptacle with deeply dissected carpocephalum and absence of pseudoperianth. A detailed account of the species along with photographs and SEM images of spores is provided here.

Surveying seaweeds from the Ulvales and Fucales in the world’s most frequently used artificial waterway, the Kiel Canal

Abstract The Kiel Canal is one of the world’s most frequently used inland waterways and connects the SW Baltic Sea with the Wadden Sea. At the same time, the canal is a highly eutrophicated environment that is characterized by salinities that range from 3 to 16. This brackish character could make the Kiel Canal an important stepping stone for the introductions of species into the inner Baltic Sea. It could also hinder the identification of native and introduced species, given the fact that salinity sometimes severely affects algal morphology. Here we report on a survey of introduced and native seaweed species in the canal, focusing on the dominant groups, which are Fucales and Ulvales. Of the Fucales, the introduced species Fucus evanescens was detected nearly exclusively inside the canal, while Fucus vesiculosus dominated rockweed communities directly outside the sluice gates. Morphological analysis and genetic barcoding distinguished three species of Ulvales, Ulva linza, Ulva intestinalis and an unknown and possibly introduced species of the genus Blidingia. Species distributions and – in the case of U. intestinalis – branching patterns were clearly affected by salinity, while thallus sizes appeared to be affected by the specific eutrophication status of sites within the canal.

Under pressure: biomechanical limitations of developing pneumatocysts in the bull kelp (Nereocystis luetkeana, Phaeophyceae).

Maintaining buoyancy with gas-filled floats (pneumatocysts) is essential for some subtidal kelps to achieve an upright stature and compete for light . However, as these kelps grow up through the water column, pneumatocysts are exposed to substantial changes in hydrostatic pressure, which could cause complications as internal gases may expand or contract, potentially causing them to rupture, flood, and lose buoyancy. In this study, we investigate how pneumatocysts of Nereocystis luetkeana resist biomechanical stress and maintain buoyancy as they develop across a hydrostatic gradient. We measured internal pressure, material properties, and pneumatocyst geometry across a range of thallus sizes and collection depths to identify strategies used to resist pressure-induced mechanical failure. Contrary to expectations, all pneumatocysts had internal pressures less than atmospheric pressure, ensuring that thalli are always exposed to a positive pressure gradient and compressional loads, indicating that they are more likely to buckle than rupture at all depths. Small pneumatocysts collected from depths between 1 and 9m (inner radius=0.4-1.0cm) were demonstrated to have elevated wall stresses under high compressive loads and are at greatest risk of buckling. Although small kelps do not adjust pneumatocyst material properties or geometry to reduce wall stress as they grow, they are ~3.4 times stronger than they need to be to resist hydrostatic buckling. When tested, pneumatocysts buckled around 35m depth, which agrees with previous measures of lower limits due to light attenuation, suggesting that hydrostatic pressure may also define the lower limit of Nereocystis in the field.

Epibionts on Turbinaria ornata, a secondary foundational macroalga on coral reefs, provide diverse trophic support to fishes

Worldwide, many coral reef ecosystems have shifted from coral to algal dominance, yet the ecological function of these emergent communities remains relatively unknown. Turbinaria ornata, a macroalga with a rapidly expanding range in the South Pacific, forms dense stands on hard substrate, likely providing ecological services unique from corals. While generally unpalatable, T. ornata can function as a secondary foundation species and hosts an epibiont community that may provide overlooked trophic resources in phase shifted reef ecosystems. Results from video recorded field experiments designed to quantify consumer pressure on T. ornata epibionts showed that both consumer pressure and epibiont cover increased with thallus size. Additionally, most fish species, including herbivores, omnivores, and detritivores, exhibited higher bite rates on thalli with epibionts compared to thali with epibionts experimentally removed. Juvenile parrotfishes were responsible for 50% of total bites recorded and also had the highest bite rates. Results indicate that epibionts, particularly on large T. ornata, are a food resource for a diversity of fishes, representing a previously undescribed function of this macroalga in coral reef ecosystems. Exploring the functions of macroalgal dominated reef communities will be increasingly important as reefs continue to phase shift toward macroalgal dominance in the Anthropocene.

Photosynthetic rate and thallus size are not related in alpine yellow-green Rhizocarpon crustose lichens: Implications for lichenometry and growth

Lichenometry, first proposed at the beginning of the XXth century, is a technique that uses growth rates of saxicolous crustose lichens to date exposed surfaces over an age range of 500 years from present. Despite of the wide use of the methodology, it has been strongly criticized by several authors who consider that biological aspects involved in growth rates of lichens used are not sufficiently considered and that this will contribute to uncertainty in the final surface dating. The assumption, important for direct lichenometric measurements, that crustose lichens have constant growth rates through all their life span, is controversial, with some works pointing to a change in growth rate with thallus size in yellow-green Rhizocarpon sp. samples, the most widely lichens used for surface dating. This change in growth rate with size would contribute to inaccuracy in dating. In this work, we contribute to the discussion with a novel approach in which we measure several physiological parameters linked to carbon gain in 42 Rhizocarpon sp. samples of different sizes collected in the same locality. We found no significant relationship between thallus photosynthetic rate and thallus size indicating that possible growth rate variations over Rhizocarpon life span are not linked to carbon gain. The experiment performed provides a new data set to include in lichenometric modelling with the aim to obtain a better understanding of crustose lichens biology before attempting more feasible and accurate surface dating strategies. Measurements of length of activity periods and carbon allocation in Rhizocarpon are recommended to achieve future improvements in this direction.

Mitogenome analysis of a green tide forming Ulva from California, USA confirms its identity as Ulva expansa (Ulvaceae, Chlorophyta)

An unknown species of marine sea lettuce was observed forming green tides consecutive years from 2014 to 2016 in Seaside, California. This Ulva sp. was similar in thallus size and shape to U. expansa. To confirm this identification, whole genome sequencing was performed on the bloom-forming species of Ulva and the holotype specimen of U. expansa. The complete green tide Ulva mitogenome is 64,143 bp in length, contains 65 genes, and displays high gene synteny with U. pertusa Kjellman. The mitogenome was incomplete for the holotype of U. expansa, but the analysis yielded the mitoexome, plastid, and nuclear genetic markers. These data verify that the native U. expansa is responsible for the blooms in central California.

Influence of isotope fractionation on the nitrogen isotope composition of the brown macroalga Undaria pinnatifida

SUMMARYAlthough the stable nitrogen isotope ratio (δ15N value) of macroalgae is used to indicate sources of dissolved inorganic nitrogen in coastal marine environments, little is known about the effects of isotope fractionation on the δ15N value. We compared the δ15N values of Undaria pinnatifida cultivated at an inner bay and an exposed site (sites A and B, respectively) in Shizugawa Bay on the northeastern Pacific coast of Japan. Values for algal growth indicators, including thallus length, area, and weight, were higher at site B than at site A. The δ15N value of U. pinnatifida was significantly lower at site B (4.7 ± 1.0‰; mean ± standard deviation) than at site A (6.7 ± 0.4‰). This difference in the δ15N value of U. pinnatifida could not be explained by the δ15N value of seawater NO3 (source isotopic signature), which was similar at the two sites (7.5 ± 0.3‰). The pooled data from the two sites indicated that the δ15N value of U. pinnatifida decreased with increasing thallus size. The results suggest that the δ15N value of U. pinnatifida is influenced by isotope fractionation.

A newBunodophoronspecies (Sphaerophoraceae,Lecanorales) from the Neotropics

AbstractThis is the first part of an ongoing taxonomic treatment ofBunodophoron(Sphaerophoraceae,Lecanorales) in the Neotropics, based on the molecular phylogenetic analysis of three markers together with studies of morphology and chemistry, and using the general mixed Yule coalescence (GMYC) method to delimit species boundaries. In the Neotropics, species in this genus grow on the ground or on shrubs in the páramos, and as epiphytes in the montane rainforests. We describe here a new species from the páramos of Colombia,Bunodophoron crespoaeSoto, M. Prieto & Wedin sp. nov., and discuss its distinction from another large and common páramo speciesBunodophoron flabellatum(Hue) Soto, M. Prieto & Wedin comb. nov. Both species are primarily terrestrial in the páramos, althoughB. flabellatummay occasionally also grow as an epiphyte.Bunodophoron crespoaeis characterized by the white,c. 10–13 cm long, subterete to narrowly flattened, main branches. It differs from the otherwise similarB. flabellatumby being distinctly subterete, more abundantly branched, and by having smaller ascospores. Both are distinguished from the primarily epiphyticB. melanocarpumby the considerably larger thallus size, with the main branches ofB. melanocarpumrarely exceeding 3·5 cm in length and 2 mm in width.

Amplificación del gen rbc L revela primer registro de Porphyra mumfordii (Bangiales, Rhodophyta) en la bahía de Valparaíso, Chile central

The genus Porphyra , commonly found along Chilean coast, is an important commercial algal food known as nori. There are some specimens of Porphyra found at Montemar Marine Protected Area (32°57’S; 71°33’W) in Valparaiso Bay, which taxonomical identity is not clear. This study aimed to identify long-bladed specimens of Porphyra inhabiting the upper, mid and lower intertidal in Valparaiso Bay. To do so, 30 thalli of Porphyra were collected during the winter of 2015 in Montemar. All samples were subjected to rbc L molecular marker amplification. However, only 18 individuals were sequenced successfully. In addition, a morphological description was conducted. The rbc L analysis showed that the longbladed specimens inhabiting Valparaiso Bay correspond to Porphyra mumfordii . Differences in size and color of thalli found on each intertidal level were recorded. This research constitutes the first record of P. mumfordii for central Chile.

Depth influence on biochemical performance and thallus size of the red alga Gelidium corneum

AbstractRecent declines in the foundation species Gelidium corneum (Hudson) J.V. Lamouroux in pristine coastal stretches of the Southeastern Bay of Biscay have been documented in which individuals showed a high bleaching incidence. Among the potential factorsthat may be responsible for these changes in G. corneum abundance, it has been suggested that high levels of irradiance may be imposing stressful environmental conditions for shallow populations. Given that light is reduced exponentially as depth increases, in this investigation we examined the influence of depth on the stress of G. corneum by comparing the performance of several biochemical parameters and thallus size in plants collected at three different depths. The results revealed that plants growing in shallow waters showed lower antioxidant activity, lower concentrations of photosynthetic pigments (Chl‐a, total carotenoids, phycoerythrin), lower internal nitrogen levels (higher C:N ratio) and shorter thallus lengths than those living in deeper waters. Some of these results may be an expression of photoacclimation, but the low antioxidant activity and total carotenoid concentration detected in plants living in shallow waters suggest that the photoprotective mechanisms of G. corneum may have failed to offset photodamage. If so, our findings support previous studies that suggest that recent changes in solar radiation are partially responsible for G. corneum declines. Nevertheless, given that the metabolic performance of macroalgae is determined by the inter‐play of irradiance with other factors, including temperature and nutrient availability, further research is needed to reach a better understanding of the factors causing stress in G. corneum.

Hydration traits in cephalolichen members of the epiphytic old forest genusLobaria(s. lat.)

AbstractThis study aims to quantify the size-dependency of important hydration traits inLobaria amplissima, L. pulmonariaandL. virenssampled in sympatric populations on deciduous tree trunks in southern Norway, and to discuss possible implications of species-specific traits for the ecological niches of these old forest cephalolichens. Traits measured were thallus size (area and mass), specific thallus mass (STM), internal (WHCinternal) and external water-holding capacity (WHCexternal), and water content (WC) after shaking and after blotting.Lobaria amplissimahad the highest WHCinternal, 2·6 times higher thanL. pulmonariawith the lowest WHCinternal. WHCinternal, driven by STM, strongly depended on size. WHCexternalwas 28% (L. virens)to 47% (L. pulmonaria) of the WHCinternal. Unlike WHCinternal, WHCexternaldid not depend on thallus area, meaning that WHCexternalis proportionally higher for smaller compared with larger thalli. The most widespread species,L. pulmonaria, benefits from a flexible hydration strategy due to low STM, facilitating the use of more diverse water sources than the other two species that depend more on rain, particularlyL. amplissimawith the highest STM and thus relatively high WHCinternal. ForL. virens, a combination of less specialized hydration traits and a low tolerance to higher light intensity probably jeopardizes its survival outside rainforest habitats.

Thallus variation of Sargassum polycystum from Central Java, Indonesia

Widyartini DS, Widodo P, Susanto AB. 2017. Thallus variation of Sargassum polycystum from Central Java, Indonesia. Biodiversitas 18: 1004-1011. Sargassum polycystum C. Agardh has characteristic of highly polymorphic thalli among individuals in its population. The shape and size of thalli are variable among the individuals. The study was aimed at knowing the variation of S. polycystum thalli grown in different beaches of Central Java. The data showed that the thalli of the S. polycystum grown in the Karimunjawa had more variation than those grown in the Menganti Beach, as obviously seen on its leaves and vesicles. There was a relationship between salinity and pH of the Menganti Beach of the length of vesicle S. polycystum, but nitrate and phosphate content weren't correlated of variation thalli. Temperature, salinity, and pH of Karimunjawa Beach were correlated with the vesicle's color, nitrate and phosphate correlated of the width as well as upper and lower surfaces of the leaves, phosphate also correlated with the leaf edge and vesicle diameter. Environmental factors in the Menganti Beach had not significantly affected the thalli variation, but salinity and pH had an effect on vesicle length. Temperature, salinity, and pH in the Karimunjawa Beach significantly affected the vesicle color, nitrate and phosphate affect the width of leaves as well as their upper and lower surface. Phosphate also affected the edge of the leaf and the vesicle diameter

Functional traits prolonging photosynthetically active periods in epiphytic cephalolichens during desiccation

By fluorescence imaging, we quantified how hydration traits and thallus size determine the duration of photosynthetic activity during drying in light and darkness for sympatric populations of three epiphytic old forest cephalolichens differing in specific thallus mass (STM) and growth form. Maximal PS II efficiency (FV/FM) during drying in darkness and effective PSII yield (ФPSII) during drying in light (200μmolm−2s−1) was repeatedly monitored in lichens under controlled conditions, using chlorophyll fluorescence imaging with simultaneous recordings of wet mass. STM shaping the water holding capacity (WHC), water content in percent at start (WC), and water loss rate per thallus area (WLR) in combination determined the duration of active periods during desiccation between (r2adj=0.86) and within species (r2adj=0.60–0.92). Lobaria pulmonaria with the lowest STM and the most upright growth form experienced the shortest active periods, Lobaria amplissima with the highest STM and the most prostrate growth form had the longest periods. Across the more compact species (L. amplissima, L. virens), WLR strongly declined with increasing thallus size, but weakly in the loosely attached L. pulmonaria. ФPSII, a proxy of photosynthesis, exhibited suprasaturation depression at maximal hydration. Fluorescence imaging allowed a rapid, non-invasive evaluation of the contribution of various functional traits to active periods in entire thalli during desiccation. A thick, prostrate growth form prolongs active periods by maximizing water storage, whereas a thin, loosely attached growth form uses a more flexible hydration strategy with shorter active periods.

Екологічні особливості поширення лишайників антропогенно трансформованих територій Криворіжжя

<p>We observed the diversity of lichens at different levels of aerotechnogenic pollution of industrial mining – metallurgical complex of Krivyi Rig Basin. We registered the reduction of vegetation cover and reducing of the size of thallus in the highway area with undesirable living conditions to lichens<strong>.</strong><strong> </strong>We suggested<strong> </strong>that specific composition and distribution patterns of cladinas is subject of the industrial loading. Most of lichen species are tolerant to urbanization and widely distributed in the built-up area of industrial mining, whereas only a few lichens sensitive to urbanization were recorded. We registered the domination of crustaceous cladinas with insignificant participation of fissile cladinas and complete absence of bushy forms that could be possible consequence of industrial influence.</p>

Phenology of Chondracanthus tenellus (Rhodophyta) in the central Pacific coast of Honshu, Japan

This study reports variations in thallus size, density, and reproductive phenology in a natural population of underutilized seaweed, Chondracanthus tenellus, at Tateyama in the Pacific coast of Japan from August 2014 to October 2016. Growth was monitored by measuring the length and weight of thalli. In addition, a number of thalli fouled by spirobids, coralline, and/or fleshy epiphyte and/or grazing marks were also recorded from February 2015 to October 2016. Density was monitored by counting the number of clumps along a subtidal fixed line of 30 m long (3–4 m in depth). Reproductive efforts were evaluated by counting the number of cystocarps per thallus and carpospores per cystocarp. Average thallus length sharply dropped in October and gradually increased from November to record the maximum length (73.5 ± 2.7 mm) in April 2016. After the season, thalli were damaged by epiphytic algae, spirobids, and bite marks with increase in temperature. It was suggested that high temperature and low nutrition in summer probably induce weakening of thallus and promote fouling by spirobids and epiphyte which accelerate depletion of C. tenellus together with herbivorous animals. Although the population showed a large gametophytic dominance, carposporic gametophytes were found in August to September and May to July with the highest value (20.6%) in September. Maturation of gametophytes probably was induced by long day length or a transition from short to long day. Number of carpospores per cystocarp was 72 ± 15 × 103 in January. Considering the size and cleanliness of thallus, it is preferable to harvest in April.