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opencc-zeroDec 2021View details →

Evolutionary responses of energy metabolism, development, and reproduction to artificial selection for increasing heat tolerance in Drosophila subobscura

Adaptation to warming conditions involves increased heat tolerance and metabolic changes to reduce maintenance costs and maximize biological functions close to fitness. Evidence shows that energy metabolism evolves in response to warming conditions, but we know little about how heat stress intensity determines the evolutionary responses of metabolism and life history traits. Here, we evaluated the evolutionary responses of energy metabolism and life-history traits to artificial selection for increasing heat tolerance in Drosophila subobscura, using two protocols to measure and select heat tolerance: slow and fast ramping protocols. We found that the increase in heat tolerance was associated with reduced activity of the enzymes involved in the glucose-6-phosphate branchpoint, but no changes in the metabolic rate in selected lines. We also found that the evolution of increased heat tolerance increased the early fecundity in selected lines and increased the egg-to-adult viability only in the slow-ramping selected lines. This work shows heat tolerance can evolve under different thermal scenarios but with different evolutionary outcomes on associated traits depending on the heat stress intensity. Therefore, spatial and temporal variability of thermal stress intensity should be taken into account to understand and predict the adaptive response to ongoing and future climatic conditions.

opencc-zeroNov 2022View details →

High thermal tolerance in high elevation species and laboratory-reared colonies of tropical bumble bees

Bumble bees are key pollinators with some species reared in captivity at a commercial scale, but with significant evidence of population declines and with alarming predictions of substantial impacts under climate change scenarios. While studies on the thermal biology of temperate bumble bees are still limited, they are entirely absent from the tropics where the effects of climate change are expected to be greater. Herein we test if bees' thermal tolerance decreases with elevation and if the stable optimal conditions used in laboratory-reared colonies reduces their thermal tolerance. We assessed changes in the lower (CTMin) and upper (CTMax) critical thermal limits of four species at two elevations (2600 and 3600 m) in the Colombian Andes, examined the effect of body size, and evaluated the thermal tolerance of wild caught and laboratory-reared individuals of B. pauloensis. We also compiled information on bumble bees' thermal limits and assessed potential predictors for broad-scale patterns. We found that CTMin decreased with increasing elevation while CTMax was similar between elevations. CTMax was slightly higher (0.84 °C) in laboratory-reared than in wild-caught bees while CTMin was similar, and CTMin decreased with increasing body size while CTMax did not. Latitude is a good predictor for CTMin only while annual mean temperature, maximum and minimum temperatures of the warmest and coldest months are good predictors for both CTMin and CTMax. The stronger response in CTMin with increasing elevation, and similar CTMax, supports Brett's heat-invariant hypothesis, which has been documented in other taxa. Andean bumble bees appear to be about as heat tolerant as those from temperate areas, suggesting that other aspects besides temperature (e.g., water balance) might be more determinant environmental factors for these species. Laboratory-reared colonies are adequate surrogates for addressing questions on thermal tolerance and global warming impacts.

opencc-zeroDec 2022View details →

Testing the chilling: Before drought-tolerance hypothesis in Pooideae grasses

Temperate Pooideae are a large clade of economically important grasses distributed in some of the Earth's coldest and driest terrestrial environments. Previous studies have inferred that Pooideae diversified from their tropical ancestors in a cold montane habitat, suggesting that above-freezing cold (chilling) tolerance evolved early in the subfamily. By contrast, drought tolerance is hypothesized to have evolved multiple times independently in response to global aridification that occurred after the split of Pooideae tribes. To independently test predictions of the chilling before-drought hypothesis in Pooideae, we assessed the conservation of whole plant and gene expression traits in response to chilling versus drought. We demonstrated that both trait responses are more similar across tribes in cold as compared to drought, suggesting that chilling responses evolved before, and drought responses after, tribe diversification. Moreover, we found significantly more overlap between drought and chilling-responsive genes within a species than between drought-responsive genes across species, providing evidence that chilling tolerance genes acted as precursors for the novel acquisition of increased drought tolerance multiple times independently, partially through the cooption of chilling responsive genes.

opencc-zeroDec 2022View details →

Data for: Effects of parental age on salt stress tolerance in an aquatic plant

Parental age influences components of offspring fitness in many species. The ability to tolerate stress also affects fitness, but less is known regarding changes in offspring stress tolerance with increasing parental age, especially in plants. We examined first and fifth-born clonal offspring (using birth order as a proxy for parental age), and compared their fitness in several sub-lethal concentrations of salt (NaCl), to investigate the interactive effects of birth order and salt stress on the offspring of the aquatic plant Lemna minor L. We found that increasing salt concentration reduced reproduction particularly at early ages, which detrimentally affected fitness, as measured by the intrinsic rate of natural increase. Fifth offspring had greater fitness than first offspring, potentially due to the hump-shaped relationship between offspring fitness and birth order observed in other studies on Lemna, with fifth offspring near the peak of the hump. We found no interactive effect of birth order and salt concentration on offspring fitness; however, there were interactive effects on the time to first reproduction and the size of fronds. Specifically, first offspring exposed to increasing salt concentrations exhibited longer delays to first reproduction and grew to a greater size, while fifth offspring showed little change in either variable with increasing salt concentration. Thus, variation in birth order affected offspring response to salt stress, although not in terms of fitness. These results help illuminate factors impacting the age-specific strength of natural selection and stress responses, and may be environmentally relevant in the context of environmental salinization.

opencc-zeroJan 2023View details →

Barn Swallows (Hirundo rustica) in East Asia show strong tolerance against nest mites

Hosts defend against parasites through two main strategies: resistance and tolerance. These two strategies have different influences on the parasite load of the host and so have variable effects on the fitness of parasites. In this study, we investigated the strategy used by the Barn Swallow (Hirundo rustica) to defend against a haematophagous mite (Dermanyssus hirundinis) in East Asia. For the first time, we found Barn Swallows were parasitized by D. hirundinis in Asia and, in north-eastern China, over 60% of broods were parasitized. We examined whether swallows with better body condition or the males with more extravagant sexually selected traits (longer tail streamers or redder ventral plumage) showed stronger resistance or tolerance to these parasites. The results showed that males with more extravagant sexual ornaments had weaker resistance (heavier brood mite load) than males with less extravagant traits; however, the heavier mite load was likely to be a by-product of larger brood size. Moreover, the Barn Swallow population showed strong tolerance against the parasite, i.e., the brood mite load did not affect brood size or nestling body mass. It is worth noting that male swallows with redder breast plumage showed weaker tolerance (lower seasonal reproductive success) than other males. Further, we conducted experimental ectoparasite manipulations in 46 Barn Swallow nests to verify the potential effect of these parasites on the body condition of nestlings. Neither the body mass nor the hemoglobin concentration differed between the designed groups; however, the hemoglobin concentration of nestlings decreased with mite load. We concluded that the Barn Swallow population in East Asia used the tolerance strategy, rather than resistance to defend against nest mites, and the ectoparasitic mite did not appear to be a strong selective factor for reproductive success owing to the strong tolerance of the studied Barn Swallow population.

opencc-zeroJan 2023View details →

Data for: Tolerance of high temperature and associated effects on reproduction in euedaphic Collembola

Tropical and mid-latitude terrestrial ectotherms are more vulnerable to global warming than species from high latitudes. However, thermal tolerance studies from these regions still lack soil invertebrates. We chose six euedaphic species of Collembola (of the genera Onychiurus and Protaphorura) sampled across latitudes ranging from 31°N to 64°N and first determined their upper thermal limit (UTL) by static assays. Based on their upper thermal limit, we exposed them to temperatures 32 ℃ to 36 ℃ depending on the species for 1-2 hours, which was expected to result in heat-induced injuries equivalent to 5% to 30% mortality within each species. Survivors from this series of increasing heat injuries were used to determine the time to first oviposition, and the number of eggs produced following heat exposure. We found the upper thermal limit positively correlates to the soil temperature of the sampling site. Species have different UTL60 indicating they have different tolerance to high temperatures. The species order from most heat tolerance to least was O. yodai > P. fimata > P. armata ≈ P. tricampata > P. macfadyeni > P. pseudovanderdrifti. Heat stress inflicted on survivors can prolong the time to regain reproduction in all species, and two species showed reduced egg production. The most heat-tolerant species did not have advantages over the least heat-tolerant species under a sublethal heat exposure scheme (where predicted survival was less than 30%) in terms of their recovery of reproduction. We provided evidence for a potential long-term effect of high temperature on euedaphic species of Collembola and highlighted the need for further studies on the effects of global warming on soil living organisms.

opencc-zeroJan 2023View details →

Postrelease exploration and stress tolerance of landlocked and anadromous Atlantic salmon and their hybrids

Background We studied postrelease explorative behavior and stress tolerance of Landlocked and anadromous Atlantic salmon and their hybrids. For the research, we hybridized the Landlocked salmon of Lake Saimaa with a Baltic anadromous salmon from River Kymijoki, Southern Finland (strain originally from River Neva, Russia). These fish were hybridized in November 2017 and October 2018 in the Kainuu Fisheries Research Station, Paltamo, Kainuu, Finland (kfrs.fi). In the data the fish are treated as four strains (column 'Strain'): 1. Purebred Landlocked salmon (LLxLL), 2. hybrids, where the maternal population was landlocked salmon (LLxBA), 3. hybrids, where the maternal population was Baltic anadromous salmon (BAxLL) and 4. purebred Baltic anadromous salmon (BAxBA). Experiment 1. Post-release exploration In experiment 1., consisting of two separate trials, we studied post-release exploration of the fish in four circular seminatural streams. (Datasets 'Exploration2018.csv' & 'Exploration2020.csv', see also 'Figure_1.jpg') Experiment 2. Stress tolerance In experiment 2. we studied the stress response and recovery of the fish (Dataset 'Stress_BernoulliData.csv', see also 'Figure_2.jpg').

opencc-zeroFeb 2023View details →

SO2 and copper tolerance exhibit an evolutionary trade-off in Saccharomyces cerevisiae

Copper tolerance and sulfite tolerance are two well-studied phenotypic traits of Saccharomyces cerevisiae. The genetic bases of these traits are derived from allelic expansion at the CUP1 locus and reciprocal translocation at the SSU1 locus, respectively. Previous work identified a negative association between sulfite and copper tolerance in S. cerevisiae wine yeasts. Here we probe the relationship between sulfite and copper tolerance and show that an increase in CUP1 copy number does not impart copper tolerance in all S. cerevisiae wine yeast. Bulk-segregant QTL analysis was used to identify variance at SSU1 as a causative factor in copper sensitivity, which was verified by reciprocal hemizygosity analysis in a strain carrying 20 copies of CUP1. Transcriptional and proteomic analysis demonstrated that SSU1 over-expression did not suppress CUP1 transcription or constrain protein production but suggested that SSU1 overexpression induced sulfur limitation during exposure to copper. Finally, an SSU1 over-expressing strain exhibited increased sensitivity to moderately elevated copper concentrations in sulfur-limited medium, demonstrating that SSU1 over-expression burdens the sulfate assimilation pathway. Over-expression of MET 3/14/16, genes upstream of H2S production in the sulfate assimilation pathway increased the production of SO2 and H2S but did not improve copper sensitivity in an SSU1 overexpressing background. We conclude that copper and sulfite tolerance are conditional traits in S. cerevisiae and provide evidence of the metabolic basis for their mutual exclusivity. These findings suggest an evolutionary basis for the extreme amplification of CUP1 observed in some yeasts.

opencc-zeroMar 2023View details →

Data from: Enhanced climate tolerance for trees derived from microbial communities

Changing climates are pushing species outside of their evolved tolerances; populations must acclimate or adapt to the new conditions or migrate to avoid extinction. However, because plants associate with diverse microbial communities that shape their phenotype, shifts in microbial associations may provide an alternative source of novel climate tolerance. Here we show that tree seedlings inoculated with microbial communities sourced from drier, warmer, or colder sites displayed higher survival when facing drought, heat, or cold stress, respectively. Microbially mediated drought tolerance was associated with increased diversity of arbuscular mycorrhizal fungi, while cold tolerance was related to reduced diversity of non-adapted taxa. Understanding microbially mediated climate tolerance may enhance our ability to predict and manage the adaptability of forest ecosystems to changing climates.

opencc-by-4.0May 2023View details →

GWAS analysis for tolerance to heat stress and milk production, in subtropical Egyptian goats

The conservation of local Egyptian goat genotypes has been a national program since 2009. Our study investigated different subtropical Egyptian goat populations (367 does) from different harsh ecological zones(hot Upper Egypt, Coastal Zone of Western Desert, and Wahati Desert Oasis) to identify genes associated with heat stress. We examined the physiological response of animals that were exposed to simulated summer grazing conditions, and physiological parameters including respiration rate, gas volume, rectal temperature, and skin temperature. Temperature Humidity Index ranged from 98.6 to 109.3, indicating that the animals were under severe heat stress. Results showed significant differences between the populations in their tolerance to heat stress, with Saidi goats being the most adapted to hot-dry conditions. Respiration rate was found to be the most reliable physiological trait for differentiating between animals in their tolerance to heat stress. The GWAS analysis involved 157 genotypes and 54,032 marker-SNPs, revealing 90 SNPs associated with heat stress and 70 SNPs associated with milk production. Chromosome 1 had the highest number of SNPs associated with heat-stress traits, while chromosome 4 exhibited the largest number of significant SNPs associated with milk production. Several genes were associated with heat stress response and tolerance in goats. These genes were categorized into three main groups: heat stress response, stress response, and reproduction, and feed intake. The first group includes USP54, KDM6A, and ETNPPL, which have multiple functions, such as steroid biosynthesis, metabolism, and stress response, and are also involved in key biological processes such as reproduction, immune response, and metabolism. The second group, which is mostly associated with immune response, includes GLTSCR2 and NAALADL2. Finally, a group of genes that may control animal feed intakes, including TRPM3 and ZBTB8A. Additionally, several genes, such as FHIT, GALNT18, RAPGEF5, and RBFOX1, linked to milk production, are known to be linked fertility, and fatty acid composition. These findings provide insights into the potential roles of these genes in heat stress response and tolerance in goats, and the genetic basis for improved milk production and animal welfare and can be used as selection markers in the ongoing breeding programs.

opencc-by-4.0Mar 2023View details →

Upper thermal tolerance of grassland vipers (Vipera spp.): environmental drivers and local adaptation

The thermal tolerance of ectotherms is a critical factor that influences their distribution, physiology, behaviour, and ultimately survival. Understanding the factors that shape thermal tolerance in these organisms is therefore of great importance for predicting their responses to forecasted climate warming. Here, we investigated the voluntary thermal maximum (VTmax) of nine grassland viper taxa and explored the factors that influence this trait. The small size of these vipers and the open landscape they inhabit renders them particularly vulnerable to overheating and dehydration. We found that the VTmax of grassland vipers is influenced by environmental temperature, precipitation, shortwave flux, and individual body size, rather than by phylogenetic relatedness. Vipers living in colder environments exhibited a higher upper thermal tolerance, contradicting the hypothesis that environmental temperature is positively related to VTmax. Our findings emphasise the importance of considering local to regional adaptation and environmental conditions when studying thermal physiology and the evolution of thermal tolerance in ectotherms.

opencc-zeroMay 2023View details →

Interspecific differences in thermal tolerance landscape explain aphid community abundance under climate change

A single critical thermal limit is often used to explain and infer the impact of climate change on geographic range and population abundance. However, it has limited application in describing the temporal dynamic and cumulative impacts of extreme temperatures. Here, we used a thermal tolerance landscape approach to address the impacts of extreme thermal events on the survival of co-existing aphid species (Metopolophium dirhodum, Sitobion avenae and Rhopalosiphum padi). Specifically, we built the thermal death time (TDT) models based on detailed survival datasets of three aphid species with three ages across a broad range of stressful high (34–40 ÅãC) and low (−3∼-11 ÅãC) temperatures to compare the interspecific and developmental stage variations in thermal tolerance. Using these TDT parameters, we performed a thermal risk assessment by calculating the potential daily thermal injury accumulation associated with the regional temperature variations in three wheat-growing sites along a latitude gradient. Results showed that M. dirhodum was the most vulnerable to heat but more tolerant to low temperatures than R. padi and S. avenae. R. padi survived better at high temperatures than Sitobion avenae and M. dirhodum but was sensitive to cold. R. padi was estimated to accumulate higher cold injury than the other two species during winter, while M. dirhodum accrued more heat injury during summer. The warmer site had higher risks of heat injury and the cooler site had higher risks of cold injury along a latitude gradient. These results support recent field observations that the proportion of R. padi increases with the increased frequency of heat waves. We also found that young nymphs generally had a lower thermal tolerance than old nymphs or adults. Our results provide a useful dataset and method for modelling and predicting the consequence of climate change on the population dynamics and community structure of small insects.

opencc-by-4.0Mar 2023View details →

Naturally segregating variants contributing to thermal tolerance in a D. melanogaster model system.

Main_Incapacitation.zip and Incapacitation_founders.zip contain raw thermal tolerance scores for individuals measured within the heat box. Each folder is labeled with the RIL or founder ID and replicates within each file are labeled with group numbers.   RNAi_files_to_tar.txt contains the metadata for the Combined_tracks_RNAi_1.Rds.zip and  Combined_tracks_RNAi_2.Rds.zip. Combined_tracks_RNAi_1.Rds.zip and Combined_tracks_RNAi_2.Rds.zip. contains raw data for RNAi lines measured on the heat plate.  plate_finder-kinglab-2021-05-02.zip contains the DeepLabCut model used for finding the corners of aluminum mounting plate used to hold the fly vials for thermal sensitivity testing. This directory contains the training data as well as the trained and evaluated model. No retraining should be necessary for use. fly_tracker_2-king-2021-09-27.zip contains the DeepLabCut model used for tracking individual flies during thermal sensitivity testing. This directory contains the training data as well as the trained and evaluated model. No retraining should be necessary for use. fly_tracker_batch.py is a python (>= 3.0) script that processes the raw movie files collected via the Raspberry Pi. This script uses the plate finder DeepLabCut model to find the corners of the plate, rotate and crop the images, and output movie files for individual flies. It then uses the fly tracker DeepLabCut model to track the flies and output the data for subsequent processing in R.

opencc-zeroJun 2023View details →

Trans-acting genotypes drive mRNA expression affecting metabolic and thermal tolerance traits

Evolutionary processes driving physiological trait variation depend on the underlying genomic mechanisms. Evolution of these mechanisms depends on whether traits are genetically complex (involving many genes) and how gene expression that impacts the traits is converted to phenotype. Yet, genomic mechanisms that impact physiological traits are diverse and context-dependent (e.g., vary by environment or among tissues), making them difficult to discern. Here we examine the relationships between genotype, mRNA expression, and physiological traits to discern the genetic complexity and whether the gene expression affecting the physiological traits is primarily cis or trans-acting. We use low-coverage whole genome sequencing and tissue-specific mRNA expression among individuals to identify polymorphisms directly associated with physiological traits and expressed quantitative trait loci (eQTL) driving variation in six temperature-specific physiological traits (standard metabolic rate, thermal tolerance, and four substrate-specific cardiac metabolic rates). Not surprisingly, there were few, only five, SNPs directly associated with physiological traits. Yet, by focusing on a select set of mRNAs belonging to co-expression modules that explain up to 82% of temperature specific (12°C or 28°C) metabolism and thermal tolerance, we identified hundreds of significant eQTL for mRNA whose expression affects physiological traits. Surprisingly, most eQTL (97.4% for heart and 96.7% for brain) of eQTL were trans-acting. This could be due to higher effect size or greater importance of trans versus cis-acting eQTLs for mRNAs that are central to co-expression modules. That is, we may have enhanced the identification of trans-acting factors by looking for SNPs associated with mRNAs in co-expression modules that are known to be correlated with the expression of 10s or 100s of other genes, and thus have identified eQTLs with widespread effects on broad gene expression patterns. Overall, these data indicate that the genomic mechanism driving physiological variation across environments is driven by trans-acting tissue-specific mRNA expression.

opencc-zeroAug 2023View details →

High tolerance to zinc but no evidence for local adaptation in the aquatic plant Lemna minor

Duckweeds are a widely distributed and economically important aquatic plant family that have high potential for phytoremediation of polluted water bodies. We collected four ecotypes of the common duckweed (Lemna minor) from the four corners of Switzerland and assessed how their home vs. away environments influenced their growth. Additionally, we investigated their response to a metal pollutant (Zn) in both their home and away environments. Zn is found in freshwater systems and can become harmful to plants at elevated concentrations. We hypothesized that growing in their home environment would help the plants buffer the negative effect of the metal pollutant. To test this, we measured Lemna growth in a common garden experiment in a glasshouse where the four ecotypes were grown in each of the water environments, as well as in three different concentrations of Zn. To investigate whether interactions between Lemna and their microbial community can enhance or reduce tolerance to heavy metal pollution, we sampled chlorophyll-a as a proxy for algal biomass. Finally, we measured total nitrogen and total organic carbon to describe the abiotic environment in more detail. The four Lemna ecotypes exhibited significantly different growth rates across the water treatments. This difference in fitness was matched with DNA sequencing revealing genetic differentiation between the four ecotypes. However, the effect of the water and zinc treatment on Lemna growth was the same for all ecotypes. We did not find evidence for local adaptation; instead, we observed strong plastic responses. Lemna growth rates were higher under higher Zn concentrations. This positive effect of Zn on Lemna growth could be in part due to reduced competition with algae. We conclude that L. minor ecotypes may exhibit large differences in growth rate, but that the species overall have a high Zn tolerance and strong plastic adaptive potential in novel environments.

opencc-zeroSep 2023View details →

Data from: The biogeographic history of neosuchian crocodiles and the impact of saltwater tolerance variability

Extant neosuchian crocodiles are represented by only 24 taxa that are confined to the tropics and subtropics. However, at other intervals during their 200 million-year evolutionary history, the clade reached considerably higher levels of species-richness, matched by more widespread distributions. Neosuchians have occupied numerous habitats and niches, ranging from dwarf riverine forms to large marine predators. Despite numerous previous studies, several unsolved questions remain with respect to their biogeographic history, including the geographic origins of major groups, e.g., Eusuchia and Neosuchia itself. We carried out the most comprehensive biogeographic analysis of Neosuchia to date, based on a multivariate K-means clustering approach followed by the application of two ancestral area estimation methods (BioGeoBEARS and Bayesian Ancestral Location Estimation) applied to two recently published phylogenies. Our results placed the origin of Neosuchia in north-western Pangaea, with subsequent radiations into Gondwana. Eusuchia probably emerged in the European archipelago during the Late Jurassic/Early Cretaceous, followed by dispersal to the North American and Asian landmasses. We show that putative transoceanic dispersal events are statistically significantly less likely to happen in alligatoroids. This finding is consistent with the saltwater intolerant physiology of extant alligatoroids, bolstering inferences of such intolerance in their ancestral lineages.

Fig. 1a–d in Description of Epistylis camprubii n. sp., a Species Highly Tolerant to Ammonium and Nitrite

Fig. 1a–d. Morphological and oral infraciliature details of Epistylis camprubii. a – detail of the zooid. PD – peristomial disk; PL – peristomial lip; CV – contractile vacuole; Ma – macronucleus; Mi – micronucleus; b – scheme of a colony; c – oral infraciliature. Pk – polykinety; H – haplokinety; G – germinal kinety; P1 – polykinety 1; P2 – polykinety 2; P3 – polykinety 3; d – morphological characteristics measured. PDd – peristomial disk diameter; PLw – peristomial lip width; PLh – peristomial lip height; Zl – zooid lenght; Zw – zooid width; Sw – stalk width. Scale bars: 25 µm.

Fig. 5. The preferred 18s in Description of Epistylis camprubii n. sp., a Species Highly Tolerant to Ammonium and Nitrite

Fig. 5. The preferred 18s rRNA tree under maximum likelihood (ML). Rectangles on branches denote the support recovered in analyses under alternative inference methods. Left rectangle refers to maximum likelihood (ML), the middle one to Bayesian inference (BI) and the right one to maximum parsimony (MP). Black coloured rectangle indicates bootstrap support> 80 or posterior probability> 0.95, grey rectangle indicates clade recovered but with lower support than the former values, and white rectangle indicates the clade was not recovered. Main Epistylis clade boxed.

Fig. 4 in Description of Epistylis camprubii n. sp., a Species Highly Tolerant to Ammonium and Nitrite

Fig. 4. Frequency of the number of zooids per colony observed in Epistylis camprubii colonies (number of analyzed colonies = 71).

Fig. 3a–e in Description of Epistylis camprubii n. sp., a Species Highly Tolerant to Ammonium and Nitrite

Fig. 3a–e. Images of Epistylis camprubii, after silver staining method. a – view of the longitudinal fibers and the oral infraciliature; b – detail of the aboral trochal band of a feeding zooid; c – aboral trochal band of a zooid during swimmer formation; d–e – oral infraciliature details. H – haplokinety; G – germinal kinety; Pk – polykinety; P1 – polykinety 1; P2 – polykinety 2; P3 – polykinety 3. Scale bars: 15 µm.