Leaf size in mosses is structurally constrained by cell dimensions and genome size

Leaf size in mosses is structurally constrained by cell dimensions and genome size | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 3 July 2025 V1 Latest version Share on Leaf size in mosses is structurally constrained by cell dimensions and genome size Authors : Pere Miquel Mir-Rosselló 0000-0002-0004-1916 [email protected] , Jaume Flexas 0000-0002-3069-175X , and Marc Carriquí Alcover 0000-0002-0153-2602 Authors Info & Affiliations https://doi.org/10.22541/au.175152808.82476401/v1 Published Plant Biology Version of record Peer review timeline 461 views 317 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Leaf anatomy is a key factor determining plant ecology. Cell size and number are related to leaf size in tracheophytes, but this has been poorly studied in bryophytes, which never reach large leaf sizes. In this study, we aimed to study the main anatomical factors determining leaf size in mosses, and how this is related to their ecology. We measured cell and leaf dimensions in 287 moss species, as well as cell density, cell wall thickness and midrib length. These measurements were contrasted against different traits, highlighting growth form and genome size, and correlations among them were also performed. Leaf size positively correlated with cell size in mosses, while it negatively correlated with cell density. The longest moss leaves were always supported by midribs reaching or surpassing the leaf apex. Genome size positively correlated with cell and especially leaf size. All these relationships were stronger for acrocarpous mosses. Leaf size in mosses is limited by the mechanical support provided by cell turgor and midribs. Both the mechanical support and the effect of genome size were more important in acrocarpous mosses. Our findings present anatomy as a key linking factor between genome size and plant ecology. Leaf size in mosses is structurally constrained by cell dimensions and genome size Pere Miquel Mir-Rosselló 1,2,3 , Jaume Flexas 1 , Marc Carriquí 1 1 Research Group on Plant Biology under Mediterranean Conditions, Universitat de les Illes Balears (UIB)‐ Institut d’Investigacions Agroambientals i d’Economia de l’Aigua (INAGEA), Palma, Spain 2 Interdisciplinary Ecology Group, Department of Biology, Universitat de les Illes Balears, Palma, Spain 3 Botany on Mediterranean Islands Research Group, Department of Biology, Universitat de les Illes Balears, Palma, Spain Corresponding authors: Pere Miquel Mir-Rosselló ( [email protected] ); Marc Carriquí ( [email protected] ) Abstract Leaf anatomy is a key factor determining plant ecology. Cell size and number are related to leaf size in tracheophytes, but this has been poorly studied in bryophytes, which never reach large leaf sizes. In this study, we aimed to study the main anatomical factors determining leaf size in mosses, and how this is related to their ecology. We measured cell and leaf dimensions in 287 moss species, as well as cell density, cell wall thickness and midrib length. These measurements were contrasted against different traits, highlighting growth form and genome size, and correlations among them were also performed. Leaf size positively correlated with cell size in mosses, while it negatively correlated with cell density. The longest moss leaves were always supported by midribs reaching or surpassing the leaf apex. Genome size positively correlated with cell and especially leaf size. All these relationships were stronger for acrocarpous mosses. Leaf size in mosses is limited by the mechanical support provided by cell turgor and midribs. Both the mechanical support and the effect of genome size were more important in acrocarpous mosses. Our findings present anatomy as a key linking factor between genome size and plant ecology. Keywords Anatomy, bryophytes, cell size, ecology, genome size, leaf size, mechanical support, mosses Introduction Leaf anatomy is a key factor that highly determines plant biology (Bolhàr-Nordenkampf & Draxler 1993; Wright et al. 2017; Oguchi et al. 2018). Cell and leaf size, cell wall thickness and leaf structure are among the key anatomical factors determining the performance and ecology of land plants (Cutler et al. 1977; Šímová & Herben 2012; Tomás et al. 2013; Carriquí et al. 2019; Flexas et al. 2021; Xue et al. 2023). Cell size determines cell arrangement within tissues and the response and tolerance to environmental stressors, consequently influencing leaf productivity (Cutler et al. 1977; Drake et al. 2013; Brodribb et al. 2013; Théroux-Rancourt et al. 2021; Borsuk et al. 2022). In turn, leaf size and morphology are highly related to plant growth capacity, water use efficiency, stress tolerance and distribution (Knight & Ackerly 2003; Wright et al. 2017; Conesa et al. 2019). Because of this, understanding leaf dimensions and their main constraints is essential to understand plant biology. Previous studies demonstrate that cell and leaf dimensions are related in tracheophytes (Brodribb et al. 2013; John et al. 2013; Ma et al. 2025). In these plants, large leaves are typically built by increasing the number of cells at the expense of reducing their size (Gonzalez et al. 2010; Ma et al. 2025). However, while smaller cells in larger numbers increase leaf productivity (Théroux-Rancourt et al. 2021), cell size is important to maintain leaf position and shape through the physical effect of cell turgor (Trinh et al. 2021; Ali et al. 2023; Zhang et al. 2024). Thus, this trade-off between mechanical integrity and physiological performance suggests a double constraint to leaf size by both cell size and number. In tracheophytes, leaves are mechanically supported by differentiated tissues and structures such as venation and petioles (Niklas 1999; Niinemets & Fleck 2002; Kawai & Okada 2016). This makes viable the strategy of having more and smaller cells to increase plant productivity, since more complex leaves would not essentially rely on cells as mechanical units. Additionally, some components of cell walls, particularly cellulose and lignin, would also assist this mechanical support (Li et al. 2001; Hayashi et al. 2005; Weng & Chapple 2010). This grants tracheophytes a certain degree of flexibility to adjust their cell numbers and sizes across diverse environments. Bryophytes are a group of poikilohydric land plants with c. 20000 species worldwide with three main lineages: hornworts, liverworts and mosses (Vanderpoorten & Goffinet 2009). From an anatomical and structural point of view, mosses are the most similar to tracheophytes, since their gametophytes have differentiated phyllids (leaves), caulidia (stems) and rhizoids. There are three main growth forms in mosses, depending on the position of the sporophyte and the general growth of the gametophyte: acrocarpous, pleurocarpous and cladocarpous (Vanderpoorten & Goffinet 2009; Glime 2017). The stems of acrocarpous mosses are generally erect, with sporophytes growing from the apex, while pleurocarpous mosses tend to grow horizontally, with more ramified stems with sporophytes growing from branch axils. Cladocarpous mosses are a special case almost exclusive of Sphagnum species, presenting prostrate growths similar to pleurocarpous mosses, but with sporophytes growing on the apex of branches as in acrocarpous mosses. Although not being monophyletic characters, growth forms tend to have different ecological preferences and tolerance levels (Gimingham & Birse 1957; Mägdefrau 1982; Wang et al. 2016). Moss leaf morphology, anatomical structures and plant macrostructure are essential for their ecology (Proctor 1990; Hedenäs 2001; Pan et al. 2016; Niinemets & Tobias 2019; Turberville et al. 2021). Regarding moss leaves, although some acrocarpous mosses have developed structures that functionally parallel some tracheophyte anatomical characters (such as pseudo-mesophylls formed by lamellae or advanced conduction systems; (Proctor 2005; Proctor & Bates 2018; Brodribb et al. 2020; Bok et al. 2022), there is a clear limitation on their leaf size. Most moss species have small, unistratose leaves with limited internal differentiation, lacking vascular tissues and never reaching the complexity of tracheophyte mesophylls (Carriquí et al. 2019). For this reason, their leaf mechanical properties and function may depend directly on cell arrangement, with little anatomical flexibility for interactions between cell size and number. Because of this, leaf size in bryophytes would be expected to rely on the mechanical support of cell turgor, which may be assisted by anatomical structures such as midribs (Frahm 1990; Huttunen et al. 2018) or their notoriously thick cell walls (Carriquí et al. 2019; Flexas & Carriquí 2020). An important trait influencing both plant anatomy and ecology is genome size (Knight & Beaulieu 2008; Leitch & Leitch 2012; Roddy et al. 2020; Bhadra et al. 2023). It is generally accepted that genome size determines the minimum cell size of a species (Gregory 2001; Beaulieu et al. 2008). In tracheophytes, species with smaller genomes (and consequently smaller cells) tend to display higher photosynthetic capacities through the optimisation of cell arrangement in photosynthetic tissues (Théroux-Rancourt et al. 2021). This effect of genome size on tissue packaging is in concordance with the trade-off between cell number and size in tracheophytes (Ma et al. 2025). A positive relationship between genome and cell size in moss gametophytes has been previously reported (Mir‐Rosselló et al. 2025). If a relationship exists between cell size and leaf dimensions in mosses, genome size could be a key determinant of their leaf size. Moreover, given that the link between genome and cell size appears to respond to environmental pressures (Faizullah et al. 2021; Mir‐Rosselló et al. 2025), mosses as highly tolerant and anatomically simple plants represent an excellent model system for studying the interplay between genome size, anatomy and ecology. In this study, we aimed to identify the main anatomical factors that determine leaf size in mosses and interpret them from a mechanical perspective. To this end, we measured different cell and leaf dimensions to analyse their interrelationships. We also contrasted these anatomical measurements with traits of ecological relevance, highlighting growth form and genome size. We hypothesise that leaf cells and other anatomical structures serve as a mechanical support of moss leaves, for which cell and leaf dimensions will be positively correlated among them. We also hypothesise that these correlations will vary depending on growth form and genome size. Finally, we discuss the potential ecological implications of our findings. Materials and Methods Plant material and species traits data Cell measurements were obtained from orthogonal microphotographs of 287 moss species (157 acrocarpous mosses and 130 pleurocarpous mosses) available in Lüth (2019) Mosses of Europe: a photographic flora (Supporting Information Dataset S1). This source allowed to measure a wide diversity of species representing the main growth forms and leaf anatomical variations (Figure 1). Sphagnum species were excluded due to their highly specialised leaf and cell structure, as well as their particular cladocarpous growth, which differs markedly from that of other mosses (Jassey & Signarbieux 2019; Oke et al. 2020). In addition to leaf and cell measurements, the other moss traits considered in the study were growth form, water conduction system, life form, leaf stratification and genome size. These traits were selected based on their extensively reported relevance to bryophyte ecology (Mägdefrau 1982; Proctor 2000; Glime 2017). Growth form and life form data were obtained from the Bryophytes of Europe Traits dataset (van Zuijlen et al. 2023), which follows the classifications of Glime (2017) and Mägdefrau (1982). Water conduction system was assigned to properly characterised ectohydric or endohydric species (Ligrone et al. 2000; Proctor 2000). Due to strong associations between certain traits and growth form (for example, all the endohydric species except one, and all the species with bi- to pluristratose leaves, were acrocarpous), comparative analyses were primarily centred on differences between acrocarpous and pleurocarpous mosses. A subset of 133 species with genome size (1C-values in pg) data available at the Kew Plant DNA C-values Database (Leitch et al. 2019) was compiled. Cell and leaf dimensions measurements Leaf width, length and area were measured from leaf photographs using ImageJ software (Schneider et al. 2012). Midrib length was also measured when present. In species with an excurrent nerve, its length was considered equivalent to total leaf length. Cell width and length, as well as cell wall thickness were also measured. Each cellular dimension was measured in five photosynthetic laminal cells per species, and the mean value was used for analyses. Cells were considered shaped as spheroids, from which cell volume was calculated as in the equation: \begin{equation} C_{V}=\frac{\pi\times{C_{w}}^{2}\times C_{l}}{6}\times T\nonumber \\ \end{equation} Where C V is the cell volume, C w the cell width and C l the cell length, and T is the corresponding Thain (1983) curvature correction factor. Sections of 1000 µm 2 were selected, and the number of cells in that area were counted as a proxy of cell density. Data analyses Data analyses were conducted with the RStudio software ver. 4.5.0 (R Core Team 2025). To contrast cell and leaf dimensions among growth forms, water conduction types, leaf stratification and life forms, Kruskal-Wallis tests were performed since the assumption of normality of residuals was not fulfilled (Okoye & Hosseini 2024), followed by the corresponding post-hoc Dunn test when required (Dinno 2024). To assess the relationship among leaf dimensions, between cell and leaf dimensions, and between genome size and cell and leaf dimensions, Pearson and Spearman (when normality assumptions were not fulfilled) correlations were performed among the mentioned variables for the whole dataset and for acrocarpous and pleurocarpous mosses separately. Data visualization was conducted with the ggplot2 package (Wickham 2016). Anatomical dimensions in moss gametophytes The mean ± standard error values of the cell dimensions measured from leaf photosynthetic laminal cells were 9.02 ± 0.39 µm for cell width (ranging from 2.69 to 47.10 µm), 39.78 ± 1.53 µm for cell length (ranging from 4.53 to 136.39 µm) and 4.60 x 10 3 ± 0.76 x 10 3 µm 3 for cell volume (ranging from 0.08 x 10 3 to 146.12 x 10 3 µm 3 ). For leaf dimensions, the values obtained were 0.80 ± 0.04 mm for leaf width (ranging from 0.08 to 5.47 mm), 2.49 ± 0.11 mm for leaf length (ranging from 0.21 to 11.87 mm) and 1.73 ± 0.19 mm 2 for leaf area (ranging from 0.01 to 30.85 mm 2 ). Cell and leaf dimensions in moss gametophytes significantly varied, in almost all cases, among growth forms, water conduction systems, life forms and leaf stratification degrees (Table 1, Figure 2, Supporting Information Figures S1-4). Acrocarpous mosses showed larger median cell widths (11.96 ± 0.62 µm for acrocarpous mosses vs. 5.95 ± 0.23 µm for pleurocarpous mosses, P < 0.001; Figure 2a ) and volumes (7.37 x 10 3 ± 1.38 x 10 3 µm 3 for acrocarpous mosses vs. 1.73 x 10 3 ± 0.29 x 10 3 µm 3 for pleurocarpous mosses, P < 0.01; Figure 2c ), while pleurocarpous mosses showed significantly longer cells (32.68 ± 1.80 µm for acrocarpous mosses vs. 47.16 ± 2.38 µm for pleurocarpous mosses, P < 0.001; Figure 2b ). Regarding leaf dimensions, no significant differences were detected for leaf width (0.86 ± 0.06 mm for acrocarpous mosses vs. 0.73 ± 0.03 mm for pleurocarpous mosses; Figure 2d ), and acrocarpous mosses showed significantly larger leaf lengths (3.05 ± 0.18 mm for acrocarpous mosses vs. 1.80 ± 0.08 mm for pleurocarpous mosses, P < 0.001; Figure 2e ) and areas (2.35 ± 0.33 mm 2 for acrocarpous mosses vs. 0.98 ± 0.08 mm 2 for pleurocarpous mosses, P < 0.001; Figure 2f ). All the measured leaves were longer than wide, with leaf width:length ratios ranging from 0.09 to 0.89. Leaf width and length were both strongly positively correlated with leaf area (Figure 3). The strength of these correlations varied slightly between growth forms: when considered independently, the proportion of explained variance for leaf width increased (Figure 3a ), while it decreased in the case of leaf length (Figure 3b ). Acrocarpous mosses tended to have longer midribs than pleurocapous species (mean ± standard error: 2.96 ± 0.19 mm in acrocarpous vs. 0.82 ± 0.07 mm in pleurocarpous; Kruskal-Wallis test, n= 287: χ 2 = 103.8, P < 0.001). The proportion of lamina length occupied by the midrib was variable (particularly among pleurocarpous species) up to a leaf length of around 4.8 mm. Beyond this threshold: 1) only acrocarpous mosses surpassed this leaf length, and 2) the midrib reached or surpassed the leaf apex in almost all cases (Figure 4). Relationship between cell and leaf dimensions in moss gametophytes Many of the cell dimensions measured showed significant positive correlations with leaf dimensions (Figure 5, Supporting Information Table S1). In acrocarpous mosses, cell width correlated with leaf width (Figure 5a ) and leaf area (Figure 5b ), as well as cell volume correlated with leaf width (Figure 5c ). In pleurocarpous mosses, only cell and leaf lengths correlated (Figure 5d ). Although some of them were statistically significant, the correlation coefficients of cell wall thickness with leaf dimensions were always weak (R 2 < 0.10; Supporting Information Table S1). Cell density negatively correlated with leaf width in all cases (Figure 6a ), with leaf length for pleurocarpous mosses (Figure 6b ) and with leaf area for the whole dataset and for pleurocarpous mosses (Figure 6c ). Relationship between genome size and anatomical dimensions Genome size significantly positively correlated with many of the cell and leaf dimensions measured (Figure 7, Supporting Information Table S2). Genome size correlated with cell width and length in acrocarpous mosses (Figure 7a,b ), and with cell volume in acrocarpous mosses and for the whole dataset (Figure 7c ). It also correlated with leaf width and length in acrocarpous mosses (Figure 7d,e ), and with leaf area in acrocarpous mosses and for the whole dataset (Figure 7f ). Although genome size significantly correlated with cell width and volume in pleurocarpous mosses, the correlation was very weak (R 2 < 0.20; Supporting Information Table S2). Discussion Anatomy is a key factor to understand the interaction of plants with their environment (Bolhàr-Nordenkampf & Draxler 1993; Oguchi et al. 2018). This is especially remarkable for bryophytes, for which their ecology is directly linked to their structure, morphology and anatomy (Mägdefrau 1982; Glime 2017; Niinemets & Tobias 2019). In this study, we aimed to explain the mechanical basis of leaf anatomy in mosses, and its potential implications on how leaf anatomy is related to plant ecology. Our results suggest that cell size is an important determinant of leaf size in mosses, being this influenced by different traits of ecological relevance, especially growth form and genome size. Anatomical determinants of leaf size in mosses Moss leaves, including those measured in this study, are mainly simple, undivided, and longer than wide. This results in a morphological variability much lower than that of tracheophytes (Traiser et al. 2005). This results in strong, constant correlations among leaf dimensions (Figure 3). The generally unistratose leaves of mosses, without internal vascular differentiation, result in an important mechanical limitation to leaf size, preventing them to reach the mean and maximum leaf sizes of tracheophytes, which are many orders of magnitude larger (Wright et al. 2017). Our results suggest that moss leaf size is mainly dependent on cell size and midrib length. Positive correlations were found between cell and leaf dimensions, especially for cell width and volume with leaf width and area (Figure 5, Supporting Information Table S1). This suggests that leaf size in mosses is dependent on cell turgor as a mechanical support (Trinh et al. 2021; Ali et al. 2023; Zhang et al. 2024). Despite the simplified anatomy of mosses, there is a trade-off between cell size and number (approached through cell density, Figure 6) similar to the one described for tracheophytes (Ma et al. 2025). However, in contrast to tracheophytes, which tend to expand leaf area by increasing the number of smaller cells, mosses appear to favour fewer, larger cells. This reflects their dependence on individual cells for structural support. In our set of species, leaf widths and leaf lengths when lacking a midrib reached their maximum values below 5 mm (Figure 4), suggesting that this may represent the upper limit for the mechanical support provided by cells. Hence, to exceed this threshold additional anatomical structures are likely required. Midribs are pluristratose structures usually present in moss leaves, and may have specialised cells such as hydroids and leptoids for water and nutrient conduction, and stereids for mechanical support (Frahm 1990; Ligrone et al. 2000; Huttunen et al. 2018). Midrib were always percurrent to excurrent (almost reaching or extending beyond the leaf apex, respectively) in leaves exceeding 4.8 mm in length (Figure 4), indicating that this structure is required for developing longer leaves in mosses. Midribs being placed longitudinally and not ramifying could explain why they enable moss leaves to grow longer but not wider, since increasing leaf width would likely require a more complex ramification pattern such as the reticulate venation of tracheophyte leaves (Niklas 1999; Niinemets & Fleck 2002; Kawai & Okada 2016). Regarding cell walls, despite being significantly thicker in bryophytes than in tracheophytes (Carriquí et al. 2019; Flexas & Carriquí 2020), we found no evidence of this influencing leaf dimensions (Supporting Information Table S1). Bryophyte cell walls lack lignin, an important structural polymer in tracheophytes that contributes to their ability to achieve larger dimensions (Bateman et al. 1998; Weng & Chapple 2010). As a result, bryophyte cell walls are more elastic, for which they would remain involved in other functions (e.g., stress tolerance or water retention and transport; Schofield 1981; Clarke & Robinson 2008; Coe et al. 2019; Perera‐Castro & Flexas 2022) more than in mechanical support. Moss leaf allometry varies depending on growth form Acrocarpous mosses exhibited larger laminal cells in terms of width and volume compared to pleurocarpous mosses (Figure 2a,c ). These cell dimensions were positively correlated with their leaf width and area (Figure 5a-c ). This relationship may be explained through a mechanical point of view, since cell turgor supports leaves physically (Ali et al. 2023; Zhang et al. 2024). Moreover, the longest leaves among mosses are achieved by acrocarpous mosses (Figure 2e ), always presenting excurrent to percurrent midribs (Figure 4), indicating that midribs function as their primary longitudinal mechanical support (Frahm 1990; Huttunen et al. 2018). Together, cell turgor and the presence of midribs likely enable the typical spreading position of leaves in hydrated acrocarpous mosses. This architecture, combined with pigment and chloroplast distribution adjustments, allows these erect mosses to optimise light interception by their canopies (Wang et al. 2016; Niinemets & Tobias 2019). Moreover, these anatomical reinforcements may be a prerequisite for the development of bi- to pluristratose leaves and specialised structures such as lamellae to increase their photosynthetic capacities (Proctor 2005; Proctor & Bates 2018). This could explain why these characters are more common in acrocarpous mosses. In addition, the most advanced endohydric conduction systems described in mosses are found within acrocarpous species (Ligrone et al. 2000; Brodribb et al. 2020). All of this suggests that anatomical specialization of acrocarpous mosses is the basis for their relatively higher performance through evolutionary convergences with tracheophyte anatomy (Bok et al. 2022). Pleurocarpous mosses tend to have notoriously elongated laminal cells (Fig 2b ). Most of these species are ectohydric and develop numerous anatomical structures that facilitate external water transport (e.g., paraphyllia and pseudoparaphyllia, leaf denticulation and plicae, inflated alar cells; Hedenäs 2001; Huttunen et al. 2018). It has previously been suggested that elongated cells may be implicated in facilitating water transport across leaves (Schöfield 1981), for which they could assist the ectohydric water transport. In this sense, longer cells could help maintaining the metabolism in longer leaves, which could explain the positive correlation observed between these cell length and leaf length in pleurocarpous mosses (Fig 5d ). Thus, unlike acrocarpous mosses, pleurocarpous species appear to rely less on the mechanical support from cell turgor or long midribs (Turberville et al. 2021). Their prostrate caulidia and overlapping leaf arrangement likely provide sufficient structural support to their leaves. As a result, pleurocarpous mosses may have evolved anatomical traits primarily related to maximizing water interception and retention (such as leaf imbrication and plication, paraphyllia and pseudoparaphyllia, etc.) rather than increasing their photosynthetic performance at the shoot level. Genome size influence on moss leaf anatomy Genome size has been demonstrated to be a relevant trait related to plant ecology (Leitch & Leitch 2012; Faizullah et al. 2021; Schley et al. 2022). Since genome and cell size are related in moss gametophytes (Mir‐Rosselló et al. 2025), bryophytes provide an excellent model system for investigating whether anatomy mediates as the link between genome size and plant ecology. Unexpectedly, many of the correlations conducted in this study for genome size were generally stronger for leaf dimensions than for cell dimensions (Figure 7, Supporting Information Table S2). This contrasts with previous studies suggesting that the influence of genome size decreases at higher phenotypical stages (Knight & Beaulieu 2008). One possible explanation is that our measurements were restricted to laminal photosynthetic cells, while genome size is thought to determine the minimum cell size among all cell types within a species (Gregory 2001; Beaulieu et al. 2008; Roddy et al. 2020). Therefore, the observed patterns may be explained by the overall effect of genome size on all the cells across the entire leaf, including those different from laminal photosynthetic cells. To our knowledge, this is the first study contrasting genome size and growth form in bryophytes. While genome size positively correlated with both cell and leaf dimensions (especially width and area) in acrocarpous mosses, these correlations for pleurocarpous mosses were notoriously weak (Figure 7, Supporting Information Table S2). Previous studies suggest that the relationship between genome size and anatomical and functional traits (including growth form) is related to environmental pressure (Faizullah et al. 2021; Bhadra et al. 2023; Mir‐Rosselló et al. 2025). In the case of bryophytes, acrocarpous mosses tend to display higher tolerance levels, while tending tend to inhabit harsher environments, and to showing more marked colonising character than pleurocarpous mosses (Birse & Gimingham 1955; Gimingham & Birse 1957; Mägdefrau 1982; Esposito et al. 1999; Glime 2017). It could be then hypothesised that the stronger environmental pressures experienced by acrocarpous mosses contribute to a tighter relationship between genome size and anatomical dimensions in this group. Concluding remarks Our results suggest that moss leaf size is highly dependent on mechanical supports apported by their anatomy. Cell turgor appears to be the main factor for supporting leaf width and area, while midrib length allows mosses to develop longer leaves. Although the trade-off between cell size and number previously described in tracheophytes is maintained in mosses, it operates in the opposite direction. Whereas tracheophytes tend to increase the number of smaller cells to increase leaf performance, mosses increase their leaf sizes by having fewer but larger cells. This suggests that the reliance on cells as units of mechanical support may represent a constraint on moss leaf size and could explain why they never develop large leaves. The mechanical roles of cell turgor and midribs are more important in acrocarpous mosses, which may help explain their potential to develop more complex leaf anatomies. On the contrary, pleurocarpous mosses appear to be less dependent on such mechanical support and instead display anatomical traits more closely related to stable habitats and water retention. In turn, genome size being related to leaf anatomy in acrocarpous mosses may reflect their adaptation to higher environmental pressures. Our study proves that plant anatomy is an important factor linking genome size and plant ecology. Acknowledgements P.M.M.R. work was funded by the FPU 2021 grant (FPU21/03500), Ministerio de Ciencia, Innovación y Universidades, Gobierno de España. M.C. was supported by a Vicenç Mut 2022 postdoctoral fellowship (PD‐047‐2022) funded by Conselleria de Fons Europeus, Universitat i Cultura from Govern de les Illes Balears. This study was funded by project POPEYE (PGC2018‐093824‐B‐C41) funded by Ministerio de Ciencia, Innovación y Universidades, Gobierno de España, and the ERDF (FEDER). Competing interest None declared. Author contributions P.M.M.R. compiled the data, conducted the statistical analyses and prepared the initial draft of the manuscript. All authors contributed to conceive the idea, to revise the data and the manuscript, and to prepare the final version of the manuscript. Data availability The data that supports the findings of this study are available in the supplementary material of this article (Supporting Information Dataset S1). References Ali O., Cheddadi I., Landrein B. & Long Y. (2023) Revisiting the relationship between turgor pressure and plant cell growth. New Phytologist 238 , 62–69.Bateman R.M., Crane P.R., DiMichele W.A., Kenrick P.R., Rowe N.P., Speck T. & Stein W.E. (1998) Early evolution of land plants: Phylogeny, physiology, and ecology of the primary terrestrial radiation. Annual Review of Ecology and Systematics 29 , 263–292.Beaulieu J.M., Leitch I.J., Patel S., Pendharkar A. & Knight C.A. 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Tables and Figures Table 1. Cell and leaf dimension differences (Kruskal-Wallis test) among different growth forms (cell dimensions: n= 135 acrocarpous mosses, n= 130 pleurocarpous mosses; leaf dimensions: n= 157 acrocarpous mosses, n= 130 pleurocarpous mosses), water conduction systems (cell dimensions: n= 76 ectohydric, n= 22 endohydric; leaf dimensions: n= 78 ectohydric, n= 41 endohydric), life forms (cell dimensions: n= 4 annuals, n= 11 cushions, n= 8 dendroids, n= 30 mats, n= 142 turfs, n=20 wefts; leaf dimensions: n= 4 annuals, n= 11 cushions, n= 8 dendroids, n= 31 mats, n= 159 turfs, n=20 weft) and leaf stratification levels (cell dimensions: n= 254 unistratose, 10 bistratose, 1 pluristratose; leaf dimensions: n= 262 unistratose, 12 bistratose, 11 pluristratose) for the measured species. Cell dimensions Width χ 2 = 83.12, P < 0.001 χ 2 = 17.91, P < 0.001 χ 2 = 40.38, P < 0.001 χ 2 = 3.57, P = 0.168 Length χ 2 = 21.62, P < 0.001 χ 2 = 8.48, P < 0.01 χ 2 = 9.40, P = 0.094 χ 2 = 11.14, P < 0.01 Volume χ 2 = 10.46, P < 0.01 χ 2 = 24.90, P < 0.001 χ 2 = 31.65, P < 0.001 χ 2 = 12.40, P < 0.01 Leaf dimensions Width χ 2 = 0.57, P = 0.450 χ 2 = 13.62, P < 0.001 χ 2 = 31.67, P < 0.001 χ 2 = 9.46, P < 0.01 Length χ 2 = 25.24, P < 0.001 χ 2 = 26.07, P < 0.001 χ 2 = 23.50, P < 0.001 χ 2 = 33.27, P < 0.001 Volume χ 2 = 11.15, P < 0.001 χ 2 = 26.47, P < 0.001 χ 2 = 24.54, P < 0.001 χ 2 = 20.09, P < 0.001 Figure 1. Anatomical variability of mosses considered in this study. The two main growth forms in mosses are pleurocarpous, which generally grow prostate and highly ramified (A: Thuidium tamariscinum (Hedw.) Schimp.), and acrocarpous, with erect, usually less ramified stems (B: Polytrichum commune Hedw.). Some species lack midribs (C: Fontinalis antipyretica Hedw.), but most mosses have short to long midribs (D: Rhynchostegium megapolitanum (Blandow ex F.Weber & D.Mohr) Schimp.), in some cases reaching or surpassing the leaf apex (E: Tortula muralis Hedw.). Although most moss leaves are unistratose (F: Pseudoscleropodium purum (Hedw.) M.Fleisch.), some species have bistratose leaves (G: Ceratodon purpureus (Hedw.) Brid.) and some groups can develop even more complex, pluristratose leaves (H: Polytrichum formosum Hedw.). Figure 2. Differences in leaf and cell dimensions between growth forms (acrocarpous: n= 135 for cell dimensions, n= 157 for leaf dimensions; pleurocarpous: n= 130 for both cell and leaf dimensions) for the measured mosses. Kruskal-Wallis tests: **, P < 0.01; ***, P < 0.001 (see Table 1 for further statistical details). Figure 3. Correlations among leaf dimensions in the measured mosses, in logarithmic scale. Solid black lines correspond to the correlation for all species combined (All: Pearson, n= 287, P < 0.001 for both leaf width and length). Orange circles and dashed lines correspond to acrocarpous mosses (Acro: Pearson, n= 157, P < 0.001 for both leaf width and length), and blue triangles and dotted lines correspond to pleurocarpous mosses (Pleu: Spearman, n= 130, P < 0.001 for both leaf width and length). Figure 4. Comparison between midrib length and total leaf length in the measured species (n= 287). The diagonal solid line represents the 1:1 intersection, and the horizontal dashed line indicates the leaf length threshold above which all species exhibit a percurrent or excurrent midrib. Orange circles correspond to acrocarpous species (n= 157) and blue triangles correspond to pleurocarpous species (n= 130). Figure 5. Correlations between cell and leaf dimensions in the measured mosses, in logarithmic scale. Orange circles and dashed lines correspond to acrocarpous species (Acro: Pearson, n = 135), and blue triangles and dotted lines correspond to pleurocarpous species (Pleu: Spearman, n= 130). Only significant correlations with R 2 > 0.20 are shown ( P < 0.001; see Supporting Information: Table S1 for further statistical details). Figure 6. Correlations between cell density (number of cells in 1000 µm 2 ) and leaf dimensions in the measured mosses. Orange circles and dashed lines correspond to acrocarpous species (Acro: Pearson, n = 125), and blue triangles and dotted lines correspond to pleurocarpous species (Pleu: Spearman, n= 72). Only significant correlations with R 2 > 0.20 are shown ( P < 0.001; see Supporting Information: Table S1 for further statistical details). Figure 7. Correlations between genome size (1C-value in pg) and cell and leaf dimensions in the measured mosses. Solid black lines correspond to correlations for all species combined (All: Spearman, n= 128 for cell dimensions; Spearman, n= 133 for leaf dimensions). Orange circles and dashed lines correspond to acrocarpous species (Acro: Spearman, n= 47 for cell dimensions; Pearson, n= 52 for leaf dimensions), and blue triangles and dotted lines correspond to pleurocarpous species (Pleuro: Spearman, n= 81 for both cell and leaf dimensions). Only significant correlations with R 2 > 0.20 are shown ( P < 0.001; see Supporting Information: Table S2 for further statistical details). Information & Authors Information Version history V1 Version 1 03 July 2025 Peer review timeline Published Plant Biology Version of Record 8 Dec 2025 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords anatomy bryophytes cell size ecology genome size growth leaf size mechanical support mosses Authors Affiliations Pere Miquel Mir-Rosselló 0000-0002-0004-1916 [email protected] Universitat de les Illes Balears Facultat de Ciencies View all articles by this author Jaume Flexas 0000-0002-3069-175X Universitat de les Illes Balears Facultat de Ciencies View all articles by this author Marc Carriquí Alcover 0000-0002-0153-2602 Universitat de les Illes Balears Facultat de Ciencies View all articles by this author Metrics & Citations Metrics Article Usage 461 views 317 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Pere Miquel Mir-Rosselló, Jaume Flexas, Marc Carriquí Alcover. Leaf size in mosses is structurally constrained by cell dimensions and genome size. Authorea . 03 July 2025. 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