DOI: 10.5061/dryad.tx95x6b6s
title: The smallest Zosterophyllum plant from the Lower Devonian of South China and the divergent life history strategies in zosterophyllopsids

description: Plants have evolved different life history strategies to overcome the
 limited amount of available resources; however, when and how divergent
 strategies of sexual reproduction evolved in early land plants is not well
 understood. As one of the notable and vital components of early
 terrestrial vegetation, the Zosterophyllopsida and its type
 genus Zosterophyllum, reached maximum species diversity during
 the Pragian (Early Devonian; ca. 410.8–407.6 million years ago). Here we
 describe a new species, Z. baoyangense sp. nov., based on well-preserved
 specimens from the Pragian-aged Mangshan Group of Duyun, Guizhou Province,
 China. The new plant is characterized by its small size, K-shaped
 branching, and tiny spikes with five to ten sporangia. This plant is most
 likely r-selected, completing its whole lifespan in a short time, and such
 a strategy contributes to reproduction in a suitable window time. By
 contrast, most other species of Zosterophyllum and the zosterophyllopsids
 on a broader scale are larger in body size and have greater investments in
 fertile tissues reflected in the size and total number of sporangia. We
 argue that the zosterophyllopsids probably benefited from the divergence
 of various life history strategies and thus constituted a major part of
 the Early Devonian floras.
description: (a) Morphological measurements of
 <em>Zosterophyllum</em> species The
 morphological descriptors of <em>Zosterophyllum</em> species
 are shown in Figure S2, including the width and length of spikes, width,
 and length of axes, sporangial height and width, etc. These descriptors
 are commonly used in the documentation of fossil plants and can be easily
 obtained. Compared to extant plants, the fertile
 investment involving the total number of spores and survival rate of
 spores cannot be obtained from plant fossils. Here, we define a new
 descriptor, total sporangial accommodation (TSA), to evaluate the mass or
 energy investment for spore production in each plant. The TSA is estimated
 by the following formula for each fertile axis: TSA =
 <em>n</em>·(<em>h</em>·<em>w</em>·<em>t</em>) where the total number of visible sporangia in a spike or fertile region is symbolized as <em>n</em>, and the sporangial height, width, and thickness are symbolized as <em>h</em>, ,<em>w</em> and <em>t</em> respectively. The sporangial thickness is rarely observed, but it is seemingly stable as observed in a few records (such as Hao, 1992; Hao et al., 2010). Here we assume a constant value for the thickness of sporangia (0.6 mm, as measured in the well-preserved specimens of <em>Zosterophyllum shengfengense</em> (Hao et al., 2010); that is, it is assumed that <em>t</em> = 0.6 mm for all plants). For example, the spike of <em>Z</em>. <em>qujingense</em> consists of 3–8 sporangia, which are 2.5–4.5 mm high and 3.1–4.7 mm wide, and therefore the value of TSA is 14.0–101.5 mm<sup>3</sup>. When the number of visible sporangia or estimated sporangia is not documented in species descriptions, we obtained an estimation from the published illustrations in the original literature. In addition to our plant, data on other <em>Zosterophyllum</em> species were collected from descriptions in the original literature or were directly measured by the authors based on the published illustrations. The data sources include those of Cascales-Miñana and Meyer-Berthaud (2014) and recent new works, such as Edwards et al. (2015), Edwards and Li (2018a), and Wang et al. (2018), spanning from Silurian to Early Devonian. However, only species of <em>Zosterophyllum</em> that have been well-studied or well-illustrated are included, while species with open nomenclature such as those with sp. and aff. are excluded, except for the earliest record <em>Zosterophyllum</em> sp. from the Silurian of Canada (Kotyk et al., 2002). The exact assignment of species with such open nomenclature requires further work. The same species from different localities are regarded as different data points. Plant height cannot be measured directly for most species of <em>Zosterophyllum</em>, but we could assess their possible height, or compare with each other for specimens with a similar preservation condition. An exceptional specimen of <em>Z. shengfengense</em> provides a good illustration that the plants of this group can be divided into four parts: the subterraneous fibrous root-like axes, the semi-below or below-ground rhizome, the basal aerial axes with K- or H-shaped branching, and the main erect part containing fertile regions or spikes (Hao et al., 2010; Hao and Xue, 2013). Hence, the combination of lengths of the latter two parts is regarded as representing the majority of the plant height. Here, the plant remains of <em>Zosterophyllum</em> were divided into three groups, based on their preservation status. Most specimens are only preserved in the middle to upper fertile parts (Group I); some specimens exhibit fertile axes with K- or H-shaped branching (Group II), such as <em>Z</em>. <em>qujingense</em>, Z. <em>minorstachyum,</em> and our plant; and a few other specimens show complete plants, such as <em>Z</em>. <em>shengfengense</em>, here considered as Group III. If rhizomes and aerial fertile parts of a species are found, although they may not be directly connected, such as in <em>Z</em>. <em>xishanense</em>, <em>Z</em>. <em>sinense,</em> and <em>Z</em>. <em>myretonianum</em>, such species are also included in Group III. By following the above methods, we compiled a new dataset of morphological measurements of <em>Zosterophyllum</em> species (see electronic supplementary material, table S3). The genus <em>Zosterophyllum</em> is probably not monophyletic based on phylogenetic analyses (Kenrick and Crane, 1997; Hao and Xue, 2013; Nibbelink and Tomescu, 2022). However, it is noted here that cladistic analyses of early land plants usually suffer from limited character sampling; for example, most <em>Zosterophyllum</em> species lack features of anatomy and spores. At present, we consider whether or not <em>Zosterophyllum</em> is monophyletic remains an unresolved issue. Nevertheless, we extended the scope of our analysis to consider other zosterophyllopsids, in order to provide (1) an extrapolation of our results to a higher-level category and (2) a discussion of character evolution within a (proposed) monophyletic group. (b) Measurements of zosterophyllopsids (and early lycopsids) Two different phylogenetic frameworks were employed herein (see electronic supplementary material, figure S5); the first recognized an inclusive clade, the Lycophytina sensu Kenrick et Crane (L-KC for short), which includes the Zosterophyllopsida sensu Kenrick et Crane, lycopsids, and some other taxa (Kenrick and Crane, 1997) and the second recognized the Zosterophyllopsida sensu Hao et Xue (Z-HX for short) as a monophyletic clade (Hao and Xue, 2013). By using the morphological descriptors shown in Figure S2, we obtained measurements of the members of L-KC and Z-HX. In fact, numerous zosterophyllopsids were not sampled in phylogenetic analyses (Gensel, 1992; Kenrick and Crane, 1997; Hao and Xue, 2013; Nibbelink and Tomescu, 2022), including some taxa that were recently described such as <em>Baoyinia</em> (Edwards and Li, 2018b). For those that have not been included in phylogenetic analyses, their assignment to the L-KC and the Z-HX was based on the authors’ own judgment. The dataset is shown in Table S4. The species with open nomenclature were excluded, except for those from the Ludlow. The sporangiotaxis and sporangium orientation of other zosterophyllopsids are more divergent than in <em>Zosterophyllum</em>. For TSA, three types of taxa were not considered: (1) some taxa show sparsely attached sporangia, such as <em>Danziella artesiana</em>, <em>Forania</em>, <em>Sawdonia</em>, etc.; (2) sporangia in some taxa are scattered distributed on axes, and thus it is hard to discern a spike or fertile region, such as <em>Deheubarthia</em>, <em>Gosslingia</em>, etc.; (3) the fertile regions of some taxa are not complete, such as <em>Crenaticaulis</em>, <em>Ventarura</em> etc. However, other morphological parameters of all these taxa, such as the width of axes and sporangial size, were included in the analyses. (c) Data analyses We assigned the sampled taxa to five-time bins, based on the international chronostratigraphic chart v 2023/09 (https://stratigraphy.org/chart#latest-version), that is, the Ludlow, Pridoli, Lochkovian, Pragian, and Emsian in ascending order. The majority of <em>Zosterophyllum</em> species are usually confined within a single-time bin (singletons). The species that extend across two or more time bins are rare, and our treatments are as follows. We treated the <em>Zosterophyllum</em> species from the Xujiachong Formation as Pragian age; the reason is that most plants occur in the middle to upper part of the Xujiachong Formation, although the upper part of this formation is regarded as early Pragian to ?earliest Emsian in age based on spore assemblages (Wellman et al., 2012). <em>Z</em>.<em> rhenanum</em> and <em>Z</em>. <em>confertum</em> were suggested to be derived from Lochkovian to Emsian or Pragian to Emsian deposits, but the occurrences of both species are mainly distributed in the Pragian (Schweitzer, 1979; Gossmann et al., 2022), and thus a Pragian age was assigned to them in our dataset. For the species in the L-KC and the Z-HX, we followed the age assignments in the original literature and used the range-through method for their statistics, that is, if one species was recorded from the Lochkovian to Emsian, then it was counted once separately in the Lochkovian, Pragian, and Emsian. Our dataset was analyzed and visualized by using R v.4.3.2 (RStudio/2023.09.1+494) with packages ggplot2 v.3.4.2 and ggsci v. 3.0.0. <strong>References</strong> Cascales-Miñana B, Meyer-Berthaud B. 2014 Diversity dynamics of Zosterophyllopsida. <em>Lethaia</em> <strong>47</strong>, 205–215. (doi: 10.1111/let.12051) Edwards D, Li CS. 2018a Diversity in affinities of plants with lateral sporangia from the Lower Devonian of Sichuan Province, China. <em>Rev. Palaeobot. Palynol.</em> <strong>258</strong>, 98–111. (doi: 10.1016/j.revpalbo.2018.07.002) Edwards D, Li CS. 2018b Further insights into the Lower Devonian terrestrial vegetation of Sichuan Province, China. <em>Rev</em>. <em>Palaeobot</em>. <em>Palynol</em>. <strong>253</strong>, 37–48. (doi: 10.1016/j.revpalbo.2018.03.004) Edwards D, Yang N, Hueber FM, Li CS. 2015 Additional observations on <em>Zosterophyllum yunnanicum</em> Hsü from the Lower Devonian of Yunnan, China. <em>Rev. Palaeobot. Palynol</em>. <strong>221</strong>, 220–229. (doi: 10.1016/j.revpalbo.2015.03.007) Hao SG, Xue JZ, Guo DL, Wang DM. 2010 Earliest rooting system and root: shoot ratio from a new <em>Zosterophyllum</em> plant. <em>New Phytol</em>. <strong>185</strong>, 217–225. (doi: 10.1111/j.1469-8137.2009.03056.x) Hao SG, Xue JZ. 2013 <em>The Early Devonian Posongchong flora of Yunnan</em>. Beijing: Science Press. Hao SG. 1992 Some observations on <em>Zosterophyllum australianum</em> Land & Cookson from the Lower Devonian of Yunnan, China. <em>Bot. J. Linn. Soc.</em> <strong>109</strong>, 189–202. (doi: 10.1111/j.1095-8339.1992.tb00265.x) Kenrick P, Crane PR. 1997 <em>The </em><em>o</em><em>rigin and </em><em>e</em><em>arly </em><em>d</em><em>iversification of </em><em>l</em><em>and </em><em>p</em><em>lants, a </em><em>c</em><em>ladistic </em><em>s</em><em>tudy</em>. Washington: Smithsonian Institution Press. Kotyk ME, Basinger JF, Gensel PG, de Freitas TA. 2002 Morphologically complex plant macrofossils from the late Silurian of Arctic Canada. <em>Am. J. Bot.</em> <strong>89</strong>, 1004–1013. (doi: 10.3732/ajb.89.6.1004) Nibbelink M, Tomescu AMF. 2022 Exploring zosterophyll relationships within a more broadly sampled character space: a focus on anatomy. <em>Int</em>. <em>J</em>.<em> Plant Sci</em>. <strong>183</strong>, 535–547. (doi: 10.1086/720384) Schweitzer HJ. 1979 Die Zosterophyllaceae des rheinischen Unterdevons. <em>Bonner Paläobotanische Mitteilungen</em> <strong>3</strong>, 1–32. Wang Y, Xu HH, Wang Y, Qiang F. 2018 A further study of <em>Zosterophyllum sinense</em> Li and Cai (Zosterophyllopsida) based on the type and the new specimens from the Lower Devonian of Guangxi, southwestern China. <em>Rev</em>. <em>Palaeobot</em>. <em>Palynol</em>. <strong>258</strong>, 112–122. (doi: 10.1016/j.revpalbo.2018.05.008) Wellman CH, Zhu HC, Marshall JEA, Wang Y, Berry CM, Xu HH. 2012 Spore assemblages from the Lower Devonian Xujiachong Formation from Qujing, Yunnan, China. <em>Palaeontology</em> <strong>55</strong>, 583–611. (doi: 10.1111/j.1475-4983.2012.01143.x)
description: # The smallest *Zosterophyllum* plant from the Lower Devonian of South
 China and the divergent life history strategies in zosterophyllopsids ##
 **Authors** Pu Huang [puhuang@nigpas.ac.cn](mailto:puhuang@nigpas.ac.cn)
 Jia-Shu Wang [jiashuwang1996@163.com](mailto:jiashuwang1996@163.com) 
 Yi-Ling Wang [wyl89@pku.edu.cn](mailto:wyl89@pku.edu.cn) Lu
 Liu [liulu@nnhm.org.cn](mailto:liulu@nnhm.org.cn) Jing-Yu
 Zhao [88zhaojy@163.com](mailto:88zhaojy@163.com) Jin-Zhuang
 Xue [pkuxue@pku.edu.cn](mailto:pkuxue@pku.edu.cn) ## **File list**
 Huang_tables_figures_ESM.pdf source_data_of_figure_2.zip
 source_data_of_figure_3.zip The_rscript_Zosterophyllum.R ## **File
 descriptions** **Huang_tables_figures_ESM.pdf** — The file contains
 supplementary figures S1–S5 and supplementary tables S1–S4, including
 traffic map of the fossil locality and lithological column of the Mangshan
 Group at the Baoyang section with plant-bearing bed (figure S1); sketch
 illustrations of morphological descriptors of *Zosterophyllum* species and
 other related plants (figure S2); morphology of specimen PB203563 of
 *Zosterophyllum* *baoyangense *sp. nov.* *(figure S3); interpretative line
 drawings of the spikes of *Zosterophyllum* *baoyangense* sp. nov., based
 on specimens PB203562 and PB203563 respectively (figure S4); phylogenetic
 frameworks of the Lycophytina sensu Kenrick et Crane and the
 Zosterophyllopsida sensu Hao et Xue (figure S5); morphological comparisons
 between *Zosterophyllum* *baoyangense* sp. nov. and other related plants
 (table S1); morphological comparisons among the species of
 *Zosterophyllum*, including *Z. baoyangense* sp. nov. (table S2); and raw
 data of morphological measurements of *Zosterophyllum* species (table S3)
 and the members of the Lycophytina sensu Kenrick et Crane and the
 Zosterophyllopsida sensu Hao et Xue (table S4) from late Silurian to Early
 Devonian. **source_data_of_figure_2.zip** —The file contains source data
 of Figure 2, which are shown in six tables (all in CSV format), including
 Diversity-Zosterophyllum.csv, Length of axis-Zosterophyllum.csv, Width of
 axis-Zosterophyllum.csv, TSA-Zosterophyllum.csv,
 Sporangia-Zosterophyllum.csv, and TSA~Width of axis-Zosterophyllum.csv.
 Units for the variables in this source data: mm for “Length of axis”,
 “Width (min)”, “Width (max)”, “Width (mean)”, “Height (min)” and “Height
 (max)”; and mm3 for “TSA (min)”, “TSA (max)” and “TSA (mean)”.
 **“Diversity-Zosterophyllum.csv”** includes species number of
 *Zosterophyllum* in different periods from late Silurian to Early
 Devonian, based on this study and Cascales-Miñana and Meyer-Berthaud,
 (2014, Diversity dynamics of Zosterophyllopsida. Lethaia 47, 205–215).
 **“Length of axis-Zosterophyllum.csv”** includes maximum length of axes of
 *Zosterophyllum* species in different periods from late Silurian to Early
 Devonian; “Group I” indicates the specimens only preserved in the middle
 to upper fertile parts, “Group II” indicates the specimens showing fertile
 axes with K- or H-shaped branching, and “Group III” indicates the
 specimens with a complete morphology; if rhizomes and aerial fertile parts
 of a species are found, although they may not be directly connected, such
 species are also included in “Group III.” **“Width of
 axis-Zosterophyllum.csv”** includes the width of axes of *Zosterophyllum
 *species in different periods from late Silurian to Early Devonian.
 “x(left)” and “x(right)” indicates the position of left and right sides,
 respectively, of the rectangles in the plot of axial width for each time
 bin (figure 2c), and each rectangle includes the range of axial width
 values of *Zosterophyllum* species in different periods, that is, minimum
 through maximum value; “Width (min)”, “Width (max)” and “Width (mean)”
 indicate the minimum, maximum, and mean values of axial width of measured
 species. **“TSA-Zosterophyllum.csv”** includes total sporangial
 accommodation (TSA) of *Zosterophyllum* species in different periods from
 late Silurian to Early Devonian; the TSA is estimated by the following
 formula for each fertile axis: TSA = *n*·(*h*·*w*·*t*), where, the total
 number of the visible sporangia in a spike or fertile region is symbolized
 as *n*, and the sporangial height, width, and thickness are symbolized as
 *h*, *w* and *t* respectively; “x(left)” and “x(right)” indicates the
 position of left and right sides, respectively, of the rectangles in the
 plot of TSA for each time bin (figure 2d); “TSA (min)”, “TSA (max)” and
 “TSA (mean)” indicate the minimum, maximum, and mean values of TSA of
 measured species. **“Sporangia-Zosterophyllum.csv”** includes measurements
 of sporangia of *Zosterophyllum* species of different time bins; “Width
 (min)”, “Width (max)”, “Height (min)” and “Height (max)” indicate minimum
 width, maximum width, minimum height, and the maximum height of sporangia
 in different *Zosterophyllum* species. **“TSA~Width of
 axis-Zosterophyllum.csv”** includes TAS and width of axes of
 *Zosterophyllum* species; “Width (min)” and “Width (max)” indicate minimal
 and maximal value, respectively, of axial width of all measured
 *Zosterophyllum* species; “TSA (min)” and “TSA (max)” indicate minimal and
 maximal value, respectively, of TAS of all measured *Zosterophyllum*
 species. **source_data_of_figure_3.zip**—The file contains source data of
 figure 3, and includes ten tables in CSV format, including: Length of
 axis-Lycophytina.csv, Width of axis-Lycophytina.csv, TSA-Lycophytina.csv,
 Sporangia-Lycophytina.csv; TSA Width of axis-Lycophytina.csv, Length of
 axis- Zosterophyllopsida.csv, Width of axis-Zosterophyllopsida.csv; TSA-
 Zosterophyllopsida.csv, Sporangia-Zosterophyllopsida.csv, and TSA Width of
 axis-Zosterophyllopsida.csv. Units for the variables in this source data:
 mm for “Length of axis”, “Width (min)”, “Width (max)”, “Width (mean)”,
 “Height (min)” and “Height (max)”; and mm3 for “TSA (min)”, “TSA (max)”
 and “TSA (mean).” **“Length of axis-Lycophytina.csv”** includes the
 maximum preserved length of axes of the members in the Lycophytina sensu
 Kenrick et Crane in different periods from late Silurian to Early
 Devonian. **“Width of axis-Lycophytina. csv”** includes the width of axes
 of the members in the Lycophytina sensu Kenrick et Crane in different
 periods from late Silurian to Early Devonian; “x(left)” and “x(right)”
 indicate the position of left and right sides, respectively, of the
 rectangles in the plot of the width of axis for each time bin; “Width
 (min)” and “Width (max)” indicate the minimal and maximal value,
 respectively, of axial width of all measured members of the Lycophytina
 sensu Kenrick et Crane. **“TSA-Lycophytina.csv”** includes the TSA of the
 members in the Lycophytina sensu Kenrick et Crane in different periods
 from late Silurian to Early Devonian; “x(left)” and “x(right)” indicate
 the position of left and right sides, respectively, of the rectangles in
 the plot of TSA for each time bin; “TSA (min)” and “TSA (max)” indicate
 the minimal and maximal value, respectively, of TSA of all measured
 members of the Lycophytina sensu Kenrick et Crane.
 **“Sporangia-Lycophytina.csv”** includes sporangial measurements of the
 members in the Lycophytina sensu Kenrick et Crane of different time bins;
 “Width (min)”, “Width (max)”, “Height (min)”, and “Height (max)” indicate
 minimum width, maximum width, minimum height, and maximum height,
 respectively, of sporangia in the members of the Lycophytina sensu Kenrick
 et Crane. **“TSA~Width of axis-Lycophytina.csv”** include axial width and
 TSA of all sampled members in the Lycophytina sensu Kenrick et Crane;
 “Width (min)”, “Width (max)” and “Width (mean)” indicate minimum width,
 maximum width, and mean width of axes, respectively; “TSA (min)”, “TSA
 (max)” and “TSA (mean)” indicate minimum, maximum, and mean values of TSA,
 respectively. **“Length of axis- Zosterophyllopsida.csv”** includes the
 maximum preserved length of axes of the members in the Zosterophyllopsida
 sensu Hao et Xue from late Silurian to Early Devonian. **“Width of
 axis-Zosterophyllopsida.csv”** includes the width of axes of the members
 in the Zosterophyllopsida sensu Hao et Xue from late Silurian to Early
 Devonian; “x(left)” and “x(right)” indicates the position of left and
 right sides, respectively, of the rectangles in the plot of axial width
 for each time bin; “Width (min)” and “Width (max)” indicate the minimal
 and maximal value, respectively, of axial width of all measured members of
 the Zosterophyllopsida sensu Hao et Xue. **“TSA-Zosterophyllopsida.csv”**
 includes TSA of the members in the Zosterophyllopsida sensu Hao et Xue
 from late Silurian to Early Devonian; “x(left)” and “x(right)” indicate
 the position of left and right sides, respectively, of the rectangles in
 the plot of TSA for each time bin; “TSA (min)” and “TSA (max)” indicate
 the minimal and maximal value, respectively, of TSA of all measured
 members of the Zosterophyllopsida sensu Hao et Xue.
 **“Sporangia-Zosterophyllopsida.csv”** includes sporangial width and
 height of members in the Zosterophyllopsida sensu Hao et Xue in different
 time bins; “Width (min)”, “Width (max)”, “Height (min)”, and “Height
 (max)” indicate minimum width, maximum width, minimum height, and maximum
 height, respectively, of sporangia in the members of the
 Zosterophyllopsida sensu Hao et Xue. **“TSA~Width of
 axis-Zosterophyllopsida.csv”** includes TAS and axial width for all
 sampled members in the Zosterophyllopsida sensu Hao et Xue; “Width (min)”,
 “Width (max)” and “Width (mean)” indicate minimum width, maximum width,
 and mean width of axes, respectively; “TSA (min)”, “TSA (max)” and “TSA
 (mean)” indicate minimum, maximum, and mean values of TSA, respectively.
 **The_rscript_Zosterophyllum.R**—The file contains the R scripts that
 employ packages ggplot2 v.3.4.2 and ggsci v. 3.0.0. to analyze and
 visualize the data of files 2 and 3. ## **Files and variables** **File:
 source data of figure 2.rar** **Description:**
 Diversity-Zosterophyllum.csv (for figure 2a); Length of
 axis-Zosterophyllum.csv (for figure 2b); Width of axis-Zosterophyllum.csv
 (for figure 2c); TSA-Zosterophyllum.csv (for figure 2d);
 Sporangia-Zosterophyllum.csv (for figure 2e); TSA~Width of
 axis-Zosterophyllum.csv (for figure 2f, g); units for the variables in
 this source data: mm for “Length of axis”, “Width (min)”, “Width (max)”,
 “Width (mean)”, “Height (min)” and “Height (max)”; and mm3 for “TSA
 (min)”, “TSA (max)” and “TSA(mean).” **File: source data of figure 3.rar**
 **Description:** Length of axis-Lycophytina.csv (for figure 3a); Width of
 axis-Lycophytina.csv (for figure 3b); TSA-Lycophytina.csv (for figure 3c);
 Sporangia-Lycophytina.csv (for figure 3d); TSA Width of
 axis-Lycophytina.csv (for figure 3e, f); Length of
 axis-Zosterophyllopsida.csv (for figure 3g); Width of
 axis-Zosterophyllopsida.csv (for figure 3h); TSA-Zosterophyllopsida.csv
 (for figure 3i); Sporangia-Zosterophyllopsida.csv (for figure 3j); TSA
 Width of axis-Zosterophyllopsida.csv (for figure 3k, l); units for the
 variables in this source data: mm for “Length of axis”, “Width (min)”,
 “Width (max)”, “Width (mean)”, “Height (min)” and “Height (max)”; and mm3
 for “TSA (min)”, “TSA (max)” and “TSA (mean).” ## **Code/software** Our
 dataset was analyzed and visualized by using R v.4.3.2
 (RStudio/2023.09.1+494) with packages ggplot2 v.3.4.2 and ggsci v. 3.0.0.