Wood anatomy of Tabaroa, a monotypic papilionoid legume genus narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests

Wood anatomy of Tabaroa, a monotypic papilionoid legume genus narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Wood anatomy of Tabaroa, a monotypic papilionoid legume genus narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests Marcelo dos Santos Silva, Daisy Burris, Cássia Sacramento, Lazaro Benedito da Silva, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3921723/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 6 You are reading this latest preprint version Abstract Wood anatomy can serve as a source of phylogenetically informative characteristics, as well as a model of how more plastic characters have evolved over time in plants, e.g., through ancestral reconstructions of wood anatomical traits across well-resolved phylogenies. However, the evolution of wood anatomy is largely unexplored within a phylogenetic context due to limited availability of anatomical data across taxa. When compared with other angiosperm families, Leguminosae is relatively well-documented, yet it still lacks comprehensive wood anatomical information, particularly in undersampled papilionoid clades. In order to contribute to the understanding of micromorphological diversity across papilionoid legumes, we newly characterize the wood anatomy of Tabaroa caatingicola , a papilionoid species narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests, that has been molecularly placed in the poorly anatomically studied tribe Brongniartieae. Optical histology and scanning electron microscopy (SEM) were used to examine and describe the wood anatomy of T. caatingicola and compare it with six phylogenetically related Brongniartieae genera: Amphiodon , Behaimia , Haplormosia , Harpalyce , Limadendron , and Poecilanthe . Wood anatomy of Tabaroa suggests adaptations to the irregular rainfall of the harsh Caatinga environment, featuring distinct growth rings, prone to semi-ring-porous wood, multiple narrow vessels, simple perforation plates, and small and vestured pits. These traits increase water flow during abundance and ensure hydraulic safety during scarcity, minimizing embolism formation and spread. By focusing on the genus Tabaroa , an ecologically distinctive and evolutionarily isolated lineage, this study contributes to the understanding of the systematic and functional wood anatomy variation in the papilionoid legume tribe Brongniartieae. Brongniartieae ecological wood anatomy Fabaceae functional wood anatomy systematic wood anatomy vestured pits Figures Figure 1 Figure 2 Figure 3 Figure 4 1 Introduction Wood anatomical features have consistently been recognized to provide valuable sources of phylogenetic information (Baas et al. 2000 ; Carlquist 2013 ; Herendeen and Miller 2000 ; Olson 2005 ), as well as a model of how characters more plastic have evolved over time in plants, through ancestral character reconstructions in well-resolved phylogenies ( e.g. Pace et al. 2022 ; Silva et al. 2021 ). Wood anatomical features are able to shed light on obscure relationships at deeper nodes (Lens et al. 2007 ). However, the evolution of wood anatomy is still poorly unexplored within a phylogenetic context, perhaps due to the large knowledge gaps of microscopic wood structures across clades (Lens et al. 2007 ). Wood anatomy has been instrumental for delimitating several flowering plant families including the Marcgraviaceae, Tetrameristaceae (Lens et al. 2005 ), and Sapotaceae (Kukachka 1980 ). Likewise, the wood anatomy of the ecologically and economically important family Leguminosae is relatively well documented, when compared with other angiosperm families. (Gasson 1996 ; Gasson et al. 2004 ; Stepanova et al. 2013 ). However, most tribes within the Papilionoideae are yet to be anatomically studied. Describing wood anatomical features of endemic, rare, monospecific or poorly diverse genera is therefore critical for understanding evolutionary patterns of the micromorphological diversity across legumes. This is particularly critical in understanding how tree species can behave in the face of climate change, especially in relation to temperature and precipitation variations that directly affect xylem transport and the survival capacity of plants (Anfodillo and Olson 2021 ; Breshears et al. 2018 ; Fontes et al. 2022 ; Hajek et al. 2016 ) The papilionoid legume genus Tabaroa and its only known species T. caatingicola L.P.Queiroz, G.P.Lewis & M.F.Wojc. of the tribe Brongniartieae (Fig. 1 ) have been described more than ten years ago (Queiroz et al. 2010 ), yet wood anatomy in this phylogenetically important branch of the tribe remains fully unknown. T. caatingicola is narrowly endemic to a small area of seasonally dry tropical forest and woodland (SDTFW) on sandy soils of the Brazilian Caatinga domain, in southwestern Bahia (Fig. 2 a), whereas its most closely related species Amphiodon effusus Huber is ecologically confined to the Amazon rainforest (Cardoso et al. 2013 ). This clade, in turn, is sister to the genus Harpalyce , resulting in the topology ( Harpalyce ( Tabaroa , Amphiodon )) that has been corroborated in several publications involving phylogenetic analyses of nuclear and plastid DNA sequence data (Cardoso et al. 2012 , 2013 , 2017 ; Meireles et al. 2014 ; Queiroz et al. 2010 , 2017 ) (Fig. 1 ). Brongniartieae is a taxonomically intriguing tribe within the Papilionoideae subfamily of the Leguminosae, comprising 15 genera, mostly monospecific or with few species, totalling approximately 180 species (POWO 2024; WFO 2024 ). Despite the modest number of species, the Brongniartieae tribe displays significant morphological diversity, and a wide geographic distribution across continents and biomes, with marked variation in ecological preferences. For this reason, many of its morphologically disparate genera were previously placed in at least four distantly related tribes (Cardoso et al. 2017 ; Queiroz et al. 2017 ). Caatinga is an indigenous Tupi word that means “white forest”, describing the grey and light aspect during the dry season, when the majority of trees and shrubs are devoid of leaves, allowing light to reach the ground, consists predominantly of xeric shrub, thorn and drought resistant species, dominated by legume trees (Giulietti et al. 2004 ; Queiroz 2009 ). The region is characterized by an extended dry season and erratic rainfall, resulting in sparse foliage and undergrowth (Leal et al. 2005 ) (Fig. 1 ). It harbours numerous endemic species, with 34% of the flora exclusive to the area, many featuring unique physiological adaptations to mitigate water loss during prolonged drought periods (Giulietti et al. 2004 ). Although relatively common locally, T. caatingicola only occupies a restricted geographic area of about 12 Km 2 and has been classified as Critically Endangered (IUCN 2001). Here we newly present wood anatomical description for the ecologically distinctive, phylogenetically isolated, and highly threatened species T. caatingicola and compares it with six phylogenetically related genera within the Brongniartieae, for which any wood anatomical data are available: Amphiodon , Behaimia , Haplormosia , Harpalyce , Limadendron , and Poecilanthe (Fig. 2 ). Furthermore, we analyze the wood anatomy of T. caatingicola in light of the most modern theories and functional hypotheses of secondary xylem. 2 Material and methods 2.1 Wood sampling and anatomical characterization – Wood samples from three trees were collected from the trunk at diameter at breast height (DBH = 1.30 m), in trees with apparent health and straight stem in a population of Tabaroa caatingicola in the municipality of Dom Basílio, state of Bahia, northeastern Brazil (S 13º47'19"; W 41º30'04") (Fig. 3 a). Non-destructive wood collection followed the procedure described by Silva et al. ( 2022b ). Samples were vouched, recorded, and deposited in the Herbarium and Xylotheque of the Instituto de Biologia of the Universidade Federal de Pelotas (Table 1 ). The climate from the collection area is dry, semi-arid of the low latitude and altitude (BSh), according to Köppen-Geiger’s classifications, with annual mean precipitation between 588–619 mm, and temperature 23.3–24.0 ºC, with great thermal amplitude throughout the year, 14.7–31.8 ºC (Fick and Hijmans 2017 ). The main biome in the Caatinga phytogeographical domain is the seasonally dry tropical forest and woodland (Queiroz et al. 2017 ), which can be characterized as a xerophytic vegetation across most of the semi-arid region of Northeastern Brazil, including mostly sparse vegetation that covers massifs and plateaus where rivers are usually seasonal. Leguminosae species are among the most dominating flowering plants of the Caatinga (Queiroz 2006 ; Queiroz et al. 2017 ). Despite comprising almost 10% of Brazil's territory, the Caatinga is inadequately explored, with only 1% designated as a Conservation Protection Area. The region faces threats from deforestation and unsustainable agricultural practices, such as agriculture and cattle ranching, leading to soil salinization (Leal et al. 2005 ; Silva et al. 2004). Table 1 Diameter at breast height, height and quantitative data on the wood anatomy of Tabaroa caatingicola (Leguminosae) for the three tree samples analyzed, recorded, and deposited in the xylotheque of the Instituto de Biologia of the Universidade Federal de Pelotas (PELw). Additionally, vulnerability and mesomorphy indices (sensu Carlquist 1977 ) are provided. Herbarium voucher: PEL 27325. Morphological and anatomical parameters / Samples PELw 01 PELw 02 PELw 03 Diameter at breast height (1.30 m) (cm) 5.1 3.5 4.8 Height (m) 4.5 3.0 4.0 Vessels per square millimeter 119 ± 32 129 ± 11 147 ± 17 Vessel tangential diameter (µm) 44.0 ± 23.0 49.5 ± 16.2 63.1 ± 20.9 Vessel element length (µm) 163 ± 64 180 ± 31 197 ± 26 Intervessel/vessel-ray pit size (µm) 4.0 ± 1.5 4.4 ± 1.1 4.3 ± 1.2 Fibre length (µm) 727 ± 255 709 ± 140 799 ± 128 Fibre diameter (µm) 13.4 ± 3.7 13.1 ± 2.0 14.0 ± 2.1 Fibre wall thickness (µm) 4.7 ± 1.8 5.0 ± 0.8 5.4 ± 1.0 Fibre pit size (µm) 3.7 ± 1.3 2.8 ± 0.9 2.5 ± 0.5 Rays/mm 9.7 ± 2.3 11.4 ± 1.3 10.4 ± 1.1 Ray width (µm) 14.0 ± 4.3 18.2 ± 3.6 22.6 ± 4.1 Ray width (cell numbers) 2.0 ± 0.5 2.1 ± 0.4 2.5 ± 0.5 Ray height (µm) 126 ± 22 117 ± 15 129 ± 18 Ray height (cell numbers) 6.9 ± 1.3 6.1 ± 0.9 7.2 ± 1.0 Vulnerability index 0.4 ± 0.2 0.4 ± 0.1 0.4 ± 0.1 Mesomorphy index 68.2 ± 51.8 68.9 ± 25.5 83.1 ± 27.0 The preparation of histological slides followed the usual plant anatomical methods described by Johansen ( 1940 ) and Sass ( 1951 ). Histological sections between 18–30 µm in thickness were made using a Leica© sliding microtome. Sections of each sample were clarified with sodium hypochlorite (50%), coloured with 1% alcoholic safranin, dehydrated in a 50–100% alcoholic series and mounted in synthetic Canada Balsam© or Entellan©. The method proposed by Franklin ( 1945 ), modified by Kraus and Arduin ( 1997 ), was followed to analyze dissociated cell elements. For the Scanning Electron Microscopy (SEM) analyses, longitudinal wood sections, 18–30 µm thick were dehydrated in an alcoholic series, dried in a dry chamber (60°C ~ 12hs), placed on a stub with a double-faced carbon label and metallized with gold. The observation of the samples was carried out with a SEM JEOL 6390LV© in the Instituto Gonçalo Moniz (Fundação Oswaldo Cruz – FIOCRUZ). The following quantitative wood anatomy characters were measured: vessels (vessels per square millimeter, tangential diameter, length, intervessel/vessel-ray pit outer aperture diameter); fibres (length, width, and wall thickness); and rays (rays/mm, width, and height). Indices of vulnerability (vessel diameter / vessels per square millimeter) and mesomorphy (indices of vulnerability × vessel element length) were calculated according to Carlquist ( 1977 ). The measurements of vessels per square millimeter were obtained with ANATI QUANTI© software (Aguiar et al. 2007 ). The other measurements were carried out with an Olympus CX40© microscope coupled with a micrometric lens and the factors obtained were converted to µm through a conversion factor. The terminology used in the anatomical descriptions is in accordance with the IAWA Committee ( 1989 ). The measurement of the anatomical parameters n = 30 was fixed. 2.2 Mapping distribution across geographic and climatic spaces – To assess the distribution range across geographic and climatic spaces, we used built a taxonomically verified specimen record data from the analyses of herbarium collections (ALCB, CEPEC, CEN, HUEFS, MO, MBM, NY, RB, RON, SP, SPF, US); acronyms according to Thiers 2022 ) as available in the online databases of speciesLink ( https://specieslink.net/ ) and Reflora ( https://reflora.jbrj.gov.br/reflora/PrincipalUC/PrincipalUC.do ). Any erroneous georeferenced records were filtered with the R package coordinateCleaner (Zizka et al. 2019 ), but whenever possible these specimens and others without geographic coordinates were included based on the closest georeferenced locality accessed by comparing with localities of other plant specimens in the speciesLink database. From the latitude and longitude of each herbarium collection (a total of 153 specimens recorded, Appendix S1), the variables BIO12 (Annual Precipitation) and BIO4 (Temperature Seasonality) were extracted from the WorldClim v.2.0 model layers (WGS84 projection; Fick and Hijmans 2017 ) using the R library raster (Hijmans 2024 ). Bioclimatic variables (O’Donnell and Ignizio 2012 ) were derived from these climate models using the extract function in the raster library. We mapped the range distribution of Tabaroa and its most phylogenetically closely related genus Amphiodon against the BIO12 bioclimatic variable, by using the R packages raster , ggmap (Kahle and Wickham 2013 ), ggplot2 (Wickham et al. 2016), ggspatial (Dunnington 2023 ), and rnaturalearth (South 2017 ) as implemented in RStudio ( 2022 ). The built a scatterplot to show the bioclimatic space of Tabaroa and Amphiodon across the BIO12 and BIO4 axes. 3 Results Here, we provide a full description of the wood anatomy of Tabaroa caatingicola . This species exhibits “bark light grey with darker wavy stripes that interlink and cross over to form a pattern suggestive of crocodile skin, inner bark dark green” (Fig. 3 b), hence the vernacular name pau-jacaré (crocodile wood) because of the resemblance of the bark to crocodile skin, as described by Queiroz et al. ( 2010 , p. 199). 3.1 General, organoleptic, and macroscopic wood anatomical characters – Wood with a brown heartwood and yellowish sapwood (Fig. 3 c-d), highly dense, and resistant to cutting. Odorless; straight grain; fine texture; gloss present. Distinct growth ring, macroscopically demarcated by fibrous zone (Fig. 3 c-d). Diffuse porosity, tending towards semi-ring porosity. Vessels arranged in a radial pattern; solitary and multiple radial vessels of 2–5, less frequently 6–10; noticeable only under magnification; small in tangential diameter; very numerous; some vessels obstructed by deposits in the heartwood. Axial parenchyma indistinct even under magnification. Rays noticeable only under magnification, storied. 3.2 Microscopic anatomical description of wood – Growth rings are well defined, marked by thick-walled and radially flattened fibres and distended rays, prone to semi-ring-porous wood (Fig. 4 a-b). Vessels are wood diffuse-porous with a tendency to semi-porous (Fig. 4 a-b). Vessel arrangement in radial pattern (Fig. 4 a-b). Vessel groupings: solitary and radial multiples of 2–5 (solitary 20–29%, radial multiples of two 26–29%, of three 18–27%, of four 10–15%, of five 4–7%) occurring less frequently radial multiples of 6–10, 4–12% (Fig. 4 a-b); simple perforation plates (Fig. 4 f); intervessel pits alternate, small and vestured (Fig. 4 g); vessel-ray pits with distinct borders, similar to intervessel pits in size and shape throughout the ray cell (Fig. 4 d). Deposits in heartwood vessels. Fibres with simple to minutely bordered pits, few, common in both radial and tangential walls. Nonseptate fibres present. Fibres very thick-walled (Fig. 4 b). Axial parenchyma diffuse and paratracheal, both scanty (Fig. 4 a-b). Fusiform and two cells per parenchyma strand (Fig. 4 c), less often 3–4 cells per parenchyma strand. Rays width 1 to 3 cells (Fig. 4 c). Cellular composition of rays: homocellular, all ray cells procumbent (Fig. 4 d); and heterocellular, body ray cells procumbent with one or more row of procumbent cells more high, upright and/or square marginal cells (Fig. 4 e). Perforated ray cells present, rare. Storied structure includes rays, axial parenchyma and vessel elements that are storied or irregularly storied (4c). Mineral inclusions include prismatic crystals in chambered axial parenchyma cells (Fig. 4 c, e), and in procumbent, upright and square ray cells (Fig. 4 e), less often in chambered upright and/or square ray cells. The anatomical quantitative features of the wood are described in Table 1 . Data on wood anatomy for the tribe Brongniartieae is largely scarce. Among the 15 genera and approximately 180 species constituting this tribe, only six genera and merely eight species, about 4%, have any data on wood anatomy. Yet, some species such as A. effusus and Limadendron amazonicum (Ducke) Meireles & A.M.G.Azevedo only have quite incomplete anatomical information (Table 2 ). Table 2 Comparative wood anatomy of Tabaroa caatingicola with eight species from the Brongniartieae tribe for which some wood anatomy data is available. For quantitative data, the classes indicated by the IAWA Committee ( 1989 ) are presented, and when available, the range of variation (minimum – maximum), mean, and/or mean ± standard deviation. (-) Not applicable; (?) Information is missing. The list of anatomical character follows the IAWA Committee ( 1989 ). Anatomical character / Species Tabaroa caatingicola L.P.Queiroz, G.P.Lewis & M.F.Wojc. Amphiodon effusus Huber Behaimia cubensis Griseb. Harpalyce formosa DC. Harpalyce arborescens A.Gray Haplormosia monophylla (Harms) Harms Limadendron amazonicum (Ducke) Meireles & A.M.G.Azevedo Limadendron hostmannii (Benth.) Meireles & A.M.G.Azevedo Poecilanthe parviflora Benth. Growth rings Present Absent Absent Present Present Absent ? Present Present Growth rings: anatomical markers Thick-walled and radially flattened fibres and distended rays, prone to semi-ring-porous wood - - Marginal parenchyma Marginal parenchyma, semi-ring-porous and thick-walled and/or radially flattened latewood fibres - ? Marginal parenchyma Thick-walled and/or radially flattened latewood fibres and marginal parenchyma Porosity Diffuse-porous with a tendency to semi-porous Diffuse-porous Diffuse-porous Diffuse-porous Diffuse-porous with tendency to semi-ring porosity Diffuse-porous ? Diffuse-porous Diffuse-porous Vessel arrangement Vessels in radial pattern No specific arrangement No specific arrangement No specific arrangement No specific arrangement No specific arrangement ? No specific arrangement No specific arrangement Vessel groupings Solitary and radial multiples of 2–5, less common radial multiples of 6–10 ? Vessels in radial multiples of 4 or more common Vessels in radial multiples of 4 or more common Vessels in radial multiples of 4 or more common Solitary and radial multiples of 2–3 ? Solitary and radial multiples of 2–3, less common radial multiples of 4–5 Solitaires and radial multiples of 2, radial multiples of 3–4 Perforation plate Simple ? Simple Simple Simple Simple Simple Simple Simple Intervessel pits arrangement Alternate ? Alternate, polygonal Alternate, polygonal Alternate Alternate, polygonal Alternate Alternate, polygonal Alternate Intervessel pits size Small ? Small Small ? Medium to large ? Medium Small to Medium Vestured pit Present ? Present Present Present Present ? Present Present Vessel–ray pits Similar to intervessel pits in size and shape throughout the ray cell ? Similar to intervessel pits in size and shape throughout the ray cell Similar to intervessel pits in size and shape throughout the ray cell ? Similar to intervessel pits in size and shape throughout the ray cell Similar to intervessel pits in size and shape throughout the ray cell Similar to intervessel pits in size and shape throughout the ray cell Similar to intervessel pits in size and shape throughout the ray cell Mean tangential diameter of vessel (µm) ≤ 50 to 50–100 (44.0–63.1) 50–100 (50–60) 50–100 50–100 (61) 50–100 (61.16 ± 10.90) 100–200 ? 50–200 (100) 50–100 (40–100) Vessels per square millimeter ≥ 100 (119–147) 5–20 (8.3) ? 40–100 (68) 40–100 (45 ± 15; 29–66) ≤ 5 to 5–20 ? 5–20 (10–15) 5–40 (11–38) Mean vessel element length (µm) ≤ 350 (163–197) ? ? <= 350 (208; (173–270)) 350–800 (187.06 ± 20.66) ? ? ? ? Tyloses Absent ? Absent Absent Absent Absent Present Absent Present Deposits in heartwood vessels Present ? Absent Absent Present Present ? Present Present, oleoresin Ground tissue fibres Fibres with simple to minutely bordered pits ? Fibres with simple to minutely bordered pits Fibres with simple to minutely bordered pits Fibres with simple to minutely bordered pits Fibres with simple to minutely bordered pits ? Fibres with simple to minutely bordered pits Fibres with simple to minutely bordered pits Septate and nonseptate fibres Nonseptate fibres ? Nonseptate fibres Nonseptate fibres Nonseptate fibres Nonseptate fibres ? Nonseptate fibres ? Fibre wall thickness Very thick-walled ? Very thick-walled Very thick-walled Thin- to thick-walled and very thick-walled Very thick-walled ? Very thick-walled Very thick-walled Fibre diameter (µm) 13.1–14.0 ? ? 12 13.51 ± 1.77 ? ? ? ? Fibre wall thickness (µm) 4.7–5.4 ? ? 4 3.13 ± 0.74 ? ? ? ? Fibre length (µm) ≤ 900 (709–799) ? ? 900–1600 (947; 799–1021)) ≤ 900 (270.30 ± 43.14) 900–1600 ? ? ? Axial parenchyma Diffuse and paratracheal, both scanty Bands more than three cells wide Diffuse and paratracheal, both scanty and marginal Scanty paratracheal, vasicentric, confluent and marginal Unilateral paratracheal, lozenge-aliform, confluent, bands more than three cells wide and marginal Scanty paratracheal and bands with 2–6 cells wide ? Scanty paratracheal, bands with 2–4 cells wide and marginal Scanty paratracheal and in narrow bands or lines up to three cells wide Axial parenchyma cell type / Strand length Fusiform and two cells, less often 3–4 cells ? 2–4 cells 1–2 cells 2 cells 3–4 cells 2–4 cells 2–4 cells 3–4 cells Ray width 1–3 cells ? 1–3 cells 2 cells 1–3 cells 1–3 cells 1–4 cells 1–3 cells 2–3 cells Rays with multiseriate portion(s) as wide as uniseriate portions Absent ? Absent Absent Absent Absent Present Absent Absent Ray height 14.0–22.6 µm; 6.1–7.2 cells ? 7–10 cells 230 µm 178.68 ± 31.89 µm ? ? ? 120–240 µm Rays: cellular composition All ray cells procumbent and body ray cells procumbent with one or more row of procumbent cells more high, upright and/or square marginal cells ? All ray cells procumbent and body ray cells procumbent with one row of upright and/or square marginal cells Body ray cells procumbent with one row of upright and/or square marginal cells All ray cells procumbent All ray cells procumbent and body ray cells procumbent with one or more row of upright and/or square marginal cells Body ray cells procumbent with one row of upright and/or square marginal cells Body ray cells procumbent with one row of upright and/or square marginal cells Homocellular Sheath cells Absent ? Absent Absent Absent Absent Absent Absent Absent Perforated ray cells Present, rare ? Absent Absent Absent Absent ? Absent Absent Rays per millimeter 9.7–11.4 ? 4–12 4–12 (12) 4–12 (4 ± 1; 2–6) 4–12 to ≥ 12 ? 4–12 (6–7) 4–12 to ≥ 12 (8–14) Storied structure Rays, axial parenchyma and vessel elements ? Rays, axial parenchyma and vessel elements Rays, axial parenchyma and vessel elements Rays, axial parenchyma and vessel elements Rays, axial parenchyma and vessel elements Absent Absent Rays, axial parenchyma and vessel elements Mineral inclusions Prismatic crystals in chambered axial parenchyma cells, and in procumbent, upright and square ray cells, less often in chambered upright and/or square ray cells. ? Prismatic crystals in upright and/or square ray cells and in chambered axial parenchyma cells Prismatic crystals in upright, square and procumbent ray cells and in chambered axial parenchyma cells Absent Prismatic crystals in chambered axial parenchyma cells ? Prismatic crystals in procumbent ray cells Absent Reference This paper Tsuchiya et al. ( 2002 ) Gasson et al. ( 2004 ) Bonilla et al. ( 2004 ) Ramírez-Martíne et al. (2017) Loupe et al. (2008) InsideWood ( 2004 -onwards) Detienne and Jacquet (1983) Manieri and Chimelo (1989) 4 Discussion External characteristics of the bark, as well as the general, organoleptic, and macroscopic anatomical features of wood, can assist regulatory bodies in wood identification and in combating illegal exploitation and trade. This information can thereby contribute to the conservation efforts for this species (Silva et al. 2022b ). 4.1 Systematic wood anatomy – The wood anatomy of Tabaroa caatingicola differs from other Brongniartieae genera by having more than 100 vessels.mm − 2 , the vessels arranged in a radial pattern and perforated ray cells. Although comparing T. caatingicola with its closest genus, Amphiodon , is impeded by limited anatomical data for the latter, distinctions arise in growth rings, porosity, and axial parenchyma type: Tabaroa exhibits growth rings, semi-porous wood, and diffuse and scanty paratracheal, while Amphiodon lacks growth rings, displays diffuse-porous wood, with paratracheal parenchyma bands more than three cells wide (Tsuchiya et al. 2002 ). With the current data, no anatomical feature of the wood is exclusive to these two sister genera. A comprehensive wood anatomical description is advised for Amphiodon effusus . Most wood anatomical features observed in Tabaroa and all other Brongniartieae genera are also widely observed in woody angiosperms: wood diffuse-porous (some tend to semi-porosity), vessels multiples, simple perforation plates, intervessel pits alternate, vessel–ray pits with distinct borders similar to intervessel pits in size and shape, libriform fibres with very thick-walled and non-septate, 4–12 rays.mm − 2 and gums and other deposits in heartwood vessels (except Behaimia ) (Carlquist 2001 ; Wheeler and Baas 1991 ; Wheeler et al. 2007 ); or in Leguminosae wood: vestured pits, 1–4 cells per parenchyma strand, and all rays, axial parenchyma and vessel elements storied (except Limadendron ) (Carlquist 2001 ; Gasson 1996 ; Gasson 1999 ; Höhn 1999 ; Pernía and Melandri 2006 ; Ramanantsialonina et al. 2022 ; Stepanova et al. 2013 ). Hence, considering the limited data available among species of the Brongniartieae tribe, no anatomical trait can be inferred as synapomorphic of this clade. However, the composition of ray cells, diffuse and/or paratracheal scanty axial parenchyma (except Amphiodon effusus and Harpalyce arborescens ), and the presence of prismatic crystals (except H. arborescens and Poecilanthe parviflora ) appear to be very characteristic of the tribe. The anatomical markers semi-ring-porous wood and thick-walled and radially flattened fibres are reported for Harpalyce arborescens A.Gray (Loupe et al. 2008), the latter also being described for Poecilanthe parviflora Benth. (Manieri and Chimelo 1989). The other Brongniartieae species either lack growth rings or exhibit marginal parenchyma, a more common anatomical marker in Leguminosae, also associated with deciduous species (Carlquist, 2001 ; Höhn 1999 ; Pernía and Melandri 2006 ; Silva 2006 ; Stepanova et al. 2013 ; Worbes and Fichtler 2010), although it does not occur in Tabaroa . Tabaroa caatingicola shares with its phylogenetically related genus Harpalyce tendency to semi-ring-porous wood and short vessel element length (≤ 350 µm) (Bonilla et al. 2004 ; Ramírez-Martíne et al. 2017). Other anatomical similarities can be noted among the other Brongniartieae genera. For example, marginal parenchyma is characteristic of Harpalyce , Limadendron , and Poecilanthe (Bonilla et al. 2004 ; Detienne and Jacquet 1983; Manieri and Chimelo 1989; Ramírez-Martínez et al. 2017 ), being a type of parenchyma widely distributed across Leguminosae species (Carlquist 2001 ; Gasson 1996 ; Gasson 1999 ; Höhn 1999 ; Pernía and Melandri 2006 ; Ramanantsialonina et al. 2022 ; Stepanova et al. 2013 ). The polygonal intervessel pits are characteristic of Behaimia , Haplormosia , Harpalyce , and Limadendron (Bonilla et al. 2004 ; Detienne and Jacquet 1983; Gasson et al. 2004 ; Loupe et al. 2008). Tyloses is present in Limadendron and Poecilanthe (InsideWood 2004 -onwards; Manieri and Chimelo 1989), it goes against the data which indicate that species with vessel–ray pits similar to intervessel pits, normally with small diameter, have a low probability of having tyloses (Bonsen and Kučera 1990 ; De Micco et al. 2016 ). The absence of deposits in heartwood vessels is indicated for Behaimia and Harpalyce (Bonilla et al. 2004 ; Gasson et al. 2004 ). Rays with multiseriate portions as wide as uniseriate portions are limiting to Limadendron amazonicum (InsideWood 2004 -onwards). 4.2 Functional and ecological wood anatomy – Growth rings are present in the wood of T. caatingicola and are identified by three distinct anatomical markers: thick-walled and radially flattened fibres and distended rays, prone to semi-ring-porous wood. Anatomical markers can be characterized as histological contrasts arising from traits that vary between cells produced throughout a growing season, giving rise to early- and latewood (Silva 2023 ). Each of these variations, whether in isolation or combination, generates identifiable features in the secondary xylem, recurring across multiple growing seasons. Currently, growth rings are conceived not only as a result of the vascular cambium’s ability to respond to environmental and physiological changes but also to effect changes in the secondary xylem with significant functional implications (Carlquist 2000 ; Silva et al. 2019 , 2021 ; Silva 2023 ). Fibers with a smaller lumen and thicker walls are hypothesized to enhance the security of transport in vessel elements, representing the most widespread anatomical marker among angiosperms, particularly in tropical regions (Jacobsen et al. 2005 ; Silva et al. 2021 ). Semi-ring-porous wood is more commonly found in species distributed in subtropical and temperate regions (Silva et al. 2021 ). However, it is also present in tropical species, particularly in more seasonal habitats, like the Caatinga dry woodlands, where gradations between diffuse and semi-ring porosity can be observed (Aragão and Lisi 2019 ; Gasson et al. 2017 ; Silva 2006 ). This anatomical marker shows varying degrees of specialization in balancing the trade-off between embolism protection and efficient conduction. Broad vessels in the earlywood facilitate conduction capacity with a lower risk of embolism during periods of ample water availability. In contrast, narrower vessels in conditions of water scarcity, although less efficient for transport, ensure continuous xylem flow while minimizing the risk of embolism formation and the subsequent disruption of conduction (Baas et al. 2004 ; Sperry et al. 2006 ; Tyree and Zimmermann 2013 ). Distended rays, on the other hand, adapt to adjacent cells, preventing the formation of large intercellular spaces and, consequently, cracks in the secondary xylem (Silva et al. 2021 ; Silva 2023 ). The seasonality and irregularity of rainfall in the Caatinga explain the formation of growth rings with an annual periodicity (Mattos et al. 2015 ; Nogueira Jr et al. 2018; Silva 2006 ) and infra-annual, potentially forming multiple annual ring ( sensu Silva et al. 2019 ; Pagotto et al. 2015 ). The latter periodicity is also recognized for dry forests in East Africa (Gourlay 1995a , b ; Jacoby 1989 ). The growth rings of T. caatingicola should be studied to determine the periodicity. Given that T. caatingicola is ecologically confined to a semi-arid region, the vessel characteristics outlined here — including a degree of semi-porosity, predominantly multiple vessels, small diameter, high number of vessels per square millimeter, short vessel element length, simple perforation plates, and vestured pits — are consistent with the well-established trade-off between safety × efficiency in conduction. This trade-off is widely recognized as the primary factor influencing the evolution of secondary xylem in seed plants (Baas et al. 2004 ). While the mechanisms of freeze-induced embolism are better understood, drought-induced embolism is a theme that has gained strength in recent years (Hacke et al. 2023 ; Olson 2022 ). Failures in sap transport through the xylem are among the main causes of death or reduced productivity in plants subjected to drought (Anfodillo and Olson 2021 ; Breshears et al. 2005 , 2018 ; Brodribb and Cochard 2009 ; McDowell et al. 2018 ). So, cavitation vulnerability is a relevant hydraulic trait that determines a species’ competitiveness under water stress (Islam et al. 2018 ). Wider vessels are at a higher risk of embolism than narrower vessels within a tissue and when we compare species. Typically, wider vessels tend to be more vulnerable to drought-induced embolism, whereas embolism-resistant vessels are generally narrow (Hacke et al. 2023 ). This explanation justifies the small mean vessel diameter, as well as the slightly semi-ring porosity observed in T. caatingicola . The vessel diameter influences the variation in vessels per square millimeter. The inverse correlation between diameter and vessel frequency has been previously reported and is widely supported in the literature (Chave et al. 2009 ; Sperry et al. 2008 ). The high percentage of multiple vessels, ranging from 71–80%, stands out in the wood of T. caatingicola . The performance of different levels of vessel grouping in secondary xylem is a controversial subject in the literature since Carlquist ( 1984 ). Some authors argue that multiple vessels increase the chances of propagating cavitation of air-seeding (Jacobsen et al. 2007 ; Tyree and Zimmermann 2013 ), while others suggest that a higher connection between vessels improves hydraulic integration, providing alternative pathways when embolism blocks flow in one vessel, reducing the potential loss of water transport capacity associated with cavitation (Loepfe et al. 2007 ; Trifilò et al. 2014 ). Despite many gaps in understanding the functioning of different vessel grouping levels, the latter argument is gaining strength by providing data that cavitation can be more easily removed in grouped vessels than in solitary ones (Hölttä et al. 2009 ), and that multiple vessels are more resistant to cavitation (Lens et al. 2011 ). In any case, T. caatingicola is part of the species in dry forests that exhibit greater vessel connectivity (Apgaua et al. 2022 ; Scholz et al. 2014 ). The considerably short vessel elements described for the wood of T. caatingicola also have a functional relationship with the water-limited, arid environment in which this species grows. Carlquist ( 1975 ) hypothesized that shorter vessel elements have adaptive value, resisting mechanical deformation under negative pressure. Recently, a comparative analysis of over 1000 species confirmed Carlquist’s prediction, indicating that shorter vessel elements affect vessel resistance to deformation. Species with exceptionally short vessel elements tend to grow in arid lands, while those with exceptionally long vessel elements tend to thrive in humid climates (Echeverría et al. 2023 ). Simple perforation plates are widely found in angiosperms, prevailing in species distributed across the tropics. Simple perforation plates provide considerably less resistance to sap flow between vessel elements, being crucial, when compared to compound perforation plates, for high flow demands (Carlquist 2001 ; Wheeler and Baas 1991 ; Wheeler et al. 2007 ), notably for plants growing in environments with scarce and irregular precipitation such as the Caatinga. The vestured pits described for the vessel elements of T. caatingicola are an anatomical trait of wood with systematic and functional value. They are generally associated with simple perforation plates, bordered pits, and dry environments, characterizing taxa such as the orders Myrtales and Gentianales, and families such as Malpighiaceae and Leguminosae (Jansen et al. 2001 , 2008 ). Although vestured pits are related to safety in hydraulic conduction, reducing vulnerability to cavitation or assisting in embolism repair, the mechanisms associated with these functionalities remain poorly understood (Jansen et al. 2008 ; Rabaey et al. 2010 ). The presence of highly lignified structures within the pit chamber can influence hydraulic resistance. Vestured pits appear to facilitate embolism reversal in xeric and warmer regions with high transpiration rates (Jansen et al. 2003 ), like the Caatinga seasonally dry vegetation. The small pit apertures recorded for the vessel elements of T. caatingicola are associated with the small diameters of the vessel elements, as this trait covaries with vessel element diameter (Silva et al. 2022a ). The xylem of T. caatingicola is considered to have a high degree of xeromorphism based on the vulnerability and mesomorphy index values. Carlquist ( 1977 ) considered vulnerability index values significantly below 1.0, as observed here, to indicate a high degree of xeromorphy. Meanwhile, mesomorphy index values above 200 indicate mesophytic species. For these indices, higher values are associated with greater efficiency, while lower values are linked to greater safety (Carlquist 1977 ). The vulnerability and mesomorphy indices proposed by Carlquist ( 1977 ) help quantify the degree of mesomorphy or xeromorphy exhibited in the plant xylem, contributing to a quantitative approach to plant anatomy, particularly in the fields of functional and ecological anatomy (Ewers et al. 2023 ). Despite of the few data available for comparison, the other species of the Brongniartieae tribe exhibit a series of wood anatomy traits common to T. caatingicola . However, most of these traits are widely found in angiosperms and/or in Leguminosae. Tabaroa caatingicola has a set of wood anatomy traits that distinguishes it from other Brongniartieae genera. The wood anatomy of T. caatingicola reveals a set of adaptations to the SDTFW biome of the Brazilian Caatinga domain, where the low precipitation with irregular rains, high temperatures, and high transpiration potential increases the likelihood of drought-induced embolism formation (Hacke et al. 2023 ; Olson et al. 2023 ). These adaptations include semi-ring-porous wood with distinct growth ring boundaries, multiple narrow vessels, simple perforation plates, small and vestured pits. These traits increase flow during periods of higher water availability and ensure hydraulic safety during scarcity, while also providing mechanisms to minimize the formation and spread of embolisms. Declarations Author contributions The research project was initiated, and the study designed by M.S.S. and D.C. M.S.S. and D.C. conducted fieldwork and prepared the figures. M.S., L.B.S. and D.B. did the laboratory work and anatomical analyses. C.S. performed the SEM analyses. All authors contributed to the writing of the manuscript as well as approving the final submitted version of the paper. Statements and Declarations Conflict of interests: The authors declare that the manuscript does not present any kind of conflict of interests. The purpose and content of the work are original and not previously published. The published data are of non-financial interest. References Aguiar TV, Sant’anna-Santos BF, Azevedo AA, Ferreira RS (2007) ANATI QUANTI: software de análises quantitativas para estudos em anatomia vegetal. Planta Daninha 25:649–659. https://doi.org/10.1590/S0100-83582007000400001 Anfodillo T, Olson ME (2021) Tree mortality: testing the link between drought, embolism vulnerability, and xylem conduit diameter remains a priority. 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Acta Amazon 32:241–241. https://doi.org/10.1590/1809-43922002322256 Tyree MT, Zimmermann MH (2013) Xylem structure and the ascent of sap. Springer Science & Business Media, Berlin WFO (2024) World Flora Online. Published on the Internet; http://www.worldfloraonline.org . Accessed on 30 Jan 2024' Wheeler EA, Baas P (1991) A survey of the fossil record for dicotiledonous wood and its significance for evolutionary and ecological wood anatomy. IAWA J 12:275–318. https://doi.org/10.1163/22941932-90001256 Wheeler EA, Baas P, Rodgers S (2007) Variations in dicot wood anatomy: a global analysis based on the Insidewood database. IAWA J 28:229–258. https://doi.org/10.1163/22941932-90001638 Wickham H (2016) ggplot2 - elegant graphics for data analysis. Springer-, New York, USA Worbes M, Fichtler E (2010) Wood anatomy and tree–ring structure and their importance for tropical dendrochronology. In: Junk WJ et al (eds) Amazonian Floodplain Forests: Ecophysiology, Biodiversity and Sustainable Management. Springer, Berlin, pp 329–346 Zizka A, Silvestro D, Andermann T et al (2019) CoordinateCleaner: Standardized cleaning of occurrence records from biological collection databases. Methods Ecol Evol 10:744–751. https://doi.org/10.1111/2041-210X.13152 Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Minor revisions 26 Apr, 2024 Reviewers agreed at journal 28 Feb, 2024 Reviewers invited by journal 28 Feb, 2024 Editor invited by journal 26 Feb, 2024 Editor assigned by journal 24 Feb, 2024 First submitted to journal 22 Feb, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3921723","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":275683796,"identity":"0e9f39bb-7338-477a-8918-2ed00b72ca52","order_by":0,"name":"Marcelo dos Santos Silva","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2ElEQVRIiWNgGAWjYNCCAiBm7wEzefiI02IAUnuGgeEAkGIjXotEDlgLA0Etuu1nH374YFBnzz/z7cHHH3PsZNgYmB8+uoFHi9mZdGPJGQaHE2fczks2OLgtGegwNmPjHHxaDqSxMfMYHEhguJ1jJnFwGzNQCw+bNF4t55+xMf8BOkz+5hmQlnoitNwA2sJgwMy44QYPSMthYrQ8Y5bsAfpl45kcY4Oz247zsDET8sv5NMYPPyrq7OWOnzF8ULmt2p6fvfnhY3xasABm0pSPglEwCkbBKMACAIimRQQ//gr+AAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-4982-8808","institution":"Universidade Federal de Pelotas","correspondingAuthor":true,"prefix":"","firstName":"Marcelo","middleName":"dos Santos","lastName":"Silva","suffix":""},{"id":275683797,"identity":"db177395-8ac3-4e8e-95fb-39af5fbbb833","order_by":1,"name":"Daisy Burris","email":"","orcid":"","institution":"Imperial College London","correspondingAuthor":false,"prefix":"","firstName":"Daisy","middleName":"","lastName":"Burris","suffix":""},{"id":275683798,"identity":"785d9268-94b7-4bb3-98f2-516ccba23f66","order_by":2,"name":"Cássia Sacramento","email":"","orcid":"","institution":"Universidade Federal da Bahia","correspondingAuthor":false,"prefix":"","firstName":"Cássia","middleName":"","lastName":"Sacramento","suffix":""},{"id":275683799,"identity":"3a2f92f3-d8b6-411b-bf7c-8a2929da2d8f","order_by":3,"name":"Lazaro Benedito da Silva","email":"","orcid":"","institution":"Universidade Federal da Bahia","correspondingAuthor":false,"prefix":"","firstName":"Lazaro","middleName":"Benedito da","lastName":"Silva","suffix":""},{"id":275683800,"identity":"7a28f676-be47-4a3b-b46d-6c6d31ed0421","order_by":4,"name":"Domingos Cardoso","email":"","orcid":"","institution":"Universidade Federal da Bahia","correspondingAuthor":false,"prefix":"","firstName":"Domingos","middleName":"","lastName":"Cardoso","suffix":""}],"badges":[],"createdAt":"2024-02-02 17:20:24","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3921723/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3921723/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":52045344,"identity":"1b44e92f-fdf9-4c07-b1f5-338ed5a6985c","added_by":"auto","created_at":"2024-03-05 19:36:24","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":272197,"visible":true,"origin":"","legend":"\u003cp\u003eBayesian majority-rule consensus tree based on \u003cem\u003ematK\u003c/em\u003e sequence data (from Cardoso et al. 2017) showing the relationship of the tribe Brongniartieae with respect to the main lineages of the early branching Genistoid clade of papilionoid legumes (circular cladogram on the lower left), and a summary of the genus-level Brongniartieae phylogeny showing the placement of \u003cem\u003eTabaroa\u003c/em\u003e (cladogram on the right). Photo of the \u003cem\u003eTabaroa\u003c/em\u003e flowers by Luciano Paganucci de Queiroz.\u003c/p\u003e","description":"","filename":"Fig1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3921723/v1/fab70ba4bfebd794ebdcd5bd.jpg"},{"id":52045346,"identity":"a6ab522f-0abb-4734-acda-415a2805137b","added_by":"auto","created_at":"2024-03-05 19:36:25","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1506719,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of the phylogenetically closely related monotypic genera \u003cem\u003eTabaroa\u003c/em\u003e (focus of the current anatomical characterization) and \u003cem\u003eAmphiodon\u003c/em\u003e across geographic and climatic spaces. a) The geographical distribution of \u003cem\u003eTabaroa\u003c/em\u003eis narrowly restricted to areas with low annual precipitation in the Caatinga of the Brazilian Northeast, whereas \u003cem\u003eAmphodion\u003c/em\u003e distribution spans a wide range across wetter areas of the Amazon. b) Climate space of \u003cem\u003eTabaroa\u003c/em\u003e and \u003cem\u003eAmphiodon\u003c/em\u003e plotted using the WorldClim-derived bioclimatic variable BIO12 and BIO4 (Fick and Hijmans 2017).\u003c/p\u003e","description":"","filename":"Fig2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3921723/v1/dfbd0519fba1d3dbd002d3ce.jpg"},{"id":52045347,"identity":"f1548d9c-0573-40c8-888a-d33715cc3364","added_by":"auto","created_at":"2024-03-05 19:36:25","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1497073,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eTabaroa caatingicola\u003c/em\u003e (Leguminosae). \u003cstrong\u003ea:\u003c/strong\u003e \u003cem\u003eT. caatingicola\u003c/em\u003e, most prominent tree in the center of the image, in its natural habitat during the rainy season. \u003cstrong\u003eb:\u003c/strong\u003e Trunk showing the characteristic ‘crocodile skin’ bark pattern. \u003cstrong\u003ec-d:\u003c/strong\u003e Transversely cut stem showing the brown heartwood and yellowish sapwood, in addition to the growth rings. Scale bar – c: 1.0 cm; d: 0.5 cm. Photos: Marcelo dos S. Silva.\u003c/p\u003e","description":"","filename":"Fig3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3921723/v1/171b15b89f9fa1d6627a49ea.jpg"},{"id":52045345,"identity":"d7ec793f-51e4-4382-88fc-748e766e3a45","added_by":"auto","created_at":"2024-03-05 19:36:25","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":569338,"visible":true,"origin":"","legend":"\u003cp\u003eWood anatomy of the papilionoid legume \u003cem\u003eTabaroa caatingicola\u003c/em\u003e. \u003cstrong\u003ea – e:\u003c/strong\u003e Light microscopy. \u003cstrong\u003ef – g:\u003c/strong\u003e Scanning electron microscope. \u003cstrong\u003ea – b:\u003c/strong\u003e Cross section, black arrowhead indicates the limit of the growth rings. \u003cstrong\u003ec:\u003c/strong\u003e Tangential section, white arrowhead indicates prismatic crystals in chambered axial parenchyma cells. \u003cstrong\u003ed – e:\u003c/strong\u003e Radial section, white arrowhead indicates prismatic crystals in chambered axial parenchyma cells; white arrow identifies prismatic crystals in procumbent ray cells and black arrow indicates prismatic crystals in upright ray cells. \u003cstrong\u003ef:\u003c/strong\u003e Radial section showing a detail of vessel elements with simple perforation plate. \u003cstrong\u003eg:\u003c/strong\u003eDetail of a vessel element in tangential view showing intervessel pits alternate and vestured. Scale bar – a: 300 mm; b – c: 200 mm; d – e: 100 mm; f – g: 10 mm.\u003c/p\u003e","description":"","filename":"Fig4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3921723/v1/0e7def8dae63cef6a052aa5c.jpg"},{"id":52045575,"identity":"a3a0acc2-4279-43f2-8198-1cfe1092f951","added_by":"auto","created_at":"2024-03-05 19:44:26","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1487939,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3921723/v1/311dba47-0820-43c8-a117-a892e2fbe7d9.pdf"}],"financialInterests":"","formattedTitle":"Wood anatomy of Tabaroa, a monotypic papilionoid legume genus narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests","fulltext":[{"header":"1 Introduction","content":"\u003cp\u003eWood anatomical features have consistently been recognized to provide valuable sources of phylogenetic information (Baas et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Carlquist \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Herendeen and Miller \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Olson \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), as well as a model of how characters more plastic have evolved over time in plants, through ancestral character reconstructions in well-resolved phylogenies (\u003cem\u003ee.g.\u003c/em\u003e Pace et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Silva et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Wood anatomical features are able to shed light on obscure relationships at deeper nodes (Lens et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). However, the evolution of wood anatomy is still poorly unexplored within a phylogenetic context, perhaps due to the large knowledge gaps of microscopic wood structures across clades (Lens et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWood anatomy has been instrumental for delimitating several flowering plant families including the Marcgraviaceae, Tetrameristaceae (Lens et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), and Sapotaceae (Kukachka \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). Likewise, the wood anatomy of the ecologically and economically important family Leguminosae is relatively well documented, when compared with other angiosperm families. (Gasson \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Gasson et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Stepanova et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). However, most tribes within the Papilionoideae are yet to be anatomically studied. Describing wood anatomical features of endemic, rare, monospecific or poorly diverse genera is therefore critical for understanding evolutionary patterns of the micromorphological diversity across legumes. This is particularly critical in understanding how tree species can behave in the face of climate change, especially in relation to temperature and precipitation variations that directly affect xylem transport and the survival capacity of plants (Anfodillo and Olson \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Breshears et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Fontes et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Hajek et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2016\u003c/span\u003e)\u003c/p\u003e \u003cp\u003eThe papilionoid legume genus \u003cem\u003eTabaroa\u003c/em\u003e and its only known species \u003cem\u003eT. caatingicola\u003c/em\u003e L.P.Queiroz, G.P.Lewis \u0026amp; M.F.Wojc. of the tribe Brongniartieae (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) have been described more than ten years ago (Queiroz et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), yet wood anatomy in this phylogenetically important branch of the tribe remains fully unknown. \u003cem\u003eT. caatingicola\u003c/em\u003e is narrowly endemic to a small area of seasonally dry tropical forest and woodland (SDTFW) on sandy soils of the Brazilian Caatinga domain, in southwestern Bahia (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea), whereas its most closely related species \u003cem\u003eAmphiodon effusus\u003c/em\u003e Huber is ecologically confined to the Amazon rainforest (Cardoso et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This clade, in turn, is sister to the genus \u003cem\u003eHarpalyce\u003c/em\u003e, resulting in the topology (\u003cem\u003eHarpalyce\u003c/em\u003e (\u003cem\u003eTabaroa\u003c/em\u003e, \u003cem\u003eAmphiodon\u003c/em\u003e)) that has been corroborated in several publications involving phylogenetic analyses of nuclear and plastid DNA sequence data (Cardoso et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2013\u003c/span\u003e, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Meireles et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Queiroz et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Brongniartieae is a taxonomically intriguing tribe within the Papilionoideae subfamily of the Leguminosae, comprising 15 genera, mostly monospecific or with few species, totalling approximately 180 species (POWO 2024; WFO \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Despite the modest number of species, the Brongniartieae tribe displays significant morphological diversity, and a wide geographic distribution across continents and biomes, with marked variation in ecological preferences. For this reason, many of its morphologically disparate genera were previously placed in at least four distantly related tribes (Cardoso et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Queiroz et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eCaatinga is an indigenous \u003cem\u003eTupi\u003c/em\u003e word that means \u0026ldquo;white forest\u0026rdquo;, describing the grey and light aspect during the dry season, when the majority of trees and shrubs are devoid of leaves, allowing light to reach the ground, consists predominantly of xeric shrub, thorn and drought resistant species, dominated by legume trees (Giulietti et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Queiroz \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The region is characterized by an extended dry season and erratic rainfall, resulting in sparse foliage and undergrowth (Leal et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). It harbours numerous endemic species, with 34% of the flora exclusive to the area, many featuring unique physiological adaptations to mitigate water loss during prolonged drought periods (Giulietti et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough relatively common locally, \u003cem\u003eT. caatingicola\u003c/em\u003e only occupies a restricted geographic area of about 12 Km\u003csup\u003e2\u003c/sup\u003e and has been classified as Critically Endangered (IUCN 2001). Here we newly present wood anatomical description for the ecologically distinctive, phylogenetically isolated, and highly threatened species \u003cem\u003eT. caatingicola\u003c/em\u003e and compares it with six phylogenetically related genera within the Brongniartieae, for which any wood anatomical data are available: \u003cem\u003eAmphiodon\u003c/em\u003e, \u003cem\u003eBehaimia\u003c/em\u003e, \u003cem\u003eHaplormosia\u003c/em\u003e, \u003cem\u003eHarpalyce\u003c/em\u003e, \u003cem\u003eLimadendron\u003c/em\u003e, and \u003cem\u003ePoecilanthe\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Furthermore, we analyze the wood anatomy of \u003cem\u003eT. caatingicola\u003c/em\u003e in light of the most modern theories and functional hypotheses of secondary xylem.\u003c/p\u003e"},{"header":"2 Material and methods","content":"\u003cp\u003e \u003cb\u003e2.1 Wood sampling and anatomical characterization \u0026ndash;\u003c/b\u003e Wood samples from three trees were collected from the trunk at diameter at breast height (DBH\u0026thinsp;=\u0026thinsp;1.30 m), in trees with apparent health and straight stem in a population of \u003cem\u003eTabaroa caatingicola\u003c/em\u003e in the municipality of Dom Bas\u0026iacute;lio, state of Bahia, northeastern Brazil (S 13\u0026ordm;47'19\"; W 41\u0026ordm;30'04\") (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea). Non-destructive wood collection followed the procedure described by Silva et al. (\u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e). Samples were vouched, recorded, and deposited in the Herbarium and Xylotheque of the Instituto de Biologia of the Universidade Federal de Pelotas (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The climate from the collection area is dry, semi-arid of the low latitude and altitude (BSh), according to K\u0026ouml;ppen-Geiger\u0026rsquo;s classifications, with annual mean precipitation between 588\u0026ndash;619 mm, and temperature 23.3\u0026ndash;24.0 \u0026ordm;C, with great thermal amplitude throughout the year, 14.7\u0026ndash;31.8 \u0026ordm;C (Fick and Hijmans \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The main biome in the Caatinga phytogeographical domain is the seasonally dry tropical forest and woodland (Queiroz et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), which can be characterized as a xerophytic vegetation across most of the semi-arid region of Northeastern Brazil, including mostly sparse vegetation that covers massifs and plateaus where rivers are usually seasonal. Leguminosae species are among the most dominating flowering plants of the Caatinga (Queiroz \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Queiroz et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Despite comprising almost 10% of Brazil's territory, the Caatinga is inadequately explored, with only 1% designated as a Conservation Protection Area. The region faces threats from deforestation and unsustainable agricultural practices, such as agriculture and cattle ranching, leading to soil salinization (Leal et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Silva et al. 2004).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eDiameter at breast height, height and quantitative data on the wood anatomy of \u003cem\u003eTabaroa caatingicola\u003c/em\u003e (Leguminosae) for the three tree samples analyzed, recorded, and deposited in the xylotheque of the Instituto de Biologia of the Universidade Federal de Pelotas (PELw). Additionally, vulnerability and mesomorphy indices (sensu Carlquist \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1977\u003c/span\u003e) are provided. Herbarium voucher: PEL 27325.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMorphological and anatomical parameters / Samples\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePELw 01\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePELw 02\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePELw 03\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDiameter at breast height (1.30 m) (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHeight (m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVessels per square millimeter\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e119\u0026thinsp;\u0026plusmn;\u0026thinsp;32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e129\u0026thinsp;\u0026plusmn;\u0026thinsp;11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e147\u0026thinsp;\u0026plusmn;\u0026thinsp;17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVessel tangential diameter (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e44.0\u0026thinsp;\u0026plusmn;\u0026thinsp;23.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e49.5\u0026thinsp;\u0026plusmn;\u0026thinsp;16.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e63.1\u0026thinsp;\u0026plusmn;\u0026thinsp;20.9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVessel element length (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e163\u0026thinsp;\u0026plusmn;\u0026thinsp;64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e180\u0026thinsp;\u0026plusmn;\u0026thinsp;31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e197\u0026thinsp;\u0026plusmn;\u0026thinsp;26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eIntervessel/vessel-ray pit size (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFibre length (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e727\u0026thinsp;\u0026plusmn;\u0026thinsp;255\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e709\u0026thinsp;\u0026plusmn;\u0026thinsp;140\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e799\u0026thinsp;\u0026plusmn;\u0026thinsp;128\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFibre diameter (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e13.4\u0026thinsp;\u0026plusmn;\u0026thinsp;3.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e13.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14.0\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFibre wall thickness (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFibre pit size (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRays/mm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRay width (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.0\u0026thinsp;\u0026plusmn;\u0026thinsp;4.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18.2\u0026thinsp;\u0026plusmn;\u0026thinsp;3.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e22.6\u0026thinsp;\u0026plusmn;\u0026thinsp;4.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRay width (cell numbers)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRay height (\u0026micro;m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e126\u0026thinsp;\u0026plusmn;\u0026thinsp;22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e117\u0026thinsp;\u0026plusmn;\u0026thinsp;15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e129\u0026thinsp;\u0026plusmn;\u0026thinsp;18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRay height (cell numbers)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVulnerability index\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMesomorphy index\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e68.2\u0026thinsp;\u0026plusmn;\u0026thinsp;51.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e68.9\u0026thinsp;\u0026plusmn;\u0026thinsp;25.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e83.1\u0026thinsp;\u0026plusmn;\u0026thinsp;27.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe preparation of histological slides followed the usual plant anatomical methods described by Johansen (\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1940\u003c/span\u003e) and Sass (\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1951\u003c/span\u003e). Histological sections between 18\u0026ndash;30 \u0026micro;m in thickness were made using a Leica\u0026copy; sliding microtome. Sections of each sample were clarified with sodium hypochlorite (50%), coloured with 1% alcoholic safranin, dehydrated in a 50\u0026ndash;100% alcoholic series and mounted in synthetic Canada Balsam\u0026copy; or Entellan\u0026copy;. The method proposed by Franklin (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1945\u003c/span\u003e), modified by Kraus and Arduin (\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), was followed to analyze dissociated cell elements. For the Scanning Electron Microscopy (SEM) analyses, longitudinal wood sections, 18\u0026ndash;30 \u0026micro;m thick were dehydrated in an alcoholic series, dried in a dry chamber (60\u0026deg;C\u0026thinsp;~\u0026thinsp;12hs), placed on a stub with a double-faced carbon label and metallized with gold. The observation of the samples was carried out with a SEM JEOL 6390LV\u0026copy; in the Instituto Gon\u0026ccedil;alo Moniz (Funda\u0026ccedil;\u0026atilde;o Oswaldo Cruz \u0026ndash; FIOCRUZ).\u003c/p\u003e \u003cp\u003eThe following quantitative wood anatomy characters were measured: vessels (vessels per square millimeter, tangential diameter, length, intervessel/vessel-ray pit outer aperture diameter); fibres (length, width, and wall thickness); and rays (rays/mm, width, and height). Indices of vulnerability (vessel diameter / vessels per square millimeter) and mesomorphy (indices of vulnerability \u0026times; vessel element length) were calculated according to Carlquist (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1977\u003c/span\u003e). The measurements of vessels per square millimeter were obtained with ANATI QUANTI\u0026copy; software (Aguiar et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The other measurements were carried out with an Olympus CX40\u0026copy; microscope coupled with a micrometric lens and the factors obtained were converted to \u0026micro;m through a conversion factor. The terminology used in the anatomical descriptions is in accordance with the IAWA Committee (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1989\u003c/span\u003e). The measurement of the anatomical parameters n\u0026thinsp;=\u0026thinsp;30 was fixed.\u003c/p\u003e \u003cp\u003e \u003cb\u003e2.2 Mapping distribution across geographic and climatic spaces \u0026ndash;\u003c/b\u003e To assess the distribution range across geographic and climatic spaces, we used built a taxonomically verified specimen record data from the analyses of herbarium collections (ALCB, CEPEC, CEN, HUEFS, MO, MBM, NY, RB, RON, SP, SPF, US); acronyms according to Thiers \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) as available in the online databases of speciesLink (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://specieslink.net/\u003c/span\u003e\u003cspan address=\"https://specieslink.net/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and Reflora (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://reflora.jbrj.gov.br/reflora/PrincipalUC/PrincipalUC.do\u003c/span\u003e\u003cspan address=\"https://reflora.jbrj.gov.br/reflora/PrincipalUC/PrincipalUC.do\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). Any erroneous georeferenced records were filtered with the R package coordinateCleaner (Zizka et al. \u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), but whenever possible these specimens and others without geographic coordinates were included based on the closest georeferenced locality accessed by comparing with localities of other plant specimens in the speciesLink database.\u003c/p\u003e \u003cp\u003eFrom the latitude and longitude of each herbarium collection (a total of 153 specimens recorded, Appendix S1), the variables BIO12 (Annual Precipitation) and BIO4 (Temperature Seasonality) were extracted from the WorldClim v.2.0 model layers (WGS84 projection; Fick and Hijmans \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) using the R library \u003cem\u003eraster\u003c/em\u003e (Hijmans \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Bioclimatic variables (O\u0026rsquo;Donnell and Ignizio \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) were derived from these climate models using the \u003cem\u003eextract\u003c/em\u003e function in the \u003cem\u003eraster\u003c/em\u003e library. We mapped the range distribution of \u003cem\u003eTabaroa\u003c/em\u003e and its most phylogenetically closely related genus \u003cem\u003eAmphiodon\u003c/em\u003e against the BIO12 bioclimatic variable, by using the R packages \u003cem\u003eraster\u003c/em\u003e, \u003cem\u003eggmap\u003c/em\u003e (Kahle and Wickham \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), \u003cem\u003eggplot2\u003c/em\u003e (Wickham et al. 2016), \u003cem\u003eggspatial\u003c/em\u003e (Dunnington \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and \u003cem\u003ernaturalearth\u003c/em\u003e (South \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) as implemented in RStudio (\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The built a scatterplot to show the bioclimatic space of \u003cem\u003eTabaroa\u003c/em\u003e and \u003cem\u003eAmphiodon\u003c/em\u003e across the BIO12 and BIO4 axes.\u003c/p\u003e"},{"header":"3 Results","content":"\u003cp\u003eHere, we provide a full description of the wood anatomy of \u003cem\u003eTabaroa caatingicola\u003c/em\u003e. This species exhibits \u0026ldquo;bark light grey with darker wavy stripes that interlink and cross over to form a pattern suggestive of crocodile skin, inner bark dark green\u0026rdquo; (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb), hence the vernacular name \u003cem\u003epau-jacar\u0026eacute;\u003c/em\u003e (crocodile wood) because of the resemblance of the bark to crocodile skin, as described by Queiroz et al. (\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, p. 199).\u003c/p\u003e \u003cp\u003e \u003col\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003e \u003cb\u003e3.1 General, organoleptic, and macroscopic wood anatomical characters\u003c/b\u003e \u0026ndash; Wood with a brown heartwood and yellowish sapwood (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec-d), highly dense, and resistant to cutting. Odorless; straight grain; fine texture; gloss present. Distinct growth ring, macroscopically demarcated by fibrous zone (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec-d). Diffuse porosity, tending towards semi-ring porosity. Vessels arranged in a radial pattern; solitary and multiple radial vessels of 2\u0026ndash;5, less frequently 6\u0026ndash;10; noticeable only under magnification; small in tangential diameter; very numerous; some vessels obstructed by deposits in the heartwood. Axial parenchyma indistinct even under magnification. Rays noticeable only under magnification, storied.\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003e \u003cb\u003e3.2 Microscopic anatomical description of wood\u003c/b\u003e \u0026ndash; \u003cb\u003eGrowth rings\u003c/b\u003e are well defined, marked by thick-walled and radially flattened fibres and distended rays, prone to semi-ring-porous wood (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-b). \u003cb\u003eVessels\u003c/b\u003e are wood diffuse-porous with a tendency to semi-porous (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-b). Vessel arrangement in radial pattern (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-b). Vessel groupings: solitary and radial multiples of 2\u0026ndash;5 (solitary 20\u0026ndash;29%, radial multiples of two 26\u0026ndash;29%, of three 18\u0026ndash;27%, of four 10\u0026ndash;15%, of five 4\u0026ndash;7%) occurring less frequently radial multiples of 6\u0026ndash;10, 4\u0026ndash;12% (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-b); simple perforation plates (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef); intervessel pits alternate, small and vestured (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eg); vessel-ray pits with distinct borders, similar to intervessel pits in size and shape throughout the ray cell (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ed). Deposits in heartwood vessels. \u003cb\u003eFibres\u003c/b\u003e with simple to minutely bordered pits, few, common in both radial and tangential walls. Nonseptate fibres present. Fibres very thick-walled (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb). \u003cb\u003eAxial parenchyma\u003c/b\u003e diffuse and paratracheal, both scanty (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea-b). Fusiform and two cells per parenchyma strand (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec), less often 3\u0026ndash;4 cells per parenchyma strand. \u003cb\u003eRays\u003c/b\u003e width 1 to 3 cells (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec). Cellular composition of rays: homocellular, all ray cells procumbent (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ed); and heterocellular, body ray cells procumbent with one or more row of procumbent cells more high, upright and/or square marginal cells (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). Perforated ray cells present, rare. \u003cb\u003eStoried structure\u003c/b\u003e includes rays, axial parenchyma and vessel elements that are storied or irregularly storied (4c). \u003cb\u003eMineral inclusions\u003c/b\u003e include prismatic crystals in chambered axial parenchyma cells (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec, e), and in procumbent, upright and square ray cells (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee), less often in chambered upright and/or square ray cells.\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003c/ol\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe anatomical quantitative features of the wood are described in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Data on wood anatomy for the tribe Brongniartieae is largely scarce. Among the 15 genera and approximately 180 species constituting this tribe, only six genera and merely eight species, about 4%, have any data on wood anatomy. Yet, some species such as \u003cem\u003eA. effusus\u003c/em\u003e and \u003cem\u003eLimadendron amazonicum\u003c/em\u003e (Ducke) Meireles \u0026amp; A.M.G.Azevedo only have quite incomplete anatomical information (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eComparative wood anatomy of \u003cem\u003eTabaroa caatingicola\u003c/em\u003e with eight species from the Brongniartieae tribe for which some wood anatomy data is available. For quantitative data, the classes indicated by the IAWA Committee (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1989\u003c/span\u003e) are presented, and when available, the range of variation (minimum \u0026ndash; maximum), mean, and/or mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation. (-) Not applicable; (?) Information is missing. The list of anatomical character follows the IAWA Committee (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1989\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAnatomical character / Species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eTabaroa caatingicola\u003c/em\u003e L.P.Queiroz, G.P.Lewis \u0026amp; M.F.Wojc.\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eAmphiodon effusus\u003c/em\u003e Huber\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eBehaimia cubensis\u003c/em\u003e Griseb.\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eHarpalyce formosa\u003c/em\u003e DC.\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eHarpalyce arborescens\u003c/em\u003e A.Gray\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eHaplormosia monophylla\u003c/em\u003e (Harms) Harms\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eLimadendron amazonicum\u003c/em\u003e (Ducke) Meireles \u0026amp; A.M.G.Azevedo\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cem\u003eLimadendron hostmannii\u003c/em\u003e (Benth.) Meireles \u0026amp; A.M.G.Azevedo\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cem\u003ePoecilanthe parviflora\u003c/em\u003e Benth.\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eGrowth rings\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eGrowth rings: anatomical markers\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eThick-walled and radially flattened fibres and distended rays, prone to semi-ring-porous wood\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMarginal parenchyma\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eMarginal parenchyma, semi-ring-porous and thick-walled and/or radially flattened latewood fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMarginal parenchyma\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eThick-walled and/or radially flattened latewood fibres and marginal parenchyma\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePorosity\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDiffuse-porous with a tendency to semi-porous\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDiffuse-porous\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDiffuse-porous\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDiffuse-porous\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eDiffuse-porous with tendency to semi-ring porosity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eDiffuse-porous\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eDiffuse-porous\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eDiffuse-porous\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eVessel arrangement\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVessels in radial pattern\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eNo specific arrangement\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eVessel groupings\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSolitary and radial multiples of 2\u0026ndash;5, less common radial multiples of 6\u0026ndash;10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVessels in radial multiples of 4 or more common\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVessels in radial multiples of 4 or more common\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eVessels in radial multiples of 4 or more common\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSolitary and radial multiples of 2\u0026ndash;3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSolitary and radial multiples of 2\u0026ndash;3, less common radial multiples of 4\u0026ndash;5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eSolitaires and radial multiples of 2, radial multiples of 3\u0026ndash;4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePerforation plate\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eSimple\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eIntervessel pits arrangement\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAlternate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAlternate, polygonal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAlternate, polygonal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAlternate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAlternate, polygonal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eAlternate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAlternate, polygonal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAlternate\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eIntervessel pits size\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSmall\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSmall\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSmall\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMedium to large\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMedium\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eSmall to Medium\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eVestured pit\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eVessel\u0026ndash;ray pits\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eSimilar to intervessel pits in size and shape throughout the ray cell\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMean tangential diameter of vessel (\u0026micro;m)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026le;\u0026thinsp;50 to 50\u0026ndash;100 (44.0\u0026ndash;63.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50\u0026ndash;100 (50\u0026ndash;60)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e50\u0026ndash;100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e50\u0026ndash;100 (61)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e50\u0026ndash;100 (61.16\u0026thinsp;\u0026plusmn;\u0026thinsp;10.90)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e100\u0026ndash;200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e50\u0026ndash;200 (100)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e50\u0026ndash;100 (40\u0026ndash;100)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eVessels per square millimeter\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026ge;\u0026thinsp;100 (119\u0026ndash;147)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5\u0026ndash;20 (8.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e40\u0026ndash;100 (68)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e40\u0026ndash;100 (45\u0026thinsp;\u0026plusmn;\u0026thinsp;15; 29\u0026ndash;66)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026le;\u0026thinsp;5 to 5\u0026ndash;20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e5\u0026ndash;20 (10\u0026ndash;15)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e5\u0026ndash;40 (11\u0026ndash;38)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMean vessel element length (\u0026micro;m)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026le;\u0026thinsp;350 (163\u0026ndash;197)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;= 350 (208; (173\u0026ndash;270))\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e350\u0026ndash;800 (187.06\u0026thinsp;\u0026plusmn;\u0026thinsp;20.66)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTyloses\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eDeposits in heartwood vessels\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePresent, oleoresin\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eGround tissue fibres\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eFibres with simple to minutely bordered pits\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSeptate and nonseptate fibres\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNonseptate fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNonseptate fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNonseptate fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNonseptate fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNonseptate fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eNonseptate fibres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eFibre wall thickness\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVery thick-walled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVery thick-walled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVery thick-walled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eThin- to thick-walled and\u003c/p\u003e \u003cp\u003every thick-walled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVery thick-walled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eVery thick-walled\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eVery thick-walled\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eFibre diameter (\u0026micro;m)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e13.1\u0026ndash;14.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.51\u0026thinsp;\u0026plusmn;\u0026thinsp;1.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eFibre wall thickness (\u0026micro;m)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.7\u0026ndash;5.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.74\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eFibre length (\u0026micro;m)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026le;\u0026thinsp;900 (709\u0026ndash;799)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e900\u0026ndash;1600 (947; 799\u0026ndash;1021))\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026le;\u0026thinsp;900 (270.30\u0026thinsp;\u0026plusmn;\u0026thinsp;43.14)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e900\u0026ndash;1600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eAxial parenchyma\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDiffuse and paratracheal, both scanty\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBands more than three cells wide\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDiffuse and paratracheal, both scanty and marginal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eScanty paratracheal, vasicentric, confluent and marginal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eUnilateral paratracheal, lozenge-aliform, confluent, bands more than three cells wide and marginal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eScanty paratracheal and bands with 2\u0026ndash;6 cells wide\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eScanty paratracheal, bands with 2\u0026ndash;4 cells wide and marginal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eScanty paratracheal and in narrow bands or lines up to three cells wide\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eAxial parenchyma cell type / Strand length\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFusiform and two cells, less often 3\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1\u0026ndash;2 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eRay width\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1\u0026ndash;3 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1\u0026ndash;3 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u0026ndash;3 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1\u0026ndash;3 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1\u0026ndash;4 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1\u0026ndash;3 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e2\u0026ndash;3 cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eRays with multiseriate portion(s) as wide as uniseriate portions\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003ePresent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eRay height\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.0\u0026ndash;22.6 \u0026micro;m; 6.1\u0026ndash;7.2 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7\u0026ndash;10 cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e230 \u0026micro;m\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e178.68\u0026thinsp;\u0026plusmn;\u0026thinsp;31.89 \u0026micro;m\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e120\u0026ndash;240 \u0026micro;m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eRays: cellular composition\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAll ray cells procumbent and body ray cells procumbent with one or more row of procumbent cells more high, upright and/or square marginal cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAll ray cells procumbent and body ray cells procumbent with one row of upright and/or square marginal cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBody ray cells procumbent with one row of upright and/or square marginal cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAll ray cells procumbent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAll ray cells procumbent and body ray cells procumbent with one or more row of upright and/or square marginal cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eBody ray cells procumbent with one row of upright and/or square marginal cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eBody ray cells procumbent with one row of upright and/or square marginal cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eHomocellular\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSheath cells\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePerforated ray cells\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresent, rare\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eRays per millimeter\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.7\u0026ndash;11.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4\u0026ndash;12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4\u0026ndash;12 (12)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4\u0026ndash;12 (4\u0026thinsp;\u0026plusmn;\u0026thinsp;1; 2\u0026ndash;6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4\u0026ndash;12 to \u0026ge;\u0026thinsp;12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e4\u0026ndash;12 (6\u0026ndash;7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4\u0026ndash;12 to \u0026ge;\u0026thinsp;12 (8\u0026ndash;14)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eStoried structure\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRays, axial parenchyma and vessel elements\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRays, axial parenchyma and vessel elements\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eRays, axial parenchyma and vessel elements\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eRays, axial parenchyma and vessel elements\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eRays, axial parenchyma and vessel elements\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eRays, axial parenchyma and vessel elements\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMineral inclusions\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePrismatic crystals in chambered axial parenchyma cells, and in procumbent, upright and square ray cells, less often in chambered upright and/or square ray cells.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePrismatic crystals in upright and/or square ray cells and in chambered axial parenchyma cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePrismatic crystals in upright, square and procumbent ray cells and in chambered axial parenchyma cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePrismatic crystals in chambered axial parenchyma cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e?\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePrismatic crystals in procumbent ray cells\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAbsent\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eReference\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eThis paper\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTsuchiya et al. (\u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2002\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGasson et al. (\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2004\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBonilla et al. (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2004\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eRam\u0026iacute;rez-Mart\u0026iacute;ne et al. (2017)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eLoupe et al. (2008)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eInsideWood (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e-onwards)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eDetienne and Jacquet (1983)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eManieri and Chimelo (1989)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"4 Discussion","content":"\u003cp\u003eExternal characteristics of the bark, as well as the general, organoleptic, and macroscopic anatomical features of wood, can assist regulatory bodies in wood identification and in combating illegal exploitation and trade. This information can thereby contribute to the conservation efforts for this species (Silva et al. \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003e4.1 Systematic wood anatomy \u0026ndash;\u003c/b\u003e The wood anatomy of \u003cem\u003eTabaroa caatingicola\u003c/em\u003e differs from other Brongniartieae genera by having more than 100 vessels.mm\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e, the vessels arranged in a radial pattern and perforated ray cells. Although comparing \u003cem\u003eT. caatingicola\u003c/em\u003e with its closest genus, \u003cem\u003eAmphiodon\u003c/em\u003e, is impeded by limited anatomical data for the latter, distinctions arise in growth rings, porosity, and axial parenchyma type: \u003cem\u003eTabaroa\u003c/em\u003e exhibits growth rings, semi-porous wood, and diffuse and scanty paratracheal, while \u003cem\u003eAmphiodon\u003c/em\u003e lacks growth rings, displays diffuse-porous wood, with paratracheal parenchyma bands more than three cells wide (Tsuchiya et al. \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). With the current data, no anatomical feature of the wood is exclusive to these two sister genera. A comprehensive wood anatomical description is advised for \u003cem\u003eAmphiodon effusus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eMost wood anatomical features observed in \u003cem\u003eTabaroa\u003c/em\u003e and all other Brongniartieae genera are also widely observed in woody angiosperms: wood diffuse-porous (some tend to semi-porosity), vessels multiples, simple perforation plates, intervessel pits alternate, vessel\u0026ndash;ray pits with distinct borders similar to intervessel pits in size and shape, libriform fibres with very thick-walled and non-septate, 4\u0026ndash;12 rays.mm\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e and gums and other deposits in heartwood vessels (except \u003cem\u003eBehaimia\u003c/em\u003e) (Carlquist \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Wheeler and Baas \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Wheeler et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2007\u003c/span\u003e); or in Leguminosae wood: vestured pits, 1\u0026ndash;4 cells per parenchyma strand, and all rays, axial parenchyma and vessel elements storied (except \u003cem\u003eLimadendron\u003c/em\u003e) (Carlquist \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Gasson \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Gasson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; H\u0026ouml;hn \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Pern\u0026iacute;a and Melandri \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Ramanantsialonina et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Stepanova et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Hence, considering the limited data available among species of the Brongniartieae tribe, no anatomical trait can be inferred as synapomorphic of this clade. However, the composition of ray cells, diffuse and/or paratracheal scanty axial parenchyma (except \u003cem\u003eAmphiodon effusus\u003c/em\u003e and \u003cem\u003eHarpalyce arborescens\u003c/em\u003e), and the presence of prismatic crystals (except \u003cem\u003eH. arborescens\u003c/em\u003e and \u003cem\u003ePoecilanthe parviflora\u003c/em\u003e) appear to be very characteristic of the tribe.\u003c/p\u003e \u003cp\u003eThe anatomical markers semi-ring-porous wood and thick-walled and radially flattened fibres are reported for \u003cem\u003eHarpalyce arborescens\u003c/em\u003e A.Gray (Loupe et al. 2008), the latter also being described for \u003cem\u003ePoecilanthe parviflora\u003c/em\u003e Benth. (Manieri and Chimelo 1989). The other Brongniartieae species either lack growth rings or exhibit marginal parenchyma, a more common anatomical marker in Leguminosae, also associated with deciduous species (Carlquist, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; H\u0026ouml;hn \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Pern\u0026iacute;a and Melandri \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Silva \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Stepanova et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Worbes and Fichtler 2010), although it does not occur in \u003cem\u003eTabaroa\u003c/em\u003e. \u003cem\u003eTabaroa caatingicola\u003c/em\u003e shares with its phylogenetically related genus \u003cem\u003eHarpalyce\u003c/em\u003e tendency to semi-ring-porous wood and short vessel element length (\u0026le;\u0026thinsp;350 \u0026micro;m) (Bonilla et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Ram\u0026iacute;rez-Mart\u0026iacute;ne et al. 2017).\u003c/p\u003e \u003cp\u003eOther anatomical similarities can be noted among the other Brongniartieae genera. For example, marginal parenchyma is characteristic of \u003cem\u003eHarpalyce\u003c/em\u003e, \u003cem\u003eLimadendron\u003c/em\u003e, and \u003cem\u003ePoecilanthe\u003c/em\u003e (Bonilla et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Detienne and Jacquet 1983; Manieri and Chimelo 1989; Ram\u0026iacute;rez-Mart\u0026iacute;nez et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), being a type of parenchyma widely distributed across Leguminosae species (Carlquist \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Gasson \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Gasson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; H\u0026ouml;hn \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Pern\u0026iacute;a and Melandri \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Ramanantsialonina et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Stepanova et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The polygonal intervessel pits are characteristic of \u003cem\u003eBehaimia\u003c/em\u003e, \u003cem\u003eHaplormosia\u003c/em\u003e, \u003cem\u003eHarpalyce\u003c/em\u003e, and \u003cem\u003eLimadendron\u003c/em\u003e (Bonilla et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Detienne and Jacquet 1983; Gasson et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Loupe et al. 2008). Tyloses is present in \u003cem\u003eLimadendron\u003c/em\u003e and \u003cem\u003ePoecilanthe\u003c/em\u003e (InsideWood \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e-onwards; Manieri and Chimelo 1989), it goes against the data which indicate that species with vessel\u0026ndash;ray pits similar to intervessel pits, normally with small diameter, have a low probability of having tyloses (Bonsen and Kučera \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; De Micco et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The absence of deposits in heartwood vessels is indicated for \u003cem\u003eBehaimia\u003c/em\u003e and \u003cem\u003eHarpalyce\u003c/em\u003e (Bonilla et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Gasson et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Rays with multiseriate portions as wide as uniseriate portions are limiting to \u003cem\u003eLimadendron amazonicum\u003c/em\u003e (InsideWood \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e-onwards).\u003c/p\u003e \u003cp\u003e \u003cb\u003e4.2 Functional and ecological wood anatomy \u0026ndash;\u003c/b\u003e Growth rings are present in the wood of \u003cem\u003eT. caatingicola\u003c/em\u003e and are identified by three distinct anatomical markers: thick-walled and radially flattened fibres and distended rays, prone to semi-ring-porous wood. Anatomical markers can be characterized as histological contrasts arising from traits that vary between cells produced throughout a growing season, giving rise to early- and latewood (Silva \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Each of these variations, whether in isolation or combination, generates identifiable features in the secondary xylem, recurring across multiple growing seasons. Currently, growth rings are conceived not only as a result of the vascular cambium\u0026rsquo;s ability to respond to environmental and physiological changes but also to effect changes in the secondary xylem with significant functional implications (Carlquist \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Silva et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Silva \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFibers with a smaller lumen and thicker walls are hypothesized to enhance the security of transport in vessel elements, representing the most widespread anatomical marker among angiosperms, particularly in tropical regions (Jacobsen et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Silva et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Semi-ring-porous wood is more commonly found in species distributed in subtropical and temperate regions (Silva et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, it is also present in tropical species, particularly in more seasonal habitats, like the Caatinga dry woodlands, where gradations between diffuse and semi-ring porosity can be observed (Arag\u0026atilde;o and Lisi \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Gasson et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Silva \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). This anatomical marker shows varying degrees of specialization in balancing the trade-off between embolism protection and efficient conduction. Broad vessels in the earlywood facilitate conduction capacity with a lower risk of embolism during periods of ample water availability. In contrast, narrower vessels in conditions of water scarcity, although less efficient for transport, ensure continuous xylem flow while minimizing the risk of embolism formation and the subsequent disruption of conduction (Baas et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Sperry et al. \u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Tyree and Zimmermann \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Distended rays, on the other hand, adapt to adjacent cells, preventing the formation of large intercellular spaces and, consequently, cracks in the secondary xylem (Silva et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Silva \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe seasonality and irregularity of rainfall in the Caatinga explain the formation of growth rings with an annual periodicity (Mattos et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Nogueira Jr et al. 2018; Silva \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) and infra-annual, potentially forming multiple annual ring (\u003cem\u003esensu\u003c/em\u003e Silva et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Pagotto et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The latter periodicity is also recognized for dry forests in East Africa (Gourlay \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1995a\u003c/span\u003e, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003eb\u003c/span\u003e; Jacoby \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1989\u003c/span\u003e). The growth rings of \u003cem\u003eT. caatingicola\u003c/em\u003e should be studied to determine the periodicity.\u003c/p\u003e \u003cp\u003eGiven that \u003cem\u003eT. caatingicola\u003c/em\u003e is ecologically confined to a semi-arid region, the vessel characteristics outlined here \u0026mdash; including a degree of semi-porosity, predominantly multiple vessels, small diameter, high number of vessels per square millimeter, short vessel element length, simple perforation plates, and vestured pits \u0026mdash; are consistent with the well-established trade-off between safety \u0026times; efficiency in conduction. This trade-off is widely recognized as the primary factor influencing the evolution of secondary xylem in seed plants (Baas et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). While the mechanisms of freeze-induced embolism are better understood, drought-induced embolism is a theme that has gained strength in recent years (Hacke et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Olson \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Failures in sap transport through the xylem are among the main causes of death or reduced productivity in plants subjected to drought (Anfodillo and Olson \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Breshears et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2005\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Brodribb and Cochard \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; McDowell et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). So, cavitation vulnerability is a relevant hydraulic trait that determines a species\u0026rsquo; competitiveness under water stress (Islam et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWider vessels are at a higher risk of embolism than narrower vessels within a tissue and when we compare species. Typically, wider vessels tend to be more vulnerable to drought-induced embolism, whereas embolism-resistant vessels are generally narrow (Hacke et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This explanation justifies the small mean vessel diameter, as well as the slightly semi-ring porosity observed in \u003cem\u003eT. caatingicola\u003c/em\u003e. The vessel diameter influences the variation in vessels per square millimeter. The inverse correlation between diameter and vessel frequency has been previously reported and is widely supported in the literature (Chave et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Sperry et al. \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe high percentage of multiple vessels, ranging from 71\u0026ndash;80%, stands out in the wood of \u003cem\u003eT. caatingicola\u003c/em\u003e. The performance of different levels of vessel grouping in secondary xylem is a controversial subject in the literature since Carlquist (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). Some authors argue that multiple vessels increase the chances of propagating cavitation of air-seeding (Jacobsen et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Tyree and Zimmermann \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), while others suggest that a higher connection between vessels improves hydraulic integration, providing alternative pathways when embolism blocks flow in one vessel, reducing the potential loss of water transport capacity associated with cavitation (Loepfe et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Trifil\u0026ograve; et al. \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Despite many gaps in understanding the functioning of different vessel grouping levels, the latter argument is gaining strength by providing data that cavitation can be more easily removed in grouped vessels than in solitary ones (H\u0026ouml;ltt\u0026auml; et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), and that multiple vessels are more resistant to cavitation (Lens et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In any case, \u003cem\u003eT. caatingicola\u003c/em\u003e is part of the species in dry forests that exhibit greater vessel connectivity (Apgaua et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Scholz et al. \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe considerably short vessel elements described for the wood of \u003cem\u003eT. caatingicola\u003c/em\u003e also have a functional relationship with the water-limited, arid environment in which this species grows. Carlquist (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1975\u003c/span\u003e) hypothesized that shorter vessel elements have adaptive value, resisting mechanical deformation under negative pressure. Recently, a comparative analysis of over 1000 species confirmed Carlquist\u0026rsquo;s prediction, indicating that shorter vessel elements affect vessel resistance to deformation. Species with exceptionally short vessel elements tend to grow in arid lands, while those with exceptionally long vessel elements tend to thrive in humid climates (Echeverr\u0026iacute;a et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Simple perforation plates are widely found in angiosperms, prevailing in species distributed across the tropics. Simple perforation plates provide considerably less resistance to sap flow between vessel elements, being crucial, when compared to compound perforation plates, for high flow demands (Carlquist \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Wheeler and Baas \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Wheeler et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), notably for plants growing in environments with scarce and irregular precipitation such as the Caatinga.\u003c/p\u003e \u003cp\u003eThe vestured pits described for the vessel elements of \u003cem\u003eT. caatingicola\u003c/em\u003e are an anatomical trait of wood with systematic and functional value. They are generally associated with simple perforation plates, bordered pits, and dry environments, characterizing taxa such as the orders Myrtales and Gentianales, and families such as Malpighiaceae and Leguminosae (Jansen et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Although vestured pits are related to safety in hydraulic conduction, reducing vulnerability to cavitation or assisting in embolism repair, the mechanisms associated with these functionalities remain poorly understood (Jansen et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Rabaey et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The presence of highly lignified structures within the pit chamber can influence hydraulic resistance. Vestured pits appear to facilitate embolism reversal in xeric and warmer regions with high transpiration rates (Jansen et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), like the Caatinga seasonally dry vegetation. The small pit apertures recorded for the vessel elements of \u003cem\u003eT. caatingicola\u003c/em\u003e are associated with the small diameters of the vessel elements, as this trait covaries with vessel element diameter (Silva et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe xylem of \u003cem\u003eT. caatingicola\u003c/em\u003e is considered to have a high degree of xeromorphism based on the vulnerability and mesomorphy index values. Carlquist (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1977\u003c/span\u003e) considered vulnerability index values significantly below 1.0, as observed here, to indicate a high degree of xeromorphy. Meanwhile, mesomorphy index values above 200 indicate mesophytic species. For these indices, higher values are associated with greater efficiency, while lower values are linked to greater safety (Carlquist \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1977\u003c/span\u003e). The vulnerability and mesomorphy indices proposed by Carlquist (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1977\u003c/span\u003e) help quantify the degree of mesomorphy or xeromorphy exhibited in the plant xylem, contributing to a quantitative approach to plant anatomy, particularly in the fields of functional and ecological anatomy (Ewers et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDespite of the few data available for comparison, the other species of the Brongniartieae tribe exhibit a series of wood anatomy traits common to \u003cem\u003eT. caatingicola\u003c/em\u003e. However, most of these traits are widely found in angiosperms and/or in Leguminosae. \u003cem\u003eTabaroa caatingicola\u003c/em\u003e has a set of wood anatomy traits that distinguishes it from other Brongniartieae genera. The wood anatomy of \u003cem\u003eT. caatingicola\u003c/em\u003e reveals a set of adaptations to the SDTFW biome of the Brazilian Caatinga domain, where the low precipitation with irregular rains, high temperatures, and high transpiration potential increases the likelihood of drought-induced embolism formation (Hacke et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Olson et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). These adaptations include semi-ring-porous wood with distinct growth ring boundaries, multiple narrow vessels, simple perforation plates, small and vestured pits. These traits increase flow during periods of higher water availability and ensure hydraulic safety during scarcity, while also providing mechanisms to minimize the formation and spread of embolisms.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe research project was initiated, and the study designed by M.S.S. and D.C. M.S.S. and D.C. conducted fieldwork and prepared the figures. M.S., L.B.S. and D.B. did the laboratory work and anatomical analyses. C.S. performed the SEM analyses. All authors contributed to the writing of the manuscript as well as approving the final submitted version of the paper.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatements and Declarations\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConflict of interests: The authors declare that the manuscript does not present any kind of conflict of interests. 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Methods Ecol Evol 10:744\u0026ndash;751. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/2041-210X.13152\u003c/span\u003e\u003cspan address=\"10.1111/2041-210X.13152\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"brazilian-journal-of-botany","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"brjb","sideBox":"Learn more about [Brazilian Journal of Botany](https://www.springer.com/journal/40415)","snPcode":"40415","submissionUrl":"https://www.editorialmanager.com/brjb/default2.aspx","title":"Brazilian Journal of Botany","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Brongniartieae, ecological wood anatomy, Fabaceae, functional wood anatomy, systematic wood anatomy, vestured pits","lastPublishedDoi":"10.21203/rs.3.rs-3921723/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3921723/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWood anatomy can serve as a source of phylogenetically informative characteristics, as well as a model of how more plastic characters have evolved over time in plants, e.g., through ancestral reconstructions of wood anatomical traits across well-resolved phylogenies. However, the evolution of wood anatomy is largely unexplored within a phylogenetic context due to limited availability of anatomical data across taxa. When compared with other angiosperm families, Leguminosae is relatively well-documented, yet it still lacks comprehensive wood anatomical information, particularly in undersampled papilionoid clades. In order to contribute to the understanding of micromorphological diversity across papilionoid legumes, we newly characterize the wood anatomy of \u003cem\u003eTabaroa caatingicola\u003c/em\u003e, a papilionoid species narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests, that has been molecularly placed in the poorly anatomically studied tribe Brongniartieae. Optical histology and scanning electron microscopy (SEM) were used to examine and describe the wood anatomy of \u003cem\u003eT. caatingicola\u003c/em\u003e and compare it with six phylogenetically related Brongniartieae genera: \u003cem\u003eAmphiodon\u003c/em\u003e, \u003cem\u003eBehaimia\u003c/em\u003e, \u003cem\u003eHaplormosia\u003c/em\u003e, \u003cem\u003eHarpalyce\u003c/em\u003e, \u003cem\u003eLimadendron\u003c/em\u003e, and \u003cem\u003ePoecilanthe\u003c/em\u003e. Wood anatomy of \u003cem\u003eTabaroa\u003c/em\u003e suggests adaptations to the irregular rainfall of the harsh Caatinga environment, featuring distinct growth rings, prone to semi-ring-porous wood, multiple narrow vessels, simple perforation plates, and small and vestured pits. These traits increase water flow during abundance and ensure hydraulic safety during scarcity, minimizing embolism formation and spread. By focusing on the genus \u003cem\u003eTabaroa\u003c/em\u003e, an ecologically distinctive and evolutionarily isolated lineage, this study contributes to the understanding of the systematic and functional wood anatomy variation in the papilionoid legume tribe Brongniartieae.\u003c/p\u003e","manuscriptTitle":"Wood anatomy of Tabaroa, a monotypic papilionoid legume genus narrowly endemic to the Brazilian Caatinga seasonally dry tropical forests","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-03-05 19:36:20","doi":"10.21203/rs.3.rs-3921723/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Minor revisions","date":"2024-04-26T16:12:46+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2024-02-29T03:18:16+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-02-28T18:38:05+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Brazilian Journal of Botany","date":"2024-02-26T14:35:55+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-02-24T06:19:15+00:00","index":"","fulltext":""},{"type":"submitted","content":"Brazilian Journal of Botany","date":"2024-02-22T15:51:57+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"brazilian-journal-of-botany","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"brjb","sideBox":"Learn more about [Brazilian Journal of Botany](https://www.springer.com/journal/40415)","snPcode":"40415","submissionUrl":"https://www.editorialmanager.com/brjb/default2.aspx","title":"Brazilian Journal of Botany","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"71c51d4b-b63f-4388-9c9f-ade311aeb336","owner":[],"postedDate":"March 5th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2024-05-27T14:55:00+00:00","versionOfRecord":[],"versionCreatedAt":"2024-03-05 19:36:20","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3921723","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3921723","identity":"rs-3921723","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}