Use, management and domestication of Agave americana in Mexico

Use, management and domestication of Agave americana in Mexico | 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 Use, management and domestication of Agave americana in Mexico NADIA CAMPOS SALAS, ALEJANDRO CASAS This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9078121/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 10 You are reading this latest preprint version Abstract Background Domestication is an evolutionary process guided by humans. It operates continuously and dynamically in multiple directions, with species undergoing adaption to diverse human-influenced environments and cultural and technological contexts. Mexico and the neighbouring Mesoamerican area are among the world’s main regions of domestication. The Agave genus is widely used and managed, with at least eleven domesticated and semi-domesticated species. One species, Agave americana , comprises two subspecies and four varieties, the diversification of which is hypothetically related to human management. This study explores this hypothesis. Methods Ethnobotanical studies were conducted on the use, management, and targets of human selection, as well as the patterns of morphological variation, in different ecological and cultural contexts. Semi-structured interviews were conducted with people managing these Agave s across their distribution range, from northern to southern Mexico. Morphometric studies were also conducted on populations distributed in contrasting social and ecological contexts to explore phenotypic variation and trends in relation to selection targets. Multivariate and univariate statistical analyses were performed to analyse patterns of variation in relation to environmental conditions, geographical distribution and forms of use and management. Results A. americana is used to extract sap for preparing the fermented beverage called pulque. Its stems have been used as food since prehistoric times, when they were cooked in underground ovens. This cooked matter forms the basis of current fermentation and the production of distilled mescal. The fibre of some varieties has been used to make cords and textiles. Both subspecies are used and managed, but the subspecies protamericana clearly has wild populations. Morphometric studies confirm the presence of traits indicative of domestication syndrome in most varieties of the subspecies americana . Conclusions Phenotypic variation is clearly related to the taxonomic status and geographic distribution area of the analysed plants, but the influence of human selection favouring gigantism and reduced spines on the leaves is also evident. Further studies of population genetics and phylogeographic patterns would improve our understanding of these patterns. Agave management domestication maguey mescal Mesoamerica pulque Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Background Domestication is an evolutionary process directed by humans that provides valuable insights into general evolutionary mechanisms in organisms [ 1 – 4 ]. As the biocultural contexts in which domestication occurs are diverse and dynamic, the process operates continuously and dynamically in multiple directions. The species being tamed adapt to diverse human-influenced environments and cultural and technological contexts, enabling them to live and reproduce successfully. Domesticated organisms are shaped by morphological and physiological attributes that are markedly (though not exclusively) aligned with human requirements [ 5 , 6 ]. Although selection is the primary evolutionary force, other forces—mutation, recombination, gene flow, genetic drift, and mating systems—also influence domestication under both natural and human guidance [ 7 , 8 ]. Mesoamerica, the cultural region comprised between Mexico and Central America, is one of the world’s main domestication regions [ 9 – 12 ]. Indigenous peoples in this area domesticated more than 250 native species [ 13 ], including maize ( Zea mays L.), beans ( Phaseolus spp.), squashes ( Cucurbita spp.), agaves ( Agave spp.), prickly pears ( Opuntia spp.), chili peppers ( Capsicum annuum L.), vanilla ( Vanilla planifolia Jacks. ex Andrews), cotton ( Gossypium hirsutum L.), and other crops which are currently of worldwide relevance. The coexistence of crops and their wild relatives, together with varying intensities and directions of selection and management, produced extensive morphological and physiological diversity [ 14 – 17 ]. This diversity is reflected in the multiple varieties often found within a single species. Records from archaeological, ethnobiological, and evolutionary ecology studies have revealed that domestication began in Mesoamerica around 10 thousand years ago [ 10 , 18 – 20 ] and continues today, influencing both native and introduced species [ 4 , 18 ]. The region harbors one of the world’s richest reservoirs of agrobiodiversity [ 16 , 22 – 24 ]. Documenting how this diversity originated and identifying the factors that maintain it remain key priorities for its conservation [ 25 ]. Indigenous peoples in Mesoamerica use nearly 6,500 native plant species [ 13 ], about 20% of those recorded by botanists [ 26 ]. Among them, Agave stands out as a genus of exceptional ecological and cultural importance. Its species have been used since prehistoric times and dominate many regional landscapes, where the genus also diversified. At least twelve species have been domesticated to varying degrees: A. americana L., A. angustifolia Haw., A. atrovirens Karw. ex Salm-Dyck, A. fourcroydes Lem., A. karwinskii Zucc., A. mapisaga Trel., A. salmiana Otto ex Salm-Dyck, A. sisalana Perrine, A. tequilana FAC Weber, A. parryi Engelm., A. hookeri Jacobi, and A. inaequidens K. Koch. Others, such as A. cupreata Trel. & A. Berger, A. potatorum Zucc., A. maximiliana Baker, and A. rhodacantha Trel., have been cultivated for at least fifty years without clear signs of domestication [ 27 ]. The cultivation and management of these species, and the associated selective mechanisms, have been well documented [ 14 , 28 – 33 ]. High levels of interspecific hybridization and genetic variation have also been reported [ 30 – 36 ]. Since phenotypic changes reflect those occurring in genotypes [ 37 ], researchers commonly assess domestication by examining divergence between wild and managed phenotypes [ 38 , 39 ]. In Agave , domestication syndromes reflect use type, often involving enlargement of useful parts—stems, leaves, fibers, or sap—and reduction of defensive or undesirable features such as spines, teeth, and saponins and other secondary metabolites [ 14 , 29 – 31 ]. The interaction between natural and human selection makes Agave an excellent model for studying domestication. Mexico, with the greatest diversity and distribution of Agave s [ 40 ], offers ideal conditions for this research. Humans have managed and domesticated these plants since prehistoric times, using them for food, fiber, construction, fermented beverages, medicines, soil retention, and ceremonial purposes [ 10 , 23 , 14 , 27 ]. The case of Agave americana is of particular interest, given the extraordinary degree of diversification it has undergone during domestication. According to Gentry [ 23 ], the subspecies A. americana subsp. americana comprises four varieties. The putative wild relatives ( A. americana subsp. protamericana ) also occur in Mexico and the southern USA. Currently, all varieties of A. americana subsp. americana are found exclusively in managed populations, and their direct wild relatives are unknown. This gives rise to two hypotheses regarding their origin: (1) the species diversified prior to human management, with the ancestors of the domesticated varieties of A. a. americana subsequently becoming extinct; or (2) all varieties result from the domestication of one or few single ancestors, likely represented by the only clearly recognised wild taxon, A. a. protamericana . Testing these hypotheses requires comprehensive information on their genetic similarities and relationships, as well as on ecological aspects such as their distribution, abundance, local adaptations, and cultural aspects such as forms of management, targets and mechanisms of human selection, conditions of genetic drift associated to management, as well as gene flow influenced by translocation and other practices. The phenotypic consequences of these processes in relation to management must also be considered. This study explores aspects on management, selection, and phenotypic variation in the taxa included in this species, as well as some of its closest wild relatives. Human selection drives morphological and genetic differentiation in managed plants. A persistent challenge in evolutionary studies is determining how phenotypic traits are inherited and whether observed morphological divergence corresponds to genetic differentiation. Plant domestication is typically gradual; domesticated forms retain traits that reveal relationships with wild ancestors and display intermediate morphotypes depending on domestication degree. Examining these patterns helps reconstruct evolutionary pathways [ 9 , 41 ]. Currently, Agave americana subsp. protamericana is the only taxon of the species studied with clear wild populations, although this taxon’s populations may also be subject to some forms of management, as will be discussed below. Therefore, this taxon offers the opportunity to compare the morphology, physiology and genetics of the domesticated varieties by analysing how they diverge from other taxa and which features that are mainly affected by management. We hypothesize that these features should be related to the historical use of each taxon. We are aware of the extraordinary capacity for hybridization between varieties, subspecies, and even other closely related species within the genus. Therefore, phenotypic variation is influenced by complex factors, and such influence requires careful analysis using an integral approach to morphology, population genetics and reproductive biology. The domestication syndrome of Agave s may vary according to their use. For instance, their use as food has primarily involved the stem size, since stems have been cooked in underground ovens since prehistoric times. In addition, the inflorescences and flower buds can be consumed when roasted or boiled, respectively. Features of the domestication syndrome related to this use could include a large stem and scape, and flowers, as well as a reduction in secondary compounds that impart a bitter flavour. Agave fibres have also been used since ancient times, and gigantism expressed in leaf length, as well as the durability and strength of fibres would be expected in relation to this use. The sap of several Agave species has been used as a fresh (aguamiel) or fermented (pulque) beverage for thousands of years. To this end, selection favouring the quantity and quality of the sap (e.g. plant size, sap yield, and sugar content) would be expected. Currently, Agave s are most commonly used to produce mescal. While there is controversy surrounding pre-Columbian distillation, the material used to make mescal is the same as that used for cooking the stems. Therefore, the main attributes selected for this purpose are those preferred for preparing food in underground ovens. Gigantism of stems and leaves and the length and resistance of fibres, appear to be features shared by different species under domestication. In addition, the difficulty of managing Agave s has led people to favour plants that produce fewer and shorter spines and teeth, as well as lower amounts of saponins, which cause skin irritation in those managing them [ 14 ]. Understanding domestication thus requires examining divergence in these traits according to use and management intensity. This study aims to:1) document ethnobotanical and morphological information in order to analyse the Agave americana complex, its use and management, and the patterns and mechanisms of selection, the morphological divergence among the varieties and in relation to their putative wild ancestor A. americana subsp. protamericana , and 2) analyse the degree and patterns of divergence in relation to the targets of selection and management intensity in order to examine the role of management in morphological divergence. We expected to observe clear trends of divergence among the taxonomically defined varieties, as well as in domestication syndrome traits relative to A. a. subsp. protamericana , in order to support the hypothesis of a common ancestor of the domesticated varieties. Materials and methods Study species The Agave americana complex comprises the following two subspecies and four varieties: Agave americana subsp. protamericana , (Fig 1 G.) This predominantly wild taxon occurs along the Sierra Madre Oriental. On the coastal slopes, where rainfall is higher, rosettes are larger than on the drier continental side. Leaves are four to six times longer than wide, plane or guttered, straight or curved, and green to glaucous grey. The panicle is short, with 15–20 branches. Gentry [23] reported introgression between A. a. protamericana and A. asperrima , suggesting further research on reproductive biology and morphometrics to explore possible hybridization and its genetic and phenotypic outcomes. Agave americana subsp. americana var. americana (Fig 1 A.) This variety includes highly surculose plants with lanceolate leaves, widest near or slightly above mid-blade. Leaves are glaucous grey to light green, tapering above the base, and sometimes reflexed. The habitat is mainly anthropogenic, although Gentry [40] and Hernández-Sandoval et al. [42] reported putative wild populations near Ciudad Victoria, Tamaulipas. It is widespread in Mexico [43-45], though its wild status needs confirmation. Agave americana subsp. americana var. expansa . (Fig 1 B.) This taxon was first described by William Trelease in 1900 as var. americana . However, the large, glaucous grey, prolifically suckering rosettes and the tall, diffuse panicle resemble the taxon that occurs in Europe. The most notable characteristics are the valleculate leaves and the short, conical spines [23]. It has been reported in Chihuahua [46] and Jalisco [45]. Agave americana subsp. americana var. oaxacensis. (Fig 1 C.) It occurs in Oaxaca, where is commonly known as “maguey arroqueño”, “maguey coyote”, “maguey de Castilla”, or “maguey Sierra Negra” [24]. It has large, spreading, glaucous to whitish leaves with small teeth, producing large flowers and thick-walled capsules [23]. Because its stems accumulate high carbohydrate levels, it is used for high-quality mescal. Plants require 18–20 years to mature [45]. Historically, it was also used for pulque and fiber extraction [40]. Agave americana subsp. americana var. subtilis (Fig 1 H.) Known locally as “maguey chato” or “chato de Sahuayo,” this taxon was initially placed in the Rigidae group due to its compact rosette. Valenzuela-Zapata et al. [48] later reassigned it to the Americanae group based on inflorescence morphology. It mainly grows in Jiquilpan and Sahuayo, Michoacán. Related taxa considered Three additional species were included for comparison due to their close relationships and potential reproductive interactions with A. americana . Agave abrupta (Fig 1 D.), closely related to var. expansa , this species was first described by Trelease (1912) and later regarded by Gentry [23] as part of var. expansa . It differs mainly in rosette size, leaf shape, and terminal spine. Agave scaposa (Fig 1 F.), a large plant, with light green to glaucous leaves and tall, narrow panicles bearing small, dry bracts, aligning it with the Americanae group. Diagnostic traits include large rosettes, dark- to yellow-green undulate leaves with small marginal teeth, and tall inflorescences with small flowers [23]. According to García-Mendoza [49], A. scaposa is used for pulque production. Casas et al. [50] and Blancas et al. [51] reported its use in soil stabilization, fencing, and as a source of durable inflorescence stalks (or “quiote”) for construction. Agave asperrima subsp. Asperrima (Fig 1 E.), the most widespread Agave in northern Mexico [23], distinguished by scabrous, shorter leaves and narrow-bracted inflorescences with fewer branches. Its flowering overlaps and introgression with A. a. var. americana and A. a. subsp. protamericana have been reported [52-54]. Study area The field study sites for the analyzed taxa of the Agave americana complex are shown in Fig. 1 and Table 1. The study was conducted in three main regions of Mexico: Northeastern Mexico , covering the states of Nuevo León and Tamaulipas; The Trans-Mexican Volcanic Belt , including parts of Michoacán and Jalisco; and The Sierra Madre del Sur , represented by populations in Oaxaca. General information on the populations studied is summarized in Table 1, which includes altitude (m a.s.l.), mean annual temperature (°C), mean annual rainfall (mm), and associated habitat types. In Nuevo León, populations were examined in the localities Doctor Arroyo and Iturbide; in Tamaulipas, in San Carlos, El Tinieblo, Jaramillo, and Miquihuana; in Michoacán, in Tarímbaro, Santiago Undameo, Zacapu, Cojumatlán, and Jiquilpan; in Jalisco, in Zacoalco and Ciudad Guzmán; and in Oaxaca, in Coixtlahuaca, San Baltazar Guelavila, and San Pablo Huixtepec. These sites represent a broad range of ecological conditions, from microphyllous desert scrub and oak–pine forests in the north to agricultural plots, orchards, and intensive home-garden systems in the south. Such variation provides the environmental and management context for evaluating phenotypic differentiation and adaptation patterns within the A. americana complex. Table 1. Study sites of the analyzed Agave populations, including elevation (m a.s.l.), annual mean temperature (°C), annual rainfall (mm), and associated habitat types. These environmental variables provide the ecological context for population differentiation and habitat adaptation across the sites considered in the morphological variation analysis. Study Site Elevation (m a.s.l.) Annual Mean Temp (°c) Annual Rain Fall (mm) Habitat Dr. Arroyo, Nuevo León 1947 18.9 400 Mycrophyllous, subinerme desert scrub Iturbide, Nuevo León 1518 19 560 Oak-pine forest Orchards Miquihuana, Tamaulipas 1842 23 600 Thorny scrub San Carlos, Tamaulipas 570 22 850 Tamaulipas scrub El Tinieblo, Tamaulipas 199 22 100 Tamaulipas scrub Jaramillo, Tamaulipas 220 22 850 Tamaulipas scrub Zacapu, Michoacán. 2004 17 1069 Along the road Tarímbaro, Michoacán. 1885 18.1 609 Natural fence Santiago Undameo, Michoacán. 2050 16.2 754 Natural fence and home gardens Zacoalco, Jalisco. 1420 22.7 578.7 Agricultural field Cojumatlán, Michoacán. 1558 18 800 Orchard Jiquilpan, Michoacán. 1830 18 850 Monoculture plot Coixtlahuaca, Oaxaca. 2280 16 750 Rosetophylous scrub San Baltazar Guelavila, Oaxaca. 1570 20.7 650 Intensive Plots San Pablo Huixtepec, Oaxaca. 1480 20 1000 Intensive Plots Ethnobotanical studies A total of 14 in-depth interviews were conducted with the managers and owners of the visited sites to document traditional knowledge, uses, selection criteria, selection targets and mechanisms. These interviews were supplemented with previously collected ethnobotanical information from our team and other colleagues, including from published sources. Direct observations were also conducted on wild populations of Agave asperrima , Agave scaposa and Agave americana subsp. protamericana and subsp. americana . Ethnobotanical data were analyzed qualitatively to identify patterns of management intensity, domestication traits, and traditional selection strategies. Information was categorized according to management system (wild, promoted, managed, cultivated, and intensified plots) and cross-referenced with morphological variation patterns observed in the same populations. Studies of morphological variation Nineteen vegetative morphological traits were evaluated in field-collected plants from each population (Table 2). These characters were selected based on their ethnobotanical relevance and previous studies on domestication syndromes in Agave [14,29,31,55]. Most studied populations are cultivated or semi-cultivated, where plants are commonly harvested before flowering, making reproductive structures rarely available. To complement the data, 21 vegetative and reproductive traits were measured from herbarium specimens to characterize morphological differentiation among species, subspecies, and varieties. Preliminary analyses that included reproductive traits did not yield statistically significant differences among taxa; therefore, subsequent analyses focused on vegetative structures, following methodological precedents in ethnobotanical studies of Agave [14,29,31,55]. Table 2. Morphological characters measured in Agave populations studied. Their abbreviations and measurement units are indicated. In addition, it is indicated whether these characteristics were evaluated in herbarium specimens and/or in natural populations. Character Abbreviation Measure units Herbarium specimens Evaluated populations Plant length PL m X Stem length SL m X Diameter D m X Leaf length LL m X X Leaf width*** LW cm X X Leaf length/width ratio LL/LW cm X X Leaf/stem length ratio LL/SL m X Number of leaves LN Counted number X Terminal thorn length TTL cm X X Terminal thorn width TTW cm X X Terminal thorn length/width ratio TTL/TTW cm X X Terminal thorn/leaf length ratio TTL/LL cm X X Teeth length TEEL cm X X Teeth/leaf length ratio TEEL/LL cm X X Teeth width TEEW cm X X Teeth length/width ratio TEEL/TEEW cm X Teeth width/leaf length TEEW/LL cm X X Distance between teeth DBTEE cm X X Distance between teeth/leaf length ratio DBTEE/LL cm X X Inflorescence length IL m X Ramification number RN Counted number X Ovary length OL cm X Ovary width OW Cm X Tepal length TL cm X Filament length FL cm X Anther lenght AL cm X Stamen length SL cm X Statistical analyses of morphological characters Three multivariate analyses were performed to identify morphological grouping patterns among the studied taxa: Cluster Analysis (CA), Principal Components Analysis (PCA), and Linear Discriminant Analysis (LDA). All analyses were conducted using R software (version 4.3.2) [56]. Cluster Analysis employed Euclidean distances to estimate morphological dissimilarities among populations, generating a dendrogram with standardized average distance as weighting factor. PCA was used to assess interspecific relationships and identify the most influential variables for taxonomic differentiation. The analysis considered a matrix of 122 individuals and 19 variables, standardized by z-scores. Eigenvalues and eigenvectors were derived from Pearson’s correlation matrix, and individuals were projected onto the first two principal components. LDA was performed to classify individuals according to their taxonomic identity, management type, and geographic region. The eigenvectors from PCA and LDA analyses were compared to identify the most discriminant morphological traits. These analyses allowed evaluating whether morphological similarities correspond to management intensity, provenance, or taxonomic affinity within the A. americana complex. Heatmap and Clustering Analysis A heatmap was generated using the pheatmap package in R to visualize standardized morphological values and clustering patterns. Hierarchical agglomerative clustering with Euclidean distances and complete linkage was used to construct dendrograms for both individuals (rows) and traits (columns), revealing associations between taxa and morphological syndromes [57]. Color gradients were applied to highlight similarities and contrasts across populations, facilitating the visualization of domestication patterns that might not emerge from traditional plots. To evaluate significant differences between management categories and taxa, one-way ANOVAs were conducted for all 19 vegetative characters. Each variable was tested for normality (Shapiro–Wilk test) and homogeneity of variances (Levene’s test). Tukey’s post hoc tests were applied to identify which groups differed significantly (p < 0.05). Results Ethnobotanical uses All interviewees reported that the studied taxa are primarily used for mescal production, sap extraction for pulque, and ornamental purposes (Table 3). The var . oaxacensis is the most frequently cultivated and economically important taxon for mescal production, particularly in the Central Valleys of Oaxaca. Another highly valued variety, A. americana var. subtilis , is cultivated extensively in Jiquilpan and Sahuayo, Michoacán. Other taxa also contribute to mescal production. For example, A. americana subsp. protamericana is managed in San Carlos and Miquihuana (Tamaulipas), and A. americana var. americana is cultivated in San Carlos, Jiménez, and Jaramillo (Tamaulipas). Producers reported that plants are harvested only when the inflorescence stalk begins to develop; at this stage, the scape is cut to preserve sugars in the main stem of the Agave , which is essential for enhance fermentation and flavor quality. Pulque production is mainly associated with A. americana var. americana in Tarímbaro and Santiago Undameo (Michoacán), and with A. americana var. expansa in Ciudad Guzmán and Zacoalco (Jalisco). Additionally, A. americana var. americana is used for aguamiel extraction in Iturbide (Nuevo León). For ornamental purposes, A. abrupta is cultivated for its large, symmetrical rosette. This taxon is found along the shores of Lake Chapala in the state of Jalisco. Table 3. Study sites and Agave taxa recorded, including common names, registered uses, management types (wild or cultivated), selection criteria, and the number of populations and individuals per population evaluated. Site Taxa Common name Uses Management Selection (mechanisms, targets, reasons) Number of populations Number of individuals per population Dr. Arroyo, Nuevo León. Agave americana subsp. protamericana Maguey verde No use registered Wild 2 5 Iturbide, Nuevo León. Agave americana var americana Maguey cenizo Aguamiel extraction Cultivated The big steam its selected due to the high and constant pulque extraction. 1 Agave americana subsp. protamericana No use registered Wild 1 Agave asperrima No use registered Wild 1 Miquihuana, Tamaulipas. Agave americana subsp. protamericana Aguamiel extraction Managed in situ They select several kind of Agave taxa for the aguamiel extraction. 2 5 San Carlos, Tamaulipas. Agave americana var americana Mescal production Managed in situ This taxa are the most common and abundant in the region so people select them because it is the only option. 1 5 Agave americana subsp. protamericana Mescal production Managed in situ 1 5 Agave asperrima Maguey blanco Mescal production Managed in situ 1 5 El Tinieblo, Tamaulipas. Agave americana var. americana Maguey verde Mescal production Promoted Size of the steam 1 5 Jaramillo, Tamaulipas. Agave americana var. americana Maguey verde Mescal production Promoted Size of the steam 1 5 Zacapu, Michoacán. Agave americana var americana Maguey cenizo Ornamental Cultivated The most important is the size of the plant 1 4 Tarímbaro, Michoacán. Agave americana var americana Maguey cenizo Pulque extraction Cultivated Its selected for the quantity of sugar, the size of the steam and 1 5 Santiago Undameo, Michoacán. Agave americana var. americana Maguey cenizo Pulque extraction Cultivated 1 5 Zacoalco, Jalisco. Agave americana var. expansa Maguey cenizo Pulque extraction Cultivated It’s a taxon that produce pulque for three months 1 5 Cojumatlán, Michoacán. Agave abrupta Ornamental Cultivated The size of the plant and the few quantity of spines 1 5 Jiquilpan, Michoacán. Agave americana var. subtilis Chato de Sahuayo Mescal production Intensive Hight quantity of sugar and fast maturation 2 5 Coixtlahuaca, Oaxaca. Agave scaposa Maguey de potrero, maguey cimarrón, maguey de caballo, maguey de pitzorra o tequiol [57] Used in the construction of houses and for other local purposes (extraction of grubsfrom the plant), and occasionally for cooking. It is also used as living fences and to delimit terrain (asmarkers) [58,59] Wild Its selected due to the width of the leaves and the big size of the inflorescence. 1 5 San Baltazar Guelavila, Oaxaca. Agave americana var. oaxacensis Maguey de castilla Mescal production [60] reported for fiber use Intensive Flavor, high quantity and quality of sugar 1 5 San Pablo Huixtepec, Oaxaca. Agave americana var. oaxacensis Arroqueño Coyote Mescal production [60] reported for fiber use Intensive The flavor of the mescal its unique with this taxa, the big amount of sugar and the size of the steam. 1 5 The subspecies A. americana subsp. protamericana grows wild in the San Carlos region (Tamaulipas), where informants indicated that cultivation is recent. Locals prefer plants from the surrounding hillsides, asserting that wild individuals produce mescal with superior flavor. The studied plants are believed to have originated from wild populations established for over a century, and are thus considered genuinely wild. In Tamaulipas, mescal production based on A. americana subsp. protamericana has become a significant economic activity, sustained mainly by small-scale producers. Its growing popularity in recent decades supplements local agricultural economies (maize, sorghum, and livestock). However, the number of small production units and the extraction of reproductive Agaves from the forest has increased. No evaluation has yet been conducted in this respect, but it is undoubtedly needed to establish criteria for the sustainable use of these populations. Among all taxa, A. americana var. oaxacensis stands out for its cultural and linguistic richness. It is one of the most culturally and economically important taxa in the A. americana complex. Traditionally used for fiber extraction, it is now primarily cultivated for high-quality mescal production of high economic value. The wide range of ethnotaxa (traditional varieties recognized by local people) recognized in Oaxaca reflects both morphological variation and deep cultural knowledge. Common names include maguey arroqueño, maguey ruqueño, maguey cenizo, maguey blanco (“pale Agave ”), and maguey sierrudo or maguey sierra negra (“saw-leaf Agave ”). In Mixtec, it is called yavi cuaam, in Spanish maguey coyote, maguey xolo, maguey de rayo, and in Zapotec it bears names such as doba nupy, dòb-dzìn, dòb mpiè, dobncjp, dua-bsug, dua-bzog, dua-pchez, dua-ya-do, and dua-yesh. This linguistic diversity reflects the long-standing cultural relationship and morphological differentiation of local forms, which merit focused ethnobotanical and genetic research. Management Practices The surveyed Agave populations exhibited a broad spectrum of management regimes, reflecting a domestication gradient ranging from wild to fully cultivated forms (Table 3): Wild populations – e.g., A. asperrima, A. scaposa, A. americana subsp. protamericana and subsp. americana , growing in xeric scrublands and rocky slopes. These plants regenerate naturally and are opportunistically harvested. They maintain strong defense traits (spines, thick cuticles) and high morphological uniformity. Managed in situ populations – predominantly A. americana subsp. americana and A. scaposa , are tolerated or let standing when clearing land, as well as protected in agricultural and fallow plots. Plants are not actively propagated but are occasionally transplanted from forest or disturbed areas to agricultural plots and protected, showing slight selection favoring straight and vigorous scape and variable morphology. Promoted populations – such as A. americana var. oaxacensis and certain variants of A. americana subsp . americana are composed by semi-wild individuals that are transplanted or selectively favored, particularly for pulque or fencing uses. These Agave s exhibit intermediate phenotypes, suggesting early domestication stages. Cultivated populations – composed mainly of A. americana var. americana, var. expansa , and subsp. protamericana , grown in agroforestry systems or small plantations. Propagation is primarily vegetative, and selection favors large stem size, high sap yield, and reduced thorniness. Intensively managed populations – such as A. americana var. subtilis (Michoacán) and A. americana var. oaxacensis (Oaxaca). These are maintained in monocultures and exhibit strong domestication syndromes, including increased vegetative size and reduction of defense structures. The geographic distribution of management systems correlates strongly with species identity and regional cultural traditions, emphasizing the co-evolutionary nature of human– Agave interactions throughout Mexico. Morphological patterns Reproductive and vegetative characters analysed in herbarium specimens Principal Component Analysis (PCA) (Fig 3.) revealed significant patterns of morphological variation among the studied Agave species. The first two principal components accounted for 64.6% of the total variance (PC1 = 41.9%, PC2 = 22.7%). These components explained the main axes of morphological differentiation among the taxa. The variables with the highest loadings on PC1 were terminal thorn length (TTL), teeth length (TEEL), and the ratio TEEL/LL (teeth-to-leaf length). On PC2, the variables with the greatest contributions were teeth width (TEEW), distance between teeth (DBTEE), TEEL/TEEW (teeth length-to-width ratio), DBTEE/LL (distance between teeth-to-leaf length ratio), and TTL/TTW (terminal thorn length-to-width ratio). The ordination revealed clear segregation of the taxa along PC1 and PC2 based on these morphological variables . Agave americana var. americana, A. a. var. oaxacensis, and A. a. subsp. protamericana were positioned toward the positive side of PC1, while A. asperrima and A. scaposa were located on the negative side. Along PC2, A. a. subsp. protamericana and A. a. var. oaxacensis were grouped toward the positive quadrant, whereas A. scaposa and A. asperrima were placed toward the negative quadrant. The distribution pattern indicated that PC1 represented the axis of greatest morphological differentiation among the taxa, mainly influenced by the variables TTL, TEEL, and TEEL/LL, while PC2 captured additional variation explained by TEEL/TEEW, DBTEE, and TTL/TTW. Linear Discriminant Analysis (LDA) LDA revealed clear morphological structuring among the analysed Agave taxa according to geography, management regime, and taxonomic identity (Fig. 4). When grouped by geographic origin (north, centre, and south) (Fig.4b), southern taxa—particularly A. americana var. oaxacensis —formed a distinct cluster separated from northern and central groups. Northern taxa, including A. asperrima and A. americana subsp. americana , were characterized by elevated values of terminal thorn width (TTW), teeth length (TEEL), and the ratio TTL/LL (terminal thorn-to-leaf length). Central taxa, represented mainly by A. scaposa , exhibited intermediate or overlapping morphological profiles. Discriminant analysis based on management regime showed well-defined segregation between cultivated, intensively managed, and wild populations. Individuals under intensive management, particularly those from Oaxaca, were associated with higher values of inflorescence branching and reduced spine-related variables. Wild accessions, notably A. asperrima , displayed compact rosettes and higher thorn and tooth measurements. When grouped by taxonomic identity, the discriminant functions successfully differentiated the species and varieties within the Agave americana complex. The resulting morphological clusters were consistent with their formal taxonomic assignments, confirming the discriminant capacity of the evaluated variables. HEAT MAP ANALYSIS (Fig. 5) The A. americana var. oaxacensis group exhibited moderate vegetative and defensive traits but strong reproductive investment, with medium-sized, narrow leaves and intermediate thorn and tooth dimensions. High values for inflorescence length and ovary width indicated advanced reproductive development and a clear emphasis on seed production. The A. americana subsp. protamericana group showed moderate vegetative growth, strong defensive features, and intermediate reproductive effort. Agave s of the subspecies protamericana from Zacatecas displayed the most robust morphology—large leaves, thick thorns, and broad teeth—while other populations maintained moderate defence and reproductive traits. This group is defined by structural robustness and enhanced defence, particularly in the Zacatecan populations. The A. americana group exhibited wide morphological variability, combining vigorous vegetative growth with moderate-to-strong defences and diverse reproductive investment. A. americana Saltillo1 and Coahuila showed the largest leaves and strong defences, while Coahuila also had the largest inflorescences and ovaries, marking the highest reproductive investment. Overall, the group is characterized by large plant size, strong defence, and variable reproductive effort. The A. asperrima group displayed balanced vegetative and reproductive traits with well-developed defensive morphology. A. asperrima Zacatecas had the longest and thickest thorns and teeth, representing the strongest defence, while other populations showed intermediate values. Reproductive traits were consistently moderate across all taxa, defining the group as morphologically balanced with notable defensive strength. The A. scaposa group presented moderate differentiation, with balanced vegetative growth and intermediate defence. A. scaposa Suchixtlahuaca had the largest leaves and slightly greater reproductive investment, while A. scaposa Puebla and Puebla1 maintained smaller leaves and moderate defensive traits. Overall, the group exhibits intermediate morphology with modest variability among populations. Finally, A. lurida Oaxaca and A. a. var. marginata_ CDMX showed compact vegetative forms and moderate defence and reproduction. A. lurida possessed small leaves and slender spines but stood out for its larger ovary, reflecting higher reproductive investment. A. a. var. marginata exhibited narrow leaves and intermediate structural and reproductive traits. Both represent stable morphologies within the A. americana complex, balancing defence with reproductive efficiency. The integrated multivariate analyses provided a comprehensive understanding of the morphological diversity present among Agave species herborized, populations, and management types. Principal component analysis (PCA) (Fig. 3, Table. 5) revealed that traits such as leaf width (LW), LL/SL (leaf length/stem length ratio), and the teeth length/leaf length ratio, teeth length/leaf length and teeth width/leaf length (TEEL/LL, TEEW/LL) ratios were key contributors to the main axes of morphological variation. These variables defined a spectrum ranging from robust, large-leafed taxa to smaller, more dentate forms. Taxonomically defined groups exhibited varying degrees of cohesion in PCA space. For instance, A. asperrima occupied a distinct region, whereas A. americana subsp. americana , A . americana subsp . protamericana, and A. expansa were more broadly distributed and often overlapped with each other and with other taxa. A. americana var . oaxacensis and A. scaposa displayed greater variability. Individuals of A. americana var. oaxacensis were widely dispersed in the PCA space and split across dendrogram branches, whereas populations of A . scaposa clustered more consistently. Linear discriminant analysis (LDA) (see Fig. 4) by geographic area demonstrated strong regional structuring of morphology. Samples from the southern region formed a distinct cluster, separated along both LD1 and LD2 axes, suggesting the presence of unique morphological characters. Samples from central and northern regions displayed partial overlap, albeit with distinguishable sub-clusters, indicating that geography plays a significant role in shaping the Agave americana complex, potentially through local selection pressures or environmental constraints. Analysis by management type (wild versus cultivated) suggest evidence of domestication-related morphological patterns. Cultivated individuals formed a cohesive and relatively compact cluster, indicating reduced morphological variability and a tendency towards standardized traits. These included broader leaves and lower dentition, which are favored for harvest and processing efficiency. In contrast, the trait SL (stem length) in wild specimens exhibited more dispersed pattern in LDA space, reflecting higher phenotypic variability, which is likely to result from natural selection and genetic diversity. This supports the idea of a “domestication syndrome” in Agave , whereby human influence reduces diversity in favour of functional traits. Finally, LDA based on taxonomic assignment revealed that A. asperrima exhibited significant morphological diversity, with minimal overlap with other taxa. By contrast, subsp. americana , subsp. protamericana , and var. expansa exhibited substantial overlap in the discriminant space, underscoring the limitations of morphology in resolving species boundaries within this complex. These results are consistent with previous studies suggesting extensive hybridisation, local domestication, or shared ancestry among these taxa. Overall, the combined evidence from PCA, hierarchical clustering and LDA highlights the multifactorial nature of Agave morphological diversity, which is shaped not only by taxonomy but also by geography and human management. Based on characters evaluated in herborized specimens, the cluster analysis had a cophenetic test r= 0.747. This clearly groups A. asperrima, A. scaposa, and A. lurida separated from subsp. americana , in accordance with their recognised taxonomic identity (Figure 3). The complex Agave americana can be seen to include three differentiated groups: one of them including var. americana and var. marginata with A. americana subsp. protamericana from San Luis Potosí; a second group is formed by var. americana and var. oaxacensis grouped with subsp. protamericana specimens from Nuevo León; and a third group discriminating var. oaxacensis and var. expansa . The four varieties of A. americana are not clearly discriminated. Unfortunately, this study based on herbarium specimens does not provide information on differentiation associated with management, since this information is absent from the specimen’s labels. We can only record three types of practice: wild, cultivated and intensively cultivated. Eigenvectors indicate that the characters with the highest contribution to the classification in the first principal component are the number of branches, filament of stamens, length of the ovary, the ratio of the terminal spine and the leaf length. In the second principal component the most relevant characters were the anther length, the ratio of teeth width and leaf length (WTEE/LL), the ratio of terminal thorn length and leaf length (TTL/LL), the ratio of teeth length and leaf length (LTEE/LL), the ovary width and length and number of branches. Table 4. Results of the multivariate analysis of variance (MANOVA) for the variables Area, Management, and Taxonomy. The table shows Wilks’ Lambda values, Chi-square statistics, degrees of freedom (DF), and corresponding p-values for each variable. Analyses were performed using two datasets: H, specimens measured from herbarium material, and F, specimens measured directly in the field. P-values lower than 0.05 indicate statistically significant differences among the analyzed categories. FUNCTION WILKS´LAMBDA CHI SQUARE DF P VALUE AREA H 3.5942e-05 86.98 32 0.09173 F 0.00101 615.38 76 2.2e-16 MANAGEMENT H 1.8139e-05 92.79 32 0.06601 F 0.0055 487.24 76 2.2e-16 TAXONOMY H --------------- ----------------- 64 ------------ F 0.00278 734.63 133 2.2e-16 We conducted three analyses using the LDA to examine the role of the area of distribution, the taxonomic classification of the taxa, and the associated management to each taxon. 1) The LDA analysis for area has a small lambda value of 0.0193, which is significant at a p value of 0.091. The first function explains 91.2% of the variance, whereas the second function contributes significantly less to the discrimination process. 2) The LDA analysis for management has a small lambda value of 1.8139e-05. The first function explains 96.1% of the variance, whereas the second function does not contribute significantly to the discrimination process as compared to that of the first function. 3) The taxonomy analysis for does not provide a lambda value because there are few samples to analyze. The first function explains 81.33% of the variance, while the second function contributes with the 15.05%. Vegetative characters measured in the field The Principal Component Analysis (PCA) (Fig. 6, Table. 5) revealed the main axes of morphological variation among species within the Agave americana complex, explaining 43.6% of the total variation through the first two components (PC1 = 25.3%, PC2 = 18.3%). PC1 was primarily driven by variables related to leaf shape and defensive traits, including the teeth length/leaf length ratio (TEEL/LL), leaf length/width ratio (LL/LW), and distance between teeth (DBTEE). This axis separated species with long, narrow leaves and well-developed teeth (e.g., A. a. var. expansa , A.a. var. oaxacensis y A. a. var. subtilis ) toward the positive side, from those with broader leaves and smaller teeth (e.g., A. abrupta, A. a. var . americana, A. asperrima ) toward the negative side. PC2 was mainly associated with terminal thorn length and width (TTL, TTW) and plant diameter (D). Species with longer thorns and larger diameters, such as var. oaxacensis and var. subtilis , were positioned on the positive side of the axis, while species with shorter thorns and more compact forms, such as A. a . subsp. protamericana and A. a . subsp. americana , occupied the negative side. Overall, the PCA showed a clear separation between robust, defensively adapted species ( A. asperrima, A. scaposa, A. a. subsp. protamericana ) and those with greater vegetative and reproductive investment ( A. a. var . oaxacensis, A. a. var . subtilis ,), while intermediate forms (A. a. subsp. americana , A. a. var . expansa ) clustered in central positions. The variables with the greatest discriminant weight were TEEL/LL, LL/LW, DBTEE, TTL, and TTW, confirming that leaf morphology and defensive features are the main factors structuring variability within the Agave americana complex. The heatmap based on vegetative and defensive traits (Fig. 7) revealed clear morphological differentiation among taxa of the Agave americana complex. The most influential variables included tooth length and width (TEEL, TEEW), terminal thorn dimensions (TTL, TTW), leaf proportions (LL, LW, LL/LW), plant diameter (D), and total plant length (PL), while reproductive traits were not considered in this analysis. The A. americana group exhibited moderate to strong defensive development, particularly in cultivated and managed forms, where high TEEL/LL and TTL/LL ratios reflected proportionally long teeth and terminal thorns. The A. protamericana taxon displayed the most robust morphologies, especially in wild populations, with large leaves and highly developed spines indicating strong structural defense. The var. oaxacensis and var. subtilis were characterized by elongated, narrow leaves and intermediate thorn expression, suggesting balanced vegetative and defensive investment. A. expansa showed high values for TEEL/LL and TTL/LL, emphasizing pronounced defensive structures, while A. scaposa exhibited moderate vegetative size and notable thorn development, with long terminal spines relative to leaf length. The A. asperrima , particularly wild populations, presented the most pronounced defensive traits, with long and thick thorns and high TTL/LL ratios, whereas managed individuals displayed slightly greater vegetative development. Overall, the heatmap analysis indicated that morphological variation within the Agave americana complex is primarily driven by defensive and vegetative attributes, particularly those related to spines, teeth, and leaf architecture, delineating adaptive strategies that balance protection and growth across taxa. Wild taxa exhibit the most robust defensive morphologies. Hierarchical clustering of the rows (groups) in the heatmap revealed morphologically coherent clusters, that often align with known taxonomic distinctions or management regimes. The dendrogram on the x-axis grouped variables with similar expression profiles across groups. For example, TTL (terminal thorn length) and TTW (terminal thorn width) cluster closely together, confirming their strong positive correlation, which was also observed in the PCA. Linear Discriminant Analysis (LDA) (Fig. 8) was performed to explore the morphological diversity of Agave taxa across three key factors: taxonomy, management, and region. The three separate plots reveal the following: Taxonomy (top panel). The first LDA plot (Fig. 8A) reveals clear differentiation between species based on LD1 (the horizontal axis) and LD2 (the vertical axis). The species A. americana var. oaxacensis and the wild group A. americana subsp. americana , A. americana subsp. protamericana and A. asperrima cluster in different areas of the plot, indicating distinct morphological profiles and unique trait compositions related to their vegetative structures. A. scaposa and A. expansa are also grouped separately, reflecting their particular adaptive features and differentiating them from other species. Management (middle panel) ( Fig 8B ). Management regimes are clearly distinct. Cultivated individuals (CULT) cluster in the top-right quadrant, while the intensively managed (INTEN), semi-wild (PROMOTED), and wild (WILD) groups are separated along LD1, with the wild types farthest to the left. This pattern confirms that cultivation is associated with a suite of morphological changes consistent with a domestication syndrome. Region (bottom panel) (Fig 8C). The third LDA plot shows the influence of geographic origin on the morphological variability of Agave . Northern populations (NORTH) cluster to the right along LD1, distinct from the Michoacán (MICH), Jalisco and Oaxaca samples on the left. This suggests that regional agroecological and cultural factors influence morphological divergence, potentially reinforcing localized selection practices, possibly due to environmental factors such as temperature, precipitation, and soil conditions shaping their growth and reproductive characteristics. Table 5. Characters evaluated in herbarium specimens and in the field and eigenvectors of the first and second principal components and discriminant functions. Vegetative Herbarium specimens Character PC1 PC2 DF1 DF2 PC1 PC2 DF1 DF2 Plant length -0.2166 -0.3632 0.3847 -0.3750 - - - - Diameter -0.4013 -0.1169 0.2502 -0.5270 - - - - Stem length -0.2469 -0.2997 -1.1303 0.6912 - - - - Leaf length -0.3624 -0.2564 -0.4715 3.5562 0.2854 -0.1131 -0.8631 -0.6157 Leaf width -0.1056 -0.3815 0.5092 -1.0412 0.2385 0.0654 0.4060 0.4022 Terminal thorn length 0.1354 -0.1991 -1.7469 0.2153 -0.2711 -0.0786 0.3208 -0.0574 Terminal thorn width -0.0747 -0.1933 -0.0710 0.2293 0.2468 0.2250 -1.2081 0.9294 Number of leaves -0.2946 0.1351 1.1925 0.3065 - - - - Teeth length 0.1710 -0.3112 -1.2907 -1.2840 -0.2258 0.1511 -0.4106 0.8714 Teeth width 0.0672 -0.3393 -0.2407 0.3690 0.1827 0.093 0.6154 -1.2731 Distance between teeth 0.1180 -0.0085 0.0507 0.3979 0.0133 0.2275 -0.2652 0.1553 Leaf length/width ratio -0.2529 0.1931 1.9352 -2.3732 0.1015 -0.1549 -0.7995 -1.4925 Leaf/stem length ratio -0.0578 0.1224 -0.8731 -0.3545 - - - - Terminal thorn length/width ratio 0.1959 -0.1539 -0.2083 0.8316 0.2025 0.9294 0.2025 0.9294 Terminal thorn/leaf length ratio *** 0.2124 -0.1751 1.7553 -0.9657 1.2403 -1.3071 1.2403 -1.3071 Teeth/leaf length ratio -0.2002 0.3104 3.6414 1.5015 1.0706 -1.2204 1.0706 -1.2204 Teeth width/leaf length 0.3735 -0.1594 0.7649 1.2663 0.0462 -1.3572 0.0462 -1.3572 Distance between teeth/leaf length ratio 0.31534 0.1316 -0.0611 -0.5001 -0.0111 -0.3907 -0.0111 -0.3907 Inflorescence length -0.0948 -0.1676 -0.0948 -0.1676 Ramification number -2.1962 -1.1654 -2.1962 -1.1654 Ovary length 1.36 -1.1042 1.360 -1.104 Ovary width 0.25 -1.2011 0.250 -1.2011 Tepal length -0.4911 0.3617 -0.4911 0.3617 Filament length 1.467 0.2 1.467 0.1991 Anther length 0.9401 -1.6291 0.9401 -1.6249 Stamen length -1.0104 0.8 -1.0104 0.7998 Total flower length 0.3424 -0.2506 0.3424 -0.2506 Table 6. Averages and standard errors of 19 characters that presented significant differences when comparing categories wild, promoted, managed in situ, and cultivated. Character wild promoted Managed in situ cultivated Plant height*** 1.865 ± 0.463b 2.296 ± 0.584bc 2.336 ± 0.632c 2.557± 0.782c Stem length*** 0.339 ± 0.125ab 0.406 ± 0.069b 0.406 ± 0.070b 0.634 ±0.125c Diameter*** 1.348 ± 0.526a 1.583 ± 0.276ab 1.686 ± 0.474b 2.545 ±0.445c Leaf length*** 0.884 ± 0.257a 1.210 ± 0.170b 1.231 ± 0.335b 1.637 ±0.312c Leaf width*** 0.220 ± 0.041bc 0.182 ± 0.039b 0.216 ± 0.049b 0.245 ±0.033c Leaf length/width ratio *** 4. 059 ± 1.122a 6.985 ± 2.028b 5.701 ± 1.055b 6.754 ±1.498b Leaf/stem length ratio 6.134 ± 7.505a 3.033 ± 0.504a 3.023 ± 0.652a 2.665 ±0.708a Number of leaves*** 31.407± 9.604a 38.846±10.573ab 40.896±10.896b 51.958 ±18.044b Terminal thorn length*** 0.042 ± 0.010c 0.143 ± 0.207d 0.047 ± 0.012cd 0.027 ±0.005b Terminal thorn width* 0.004± 0.001ab 0.006± 0.003ab 0.008 ± 0.008b 0.006 ±0.003b Terminal thorn length/width ratio *** 9.796 ± 2.390bc 19.961 ± 19.409c 7.942 ± 3.727b 5.360 ±3.266a Terminal thorn/leaf length ratio *** 0.051 ± 0.016b 0.113 ± 0.152b 0.040 ± 0.010b 0.017 ±0.005a Teeth length*** 0.011 ± 0.003bc 0.012 ± 0.001c 0.011 ± 0.003bc 0.009 ±0.004b Teeth/leaf length ratio *** 0.286 ± 0.720b 0.006 ± 0.001a 0.006 ± 0.002a 0.721 ±0.506b Teeth width*** 0.007 ± 0.002b 0.007 ± 0.002b 0.007 ± 0.002b 0.008 ±0.002b Teeth length/width ratio *** 0.723 ± 0.204a 0.611 ± 0.192a 0.691 ± 0.202a 1.065 ±0.453b Teeth width/leaf length *** 0.013 ± 0.006d 0.010 ± 0.001cd 0.009 ± 0.002c 0.005 ±0.001b Distance between teeth 0.018 ± 0.009a 0.024 ± 0.011a 0.019 ± 0.008a 0.018 ±0.007a Distance between teeth/leaf length ratio *** 0.023 ± 0.013b 0.019 ± 0.008b 0.017 ± 0.007b 0.012 ±0.006a Different letters indicate significant differences. Significant level 0.001***, 0.01** and 0.05* The multivariate analyses PCA (Fig.5), LDA (Fig.8), and Heatmap (Fig.7) consistently revealed clear morphological differentiation within the Agave americana complex based on vegetative and defensive traits. The PCA (Fig.6) showed that variables such as teeth length (TEEL), teeth-to-leaf length ratio (TEEL/LL), leaf length-to-width ratio (LL/LW), and terminal thorn dimensions (TTL, TTW) contributed most to the observed variation, separating taxa according to their leaf morphology and defensive structures. The LDA (Fig. 8) further distinguished individuals by taxonomy, management, and region: taxonomic groups ( A. americana var . oaxacensis, A. americana subsp . protamericana, A. asperrima, and A. scaposa ) formed distinct clusters; cultivated, managed, and wild categories were clearly separated along discriminant axes; and northern populations differed morphologically from central and southern ones. The Heatmap (Fig.7) supported these patterns, highlighting higher values of thorn and teeth ratios in wild taxa ( A. asperrima, A. a. subsp . protamericana ), while cultivated and managed populations of A. a. var. americana and A. a. var oaxacensis exhibited larger leaves and moderate defensive traits. Overall, the combined analyses indicate structured morphological variation across species, management regimes, and regions within the A. americana complex. The multivariate analysis of variance (MANOVA) revealed differences among the analyzed categories for the evaluated morphological variables (Table 4). When comparing datasets obtained from herbarium specimens (H) and field-measured specimens (F), contrasting patterns were observed. For the Area factor, herbarium specimens did not show statistically significant differences (p = 0.09173), whereas field measurements revealed highly significant differences among groups (Wilks’ λ = 0.00101; χ² = 615.38; DF = 76; p < 0.001). A similar pattern was detected for the Management factor: herbarium specimens showed marginally non-significant differences (p = 0.06601), while field measurements indicated strong statistical significance (Wilks’ λ = 0.0055; χ² = 487.24; DF = 76; p < 0.001). For the Taxonomy factor, only field-measured specimens were analyzed, revealing highly significant differences among taxa (Wilks’ λ = 0.00278; χ² = 734.63; DF = 133; p < 0.001). These results indicate that morphological variation among taxa is more clearly detected when measurements are obtained directly from field specimens. Overall, the analyses suggest that field measurements capture a greater proportion of morphological variability than herbarium-based measurements, likely due to preservation effects or measurement constraints associated with herbarium material. These results are consistent with the multivariate patterns observed in the PCA and heatmap analyses, which revealed clear grouping of species and differentiation associated with several floral and vegetative traits. Morphological variation among taxa of the Agave americana complex Mean values and standard errors of the morphological traits that showed significant differences among taxa are presented in Table 7. Nineteen characters exhibited statistically significant variation among the analyzed taxa (p < 0.001 or p < 0.01), including vegetative size variables, leaf architecture, and defensive structures. Overall, taxa belonging to the Agave americana complex showed substantial differences in plant size. A. americana var. americana exhibited the largest mean values for plant length (up to 2.922 ± 0.841) and plant diameter (2.615 ± 0.527), whereas smaller values were generally observed in A. asperrima and var. oaxacensis . Intermediate values were recorded for A. asperrima and A. americana subsp. protamericana . These patterns indicate clear morphological differentiation in overall plant size among taxa. Stem length also varied significantly among groups. The longest stems were observed in A. americana var . expansa (0.658 ± 0.079), whereas shorter stems were characteristic of var. oaxacensis and A. asperrima . Differences in stem length contributed to the variation observed in plant architecture among taxa. Leaf morphology showed marked variation as well. Leaf length ranged from 0.893 ± 0.254 in A. americana subsp . protamericana to 1.778 ± 0.324 in A. americana var . americana . Similarly, the leaf length-to-width ratio differed significantly among taxa, reflecting differences in leaf shape and proportions. Taxa such as A. oaxacensis exhibited particularly high LL/SL values, indicating relatively elongated leaves compared with other species in the complex. The number of leaves per plant also differed significantly among taxa. Higher values were recorded in A. americana var . expansa and var . oaxacensis , whereas lower values were observed in A. asperrima . These differences suggest variation in vegetative growth strategies across the complex. Defensive traits, including terminal spine length and width, showed notable variation among taxa. Prominent terminal spines were observed in A. americana var. abrupta , whereas smaller spines were characteristic of var . oaxacensis . Ratios involving terminal spine dimensions (e.g., TTL/LTL and TTL/LL) also differed significantly, indicating differences in the relative development of defensive structures. Traits related to marginal teeth—including teeth length, teeth width, and the distance between teeth—showed significant but comparatively moderate variation among taxa. Some taxa, such as A. abrupta , exhibited longer teeth relative to leaf size, whereas others presented shorter and more widely spaced teeth. Overall, these results demonstrate substantial morphological differentiation within the Agave americana complex and associated species. The significant variation observed across multiple vegetative and defensive traits supports the taxonomic distinctiveness of the analyzed taxa and reflects the morphological diversity present within the group. Significant differences were also detected among management categories (wild, promoted, managed in situ, and cultivated) for most of the evaluated morphological traits (Table 6). In general, cultivated plants exhibited larger vegetative structures than those observed in wild populations. Cultivated individuals showed the highest mean values for plant height (2.557 ± 0.782), stem length (0.634 ± 0.125), plant diameter (2.545 ± 0.445), and leaf length (1.637 ± 0.312), indicating a clear increase in overall plant size under more intensive management regimes. Intermediate values were generally observed in the promoted and managed in situ categories. Traits such as plant height and plant diameter displayed a gradual increase from wild populations to promoted and managed plants, suggesting progressive morphological modification along the management gradient. In contrast, some variables, including leaf width and the leaf/stem length ratio, showed little variation among management categories. Leaf architecture also differed among management types. Cultivated plants exhibited higher values of the leaf length-to-width ratio, indicating proportionally longer leaves, whereas wild plants generally had smaller leaves. In addition, the number of leaves was highest in cultivated individuals, suggesting that cultivation practices may promote enhanced vegetative growth. Defensive traits also varied across management categories. Terminal spine length was greatest in promoted plants (0.143 ± 0.207), whereas cultivated plants showed the lowest values (0.027 ± 0.005). Similarly, the terminal spine length-to-width ratio was highest in promoted individuals, reflecting relatively elongated spines under intermediate management conditions. By contrast, cultivated plants tended to exhibit smaller and proportionally shorter terminal spines. Traits associated with marginal teeth showed comparatively minor variation among categories. Both teeth length and teeth width remained relatively similar across management types, although slight differences were detected in the teeth-to-leaf length ratio, with cultivated plants presenting the highest values. Discussion The domestication of Agave represents a complex evolutionary process shaped by interactions among morphological traits, ecological adaptation, and human management. The morphological diversity observed across species reflects both environmental specialization and varying degrees of anthropogenic selection. This study contributes to understanding Agave domestication by integrating vegetative and reproductive morphometrics from herbarium and field specimens, revealing how natural and cultural factors have guided morphological divergence and adaptation. Multivariate analyses demonstrated consistent patterns of differentiation within the Agave americana complex. The PCA (Fig. 3 ) indicated that reproductive traits—particularly inflorescence length (IL), ovary width (OW), and stamen length (SL)—were major contributors to variation, defining a gradient from high reproductive investment to enhanced vegetative and defensive robustness. A. americana var. oaxacensis showed high reproductive development (positive PC1), while A. a. subsp. protamericana and A. asperrima aligned with greater thorn and tooth dimensions (negative PC1/PC2), suggesting stronger defensive allocation. A. a. subsp. americana occupied an intermediate position, combining large vegetative structures with moderate reproductive traits, and A. scaposa and A. expansa exhibited balanced morphologies. These patterns confirm that reproductive differentiation is a principal axis of species segregation within the complex [ 62 , 63 ]. The LDA (Fig. 4 ) reinforced these distinctions, clearly separating taxa, management regimes, and geographic regions. Cultivated and intensively cultivated individuals clustered apart from wild and managed populations, displaying larger inflorescences and ovaries but reduced thorn and tooth size—morphological signatures consistent with domestication. In contrast, wild and managed plants retained smaller reproductive organs and stronger defensive traits. Regionally, northern populations differed from those in Oaxaca, Michoacán, and Jalisco, suggesting that ecological conditions and local management practices jointly drive morphological divergence across the A. americana complex. The heatmap (Fig. 5 ) summarized these patterns, highlighting the trade-offs between reproductive investment and structural defense. A. americana var. oaxacensis exhibited the highest reproductive values (IL, OW), A. a. subsp. protamericana displayed enhanced defensive structures (long, thick terminal thorns, prominent teeth), and A. asperrima combined robust defensive and vegetative development with moderate reproduction. A. scaposa and A. expansa showed intermediate values across all traits, suggesting morphological equilibrium in these traits. An important finding of this study is the importance of reproductive traits, particularly floral morphology, in Agave species differentiation. Floral features such as stamen and filament length, ovary dimensions, and inflorescence size emerged as key discriminant variables. These results align with Trejo et al. [ 61 ], who identified floral morphology as the most effective character set for species delimitation in Agave , surpassing fruit and seed traits. The persistence of floral variability under domestication is vital for the resilience and adaptive capacity of Agave populations. Even in cultivated and clonal stands, floral diversity remains a strong axis of variation, crucial for maintaining ecological stability under pressures such as climate change and pollinator decline [ 61 , 62 ]. This reproductive diversity supports long-term adaptability and evolutionary potential within the A. americana complex [ 63 , 64 ] Complementary analyses of field-measured specimens revealed clear differentiation based on vegetative traits. The PCA (Fig. 6 ) emphasized the influence of leaf morphology, spine and tooth dimensions, and plant diameter on species segregation. PC1 captured variation associated with leaf form and defensive traits, consistent with studies identifying spines and teeth as key domestication indicators [ 65 , 66 ]. PC2, linked to terminal thorn dimensions and plant diameter, distinguished species with large, robust morphologies ( A. protamericana, A. asperrima ) from more compact, human-managed forms ( A. americana ). This trend supports domestication patterns characterized by reduced defensive traits and enhanced vegetative growth [ 65 , 66 ]. LDA results based on field specimens also demonstrated strong separation among taxa, management regimes, and regions. Cultivated individuals exhibited trait combinations characteristic of a domestication syndrome [ 20 ], while regional clustering indicated the influence of both artificial selection and local environmental factors such as temperature, precipitation, and soil conditions [ 21 ]. The heatmap corroborated these distinctions: wild species ( A. asperrima, A. protamericana ) exhibited pronounced defensive features, while cultivated and managed populations showed larger leaves and reduced spine development [ 62 , 63 ]. The morphological differences observed between species in the Agave americana complex have significant ecological and evolutionary implications. Wild species, exposed to greater herbivory pressure and environmental stressors, have developed robust defense strategies, such as long spines and thick leaves, to protect themselves from predators and competitors. In contrast, managed species, which are under human control, exhibit traits that prioritize vegetative growth and biomass accumulation, such as larger plant sizes and greater leaf production. These traits are selected during domestication to meet human needs, such as food production or fiber harvesting [ 69 , 70 ]. The human-directed evolution through management practices such as cultivation and breeding has favored traits that enhance the usefulness of these plants, but it has also shaped their genetic and morphological diversity. Wild and less-managed species have retained a repertoire of defensive characteristics that enable them to survive in more competitive natural environments. This morphological diversity could be crucial for future conservation strategies, as cultivated and managed populations may be more vulnerable to rapid environmental changes, while wild populations may offer a genetic foundation for future adaptation to new ecological conditions [ 71 , 72 ]. Integrating multivariate morphometries provides a robust framework for examining the domestication of Agave americana . By analyzing vegetative and reproductive morphological traits, clear patterns of differentiation can be established among wild, cultivated, and promoted forms. In particular, traits such as plant size, leaf characteristics, and the size of terminal spines and teeth are key indicators of human management influence. Wild plants tend to be smaller, with fewer leaves and larger terminal spines and teeth, while cultivated plants exhibit intermediate characteristics. Those managed in situ display traits that are closer to those of cultivated forms. These patterns, which have been documented in other plant species [ 65 , 66 ], reinforce the idea that domestication can occur in both cultivated plants and wild populations that have been incorporated into human-managed environments. Integrating morphological data with cultural and ecological knowledge is crucial for developing effective conservation strategies for Agave species. The study emphasizes the importance of maintaining genetic diversity through both sexual and asexual reproduction, and highlights the need to incorporate seed production into management practices. Agave populations that rely on clonal propagation, in particular may experience a reduction in genetic diversity, which could limit their ability to adapt to changing environmental conditions. Therefore, preserving sexual reproduction is essential for maintaining the long-term viability of these plants. Furthermore, research into the reproductive biology of Agave species, particularly focusing on hybridization events and gene flow between wild and cultivated populations, would provide valuable insights into the evolutionary processes shaping these plants. Overall, these results (Table 4 and Table 6 ) indicate that management intensity influences several vegetative and defensive traits. Cultivated plants generally develop larger vegetative structures and reduced spine dimensions compared with wild populations, reflecting the effects of human selection under cultivation. Recommendations for Future Research The Agave americana complex exhibits significant morphological diversity, driven by both natural and human-induced evolutionary forces. The multivariate analyses provide a deep understanding of how taxonomic factors, geographic conditions, and management regimes have shaped the vegetative, defensive and reproductive traits of these species. This knowledge is essential for sustainable management and conservation of Agave species, offering new perspectives on how these plants adapt to selective pressures and how conservation efforts can support their survival under changing conditions [ 73 , 70 ]. To advance our understanding of Agave domestication, future studies should focus on expanding the geographic and ecological scope of research to include a broader range of taxa and populations. Sampling should be increased among fertile individuals and collaboration with local communities should be established to document traditional management practices. Genetic analyses, particularly the use of next-generation sequencing techniques and phylogeographic approaches, would be crucial for answering questions about the origin and diffusion of Agave taxa. Such studies will also shed light on the evolutionary relationships between species and the diffusion routes of Agave domestication throughout history, particularly in relation to Agave cordillerensis and its connection to the A. americana complex [ 74 , 75 ] . Conclusion In summary, this study deepens our understanding of the complex processes underlying the domestication of Agave . The interplay between morphological traits, management practices, and reproductive characteristics sheds light on how these plants have evolved under both natural and anthropogenic selection pressures. Integrating morphological, ecological, and genetic data, alongside cultural knowledge, is essential to understanding the full scope of Agave domestication and its implications for conservation and sustainable management. Future research should continue to explore the genetic diversity, management practices, and reproductive biology of Agave species paying particular attention to the role of floral traits in species delimitation and ecological adaptability. Our findings complement and extend the analysis of Agave domestication of the morphological diversity [ 61 ] demonstrating that reproductive characters, particularly floral traits, play a pivotal role in the organization of Agave diversity. The persistence of floral variability under domestication, the phenotypic differentiation it enables among taxa, and its potential contribution to ecological adaptability highlight its importance as both a taxonomic tool but and a conservation priority. Together, the two datasets advocate for the continued integration of floral morphology in the study, management, and protection of Agave genetic resources. The identified patterns are not conclusive; further details of management practices are required. The people of Mexico have managed Agave s for a long time for multiple purposes, so it is important to document these details to improve our understanding of domestication. Morphometric studies should be conducted at more sites, to cover the distribution ranges of the taxa included in this analysis. While it is difficult, it is not impossible to broaden the sampling of fertile individuals is indispensable, but it requires agreements with people who make use of these Agave s. Studies of reproductive biology would help to identify hybridisation events and how this process occurs. Finally, population genetics studies are needed to evaluate the state of genetic diversity and gene flow among wild and managed populations, as well as among the different analysed taxa. Phylogeographic studies would be particularly useful for answering questions about the origin and spread of the taxa that make up Agave americana . The question of whether the domesticated taxa have one or more ancestors remains unanswered. Phylogeographic approaches using next-generation genetic markers could help to identify the evolutionary relationships among the taxa, confirm or refute their identities, and establish the possible diffusion routes throughout history. Of particular interest is the South American taxon Agave cordillerensis , which was long considered to be A. americana . This Agave species is widely distributed in the Andean region, from Colombia to Bolivia, but it does not appear to have been present for long time. The evolutionary relationship between this Agave species and the A. americana complex is an interesting question that could be answered that could be addressed through phylogeographic approaches providing valuable insights into the historical exchange of crops and domestication techniques between Mesoamerica and the Andean region. This is merely an initial study of the domestication of Agave americana complex. While we are aware of the limitations of the information current collected, it enables us to identify the relevant taxa, regions, cultures, and research approaches to answering questions and improving our understanding of this group of plants, and other Agave complexes in their main domestication setting: Mesoamerica. Declarations Ethics approval and consent to participate Not applicable. Consent for publication Not applicable. Competing interests The authors declare that they have no competing interests. Author contributions N.C.S. conceived and designed the study, conducted fieldwork and data collection, and performed the morphological measurements and statistical analyses. N.C.S. organized the datasets, interpreted the results, and wrote the initial draft of the manuscript, including the preparation of tables and figures. A.C. supervised the research as the principal doctoral advisor and contributed to the conceptual framework and scientific direction of the study. A.C. provided guidance throughout the research process, contributed to the interpretation and discussion of the results, and participated in the writing process through critical revision, editing, and improvement of the manuscript. Both authors discussed the results, reviewed the manuscript critically, and approved the final version of the manuscript. Funding This research was supported by the National Council of Science and Technology of Mexico (CONACYT) through a doctoral scholarship granted to N.C.S. Additional financial support was provided by CONACYT (research project A1-S-14306), the Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (PAPIIT) of the Dirección General de Asuntos del Personal Académico (DGAPA, UNAM, Mexico; project IN224023), and by the National Commission for the Knowledge and Use of Biodiversity (CONABIO) together with the FAO–GEF project Agrobiodiversity of Mexico (ID 9380) through project RG023. The funding bodies had no role in the design of the study, data collection, analysis, interpretation of the data, or in writing the manuscript. Availability of data and materials The datasets generated and analyzed during the current study are available from the corresponding author upon reasonable request. Acknowledgements The authors thank the National Council of Science and Technology of Mexico (CONACYT) for the doctoral scholarship granted to the first author and the Posgrado en Ciencias Biológicas at the Instituto de Investigaciones en Ecosistemas y Sustentabilidad (IIES), Universidad Nacional Autónoma de México (UNAM), campus Morelia, for academic support during the development of this research. We also acknowledge the financial support provided by CONACYT (project A1-S-14306), the Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (PAPIIT–DGAPA, UNAM; project IN224023), and the National Commission for the Knowledge and Use of Biodiversity (CONABIO) together with the FAO–GEF project Agrobiodiversity of Mexico (ID 9380) through project RG023. References Darwin C. On the origin of species by means of natural selection, or the preservation of favored races in the struggle for life. London: John Murray; 1859. Darwin, C. The variation of animals and plants under domestication. London: John Murray; 1868. Harlan JR. Crops and man. Madison: American Society of Agronomy; 1975. Harlan JR. Agricultural origins: centers and non-centers. Science. 1971;174:468–474. Vavilov NI. Origin and geography of cultivated plants. Cambridge: Cambridge University Press; 1992. Hammer K. Das domestikationssyndrom. Die Kulturpflanze. 1984;32(1):11–34. Hawkes JG. The diversity of crop plants. Cambridge: Harvard University Press; 1983. Gepts P. Crop domestication as a long-term selection experiment. Plant Breed Rev. 2004;24:1–44. Pickersgill B. Domestication of plants in the Americas. Ann Bot. 2007;100:925–940. Pickersgill B. Parallel vs convergent evolution in domestication. Front Ecol Evol. 2018;6:56. Smith BD. The initial domestication of Cucurbita pepo. Science. 1997;276:932–934. MacNeish RS. The origins of agriculture and settled life. Norman: University of Oklahoma Press; 1992. Flannery K, editor. Guilá Naquitz. New York: Academic Press; 1986. Casas, A., Caballero, J., Mapes, C. & Zárate, S.. Manejo de la vegetación, domesticación de plantas y origen de la agricultura en Mesoamérica. Boletin De La Sociedad Botanica De Mexico. 2017. Casas, A., Valiente-Banuet, A., Viveros, J. L., Caballero, J., Cortés, L., Dávila, P., Lira, R. & Rodríguez, I.. Plant resources of the Tehuacán-Cuicatlán valley, Mexico. Economic Botany. 2000;55(1). Casas, A., Otero-Arnaiz, A., Pérez-Negrón, E. & Valiente-Banuet, A.. In situ Management and Domestication of Plants in Mesoamerica. Annals of Botany. 2007;100(5). Casas A, Parra F, Aguirre-Dugua X, et al. Manejo y domesticación de plantas en Mesoamérica. In: Domesticación en el Continente Americano. Vol. 2. Morelia: UNAM; 2017. p. 69–102. Casas A, Torres I, Parra F, Torres-Guevara J. Centros de origen y diversificación de plantas cultivadas. De los cultivos nativos y el cambio del clima. Hallazgos (Huánuco y Apurímac) (pp.23-56) Lima: UNALM; 2019. p. 23–62. Blancas, J., Casas, A., Rangel-Landa, S., Moreno-Calles, A. I., Torres, I., Pérez-Negrón, E., Solís, L., Delgado-Lemus, A., Parra, F., Arellanes, Y., Caballero, J., Cortés, L., Lira, R. & Dávila, P.. Plant Management in the Tehuacán-Cuicatlán Valley, Mexico1. Economic Botany. 2010;64(4). Boege E. El patrimonio biocultural de los pueblos indígenas de México. Mexico City: INAH; 2008. Bye RA, Burgess D, Mares-Trias A. Ethnobotany of the western Tarahumara. Bot Mus Leafl Harv Univ. 1975;24:85–112. García-Mendoza AJ. Agaváceas. In: Biodiversidad de Oaxaca. México: UNAM; 2004. p. 159–169. García-Mendoza AJ. Flora del Valle de Tehuacán-Cuicatlán. Agavaceae. México: UNAM; 2011. Gentry HS. Agaves of continental North America. Tucson: University of Arizona Press; 1982. Palma-Cruz FJ. El género Agave y su distribución en Oaxaca. Tesis de licenciatura. UNAM; 1991. Palma-Cruz FJ. Agaves productores de fibras duras en Oaxaca. Bol Soc Bot Mex. 2000;66:93–102. Vargas-Ponce, O. and Zizumbo-Villarreal, D. In situ diversity and maintanenace of traditional Agave landraces used in spirits production in West-Central Mexico. Economic botany. 2006. Vargas-Ponce, O., Zizumbo-Villarreal, D., Martínez-Castillo, J., Coello-Coello, J. & Colunga-Garcíamarín, P.. Diversity and structure of landraces of Agave grown for spirits under traditional agriculture: A comparison with wild populations of A. angustifolia (Agavaceae) and commercial plantations of A. tequilana .. American-Eurasian journal of botany. 2009;96 2. Torres, I., Blancas, J., León, A. & Casas, A. TEK, local perceptions of risk, and diversity of management practices of Agave inaequidens in Michoacán, Mexico. Journal of Ethnobiology and Ethnomedicine. 2015;11(1). Torres-García, I., Rendón-Sandoval, F. J., Blancas, J., Casas, A. & Moreno-Calles, A.. The genus Agave in agroforestry systems of Mexico. Botan‪ical Sciences. 2019. Torres-García I, Delgado-Lemus AM, Casas A, Álvarez-Ríos GD, Rangel-Landa S, Martínez-Jiménez R, et al. Agave americana L., Agave angustifolia Haw., Agave atrovirens Karw. ex Salm-Dyck, Agave asperrima Jacobi, Agave bovicornuta Gentry, Agave cupreata Trel. & A. Berger, Agave hookeri Jacobi, Agave inaequidens K. Koch, Agave karwinskii Zucc., Agave kerchovei Lem., Agave lechuguilla Torr., Agave mapisaga Trel., Agave marmorata Roezl, Agave maximiliana Baker, Agave montana Villareal, Agave potatorum Zucc., Agave rhodacantha Trel., Agave salmiana Otto ex Salm-Dyck, Agave scaposa Gentry, Agave tequilana F.A.C. Weber, Agave victoriae-reginae A. Berger (Asparagaceae). In: Casas A, Blancas Vázquez JJ, editors. Ethnobotany of the Mountain Regions of Mexico. Cham: Springer International Publishing; 2022. p. 1–52. doi: 10.1007/978-3-030-99357-3_26 Clement, C., Casas, A., Parra-Rondinel, F., Levis, C., Peroni, N., Hanazaki, N., Cortés-Zárraga, L., Rangel-Landa, S., Alves, R. P., Ferreira, M. J., Cassino, M. F., Coelho, S. D., Cruz-Soriano, A., Pancorbo-Olivera, M., Blancas, J., Martínez-Ballesté, A., Lemes, G., Lotero-Velásquez, E., Bertin, V. & Mazzochini, G. G.. Disentangling Domestication from Food Production Systems in the Neotropics. Quaternary. 2021. Alducin-Martínez C, Ruiz Mondragón KY, Jiménez-Barrón O, Aguirre-Planter E, Gasca-Pineda J, Eguiarte LE, Medellin RA. Uses, Knowledge and Extinction Risk Faced by Agave Species in Mexico. Plants. 2023; 12(1):124. https://doi.org/10.3390/plants12010124. Álvarez-Ríos, G. D., Pacheco-Torres, F., Figueredo-Urbina, C. J. & Casas, A.. Management, morphological and genetic diversity of domesticated agaves in Michoacán, México.. Journal of Ethnobiology and Ethnomedicine. 2020;16. Cabrera-Toledo D, Vargas-Ponce O, Ascencio-Ramírez S, Valadez-Sandoval LM, Pérez-Alquicira J, Morales-Saavedra J and Huerta-Galván OF. Morphological and Genetic Variation in Monocultures, Forestry Systems and Wild Populations of Agave maximiliana of Western Mexico: Implications for Its Conservation. Front. Plant Sci. 2020;11:817. doi: 10.3389/fpls.2020.00817 Figueredo, C., Casas, A., González‐Rodríguez, A., Nassar, J., Colunga-Garcíamarín, P. & Rocha-Ramírez, V.. Genetic structure of coexisting wild and managed agave populations: implications for the evolution of plants under domestication. AoB PLANTS. 2015. Figueredo-Urbina CJ, Casas A, Torres-García I. Morphological and genetic divergence between Agave inaequidens, A. cupreata and the domesticated A. hookeri. Analysis of their evolutionary relationships. PLOS ONE 2017;12(11): e0187260. https://doi.org/10.1371/journal.pone.0187260 Figueredo-Urbina CJ, Casas A, Martínez-Díaz Y, et al. Domestication and saponins contents in a gradient of management intensity of agaves: Agave cupreata , A. inaequidens and A. hookeri in central Mexico. Genet Resour Crop Evol . 2018; 65 , 1133–1146 (2018). https://doi.org/10.1007/s10722-017-0601-6 Figueredo, C.J., Casas, A., Colunga-GarcíaMarín, P. et al. Morphological variation, management and domestication of ‘maguey alto’ (Agave inaequidens) and ‘maguey manso’ (A. hookeri) in Michoacán, México. J Ethnobiology Ethnomedicine 2014;10:66.. https://doi.org/10.1186/1746-4269-10-66 González Elizondo, M., R. Galván Villanueva, I. L. López Enriquez, L. Reséndiz Rojas y M. S. González Elizondo. 2009. Agaves-magueyes, lechuguillas y noas-del Estado de Durango y sus alrededores. Centro Interdisciplinario de Investigación para el Desarrollo Integral Regional-Unidad Durango, Instituto Politécnico Nacional-Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. Durango, Dgo. 163 pp. Hernández-Sandoval L, González Romo C, González Medrano F. Plantas útiles de Tamaulipas. An Inst Biol UNAM. 1991. MacNeish RS. A summary of the subsistence. Prehistory of Tehuacan Valley. 1967;1:290–309. Miller, A. J. and Schaal, B.. Domestication of a Mesoamerican cultivated fruit tree, Spondias purpurea.. Proceedings of the National Academy of Sciences of the United States of America. 2005. Parra F, Blancas JJ, Casas A. Landscape management and domestication of Stenocereus pruinosus (Cactaceae) in the Tehuacán Valley: human guided selection and gene flow. J Ethnobiol Ethnomed. 2012;14;8:32. doi: 10.1186/1746-4269-8-32. Hancock JF. Plant evolution and the origin of crop species. Cambridge: CABI; 2006. Thiede, J. Agave Agavaceae. En U. Egli & R. Nyffeler (Eds.), Illustrated handbook of succulent plants: Monocotyledons. 2020 (2nd ed., pp. 21-311). Springer-Verlag. Villaseñor José Luis. Checklist of the native vascular plants of Mexico. Rev. Mex. Biodiv. 2016; 87( 3 ): 559-902. https://doi.org/10.1016/j.rmb.2016.06.017. Trelease W. Agave in the West Indies. Whitefish: Kessinger; 1912. Valenzuela-Zapata, A. G.. Tequila: A Natural and Cultural History. . 2002. Urrutia-Cruz SP. Etnobotánica de los agaves en Oaxaca. Tesis de licenciatura. UNAM; 1986. Barbara Pickersgill, Domestication of Plants in the Americas: Insights from Mendelian and Molecular Genetics, Annals of Botany 2007;100:925–940, https://doi.org/10.1093/aob/mcm193 Smith, B. D.. The Initial Domestication of Cucurbita pepo in the Americas 10,000 Years Ago. Science 276, no. 5314 (1997): 932–34. http://www.jstor.org/stable/2893022. Rodríguez-Garay, B., Lomelí-Sención, J., Tapia-Campos, E., Gutiérrez-Mora, A., García-Galindo, J., Rodríguez-Domínguez, J., Urbina-López, D. & Vicente-Ramírez, I.. Morphological and molecular diversity of Agave tequilana Weber var. Azul and Agave angustifolia Haw. var. Lineño.. Industrial Crops and Products. 2008;29(1). Ojeda-Linares C. , Álvarez-Ríos G. , Figueredo-Urbina C. , Islas L. , Lappe-Oliveras P. , Nabhan G. et al.. Traditional Fermented Beverages of Mexico: A Biocultural Unseen Foodscape. Foods 2021;10(10):2390. https://doi.org/10.3390/foods10102390 Casas, A., Otero-Arnaiz, A., Pérez-Negrón, E. & Valiente-Banuet, A.. In situ Management and Domestication of Plants in Mesoamerica. Annals of Botany. 2007;100(5). Clément C. , Casas A. , Parra F. , Levis C. , Peroni N. , Hanazaki N. et al.. Disentangling Domestication from Food Production Systems in the Neotropics. Quaternary 2021;4(1):4. https://doi.org/10.3390/quat4010004 García-Mendoza AJ. Agavaceae of Oaxaca. Biodiversidad de Oaxaca. 2004 Blancas J. , Casas A. , Rangel-Landa S. , Moreno-Calles A. , Torres-García I. , Pérez‐Negrón E. et al.. Plant Management in the Tehuacán-Cuicatlán Valley, Mexico1. Economic Botany 2010;64(4):287-302. https://doi.org/10.1007/s12231-010-9133-0 Casas A. , Ladio A. , & Clément C.. Editorial: Ecology and Evolution of Plants Under Domestication in the Neotropics. Frontiers in Ecology and Evolution 2019;7. https://doi.org/10.3389/fevo.2019.00231 Palma-Cruz FJ. Agave in Oaxaca. Bol Soc Bot Mex. 2000;66:93–102. Trejo L. , Soriano D. , Romano-Grande E. , Sánchez-Carmona B. , & Dávila-Navarro D.. Diversity of reproductive characters, seed set, and viability of Agave seeds used for pulque production and their wild relatives in Tlaxcala, Mexico. Genetic Resources and Crop Evolution 2023;71(6):2877-2903. https://doi.org/10.1007/s10722-023-01803-5 Figueredo-Urbina C. , Casas A. , Colunga‐GarcíaMarín P. , Nassar J. , & González‐Rodríguez A.. Morphological variation, management and domestication of ‘maguey alto’ (Agave inaequidens) and ‘maguey manso’ (A. hookeri) in Michoacán, México. Journal of Ethnobiology and Ethnomedicine 2014;10(1). https://doi.org/10.1186/1746-4269-10-66 Álvarez-Ríos G. , Pacheco-Torres F. , Figueredo-Urbina C. , & Casas A.. Management, morphological and genetic diversity of domesticated agaves in Michoacán, México. Journal of Ethnobiology and Ethnomedicine 2020;16(1). https://doi.org/10.1186/s13002-020-0353-9 Klimova, A.;Ruiz Mondragón, K.Y.;Molina Freaner, F.;Aguirre-Planter, E.; Eguiarte, L.E. Genomic Analyses of Wild and Cultivated Bacanora Agave ( Agave angustifolia var. paci fi ca ) Reveal Inbreeding, Few Signs of Cultivation History and Shallow Population Structure. Plants 2022,11, 1426. https://doi.org/10.3390/plants11111426 Casas, A., S. Rangel-Landa, E. Torres-García, E. Pérez-Negrón, L. Solís, F. Parra, A. Delgado, J. Blancas J, B. Farfán, and A. I. Moreno.. In situ management and conservation of plant resources. In: De Albuquerque, U. P., and M. Alves Ramos (eds). Current Topics in Ethnobotany. Research Signpost, Kerala, India. 2008 pp: 1-25. Parra, F., N. Pérez-Nasser, R. Lira, D. Pérez- Salicrup, and A. Casas.. Population genetics, and process of domestication of Stenocereus pruinosus (Cactaceae) in the Tehuacan Valley, Mexico. J. Arid Environ. 2008; 72: 1997-2010. Casas A, Otero-Arnaiz A, Pérez-Negrón E, Valiente-Banuet A. In situ management and domestication of plants in Mesoamerica. Ann Bot. 2007;100:1101–1115. Clement CR, Casas A, Parra-Rondinel FA, et al. Disentangling domestication from food production systems in the Neotropics. Quaternary. 2021;4:4. Gepts P. Crop domestication as a long-term selection experiment. Plant Breed Rev. 2004;24:1–44. Hancock JF. Plant evolution and the origin of crop species. 2nd ed. CABI; 2006. Parra F, Blancas J, Casas A. Landscape management and domestication of Stenocereus pruinosus. J Ethnobiol Ethnomed. 2012;8:32. Trejo L, Reyes M, Cortés-Toto D, et al. Morphological diversity and genetic relationships in pulque production agaves. Front Plant Sci. 2020;11:524812. Rodríguez-Garay B, Lomelí-Sención JA, Tapia-Campos E, et al. Morphological and molecular diversity of Agave tequilana. Ind Crops Prod. 2009;29:220–228. Ojeda-Linares C, Álvarez-Ríos GD, Figueredo-Urbina CJ, et al. Traditional fermented beverages of Mexico. Foods. 2021;10:2390. Trejo L, Soriano D, Romano-Grande E, et al. Diversity of reproductive characters in Agave used for pulque. Genet Resour Crop Evol. 2024;71:2877–2903. Table Table 7. Means and standard error of 19 characters that presented significant differences when comparing the plants of Agave americana complex and associated species of Americanae Group. Character var americana var expansa A abrupta var oaxacensis var subtilis subsp protamericana A. asperrima A scaposa C M P C C I I W M W M W GROUP 1 2 3 4 5 6 7 8 9 10 11 12 Plant length*** 2.922 ± 0.841d 2.788± 0.566cd 2.296 ± 0.584bcd 1.936 ± 0.215bcd 2.158 ± 0.163bcd 1.351± 0.613a 1.674± 0.033ab 1.951± 0.441b 2.055± 0.615bc 1.450 ± 0.550ab 2.316± 0.386bcd 1.875± 0.340 abcd Stem length*** 0.620± 0.156d 0.438± 0.082c 0.406 ± 0.069bc 0.658 ± 0.079d 0.650± 0.050d 0.236 ± 0.034a 0.386 ± 0.068bc 0.394 ± 0.081bc 0.396 ± 0.069bc 0.275± 0.086ab 0.383 ± 0.040bc 0.141 ± 0.014a Diameter*** 2.615 ± 0.527g 1.966± 0.304def 1.583± 0.276cd 2.560± 0. 361fg 2.334 ± 0.189efg 1.725± 0.562bcd± 2.456± 0.228efg 1.242± 0.306ab 1.485± 0.570bc 0.850± 0.173a 1.733± 0.103cde 2.350± 0.264efg Leaf length*** 1.778 ± 0.324f 1.506± 0.222ef 1.210± 0.170de 1.356± 0.167de 1.524 ± 0.082ef 0.910± 0.202abc 1.274± 0.105cde 0.893± 0.245ab 1.030± 0.334bcd 0.627± 0.126a 1.286± 0.080de 1.10± 0.216bcde Leaf width*** 0.237 ± 0.036 0.254± 0.052 0.182± 0.039 0.237± 0.012 0.276 ± 0.020 0.092± 0.017 0.128± 0.004 0.212± 0.034 0.193± 0.042 0.192± 0.017 0.211± 0.022 0.285± 0.031 LL/LW*** 7.561 ± 1.443de 6.025± 0.928cd 6.985± 2.028cd 5.701± 0± .608bcd 5.548 ± 0.541bcd 10.109± 2.473e 9.947± 0.640e 4.253± 1.177ab 5.310± 1.170bc 3.309± 0.904a 6.129± 0.680cd 3.889± 0.871ab LL/SL*** 2.989 ± 0.761abc 3.475± 0.447cd 3.033± 0.504bc 2.068± 0.172a 2.353 ± 0.192ab 5.853± 2.198e 3.390± 0.701bcd 2.305± 0.573a 2.576± 0.560ab 2.352± 0.422ab 3.387± 0.397bc 28.108± 43.566de Number of leaf*** 41.78 ± 12.968bc 46.44± 7.55bcd 38.846± 10.573bc 53.40± 6.618cd 79 ± 1.73e 62.10± 10.64de 75.80± 6.61e 33.21± 8.935ab 39.71± 13.09bc 20.25± 5.123a 35.33± 5.60abc 34.0± 9.933abc Terminal thorn length* 0.029 ± 0.006bc 0.056± 0.014e 0.143± 0.207e 0.026± 0.006ab± 0.025 ± 0.002ab 0.013± 0.006a 0.027± 0.002bc 0.040± 0.006d 0.044± 0.007de 0.040± 0.008cde 0.045± 0.013de 0.060± 0.008e Terminal thorn width*** 0.005 ± 0.002abc 0.014± 0.014c 0.006± 0.003bc 0.006± 0.001bc 0.009 ± 0.005bc 0.003± 0.001a 0.004± 0.0008abc 0.004± 0.001ab 0.005± 0.002ab 0.004± 0.0005abc 0.007± 0.002bc 0.005± 0.0005abc TTL/TTW*** 6.524 ± 3.776abc 6.027± 3.093abc 19.961± 19.409d 4.110± 1.239ab 3.352 ± 1.318a 4.10± 1.235ab 5.890± 0.812abc 9.622± 2.583cd 9.613± 3.791cd 9.0± 2.0bcd 6.916± 3.072abcd 11.416± 0.957cd TTL/LL** 0.017 ± 0.006ab 0.037± 0.007d 0.113± 0.152d 0.019± 0.004ab 0.017 ± 0.002ab 0.013± 0.004a 0.021± 0.0005bc 0.047± 0.013d 0.045± 0.011d 0.067± 0.025d 0.035± 0.010cd 0.056± 0.015d Teeth length*** 0.008± 0.002c 0.012± 0.002cde 0.012± 0.001de 0.015± 0.001e 0.004 ± 0.0006ab 0.004± 0.001a 0.008± 0.002bcd 0.011± 0.003cd 0.010± 0.003cd 0.012± 0.002cde 0.009± 0.001cd 0.010± 0.001cde LTEE/LL*** 0.549± 0.523ab 0.005± 0.002a 0.006± 0.001a 1.318± 0.082cd 0.606 ± 0.101bc 1.444± 0.287d 1.710± 0.199d 0.009± 0.003ab 0.007± 0.003a 0.013± 0.002ab 0.005± 0.001a 1.871± 0.750d Teeth width*** 0.007± 0.001c 0.008± 0.003cd 0.007± 0.002bc 0.011± 0.0006d 0.006 ± 0.001bc 0.002± 0.0005a 0.004± 0.001ab 0.008± 0.002c 0.007± 0.001c 0.008± 0.0005cd 0.006± 0.001bc 0.006± 0.002bc LTEE/WTEE*** 0.953± 0.379b 0.698± 0.185ab 0.612± 0.192a 0.761± 0.046ab 1.684 ± 0.273c 0.726± 0.198ab 0.590± 0.060ab 0.750± 0.206ab 0.691± 0.254ab 0.732± 0.208ab 0.677± 0.083ab 0.591± 0.193ab WTEE/LL*** 0.005± 0.001ab 0.008± 0.002a 0.010± 0.001a 0.008± 0.001cd 0.004 ± 0.0005bc 0.003± 0.0008d 0.003± 0.0005d 0.013± 0.005ab 0.0008± 0.002a 0.020± 0.006ab 0.007± 0.001a 0.006± 0.003d Distance between teeth** 0.018± 0.008abc 0.015± 0.007ab 0.024± 0.011bc 0.024± 0.005bc 0.013 ± 0.003ab 0.018± 0.005abc 0.029± 0.005c 0.021± 0.009bc 0.022± 0.007bc 0.017± 0.006abc 0.019± 0.009abc 0.009± 0.001a DTEE/LL *** 0.011± 0.007a 0.010± 0.004a 0.019± 0.008b 0.018± 0.006ab 0.008 ± 0.001a 0.020± 0.006b 0.023± 0.005b 0.022± 0.005b 0.024± 0.012b 0.029± 0.012b 0.015± 0.006ab 0.008± 0.001a Different letters indicate significant differences. Significant level 0.001***, 0.01** and 0.0 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Under Review Version 1 posted Reviews received at journal 04 May, 2026 Reviewers agreed at journal 23 Apr, 2026 Reviews received at journal 08 Apr, 2026 Reviewers agreed at journal 19 Mar, 2026 Reviewers agreed at journal 18 Mar, 2026 Reviewers agreed at journal 16 Mar, 2026 Reviewers invited by journal 16 Mar, 2026 Editor assigned by journal 16 Mar, 2026 Submission checks completed at journal 16 Mar, 2026 First submitted to journal 09 Mar, 2026 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-9078121","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":608291499,"identity":"71fdc9ed-d093-4f2b-91e1-16a910cbb9da","order_by":0,"name":"NADIA CAMPOS SALAS","email":"","orcid":"","institution":"Posgrado en Ciencias Biológicas","correspondingAuthor":false,"prefix":"","firstName":"NADIA","middleName":"CAMPOS","lastName":"SALAS","suffix":""},{"id":608291500,"identity":"596e0caf-9e2c-4b4a-b8d4-678b557d4362","order_by":1,"name":"ALEJANDRO CASAS","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA0UlEQVRIiWNgGAWjYPACGwY2UrWkka7lMAlq+dsPP/7w4895eT725ocPGNvuyOs2cKdJ4NMicSbNTLK37bZhG88xYwPGtmeG2w7wbjbAa80NBjMG3obbCWwSOWwSjG2HGYFaNj7Ap0P+Bvvnj3/+nEtgk38D1mIP1LLhAD4tBjd4DKR52A4AbeEBa0kkaIvhmZwyadm2ZKBf0owNEs4dTt52mIBf5I4f3/zxzR87efn2ww8ffCg7bLvteO82vCGGChJABDPx6kfBKBgFo2AU4AAAtQJI2b+Yus4AAAAASUVORK5CYII=","orcid":"","institution":"Instituto de Investigaciones en Ecosistemas y Sustentabilidad, Universidad Nacional Autónoma de México (UNAM).","correspondingAuthor":true,"prefix":"","firstName":"ALEJANDRO","middleName":"","lastName":"CASAS","suffix":""}],"badges":[],"createdAt":"2026-03-10 02:24:33","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9078121/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9078121/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105035341,"identity":"774d4a6c-736b-498e-bcd9-ea9ed2f122e8","added_by":"auto","created_at":"2026-03-20 07:25:53","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":839503,"visible":true,"origin":"","legend":"\u003cp\u003eRepresentative morphological diversity of Agave species across different management conditions and geographic regions in Mexico. a) Agave americana subsp americana var americana. B) Agave americana subsp americana var expansa. C) Agave americana subspamericana var oaxacensis, D) Agave abrupta, E) Agave asperrima, F) Agave scaposa, G) Agave americana subsp protamericana, and H) Agave americana subsp americana var subtilis.\u003c/p\u003e","description":"","filename":"Figure1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/f507b4e9cb033de29b17918b.jpg"},{"id":104976700,"identity":"186b5123-3661-48a7-80c3-3631b3348cc8","added_by":"auto","created_at":"2026-03-19 12:21:56","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":44193,"visible":true,"origin":"","legend":"\u003cp\u003eGeographic distribution of the sampled taxa within the Mexican territory, encompassing representative populations of \u003cem\u003eA. abrupta, A. asperrima, A. scaposa,\u003c/em\u003e and the subspecies and varieties of \u003cem\u003eA. americana\u003c/em\u003e (subsp.\u003cem\u003e protamericana, \u003c/em\u003esubsp.\u003cem\u003e americana \u003c/em\u003evar.\u003cem\u003e americana, \u003c/em\u003evar.\u003cem\u003e expansa, \u003c/em\u003evar.\u003cem\u003e oaxacensis, \u003c/em\u003eand\u003cem\u003e \u003c/em\u003evar.\u003cem\u003e subtilis\u003c/em\u003e). The map also displays the major biogeographic provinces where these taxa occur: The Neovolcanic, Oaxacan, Tamaulipan, Sierra Madre Oriental, and Altiplanicie provinces.\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/8061915e96940b18234bc259.jpg"},{"id":105035033,"identity":"e050ffa6-3c91-4ab2-af07-2b55a3b3992f","added_by":"auto","created_at":"2026-03-20 07:25:19","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":21142,"visible":true,"origin":"","legend":"\u003cp\u003ePrincipal Component Analysis (PCA) biplot, showing the distribution of morphological traits across five \u003cem\u003eAgave\u003c/em\u003e species. The first two principal components (PC1 and PC2) explain 30.8% and 24.9% of the total variance, respectively. The arrows represent the loadings of each morphological variable on the two principal components, indicating the direction and strength of their contribution. Abbreviations of morphological character names are shown in Table 2.\u003c/p\u003e","description":"","filename":"Figure3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/b883c3d4839ada77c43d45b0.jpg"},{"id":105035417,"identity":"b5b8ba21-4378-4d2f-ae60-6639bafab103","added_by":"auto","created_at":"2026-03-20 07:26:03","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":17431,"visible":true,"origin":"","legend":"\u003cp\u003eLinear Discriminant Analysis (LDA) scatterplots of LD1 vs LD2 with ~95% confidence ellipses. Panel (A) shows Taxonomic separation, (B) geographic regions (Center, North, South), and (C) management categories (Intensive cultivation, wild). Dashed lines mark axis origins. Together, the plots illustrate how taxonomy, geography, and management contribute to population differentiation.\u003c/p\u003e","description":"","filename":"Figure4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/15ffc8568d0bd161e7ebe2f0.jpg"},{"id":104976706,"identity":"51bd625c-809f-4bcd-a374-17daddee3020","added_by":"auto","created_at":"2026-03-19 12:21:56","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":61855,"visible":true,"origin":"","legend":"\u003cp\u003eHeatmap showing the distribution of standardized morphological traits across \u003cem\u003eAgave\u003c/em\u003epopulations. Each row represents a population labelled by taxon, while each column corresponds to a morphological variable. The color intensity reflects the Z-score standardized values, with red indicating higher-than-average trait values and blue indicating lower-than-average values. Hierarchical clustering (Euclidean distance, complete linkage) was applied to both individuals and traits to reveal patterns of morphological similarity.\u003c/p\u003e","description":"","filename":"Figure5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/ba1ce69e0800c992963f1925.jpg"},{"id":105034939,"identity":"83fbab8a-4aae-468c-9fe1-9c2038103dcf","added_by":"auto","created_at":"2026-03-20 07:24:54","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":28821,"visible":true,"origin":"","legend":"\u003cp\u003ePrincipal Component Analysis (PCA) biplot showing the morphological traits of field specimen samples from multiple \u003cem\u003eAgave\u003c/em\u003e species and management types. The first two principal components (PC1 = 25.3%, PC2 = 18.3%) together explain 43.6% of the total morphological variance. Each point represents an individual, color-coded by species identity. Arrows indicate the direction and strength of trait contributions (loadings) to the ordination, with longer vectors representing greater influence. Abbreviations of morphological character names are shown in Table 2.\u003c/p\u003e","description":"","filename":"Figure6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/101a99c1fdee295a1975a797.jpg"},{"id":104976703,"identity":"0dbd8935-393e-4014-9163-c8b7286f60c3","added_by":"auto","created_at":"2026-03-19 12:21:56","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":61350,"visible":true,"origin":"","legend":"\u003cp\u003eHeatmap showing the relationship between different morphological traits (rows) and herbarium specimens (columns) morphological characters of \u003cem\u003eAgave\u003c/em\u003e. The colors represent the intensity of the expression of each trait, with a color scale ranging from negative values (blue tones) to positive values (red tones). Groupings were determined using hierarchical clustering analysis.\u003c/p\u003e","description":"","filename":"Figure7.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/364c68dd5ee77840f4ef1600.jpg"},{"id":105035444,"identity":"7151b900-94ff-4fe5-aa96-8e2575e89fa1","added_by":"auto","created_at":"2026-03-20 07:26:05","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":19028,"visible":true,"origin":"","legend":"\u003cp\u003eLinear Discriminant Analysis (LDA) scatterplots of the first two discriminant functions (LD1 vs. LD2), showing population differentiation of Agave. Panel (A) illustrates the separation among Taxa, panel (B) displays variation by management categories (Cultivated, Intensive, Promoted, Wild), and panel (C) shows clustering according to geographic regions. Ellipses represent ~95% confidence intervals for each group.\u003c/p\u003e","description":"","filename":"Figure8.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/e93144843c634bebc6c90a31.jpg"},{"id":105562796,"identity":"0a22d0f4-e607-4257-975c-bff53a4987eb","added_by":"auto","created_at":"2026-03-27 12:44:46","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3194746,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9078121/v1/3740bdcd-6a41-45eb-adbb-b02faed531ea.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Use, management and domestication of Agave americana in Mexico","fulltext":[{"header":"Background","content":"\u003cp\u003eDomestication is an evolutionary process directed by humans that provides valuable insights into general evolutionary mechanisms in organisms [\u003cspan additionalcitationids=\"CR2 CR3\" citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. As the biocultural contexts in which domestication occurs are diverse and dynamic, the process operates continuously and dynamically in multiple directions. The species being tamed adapt to diverse human-influenced environments and cultural and technological contexts, enabling them to live and reproduce successfully. Domesticated organisms are shaped by morphological and physiological attributes that are markedly (though not exclusively) aligned with human requirements [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. Although selection is the primary evolutionary force, other forces\u0026mdash;mutation, recombination, gene flow, genetic drift, and mating systems\u0026mdash;also influence domestication under both natural and human guidance [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eMesoamerica, the cultural region comprised between Mexico and Central America, is one of the world\u0026rsquo;s main domestication regions [\u003cspan additionalcitationids=\"CR10 CR11\" citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. Indigenous peoples in this area domesticated more than 250 native species [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e], including maize (\u003cem\u003eZea mays\u003c/em\u003e L.), beans (\u003cem\u003ePhaseolus\u003c/em\u003e spp.), squashes (\u003cem\u003eCucurbita\u003c/em\u003e spp.), agaves (\u003cem\u003eAgave\u003c/em\u003e spp.), prickly pears (\u003cem\u003eOpuntia\u003c/em\u003e spp.), chili peppers (\u003cem\u003eCapsicum annuum\u003c/em\u003e L.), vanilla (\u003cem\u003eVanilla planifolia\u003c/em\u003e Jacks. ex Andrews), cotton (\u003cem\u003eGossypium hirsutum\u003c/em\u003e L.), and other crops which are currently of worldwide relevance. The coexistence of crops and their wild relatives, together with varying intensities and directions of selection and management, produced extensive morphological and physiological diversity [\u003cspan additionalcitationids=\"CR15 CR16\" citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. This diversity is reflected in the multiple varieties often found within a single species.\u003c/p\u003e \u003cp\u003eRecords from archaeological, ethnobiological, and evolutionary ecology studies have revealed that domestication began in Mesoamerica around 10 thousand years ago [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan additionalcitationids=\"CR19\" citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e] and continues today, influencing both native and introduced species [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. The region harbors one of the world\u0026rsquo;s richest reservoirs of agrobiodiversity [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan additionalcitationids=\"CR23\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. Documenting how this diversity originated and identifying the factors that maintain it remain key priorities for its conservation [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eIndigenous peoples in Mesoamerica use nearly 6,500 native plant species [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e], about 20% of those recorded by botanists [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. Among them, \u003cem\u003eAgave\u003c/em\u003e stands out as a genus of exceptional ecological and cultural importance. Its species have been used since prehistoric times and dominate many regional landscapes, where the genus also diversified. At least twelve species have been domesticated to varying degrees: \u003cem\u003eA. americana\u003c/em\u003e L., \u003cem\u003eA. angustifolia\u003c/em\u003e Haw., \u003cem\u003eA. atrovirens\u003c/em\u003e Karw. ex Salm-Dyck, \u003cem\u003eA. fourcroydes\u003c/em\u003e Lem., \u003cem\u003eA. karwinskii\u003c/em\u003e Zucc., \u003cem\u003eA. mapisaga\u003c/em\u003e Trel., \u003cem\u003eA. salmiana\u003c/em\u003e Otto ex Salm-Dyck, \u003cem\u003eA. sisalana\u003c/em\u003e Perrine, \u003cem\u003eA. tequilana\u003c/em\u003e FAC Weber, \u003cem\u003eA. parryi\u003c/em\u003e Engelm., \u003cem\u003eA. hookeri\u003c/em\u003e Jacobi, and \u003cem\u003eA. inaequidens\u003c/em\u003e K. Koch. Others, such as \u003cem\u003eA. cupreata\u003c/em\u003e Trel. \u0026amp; A. Berger, \u003cem\u003eA. potatorum\u003c/em\u003e Zucc., \u003cem\u003eA. maximiliana\u003c/em\u003e Baker, and \u003cem\u003eA. rhodacantha\u003c/em\u003e Trel., have been cultivated for at least fifty years without clear signs of domestication [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. The cultivation and management of these species, and the associated selective mechanisms, have been well documented [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan additionalcitationids=\"CR29 CR30 CR31 CR32\" citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]. High levels of interspecific hybridization and genetic variation have also been reported [\u003cspan additionalcitationids=\"CR31 CR32 CR33 CR34 CR35\" citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eSince phenotypic changes reflect those occurring in genotypes [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e], researchers commonly assess domestication by examining divergence between wild and managed phenotypes [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e, \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]. In \u003cem\u003eAgave\u003c/em\u003e, domestication syndromes reflect use type, often involving enlargement of useful parts\u0026mdash;stems, leaves, fibers, or sap\u0026mdash;and reduction of defensive or undesirable features such as spines, teeth, and saponins and other secondary metabolites [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan additionalcitationids=\"CR30\" citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe interaction between natural and human selection makes \u003cem\u003eAgave\u003c/em\u003e an excellent model for studying domestication. Mexico, with the greatest diversity and distribution of \u003cem\u003eAgave\u003c/em\u003es [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e], offers ideal conditions for this research. Humans have managed and domesticated these plants since prehistoric times, using them for food, fiber, construction, fermented beverages, medicines, soil retention, and ceremonial purposes [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe case of \u003cem\u003eAgave americana\u003c/em\u003e is of particular interest, given the extraordinary degree of diversification it has undergone during domestication. According to Gentry [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e], the subspecies \u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eamericana\u003c/em\u003e comprises four varieties. The putative wild relatives (\u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e) also occur in Mexico and the southern USA. Currently, all varieties of \u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eamericana\u003c/em\u003e are found exclusively in managed populations, and their direct wild relatives are unknown. This gives rise to two hypotheses regarding their origin: (1) the species diversified prior to human management, with the ancestors of the domesticated varieties of \u003cem\u003eA. a. americana\u003c/em\u003e subsequently becoming extinct; or (2) all varieties result from the domestication of one or few single ancestors, likely represented by the only clearly recognised wild taxon, \u003cem\u003eA. a. protamericana\u003c/em\u003e. Testing these hypotheses requires comprehensive information on their genetic similarities and relationships, as well as on ecological aspects such as their distribution, abundance, local adaptations, and cultural aspects such as forms of management, targets and mechanisms of human selection, conditions of genetic drift associated to management, as well as gene flow influenced by translocation and other practices. The phenotypic consequences of these processes in relation to management must also be considered. This study explores aspects on management, selection, and phenotypic variation in the taxa included in this species, as well as some of its closest wild relatives.\u003c/p\u003e \u003cp\u003eHuman selection drives morphological and genetic differentiation in managed plants. A persistent challenge in evolutionary studies is determining how phenotypic traits are inherited and whether observed morphological divergence corresponds to genetic differentiation. Plant domestication is typically gradual; domesticated forms retain traits that reveal relationships with wild ancestors and display intermediate morphotypes depending on domestication degree. Examining these patterns helps reconstruct evolutionary pathways [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eCurrently, \u003cem\u003eAgave americana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e is the only taxon of the species studied with clear wild populations, although this taxon\u0026rsquo;s populations may also be subject to some forms of management, as will be discussed below. Therefore, this taxon offers the opportunity to compare the morphology, physiology and genetics of the domesticated varieties by analysing how they diverge from other taxa and which features that are mainly affected by management. We hypothesize that these features should be related to the historical use of each taxon. We are aware of the extraordinary capacity for hybridization between varieties, subspecies, and even other closely related species within the genus. Therefore, phenotypic variation is influenced by complex factors, and such influence requires careful analysis using an integral approach to morphology, population genetics and reproductive biology.\u003c/p\u003e \u003cp\u003eThe domestication syndrome of \u003cem\u003eAgave\u003c/em\u003es may vary according to their use. For instance, their use as food has primarily involved the stem size, since stems have been cooked in underground ovens since prehistoric times. In addition, the inflorescences and flower buds can be consumed when roasted or boiled, respectively. Features of the domestication syndrome related to this use could include a large stem and scape, and flowers, as well as a reduction in secondary compounds that impart a bitter flavour. \u003cem\u003eAgave\u003c/em\u003e fibres have also been used since ancient times, and gigantism expressed in leaf length, as well as the durability and strength of fibres would be expected in relation to this use. The sap of several \u003cem\u003eAgave\u003c/em\u003e species has been used as a fresh (aguamiel) or fermented (pulque) beverage for thousands of years. To this end, selection favouring the quantity and quality of the sap (e.g. plant size, sap yield, and sugar content) would be expected. Currently, \u003cem\u003eAgave\u003c/em\u003es are most commonly used to produce mescal. While there is controversy surrounding pre-Columbian distillation, the material used to make mescal is the same as that used for cooking the stems. Therefore, the main attributes selected for this purpose are those preferred for preparing food in underground ovens. Gigantism of stems and leaves and the length and resistance of fibres, appear to be features shared by different species under domestication. In addition, the difficulty of managing \u003cem\u003eAgave\u003c/em\u003es has led people to favour plants that produce fewer and shorter spines and teeth, as well as lower amounts of saponins, which cause skin irritation in those managing them [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. Understanding domestication thus requires examining divergence in these traits according to use and management intensity.\u003c/p\u003e \u003cp\u003eThis study aims to:1) document ethnobotanical and morphological information in order to analyse the \u003cem\u003eAgave americana\u003c/em\u003e complex, its use and management, and the patterns and mechanisms of selection, the morphological divergence among the varieties and in relation to their putative wild ancestor \u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e, and 2) analyse the degree and patterns of divergence in relation to the targets of selection and management intensity in order to examine the role of management in morphological divergence. We expected to observe clear trends of divergence among the taxonomically defined varieties, as well as in domestication syndrome traits relative to \u003cem\u003eA. a.\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e, in order to support the hypothesis of a common ancestor of the domesticated varieties.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e\u003cem\u003eStudy species\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe \u003cem\u003eAgave americana\u003c/em\u003e complex comprises the following two subspecies and four varieties:\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave americana\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;subsp. \u003cem\u003eprotamericana\u003c/em\u003e\u003c/strong\u003e, (Fig 1 G.) This predominantly wild taxon occurs along the Sierra Madre Oriental. On the coastal slopes, where rainfall is higher, rosettes are larger than on the drier continental side. Leaves are four to six times longer than wide, plane or guttered, straight or curved, and green to glaucous grey. The panicle is short, with 15\u0026ndash;20 branches. Gentry [23] reported introgression between \u003cem\u003eA. a. protamericana\u003c/em\u003e and \u003cem\u003eA. asperrima\u003c/em\u003e, suggesting further research on reproductive biology and morphometrics to explore possible hybridization and its genetic and phenotypic outcomes.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave americana\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;subsp. \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eamericana\u003c/em\u003e\u0026nbsp;\u003c/strong\u003e(Fig 1 A.) This variety includes highly surculose plants with lanceolate leaves, widest near or slightly above mid-blade. Leaves are glaucous grey to light green, tapering above the base, and sometimes reflexed. The habitat is mainly anthropogenic, although Gentry [40] and Hern\u0026aacute;ndez-Sandoval et al. [42] reported putative wild populations near Ciudad Victoria, Tamaulipas. It is widespread in Mexico [43-45], though its wild status needs confirmation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave americana\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;subsp. \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eexpansa\u003c/em\u003e\u003c/strong\u003e. (Fig 1 B.) This taxon was first described by William Trelease in 1900 as var. \u003cem\u003eamericana\u003c/em\u003e. However, the large, glaucous grey, prolifically suckering rosettes and the tall, diffuse panicle resemble the taxon that occurs in Europe. The most notable characteristics are the valleculate leaves and the short, conical spines [23]. It has been reported in Chihuahua [46] and Jalisco [45].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave americana\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;subsp. \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eoaxacensis.\u003c/em\u003e\u003c/strong\u003e (Fig 1 C.) It occurs in Oaxaca, where is commonly known as \u0026ldquo;maguey arroque\u0026ntilde;o\u0026rdquo;, \u0026ldquo;maguey coyote\u0026rdquo;, \u0026ldquo;maguey de Castilla\u0026rdquo;, or \u0026ldquo;maguey Sierra Negra\u0026rdquo; [24]. It has large, spreading, glaucous to whitish leaves with small teeth, producing large flowers and thick-walled capsules [23]. Because its stems accumulate high carbohydrate levels, it is used for high-quality mescal. Plants require 18\u0026ndash;20 years to mature [45]. Historically, it was also used for pulque and fiber extraction [40].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave americana\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003esubsp.\u003cem\u003e\u0026nbsp;americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;subtilis\u003c/em\u003e\u003c/strong\u003e (Fig 1 H.) Known locally as \u0026ldquo;maguey chato\u0026rdquo; or \u0026ldquo;chato de Sahuayo,\u0026rdquo; this taxon was initially placed in the Rigidae group due to its compact rosette. Valenzuela-Zapata et al. [48] later reassigned it to the Americanae group based on inflorescence morphology. It mainly grows in Jiquilpan and Sahuayo, Michoac\u0026aacute;n.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eRelated taxa considered\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThree additional species were included for comparison due to their close relationships and potential reproductive interactions with \u003cem\u003eA. americana\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave abrupta\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e(Fig 1 D.), closely related to \u003cem\u003evar. expansa\u003c/em\u003e, this species was first described by Trelease (1912) and later regarded by Gentry [23] as part of \u003cem\u003evar. expansa\u003c/em\u003e. It differs mainly in rosette size, leaf shape, and terminal spine.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003cem\u003escaposa\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e(Fig 1 F.), a large plant, with light green to glaucous leaves and tall, narrow panicles bearing small, dry bracts, aligning it with the Americanae group. Diagnostic traits include large rosettes, dark- to yellow-green undulate leaves with small marginal teeth, and tall inflorescences with small flowers [23]. According to Garc\u0026iacute;a-Mendoza [49], \u003cem\u003eA. scaposa\u003c/em\u003e is used for pulque production. Casas et al. [50] and Blancas et al. [51] reported its use in soil stabilization, fencing, and as a source of durable inflorescence stalks (or \u0026ldquo;quiote\u0026rdquo;) for construction.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAgave\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003cem\u003easperrima\u003c/em\u003e subsp. \u003cem\u003eAsperrima\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e(Fig 1 E.), the most widespread \u003cem\u003eAgave\u003c/em\u003e in northern Mexico [23], distinguished by scabrous, shorter leaves and narrow-bracted inflorescences with fewer branches. Its flowering overlaps and introgression with \u003cem\u003eA. a. var. americana\u003c/em\u003e and \u003cem\u003eA. a. subsp. protamericana\u003c/em\u003e have been reported [52-54].\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eStudy area\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe field study sites for the analyzed taxa of the \u003cem\u003eAgave americana\u003c/em\u003e complex are shown in Fig. 1 and Table 1.\u003c/p\u003e\n\u003cp\u003eThe study was conducted in three main regions of Mexico:\u003c/p\u003e\n\u003cul\u003e\n \u003cli\u003e\u003cstrong\u003eNortheastern Mexico\u003c/strong\u003e, covering the states of Nuevo Le\u0026oacute;n and Tamaulipas;\u003c/li\u003e\n \u003cli\u003e\u003cstrong\u003eThe Trans-Mexican Volcanic Belt\u003c/strong\u003e, including parts of Michoac\u0026aacute;n and Jalisco; and\u003c/li\u003e\n \u003cli\u003e\u003cstrong\u003eThe Sierra Madre del Sur\u003c/strong\u003e, represented by populations in Oaxaca.\u0026nbsp;\u003c/li\u003e\n\u003c/ul\u003e\n\u003cp\u003eGeneral information on the populations studied is summarized in Table 1, which includes altitude (m a.s.l.), mean annual temperature (\u0026deg;C), mean annual rainfall (mm), and associated habitat types.\u003c/p\u003e\n\u003cp\u003eIn Nuevo Le\u0026oacute;n, populations were examined in the localities Doctor Arroyo and Iturbide; in Tamaulipas, in San Carlos, El Tinieblo, Jaramillo, and Miquihuana; in Michoac\u0026aacute;n, in Tar\u0026iacute;mbaro, Santiago Undameo, Zacapu, Cojumatl\u0026aacute;n, and Jiquilpan; in Jalisco, in Zacoalco and Ciudad Guzm\u0026aacute;n; and in Oaxaca, in Coixtlahuaca, San Baltazar Guelavila, and San Pablo Huixtepec.\u003c/p\u003e\n\u003cp\u003eThese sites represent a broad range of ecological conditions, from microphyllous desert scrub and oak\u0026ndash;pine forests in the north to agricultural plots, orchards, and intensive home-garden systems in the south. Such variation provides the environmental and management context for evaluating phenotypic differentiation and adaptation patterns within the \u003cem\u003eA. americana\u003c/em\u003e complex.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 1. Study sites of the analyzed \u003cem\u003eAgave\u003c/em\u003e populations, including elevation (m a.s.l.), annual mean temperature (\u0026deg;C), annual rainfall (mm), and associated habitat types. These environmental variables provide the ecological context for population differentiation and habitat adaptation across the sites considered in the morphological variation analysis.\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eStudy Site\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eElevation\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;(m a.s.l.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAnnual Mean Temp (\u0026deg;c)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAnnual Rain Fall (mm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eHabitat\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eDr. Arroyo, Nuevo Le\u0026oacute;n\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1947\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e18.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e400\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eMycrophyllous, subinerme desert scrub\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eIturbide, Nuevo Le\u0026oacute;n\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1518\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e560\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eOak-pine forest\u003c/p\u003e\n \u003cp\u003eOrchards\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eMiquihuana, Tamaulipas\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1842\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e600\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eThorny scrub\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eSan Carlos, Tamaulipas\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e570\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e850\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTamaulipas scrub\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eEl Tinieblo, Tamaulipas\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e199\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTamaulipas scrub\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eJaramillo, Tamaulipas\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e220\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e850\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTamaulipas scrub\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eZacapu, Michoac\u0026aacute;n.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2004\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1069\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAlong the road\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTar\u0026iacute;mbaro, Michoac\u0026aacute;n.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1885\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e18.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e609\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eNatural fence\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eSantiago Undameo, Michoac\u0026aacute;n.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2050\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e16.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e754\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eNatural fence and home gardens\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eZacoalco, Jalisco.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1420\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e22.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e578.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAgricultural field\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eCojumatl\u0026aacute;n, Michoac\u0026aacute;n.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1558\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e800\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eOrchard\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eJiquilpan, Michoac\u0026aacute;n.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1830\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e850\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eMonoculture plot\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eCoixtlahuaca, Oaxaca.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2280\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e750\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eRosetophylous scrub\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eSan Baltazar Guelavila, Oaxaca.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1570\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e20.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e650\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eIntensive Plots\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eSan Pablo Huixtepec, Oaxaca.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1480\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1000\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eIntensive Plots\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cem\u003eEthnobotanical studies\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eA total of 14 in-depth interviews were conducted with the managers and owners of the visited sites to document traditional knowledge, uses, selection criteria, selection targets and mechanisms. These interviews were supplemented with previously collected ethnobotanical information from our team and other colleagues, including from published sources. Direct observations were also conducted on wild populations of \u003cem\u003eAgave\u003c/em\u003e \u003cem\u003easperrima\u003c/em\u003e, \u003cem\u003eAgave\u003c/em\u003e scaposa and \u003cem\u003eAgave americana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e and subsp. \u003cem\u003eamericana\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eEthnobotanical data were analyzed qualitatively to identify patterns of management intensity, domestication traits, and traditional selection strategies. Information was categorized according to management system (wild, promoted, managed, cultivated, and intensified plots) and cross-referenced with morphological variation patterns observed in the same populations.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eStudies of morphological variation\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eNineteen vegetative morphological traits were evaluated in field-collected plants from each population (Table 2). These characters were selected based on their ethnobotanical relevance and previous studies on domestication syndromes in \u003cem\u003eAgave\u003c/em\u003e [14,29,31,55]. Most studied populations are cultivated or semi-cultivated, where plants are commonly harvested before flowering, making reproductive structures rarely available. To complement the data, 21 vegetative and reproductive traits were measured from herbarium specimens to characterize morphological differentiation among species, subspecies, and varieties.\u003c/p\u003e\n\u003cp\u003ePreliminary analyses that included reproductive traits did not yield statistically significant differences among taxa; therefore, subsequent analyses focused on vegetative structures, following methodological precedents in ethnobotanical studies of \u003cem\u003eAgave\u003c/em\u003e [14,29,31,55].\u003c/p\u003e\n\u003cp\u003eTable 2. \u0026nbsp;Morphological characters measured in \u003cem\u003eAgave\u003c/em\u003e populations studied. Their abbreviations and measurement units are indicated. In addition, it is indicated whether these characteristics were evaluated in herbarium specimens and/or in natural populations.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"641\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eCharacter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eAbbreviation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eMeasure units\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eHerbarium\u003c/p\u003e\n \u003cp\u003especimens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eEvaluated populations\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003ePlant length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003ePL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003em\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eStem length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eSL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003em\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eDiameter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003em\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eLeaf length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003em\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eLeaf width***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eLeaf length/width ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLL/LW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eLeaf/stem length ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLL/SL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003em\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eNumber of leaves\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLN\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eCounted number\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTerminal thorn length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTTL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTerminal thorn width\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTTW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTerminal thorn length/width ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTTL/TTW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTerminal thorn/leaf length ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTTL/LL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTeeth length\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTEEL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTeeth/leaf length ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTEEL/LL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTeeth width\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTEEW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTeeth length/width ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTEEL/TEEW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTeeth width/leaf length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTEEW/LL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eDistance between teeth\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eDBTEE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eDistance between teeth/leaf length ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eDBTEE/LL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eInflorescence length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eIL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003em\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eRamification number\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eRN\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eCounted number\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eOvary length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eOL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eOvary width\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eOW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eCm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eTepal length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eFilament length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eFL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eAnther lenght\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eAL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 225px;\"\u003e\n \u003cp\u003eStamen length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eSL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecm\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 92px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 107px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cem\u003eStatistical analyses of morphological characters\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThree multivariate analyses were performed to identify morphological grouping patterns among the studied taxa:\u003c/p\u003e\n\u003cul class=\"decimal_type\"\u003e\n \u003cli\u003eCluster Analysis (CA),\u003c/li\u003e\n \u003cli\u003ePrincipal Components Analysis (PCA), and\u003c/li\u003e\n \u003cli\u003eLinear Discriminant Analysis (LDA).\u003c/li\u003e\n\u003c/ul\u003e\n\u003cp\u003eAll analyses were conducted using R software (version 4.3.2) [56].\u003c/p\u003e\n\u003cp\u003eCluster Analysis employed Euclidean distances to estimate morphological dissimilarities among populations, generating a dendrogram with standardized average distance as weighting factor. PCA was used to assess interspecific relationships and identify the most influential variables for taxonomic differentiation. The analysis considered a matrix of 122 individuals and 19 variables, standardized by z-scores. Eigenvalues and eigenvectors were derived from Pearson\u0026rsquo;s correlation matrix, and individuals were projected onto the first two principal components.\u003c/p\u003e\n\u003cp\u003eLDA was performed to classify individuals according to their taxonomic identity, management type, and geographic region. The eigenvectors from PCA and LDA analyses were compared to identify the most discriminant morphological traits. These analyses allowed evaluating whether morphological similarities correspond to management intensity, provenance, or taxonomic affinity within the \u003cem\u003eA. americana\u003c/em\u003e complex.\u003c/p\u003e\n\u003cp\u003eHeatmap and Clustering Analysis\u003c/p\u003e\n\u003cp\u003eA heatmap was generated using the pheatmap package in R to visualize standardized morphological values and clustering patterns. Hierarchical agglomerative clustering with Euclidean distances and complete linkage was used to construct dendrograms for both individuals (rows) and traits (columns), revealing associations between taxa and morphological syndromes [57].\u003c/p\u003e\n\u003cp\u003eColor gradients were applied to highlight similarities and contrasts across populations, facilitating the visualization of domestication patterns that might not emerge from traditional plots.\u003c/p\u003e\n\u003cp\u003eTo evaluate significant differences between management categories and taxa, one-way ANOVAs were conducted for all 19 vegetative characters. Each variable was tested for normality (Shapiro\u0026ndash;Wilk test) and homogeneity of variances (Levene\u0026rsquo;s test). Tukey\u0026rsquo;s post hoc tests were applied to identify which groups differed significantly (p \u0026lt; 0.05).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eEthnobotanical uses\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll interviewees reported that the studied taxa are primarily used for mescal production, sap extraction for pulque, and ornamental purposes (Table 3). The var\u003cem\u003e. oaxacensis\u003c/em\u003e is the most frequently cultivated and economically important taxon for mescal production, particularly in the Central Valleys of Oaxaca. Another highly valued variety, \u003cem\u003eA. americana\u003c/em\u003e var. \u003cem\u003esubtilis\u003c/em\u003e, is cultivated extensively in Jiquilpan and Sahuayo, Michoac\u0026aacute;n.\u003c/p\u003e\n\u003cp\u003eOther taxa also contribute to mescal production. For example, \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e is managed in San Carlos and Miquihuana (Tamaulipas), and \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;americana\u003c/em\u003e is cultivated in San Carlos, Jim\u0026eacute;nez, and Jaramillo (Tamaulipas). Producers reported that plants are harvested only when the inflorescence stalk begins to develop; at this stage, the scape is cut to preserve sugars in the main stem of the \u003cem\u003eAgave\u003c/em\u003e, which is essential for enhance fermentation and flavor quality.\u003c/p\u003e\n\u003cp\u003ePulque production is mainly associated with \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar. \u003cem\u003eamericana\u003c/em\u003e in Tar\u0026iacute;mbaro and Santiago Undameo (Michoac\u0026aacute;n), and with \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;expansa\u003c/em\u003e in Ciudad Guzm\u0026aacute;n and Zacoalco (Jalisco). Additionally, \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;americana\u003c/em\u003e is used for aguamiel extraction in Iturbide (Nuevo Le\u0026oacute;n).\u003c/p\u003e\n\u003cp\u003eFor ornamental purposes, \u003cem\u003eA. abrupta\u003c/em\u003e is cultivated for its large, symmetrical rosette. This taxon is found along the shores of Lake Chapala in the state of Jalisco.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 3. Study sites and \u003cem\u003eAgave\u003c/em\u003e taxa recorded, including common names, registered uses, management types (wild or cultivated), selection criteria, and the number of populations and individuals per population evaluated.\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSite\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTaxa\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCommon name\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eUses\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eManagement\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSelection (mechanisms, targets, reasons)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of populations\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of individuals per population\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eDr. Arroyo, Nuevo Le\u0026oacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey verde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eNo use registered\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eWild\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eIturbide, Nuevo Le\u0026oacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey cenizo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eAguamiel extraction\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eCultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eThe big steam its selected due to the high and constant pulque extraction.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eNo use registered\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eWild\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave asperrima\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eNo use registered\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eWild\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMiquihuana, Tamaulipas.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eAguamiel extraction\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eManaged in situ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eThey select several kind of \u003cem\u003eAgave\u003c/em\u003e taxa for the aguamiel extraction.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSan Carlos, Tamaulipas.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eManaged in situ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"3\" valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eThis taxa are the most common and abundant in the region so people select them because it is the only option.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eManaged in situ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave asperrima\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey blanco\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eManaged in situ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eEl Tinieblo, Tamaulipas.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey verde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003ePromoted\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eSize of the steam\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eJaramillo, Tamaulipas.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey verde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003ePromoted\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eSize of the steam\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eZacapu, Michoac\u0026aacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey cenizo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eOrnamental\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eCultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eThe most important is the size of the plant\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTar\u0026iacute;mbaro, Michoac\u0026aacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey cenizo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003ePulque extraction\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eCultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eIts selected for the quantity of sugar, the size of the steam and\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSantiago Undameo, Michoac\u0026aacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey cenizo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003ePulque extraction\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eCultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eZacoalco, Jalisco.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eexpansa\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey cenizo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003ePulque extraction\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eCultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eIt\u0026rsquo;s a taxon that produce pulque for three months\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCojumatl\u0026aacute;n, Michoac\u0026aacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave abrupta\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eOrnamental\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eCultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eThe size of the plant and the few quantity of spines\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eJiquilpan, Michoac\u0026aacute;n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003esubtilis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eChato de Sahuayo\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eIntensive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eHight quantity of sugar and fast maturation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCoixtlahuaca, Oaxaca.\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e scaposa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey de potrero, maguey cimarr\u0026oacute;n, maguey de caballo, maguey de pitzorra o tequiol [57]\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eUsed in the construction of houses and for other local purposes (extraction of grubsfrom the plant), and occasionally for cooking. It is also used as living fences and to delimit terrain (asmarkers) [58,59]\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eWild\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eIts selected due to the width of the leaves and the big size of the inflorescence.\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSan Baltazar Guelavila, Oaxaca.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eoaxacensis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eMaguey de castilla\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003cp\u003e[60] reported for fiber use\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eIntensive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eFlavor, high quantity and quality of sugar\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 94px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSan Pablo Huixtepec, Oaxaca.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 76px;\"\u003e\n \u003cp\u003e\u003cem\u003eAgave\u003c/em\u003e \u003cem\u003eamericana\u003c/em\u003e var. \u003cem\u003eoaxacensis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 59px;\"\u003e\n \u003cp\u003eArroque\u0026ntilde;o\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eCoyote\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 67px;\"\u003e\n \u003cp\u003eMescal production\u003c/p\u003e\n \u003cp\u003e[60] reported for fiber use\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003eIntensive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 86px;\"\u003e\n \u003cp\u003eThe flavor of the mescal its unique with this taxa, the big amount of sugar and the size of the steam.\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 77px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eThe subspecies \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e grows wild in the San Carlos region (Tamaulipas), where informants indicated that cultivation is recent. Locals prefer plants from the surrounding hillsides, asserting that wild individuals produce mescal with superior flavor. The studied plants are believed to have originated from wild populations established for over a century, and are thus considered genuinely wild.\u003c/p\u003e\n\u003cp\u003eIn Tamaulipas, mescal production based on \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e has become a significant economic activity, sustained mainly by small-scale producers. Its growing popularity in recent decades supplements local agricultural economies (maize, sorghum, and livestock). However, the number of small production units and the extraction of reproductive Agaves from the forest has increased. No evaluation has yet been conducted in this respect, but it is undoubtedly needed to establish criteria for the sustainable use of these populations.\u003c/p\u003e\n\u003cp\u003eAmong all taxa, \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e stands out for its cultural and linguistic richness. It is one of the most culturally and economically important taxa in the \u003cem\u003eA. americana\u003c/em\u003e complex. Traditionally used for fiber extraction, it is now primarily cultivated for high-quality mescal production of high economic value.\u003c/p\u003e\n\u003cp\u003eThe wide range of ethnotaxa (traditional varieties recognized by local people) recognized in Oaxaca reflects both morphological variation and deep cultural knowledge. Common names include maguey arroque\u0026ntilde;o, maguey ruque\u0026ntilde;o, maguey cenizo, maguey blanco (\u0026ldquo;pale \u003cem\u003eAgave\u003c/em\u003e\u0026rdquo;), and maguey sierrudo or maguey sierra negra (\u0026ldquo;saw-leaf \u003cem\u003eAgave\u003c/em\u003e\u0026rdquo;). In Mixtec, it is called yavi cuaam, in Spanish maguey coyote, maguey xolo, maguey de rayo, and in Zapotec it bears names such as doba nupy, d\u0026ograve;b-dz\u0026igrave;n, d\u0026ograve;b mpi\u0026egrave;, dobncjp, dua-bsug, dua-bzog, dua-pchez, dua-ya-do, and dua-yesh. This linguistic diversity reflects the long-standing cultural relationship and morphological differentiation of local forms, which merit focused ethnobotanical and genetic research.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eManagement Practices\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe surveyed \u003cem\u003eAgave\u003c/em\u003e populations exhibited a broad spectrum of management regimes, reflecting a domestication gradient ranging from wild to fully cultivated forms (Table 3):\u003c/p\u003e\n\u003cp\u003eWild populations \u0026ndash; e.g., \u003cem\u003eA. asperrima, A. scaposa, A. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;americana\u003c/em\u003e, growing in xeric scrublands and rocky slopes. These plants regenerate naturally and are opportunistically harvested. They maintain strong defense traits (spines, thick cuticles) and high morphological uniformity.\u003c/p\u003e\n\u003cp\u003eManaged in situ populations \u0026ndash; predominantly \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;americana\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. scaposa\u003c/em\u003e, are tolerated or let standing when clearing land, as well as protected in agricultural and fallow plots. Plants are not actively propagated but are occasionally transplanted from forest or disturbed areas to agricultural plots and protected, showing slight selection favoring straight and vigorous scape and variable morphology.\u003c/p\u003e\n\u003cp\u003ePromoted populations \u0026ndash; such as \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e and certain variants of \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003esubsp\u003cem\u003e. americana\u003c/em\u003e are composed by semi-wild individuals that are transplanted or selectively favored, particularly for pulque or fencing uses. These \u003cem\u003eAgave\u003c/em\u003es exhibit intermediate phenotypes, suggesting early domestication stages.\u003c/p\u003e\n\u003cp\u003eCultivated populations \u0026ndash; composed mainly of \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;americana,\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;expansa\u003c/em\u003e, and subsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e, grown in agroforestry systems or small plantations. Propagation is primarily vegetative, and selection favors large stem size, high sap yield, and reduced thorniness.\u003c/p\u003e\n\u003cp\u003eIntensively managed populations \u0026ndash; such as \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;subtilis\u003c/em\u003e (Michoac\u0026aacute;n) and A. \u003cem\u003eamericana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e (Oaxaca). These are maintained in monocultures and exhibit strong domestication syndromes, including increased vegetative size and reduction of defense structures.\u003c/p\u003e\n\u003cp\u003eThe geographic distribution of management systems correlates strongly with species identity and regional cultural traditions, emphasizing the co-evolutionary nature of human\u0026ndash;\u003cem\u003eAgave\u003c/em\u003e interactions throughout Mexico.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMorphological patterns\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eReproductive and vegetative characters analysed in herbarium specimens\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003ePrincipal Component Analysis (PCA) (Fig 3.) revealed significant patterns of morphological variation among the studied \u003cem\u003eAgave\u003c/em\u003e species. The first two principal components accounted for 64.6% of the total variance (PC1 = 41.9%, PC2 = 22.7%). These components explained the main axes of morphological differentiation among the taxa.\u003c/p\u003e\n\u003cp\u003eThe variables with the highest loadings on PC1 were terminal thorn length (TTL), teeth length (TEEL), and the ratio TEEL/LL (teeth-to-leaf length). On PC2, the variables with the greatest contributions were teeth width (TEEW), distance between teeth (DBTEE), TEEL/TEEW (teeth length-to-width ratio), DBTEE/LL (distance between teeth-to-leaf length ratio), and TTL/TTW (terminal thorn length-to-width ratio).\u003c/p\u003e\n\u003cp\u003eThe ordination revealed clear segregation of the taxa along PC1 and PC2 based on these morphological variables\u003cem\u003e. Agave americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;americana, A. a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis,\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. a.\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e were positioned toward the positive side of PC1, while \u003cem\u003eA. asperrima\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. scaposa\u003c/em\u003e were located on the negative side. Along PC2, \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e were grouped toward the positive quadrant, whereas \u003cem\u003eA. scaposa\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. asperrima\u003c/em\u003e were placed toward the negative quadrant.\u003c/p\u003e\n\u003cp\u003eThe distribution pattern indicated that PC1 represented the axis of greatest morphological differentiation among the taxa, mainly influenced by the variables TTL, TEEL, and TEEL/LL, while PC2 captured additional variation explained by TEEL/TEEW, DBTEE, and TTL/TTW.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eLinear Discriminant Analysis (LDA)\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eLDA revealed clear morphological structuring among the analysed \u003cem\u003eAgave\u003c/em\u003e taxa according to geography, management regime, and taxonomic identity (Fig. 4). When grouped by geographic origin (north, centre, and south) (Fig.4b), southern taxa\u0026mdash;particularly \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e\u0026mdash;formed a distinct cluster separated from northern and central groups. Northern taxa, including \u003cem\u003eA. asperrima\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;americana\u003c/em\u003e, were characterized by elevated values of terminal thorn width (TTW), teeth length (TEEL), and the ratio TTL/LL (terminal thorn-to-leaf length). Central taxa, represented mainly by \u003cem\u003eA. scaposa\u003c/em\u003e, exhibited intermediate or overlapping morphological profiles.\u003c/p\u003e\n\u003cp\u003eDiscriminant analysis based on management regime showed well-defined segregation between cultivated, intensively managed, and wild populations. Individuals under intensive management, particularly those from Oaxaca, were associated with higher values of inflorescence branching and reduced spine-related variables. Wild accessions, notably \u003cem\u003eA. asperrima\u003c/em\u003e, displayed compact rosettes and higher thorn and tooth measurements.\u003c/p\u003e\n\u003cp\u003eWhen grouped by taxonomic identity, the discriminant functions successfully differentiated the species and varieties within the \u003cem\u003eAgave americana\u003c/em\u003e complex. The resulting morphological clusters were consistent with their formal taxonomic assignments, confirming the discriminant capacity of the evaluated variables.\u003c/p\u003e\n\u003cp\u003eHEAT MAP ANALYSIS (Fig. 5) The \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e group exhibited moderate vegetative and defensive traits but strong reproductive investment, with medium-sized, narrow leaves and intermediate thorn and tooth dimensions. High values for inflorescence length and ovary width indicated advanced reproductive development and a clear emphasis on seed production.\u003c/p\u003e\n\u003cp\u003eThe \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e group showed moderate vegetative growth, strong defensive features, and intermediate reproductive effort. \u003cem\u003eAgave\u003c/em\u003es of the subspecies \u003cem\u003eprotamericana\u003c/em\u003e from Zacatecas displayed the most robust morphology\u0026mdash;large leaves, thick thorns, and broad teeth\u0026mdash;while other populations maintained moderate defence and reproductive traits. This group is defined by structural robustness and enhanced defence, particularly in the Zacatecan populations.\u003c/p\u003e\n\u003cp\u003eThe \u003cem\u003eA. americana\u003c/em\u003e group exhibited wide morphological variability, combining vigorous vegetative growth with moderate-to-strong defences and diverse reproductive investment. \u003cem\u003eA. americana\u003c/em\u003e Saltillo1 and Coahuila showed the largest leaves and strong defences, while Coahuila also had the largest inflorescences and ovaries, marking the highest reproductive investment. Overall, the group is characterized by large plant size, strong defence, and variable reproductive effort.\u003c/p\u003e\n\u003cp\u003eThe \u003cem\u003eA. asperrima\u003c/em\u003e group displayed balanced vegetative and reproductive traits with well-developed defensive morphology. \u003cem\u003eA. asperrima\u003c/em\u003e Zacatecas had the longest and thickest thorns and teeth, representing the strongest defence, while other populations showed intermediate values. Reproductive traits were consistently moderate across all taxa, defining the group as morphologically balanced with notable defensive strength.\u003c/p\u003e\n\u003cp\u003eThe \u003cem\u003eA. scaposa\u003c/em\u003e group presented moderate differentiation, with balanced vegetative growth and intermediate defence. \u003cem\u003eA. scaposa\u003c/em\u003e Suchixtlahuaca had the largest leaves and slightly greater reproductive investment, while \u003cem\u003eA. scaposa\u003c/em\u003e Puebla and Puebla1 maintained smaller leaves and moderate defensive traits. Overall, the group exhibits intermediate morphology with modest variability among populations.\u003c/p\u003e\n\u003cp\u003eFinally, \u003cem\u003eA. lurida\u003c/em\u003e Oaxaca and \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;marginata_\u003c/em\u003eCDMX showed compact vegetative forms and moderate defence and reproduction. \u003cem\u003eA. lurida\u003c/em\u003e possessed small leaves and slender spines but stood out for its larger ovary, reflecting higher reproductive investment. \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;marginata\u003c/em\u003e exhibited narrow leaves and intermediate structural and reproductive traits. Both represent stable morphologies within the \u003cem\u003eA. americana\u003c/em\u003e complex, balancing defence with reproductive efficiency.\u003c/p\u003e\n\u003cp\u003eThe integrated multivariate analyses provided a comprehensive understanding of the morphological diversity present among \u003cem\u003eAgave\u003c/em\u003e species herborized, populations, and management types. Principal component analysis (PCA) (Fig. 3, Table. 5) revealed that traits such as leaf width (LW),\u0026nbsp;LL/SL (leaf length/stem length ratio), and the teeth length/leaf length ratio, teeth length/leaf length and teeth width/leaf length (TEEL/LL, TEEW/LL) ratios were key contributors to the main axes of morphological variation. These variables defined a spectrum ranging from robust, large-leafed taxa to smaller, more dentate forms. Taxonomically defined groups exhibited varying degrees of cohesion in PCA space. For instance, \u003cem\u003eA. asperrima\u003c/em\u003e occupied a distinct region, whereas \u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eamericana\u003c/em\u003e, A\u003cem\u003e. americana\u0026nbsp;\u003c/em\u003esubsp\u003cem\u003e. protamericana,\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;A. expansa\u003c/em\u003e were more broadly distributed and often overlapped with each other and with other taxa. \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. oaxacensis\u003c/em\u003e and \u003cem\u003eA. scaposa\u003c/em\u003e displayed greater variability. Individuals of \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e were widely dispersed in the PCA space and split across dendrogram branches, whereas populations of\u003cem\u003e\u0026nbsp;A\u003c/em\u003e. \u003cem\u003escaposa\u003c/em\u003e clustered more consistently.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eLinear discriminant analysis (LDA) (see Fig. 4) by geographic area demonstrated strong regional structuring of morphology. Samples from the southern region formed a distinct cluster, separated along both LD1 and LD2 axes, suggesting the presence of unique morphological characters. Samples from central and northern regions displayed partial overlap, albeit with distinguishable sub-clusters, indicating that geography plays a significant role in shaping the \u003cem\u003eAgave americana\u003c/em\u003e complex, potentially through local selection pressures or environmental constraints.\u003c/p\u003e\n\u003cp\u003eAnalysis by management type (wild versus cultivated) suggest evidence of domestication-related morphological patterns. Cultivated individuals formed a cohesive and relatively compact cluster, indicating reduced morphological variability and a tendency towards standardized traits. These included broader leaves and lower dentition, which are favored for harvest and processing efficiency. In contrast, the trait SL (stem length) in\u0026nbsp;wild specimens exhibited more dispersed pattern in LDA space, reflecting higher phenotypic variability, which is likely to result from natural selection and genetic diversity. This supports the idea of a \u0026ldquo;domestication syndrome\u0026rdquo; in \u003cem\u003eAgave\u003c/em\u003e, whereby human influence reduces diversity in favour of functional traits.\u003c/p\u003e\n\u003cp\u003eFinally, LDA based on taxonomic assignment revealed that \u003cem\u003eA. asperrima\u003c/em\u003e exhibited significant morphological diversity, with minimal overlap with other taxa. By contrast, subsp. \u003cem\u003eamericana\u003c/em\u003e, subsp. \u003cem\u003eprotamericana\u003c/em\u003e, and var. \u003cem\u003eexpansa\u003c/em\u003e exhibited substantial overlap in the discriminant space, underscoring the limitations of morphology in resolving species boundaries within this complex. These results are consistent with previous studies suggesting extensive hybridisation, local domestication, or shared ancestry among these taxa. Overall, the combined evidence from PCA, hierarchical clustering and LDA highlights the multifactorial nature of \u003cem\u003eAgave\u003c/em\u003e morphological diversity, which is shaped not only by taxonomy but also by geography and human management.\u003c/p\u003e\n\u003cp\u003eBased on characters evaluated in herborized specimens, the cluster analysis had a cophenetic test r= 0.747. This clearly groups \u003cem\u003eA. asperrima,\u003c/em\u003e \u003cem\u003eA. scaposa,\u003c/em\u003e and\u003cem\u003e\u0026nbsp;A. lurida\u003c/em\u003e separated from subsp.\u003cem\u003e\u0026nbsp;americana\u003c/em\u003e, in accordance with their recognised taxonomic identity (Figure 3). The complex \u003cem\u003eAgave americana\u003c/em\u003e can be seen to include three differentiated groups: one of them including var. \u003cem\u003eamericana\u0026nbsp;\u003c/em\u003eand var.\u003cem\u003e\u0026nbsp;marginata\u0026nbsp;\u003c/em\u003ewith\u003cem\u003e\u0026nbsp;A. americana\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u0026nbsp;\u003c/em\u003efrom San Luis Potos\u0026iacute;; a second group is formed by var. \u003cem\u003eamericana\u003c/em\u003e and var. \u003cem\u003eoaxacensis\u003c/em\u003e grouped with subsp. \u003cem\u003eprotamericana\u003c/em\u003e specimens from Nuevo Le\u0026oacute;n; and a third group discriminating var. \u003cem\u003eoaxacensis\u003c/em\u003e and var. \u003cem\u003eexpansa\u003c/em\u003e. The four varieties of \u003cem\u003eA. americana\u003c/em\u003e are not clearly discriminated. Unfortunately, this study based on herbarium specimens does not provide information on differentiation associated with management, since this information is absent from the specimen\u0026rsquo;s labels. We can only record three types of practice: wild, cultivated and intensively cultivated.\u003c/p\u003e\n\u003cp\u003eEigenvectors indicate that the characters with the highest contribution to the classification in the first principal component are the number of branches, filament of stamens, length of the ovary, the ratio of the terminal spine and the leaf length. In the second principal component the most relevant characters were the anther length, the ratio of teeth width and leaf length (WTEE/LL), the ratio of terminal thorn length and leaf length (TTL/LL), the ratio of teeth length and leaf length (LTEE/LL), the ovary width and length and number of branches.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 4. Results of the multivariate analysis of variance (MANOVA) for the variables Area, Management, and Taxonomy. The table shows Wilks\u0026rsquo; Lambda values, Chi-square statistics, degrees of freedom (DF), and corresponding p-values for each variable. Analyses were performed using two datasets: H, specimens measured from herbarium material, and F, specimens measured directly in the field. P-values lower than 0.05 indicate statistically significant differences among the analyzed categories.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" align=\"\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eFUNCTION\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eWILKS\u0026acute;LAMBDA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eCHI SQUARE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eDF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eP VALUE\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003eAREA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eH\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e3.5942e-05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e86.98\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e0.09173\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e0.00101\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e615.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2.2e-16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003eMANAGEMENT\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eH\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1.8139e-05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e92.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e0.06601\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e0.0055\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e487.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2.2e-16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003eTAXONOMY\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eH\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e---------------\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e-----------------\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e------------\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e0.00278\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e734.63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e133\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2.2e-16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eWe conducted three analyses using the LDA to examine the role of the area of distribution, the taxonomic classification of the taxa, and the associated management to each taxon. 1) The LDA analysis for area has a small lambda value of 0.0193, which is significant at a \u003cem\u003ep\u003c/em\u003e value of 0.091. The first function explains 91.2% of the variance, whereas the second function contributes significantly less to the discrimination process. 2) The LDA analysis for management has a small lambda value of 1.8139e-05. The first function explains 96.1% of the variance, whereas the second function does not contribute significantly to the discrimination process as compared to that of the first function. 3) The taxonomy analysis for does not provide a lambda value because there are few samples to analyze. The first function explains 81.33% of the variance, while the second function contributes with the 15.05%.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eVegetative characters measured in the field\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe Principal Component Analysis (PCA) (Fig. 6, Table. 5) revealed the main axes of morphological variation among species within the \u003cem\u003eAgave americana\u003c/em\u003e complex, explaining 43.6% of the total variation through the first two components (PC1 = 25.3%, PC2 = 18.3%).\u003c/p\u003e\n\u003cp\u003ePC1 was primarily driven by variables related to leaf shape and defensive traits, including the teeth length/leaf length ratio (TEEL/LL), leaf length/width ratio (LL/LW), and distance between teeth (DBTEE). This axis separated species with long, narrow leaves and well-developed teeth (e.g., \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;expansa\u003c/em\u003e, \u003cem\u003eA.a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e y \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;subtilis\u003c/em\u003e) toward the positive side, from those with broader leaves and smaller teeth (e.g., \u003cem\u003eA. abrupta, A. a.\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. americana, A. asperrima\u003c/em\u003e) toward the negative side.\u003c/p\u003e\n\u003cp\u003ePC2 was mainly associated with terminal thorn length and width (TTL, TTW) and plant diameter (D). Species with longer thorns and larger diameters, such as var. \u003cem\u003eoaxacensis\u003c/em\u003e and var. \u003cem\u003esubtilis\u003c/em\u003e, were positioned on the positive side of the axis, while species with shorter thorns and more compact forms, such as \u003cem\u003eA. a\u003c/em\u003e. subsp. \u003cem\u003eprotamericana\u003c/em\u003e and \u003cem\u003eA. a\u003c/em\u003e. subsp. \u003cem\u003eamericana\u003c/em\u003e, occupied the negative side.\u003c/p\u003e\n\u003cp\u003eOverall, the PCA showed a clear separation between robust, defensively adapted species (\u003cem\u003eA. asperrima, A. scaposa, A. a.\u0026nbsp;\u003c/em\u003esubsp.\u003cem\u003e\u0026nbsp;protamericana\u003c/em\u003e) and those with greater vegetative and reproductive investment (\u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. oaxacensis, A. a.\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. subtilis\u003c/em\u003e,), while intermediate forms (A. a. subsp. \u003cem\u003eamericana\u003c/em\u003e, \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. expansa\u003c/em\u003e) clustered in central positions.\u003c/p\u003e\n\u003cp\u003eThe variables with the greatest discriminant weight were TEEL/LL, LL/LW, DBTEE, TTL, and TTW, confirming that leaf morphology and defensive features are the main factors structuring variability within the \u003cem\u003eAgave americana\u003c/em\u003e complex.\u003c/p\u003e\n\u003cp\u003eThe heatmap based on vegetative and defensive traits (Fig. 7) revealed clear morphological differentiation among taxa of the \u003cem\u003eAgave americana\u003c/em\u003e complex. The most influential variables included tooth length and width (TEEL, TEEW), terminal thorn dimensions (TTL, TTW), leaf proportions (LL, LW, LL/LW), plant diameter (D), and total plant length (PL), while reproductive traits were not considered in this analysis. The \u003cem\u003eA. americana\u003c/em\u003e group exhibited moderate to strong defensive development, particularly in cultivated and managed forms, where high TEEL/LL and TTL/LL ratios reflected proportionally long teeth and terminal thorns. The \u003cem\u003eA. protamericana\u003c/em\u003e taxon displayed the most robust morphologies, especially in wild populations, with large leaves and highly developed spines indicating strong structural defense. The \u003cem\u003evar. oaxacensis\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;var. subtilis\u003c/em\u003e were characterized by elongated, narrow leaves and intermediate thorn expression, suggesting balanced vegetative and defensive investment. \u003cem\u003eA. expansa\u003c/em\u003e showed high values for TEEL/LL and TTL/LL, emphasizing pronounced defensive structures, while \u003cem\u003eA. scaposa\u003c/em\u003e exhibited moderate vegetative size and notable thorn development, with long terminal spines relative to leaf length. The \u003cem\u003eA. asperrima\u003c/em\u003e, particularly wild populations, presented the most pronounced defensive traits, with long and thick thorns and high TTL/LL ratios, whereas managed individuals displayed slightly greater vegetative development. Overall, the heatmap analysis indicated that morphological variation within the \u003cem\u003eAgave americana\u003c/em\u003e complex is primarily driven by defensive and vegetative attributes, particularly those related to spines, teeth, and leaf architecture, delineating adaptive strategies that balance protection and growth across taxa. Wild taxa exhibit the most robust defensive morphologies. Hierarchical clustering of the rows (groups) in the heatmap revealed morphologically coherent clusters, that often align with known taxonomic distinctions or management regimes. The dendrogram on the x-axis grouped variables with similar expression profiles across groups. For example, TTL (terminal thorn length) and TTW (terminal thorn width) cluster closely together, confirming their strong positive correlation, which was also observed in the PCA.\u003c/p\u003e\n\u003cp\u003eLinear Discriminant Analysis (LDA) (Fig. 8) was performed to explore the morphological diversity of \u003cem\u003eAgave\u003c/em\u003e taxa across three key factors: taxonomy, management, and region. The three separate plots reveal the following:\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eTaxonomy (top panel).\u0026nbsp;\u003c/em\u003eThe first LDA plot (Fig. 8A) reveals clear differentiation between species based on LD1 (the horizontal axis) and LD2 (the vertical axis). The species \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e and the wild group \u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eamericana\u003c/em\u003e, \u003cem\u003eA. americana\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e and A. \u003cem\u003easperrima\u003c/em\u003e cluster in different areas of the plot, indicating distinct morphological profiles and unique trait compositions related to their vegetative structures. \u003cem\u003eA. scaposa\u003c/em\u003e and \u003cem\u003eA. expansa\u003c/em\u003e are also grouped separately, reflecting their particular adaptive features and differentiating them from other species.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eManagement (middle panel) (\u003c/em\u003eFig 8B\u003cem\u003e).\u003c/em\u003e Management regimes are clearly distinct. Cultivated individuals (CULT) cluster in the top-right quadrant, while the intensively managed (INTEN), semi-wild (PROMOTED), and wild (WILD) groups are separated along LD1, with the wild types farthest to the left. This pattern confirms that cultivation is associated with a suite of morphological changes consistent with a domestication syndrome.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eRegion (bottom panel)\u003c/em\u003e (Fig 8C). The third LDA plot shows the influence of geographic origin on the morphological variability of \u003cem\u003eAgave\u003c/em\u003e. Northern populations (NORTH) cluster to the right along LD1, distinct from the Michoac\u0026aacute;n (MICH), Jalisco and Oaxaca samples on the left. This suggests that regional agroecological and cultural factors influence morphological divergence, potentially reinforcing localized selection practices, possibly due to environmental factors such as temperature, precipitation, and soil conditions shaping their growth and reproductive characteristics.\u003c/p\u003e\n\u003cp\u003eTable 5. Characters evaluated in herbarium specimens and in the field and eigenvectors of the first and second principal components and discriminant functions.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eVegetative\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 184px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eHerbarium specimens\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCharacter\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003ePC1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003ePC2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003eDF1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003eDF2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003ePC1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003ePC2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003eDF1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003eDF2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ePlant length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.2166\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.3632\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3847\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.3750\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eDiameter\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.4013\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.1169\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2502\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.5270\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eStem length\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.2469\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.2997\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.1303\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.6912\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLeaf length\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.3624\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.2564\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.4715\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e3.5562\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2854\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.1131\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.8631\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.6157\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLeaf width\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.1056\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.3815\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.5092\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.0412\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2385\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.0654\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.4060\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.4022\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTerminal thorn length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.1354\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.1991\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.7469\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2153\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2711\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0786\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3208\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0574\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTerminal thorn width\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.0747\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.1933\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0710\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2293\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2468\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2250\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2081\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.9294\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of leaves\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.2946\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e0.1351\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.1925\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3065\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTeeth length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.1710\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.3112\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2907\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2840\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2258\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.1511\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.4106\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.8714\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTeeth width\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.0672\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.3393\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2407\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3690\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.1827\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.093\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.6154\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2731\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eDistance between teeth\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.1180\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.0085\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.0507\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3979\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.0133\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2275\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2652\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.1553\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLeaf length/width ratio\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.2529\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e0.1931\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.9352\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-2.3732\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.1015\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.1549\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.7995\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.4925\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLeaf/stem length ratio\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.0578\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e0.1224\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.8731\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.3545\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTerminal thorn length/width ratio\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.1959\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.1539\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2083\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.8316\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2025\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.9294\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2025\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.9294\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTerminal thorn/leaf length ratio ***\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.2124\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.1751\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.7553\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.9657\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.2403\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.3071\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.2403\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.3071\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTeeth/leaf length ratio\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e-0.2002\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e0.3104\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e3.6414\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.5015\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.0706\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2204\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.0706\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2204\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTeeth width/leaf length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.3735\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e-0.1594\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.7649\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.2663\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.0462\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.3572\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.0462\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.3572\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 227px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eDistance between teeth/leaf length ratio\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 74px;\"\u003e\n \u003cp\u003e0.31534\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 49px;\"\u003e\n \u003cp\u003e0.1316\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0611\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.5001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0111\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.3907\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0111\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.3907\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eInflorescence length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0948\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.1676\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.0948\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.1676\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eRamification number\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-2.1962\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.1654\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-2.1962\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.1654\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eOvary length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.1042\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.360\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.104\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eOvary width\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2011\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.250\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.2011\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTepal length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.4911\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3617\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.4911\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3617\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eFilament length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.467\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e1.467\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.1991\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAnther length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.9401\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.6291\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.9401\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.6249\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eStamen length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.0104\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-1.0104\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.7998\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" valign=\"top\" style=\"width: 443px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTotal flower length\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3424\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2506\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e0.3424\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 46px;\"\u003e\n \u003cp\u003e-0.2506\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 6. Averages and standard errors of 19 characters that presented significant differences when comparing categories wild, promoted, managed in situ, and cultivated.\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"603\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eCharacter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ewild\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003epromoted\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003eManaged in situ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003ecultivated\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003ePlant height***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.865 \u0026plusmn; 0.463b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.296 \u0026plusmn; 0.584bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.336 \u0026plusmn; 0.632c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.557\u0026plusmn; 0.782c\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eStem length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.339 \u0026plusmn; 0.125ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.406 \u0026plusmn; 0.069b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.406 \u0026plusmn; 0.070b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.634 \u0026plusmn;0.125c\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eDiameter***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.348 \u0026plusmn; 0.526a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.583 \u0026plusmn; 0.276ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.686 \u0026plusmn; 0.474b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.545 \u0026plusmn;0.445c\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eLeaf length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.884 \u0026plusmn; 0.257a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.210 \u0026plusmn; 0.170b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.231 \u0026plusmn; 0.335b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.637 \u0026plusmn;0.312c\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eLeaf width***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.220 \u0026plusmn; 0.041bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.182 \u0026plusmn; 0.039b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.216 \u0026plusmn; 0.049b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.245 \u0026plusmn;0.033c\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eLeaf length/width ratio ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e4. 059 \u0026plusmn; 1.122a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e6.985 \u0026plusmn; 2.028b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e5.701 \u0026plusmn; 1.055b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e6.754 \u0026plusmn;1.498b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eLeaf/stem length ratio\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e6.134 \u0026plusmn; 7.505a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.033 \u0026plusmn; 0.504a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e3.023 \u0026plusmn; 0.652a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e2.665 \u0026plusmn;0.708a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eNumber of leaves***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e31.407\u0026plusmn; 9.604a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e38.846\u0026plusmn;10.573ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e40.896\u0026plusmn;10.896b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e51.958 \u0026plusmn;18.044b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTerminal thorn length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.042 \u0026plusmn; 0.010c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.143 \u0026plusmn; 0.207d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.047 \u0026plusmn; 0.012cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.027 \u0026plusmn;0.005b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTerminal thorn width*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.004\u0026plusmn; 0.001ab\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.006\u0026plusmn; 0.003ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.008 \u0026plusmn; 0.008b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.006 \u0026plusmn;0.003b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTerminal thorn length/width ratio ***\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e9.796 \u0026plusmn; 2.390bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e19.961 \u0026plusmn; 19.409c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e7.942 \u0026plusmn; 3.727b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e5.360 \u0026plusmn;3.266a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTerminal thorn/leaf length ratio ***\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.051 \u0026plusmn; 0.016b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.113 \u0026plusmn; 0.152b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.040 \u0026plusmn; 0.010b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.017 \u0026plusmn;0.005a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTeeth length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.011 \u0026plusmn; 0.003bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.012 \u0026plusmn; 0.001c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.011 \u0026plusmn; 0.003bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.009 \u0026plusmn;0.004b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTeeth/leaf length ratio ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.286 \u0026plusmn; 0.720b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.006 \u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.006 \u0026plusmn; 0.002a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.721 \u0026plusmn;0.506b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTeeth width***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.007 \u0026plusmn; 0.002b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.007 \u0026plusmn; 0.002b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.007 \u0026plusmn; 0.002b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.008 \u0026plusmn;0.002b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTeeth length/width ratio ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.723 \u0026plusmn; 0.204a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.611 \u0026plusmn; \u0026nbsp; \u0026nbsp; 0.192a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.691 \u0026plusmn; 0.202a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.065 \u0026plusmn;0.453b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eTeeth width/leaf length ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.013 \u0026plusmn; 0.006d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.010 \u0026plusmn; 0.001cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.009 \u0026plusmn; 0.002c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.005 \u0026plusmn;0.001b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eDistance between teeth\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.018 \u0026plusmn; 0.009a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.024 \u0026plusmn; 0.011a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.019 \u0026plusmn; 0.008a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.018 \u0026plusmn;0.007a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 187px;\"\u003e\n \u003cp\u003eDistance between teeth/leaf length ratio ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.023 \u0026plusmn; 0.013b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.019 \u0026plusmn; 0.008b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.017 \u0026plusmn; 0.007b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e0.012 \u0026plusmn;0.006a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eDifferent letters indicate significant differences. Significant level 0.001***, 0.01** and 0.05*\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe multivariate analyses PCA (Fig.5), LDA (Fig.8), and Heatmap (Fig.7) consistently revealed clear morphological differentiation within the \u003cem\u003eAgave americana\u003c/em\u003e complex based on vegetative and defensive traits. The PCA (Fig.6) showed that variables such as teeth length (TEEL), teeth-to-leaf length ratio (TEEL/LL), leaf length-to-width ratio (LL/LW), and terminal thorn dimensions (TTL, TTW) contributed most to the observed variation, separating taxa according to their leaf morphology and defensive structures. The LDA (Fig. 8) further distinguished individuals by taxonomy, management, and region: taxonomic groups (\u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. oaxacensis, A. americana\u0026nbsp;\u003c/em\u003esubsp\u003cem\u003e. protamericana, A. asperrima, and A. scaposa\u003c/em\u003e) formed distinct clusters; cultivated, managed, and wild categories were clearly separated along discriminant axes; and northern populations differed morphologically from central and southern ones. The Heatmap (Fig.7) supported these patterns, highlighting higher values of thorn and teeth ratios in wild taxa (\u003cem\u003eA. asperrima, A. a.\u0026nbsp;\u003c/em\u003esubsp\u003cem\u003e. protamericana\u003c/em\u003e), while cultivated and managed populations of \u003cem\u003eA. a. var. americana\u003c/em\u003e and \u003cem\u003eA. a.\u0026nbsp;\u003c/em\u003evar\u003cem\u003e\u0026nbsp;oaxacensis\u003c/em\u003e exhibited larger leaves and moderate defensive traits. Overall, the combined analyses indicate structured morphological variation across species, management regimes, and regions within the \u003cem\u003eA. americana\u003c/em\u003e complex.\u003c/p\u003e\n\u003cp\u003eThe multivariate analysis of variance (MANOVA) revealed differences among the analyzed categories for the evaluated morphological variables (Table 4). When comparing datasets obtained from herbarium specimens (H) and field-measured specimens (F), contrasting patterns were observed. For the Area factor, herbarium specimens did not show statistically significant differences (p = 0.09173), whereas field measurements revealed highly significant differences among groups (Wilks\u0026rsquo; \u0026lambda; = 0.00101; \u0026chi;\u0026sup2; = 615.38; DF = 76; p \u0026lt; 0.001). A similar pattern was detected for the Management factor: herbarium specimens showed marginally non-significant differences (p = 0.06601), while field measurements indicated strong statistical significance (Wilks\u0026rsquo; \u0026lambda; = 0.0055; \u0026chi;\u0026sup2; = 487.24; DF = 76; p \u0026lt; 0.001).\u003c/p\u003e\n\u003cp\u003eFor the Taxonomy factor, only field-measured specimens were analyzed, revealing highly significant differences among taxa (Wilks\u0026rsquo; \u0026lambda; = 0.00278; \u0026chi;\u0026sup2; = 734.63; DF = 133; p \u0026lt; 0.001). These results indicate that morphological variation among taxa is more clearly detected when measurements are obtained directly from field specimens. Overall, the analyses suggest that field measurements capture a greater proportion of morphological variability than herbarium-based measurements, likely due to preservation effects or measurement constraints associated with herbarium material.\u003c/p\u003e\n\u003cp\u003eThese results are consistent with the multivariate patterns observed in the PCA and heatmap analyses, which revealed clear grouping of species and differentiation associated with several floral and vegetative traits.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMorphological variation among taxa of the Agave americana complex\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eMean values and standard errors of the morphological traits that showed significant differences among taxa are presented in Table 7. Nineteen characters exhibited statistically significant variation among the analyzed taxa (p \u0026lt; 0.001 or p \u0026lt; 0.01), including vegetative size variables, leaf architecture, and defensive structures.\u003c/p\u003e\n\u003cp\u003eOverall, taxa belonging to the Agave americana complex showed substantial differences in plant size. A. americana var. americana exhibited the largest mean values for plant length (up to 2.922 \u0026plusmn; 0.841) and plant diameter (2.615 \u0026plusmn; 0.527), whereas smaller values were generally observed in \u003cem\u003eA. asperrima\u003c/em\u003e and var. \u003cem\u003eoaxacensis\u003c/em\u003e. Intermediate values were recorded for \u003cem\u003eA. asperrima and A. americana subsp. protamericana\u003c/em\u003e. These patterns indicate clear morphological differentiation in overall plant size among taxa.\u003c/p\u003e\n\u003cp\u003eStem length also varied significantly among groups. The longest stems were observed in \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. expansa\u003c/em\u003e (0.658 \u0026plusmn; 0.079), whereas shorter stems were characteristic of var. \u003cem\u003eoaxacensis\u003c/em\u003e and \u003cem\u003eA. asperrima\u003c/em\u003e. Differences in stem length contributed to the variation observed in plant architecture among taxa.\u003c/p\u003e\n\u003cp\u003eLeaf morphology showed marked variation as well. Leaf length ranged from 0.893 \u0026plusmn; 0.254 \u003cem\u003ein A. americana\u0026nbsp;\u003c/em\u003esubsp\u003cem\u003e. protamericana\u003c/em\u003e to 1.778 \u0026plusmn; 0.324 in \u003cem\u003eA. americana\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. americana\u003c/em\u003e. Similarly, the leaf length-to-width ratio differed significantly among taxa, reflecting differences in leaf shape and proportions. Taxa such as A. oaxacensis exhibited particularly high LL/SL values, indicating relatively elongated leaves compared with other species in the complex.\u003c/p\u003e\n\u003cp\u003eThe number of leaves per plant also differed significantly among taxa. Higher values were recorded in\u003cem\u003e\u0026nbsp;A. americana\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. expansa\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;\u003c/em\u003evar\u003cem\u003e. oaxacensis\u003c/em\u003e, whereas lower values were observed in \u003cem\u003eA. asperrima\u003c/em\u003e. These differences suggest variation in vegetative growth strategies across the complex.\u003c/p\u003e\n\u003cp\u003eDefensive traits, including terminal spine length and width, showed notable variation among taxa. Prominent terminal spines were observed in \u003cem\u003eA. americana var. abrupta\u003c/em\u003e, whereas smaller spines were characteristic of var\u003cem\u003e. oaxacensis\u003c/em\u003e. Ratios involving terminal spine dimensions (e.g., TTL/LTL and TTL/LL) also differed significantly, indicating differences in the relative development of defensive structures.\u003c/p\u003e\n\u003cp\u003eTraits related to marginal teeth\u0026mdash;including teeth length, teeth width, and the distance between teeth\u0026mdash;showed significant but comparatively moderate variation among taxa. Some taxa, such as \u003cem\u003eA. abrupta\u003c/em\u003e, exhibited longer teeth relative to leaf size, whereas others presented shorter and more widely spaced teeth.\u003c/p\u003e\n\u003cp\u003eOverall, these results demonstrate substantial morphological differentiation within the Agave americana complex and associated species. The significant variation observed across multiple vegetative and defensive traits supports the taxonomic distinctiveness of the analyzed taxa and reflects the morphological diversity present within the group.\u003c/p\u003e\n\u003cp\u003eSignificant differences were also detected among management categories (wild, promoted, managed in situ, and cultivated) for most of the evaluated morphological traits (Table 6). In general, cultivated plants exhibited larger vegetative structures than those observed in wild populations. Cultivated individuals showed the highest mean values for plant height (2.557 \u0026plusmn; 0.782), stem length (0.634 \u0026plusmn; 0.125), plant diameter (2.545 \u0026plusmn; 0.445), and leaf length (1.637 \u0026plusmn; 0.312), indicating a clear increase in overall plant size under more intensive management regimes.\u003c/p\u003e\n\u003cp\u003eIntermediate values were generally observed in the promoted and managed in situ categories. Traits such as plant height and plant diameter displayed a gradual increase from wild populations to promoted and managed plants, suggesting progressive morphological modification along the management gradient. In contrast, some variables, including leaf width and the leaf/stem length ratio, showed little variation among management categories.\u003c/p\u003e\n\u003cp\u003eLeaf architecture also differed among management types. Cultivated plants exhibited higher values of the leaf length-to-width ratio, indicating proportionally longer leaves, whereas wild plants generally had smaller leaves. In addition, the number of leaves was highest in cultivated individuals, suggesting that cultivation practices may promote enhanced vegetative growth.\u003c/p\u003e\n\u003cp\u003eDefensive traits also varied across management categories. Terminal spine length was greatest in promoted plants (0.143 \u0026plusmn; 0.207), whereas cultivated plants showed the lowest values (0.027 \u0026plusmn; 0.005). Similarly, the terminal spine length-to-width ratio was highest in promoted individuals, reflecting relatively elongated spines under intermediate management conditions. By contrast, cultivated plants tended to exhibit smaller and proportionally shorter terminal spines.\u003c/p\u003e\n\u003cp\u003eTraits associated with marginal teeth showed comparatively minor variation among categories. Both teeth length and teeth width remained relatively similar across management types, although slight differences were detected in the teeth-to-leaf length ratio, with cultivated plants presenting the highest values.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe domestication of \u003cem\u003eAgave\u003c/em\u003e represents a complex evolutionary process shaped by interactions among morphological traits, ecological adaptation, and human management. The morphological diversity observed across species reflects both environmental specialization and varying degrees of anthropogenic selection. This study contributes to understanding \u003cem\u003eAgave\u003c/em\u003e domestication by integrating vegetative and reproductive morphometrics from herbarium and field specimens, revealing how natural and cultural factors have guided morphological divergence and adaptation.\u003c/p\u003e \u003cp\u003eMultivariate analyses demonstrated consistent patterns of differentiation within the \u003cem\u003eAgave americana\u003c/em\u003e complex. The PCA (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e3\u003c/span\u003e) indicated that reproductive traits\u0026mdash;particularly inflorescence length (IL), ovary width (OW), and stamen length (SL)\u0026mdash;were major contributors to variation, defining a gradient from high reproductive investment to enhanced vegetative and defensive robustness. \u003cem\u003eA. americana\u003c/em\u003e var. \u003cem\u003eoaxacensis\u003c/em\u003e showed high reproductive development (positive PC1), while \u003cem\u003eA. a.\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e and \u003cem\u003eA. asperrima\u003c/em\u003e aligned with greater thorn and tooth dimensions (negative PC1/PC2), suggesting stronger defensive allocation. \u003cem\u003eA. a.\u003c/em\u003e subsp. \u003cem\u003eamericana\u003c/em\u003e occupied an intermediate position, combining large vegetative structures with moderate reproductive traits, and \u003cem\u003eA. scaposa\u003c/em\u003e and \u003cem\u003eA. expansa\u003c/em\u003e exhibited balanced morphologies. These patterns confirm that reproductive differentiation is a principal axis of species segregation within the complex [\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe LDA (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e4\u003c/span\u003e) reinforced these distinctions, clearly separating taxa, management regimes, and geographic regions. Cultivated and intensively cultivated individuals clustered apart from wild and managed populations, displaying larger inflorescences and ovaries but reduced thorn and tooth size\u0026mdash;morphological signatures consistent with domestication. In contrast, wild and managed plants retained smaller reproductive organs and stronger defensive traits. Regionally, northern populations differed from those in Oaxaca, Michoac\u0026aacute;n, and Jalisco, suggesting that ecological conditions and local management practices jointly drive morphological divergence across the \u003cem\u003eA. americana\u003c/em\u003e complex.\u003c/p\u003e \u003cp\u003eThe heatmap (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e) summarized these patterns, highlighting the trade-offs between reproductive investment and structural defense. \u003cem\u003eA. americana\u003c/em\u003e var. \u003cem\u003eoaxacensis\u003c/em\u003e exhibited the highest reproductive values (IL, OW), \u003cem\u003eA. a.\u003c/em\u003e subsp. \u003cem\u003eprotamericana\u003c/em\u003e displayed enhanced defensive structures (long, thick terminal thorns, prominent teeth), and \u003cem\u003eA. asperrima\u003c/em\u003e combined robust defensive and vegetative development with moderate reproduction. \u003cem\u003eA. scaposa\u003c/em\u003e and \u003cem\u003eA. expansa\u003c/em\u003e showed intermediate values across all traits, suggesting morphological equilibrium in these traits.\u003c/p\u003e \u003cp\u003eAn important finding of this study is the importance of reproductive traits, particularly floral morphology, in \u003cem\u003eAgave\u003c/em\u003e species differentiation. Floral features such as stamen and filament length, ovary dimensions, and inflorescence size emerged as key discriminant variables. These results align with Trejo et al. [\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e], who identified floral morphology as the most effective character set for species delimitation in \u003cem\u003eAgave\u003c/em\u003e, surpassing fruit and seed traits. The persistence of floral variability under domestication is vital for the resilience and adaptive capacity of \u003cem\u003eAgave\u003c/em\u003e populations. Even in cultivated and clonal stands, floral diversity remains a strong axis of variation, crucial for maintaining ecological stability under pressures such as climate change and pollinator decline [\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e, \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e]. This reproductive diversity supports long-term adaptability and evolutionary potential within the \u003cem\u003eA. americana\u003c/em\u003e complex [\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e64\u003c/span\u003e]\u003c/p\u003e \u003cp\u003eComplementary analyses of field-measured specimens revealed clear differentiation based on vegetative traits. The PCA (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e) emphasized the influence of leaf morphology, spine and tooth dimensions, and plant diameter on species segregation. PC1 captured variation associated with leaf form and defensive traits, consistent with studies identifying spines and teeth as key domestication indicators [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e]. PC2, linked to terminal thorn dimensions and plant diameter, distinguished species with large, robust morphologies (\u003cem\u003eA. protamericana, A. asperrima\u003c/em\u003e) from more compact, human-managed forms (\u003cem\u003eA. americana\u003c/em\u003e). This trend supports domestication patterns characterized by reduced defensive traits and enhanced vegetative growth [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eLDA results based on field specimens also demonstrated strong separation among taxa, management regimes, and regions. Cultivated individuals exhibited trait combinations characteristic of a domestication syndrome [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e], while regional clustering indicated the influence of both artificial selection and local environmental factors such as temperature, precipitation, and soil conditions [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. The heatmap corroborated these distinctions: wild species (\u003cem\u003eA. asperrima, A. protamericana\u003c/em\u003e) exhibited pronounced defensive features, while cultivated and managed populations showed larger leaves and reduced spine development [\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe morphological differences observed between species in the \u003cem\u003eAgave americana\u003c/em\u003e complex have significant ecological and evolutionary implications. Wild species, exposed to greater herbivory pressure and environmental stressors, have developed robust defense strategies, such as long spines and thick leaves, to protect themselves from predators and competitors. In contrast, managed species, which are under human control, exhibit traits that prioritize vegetative growth and biomass accumulation, such as larger plant sizes and greater leaf production. These traits are selected during domestication to meet human needs, such as food production or fiber harvesting [\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e69\u003c/span\u003e, \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e70\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe human-directed evolution through management practices such as cultivation and breeding has favored traits that enhance the usefulness of these plants, but it has also shaped their genetic and morphological diversity. Wild and less-managed species have retained a repertoire of defensive characteristics that enable them to survive in more competitive natural environments. This morphological diversity could be crucial for future conservation strategies, as cultivated and managed populations may be more vulnerable to rapid environmental changes, while wild populations may offer a genetic foundation for future adaptation to new ecological conditions [\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e71\u003c/span\u003e, \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eIntegrating multivariate morphometries provides a robust framework for examining the domestication of \u003cem\u003eAgave americana\u003c/em\u003e. By analyzing vegetative and reproductive morphological traits, clear patterns of differentiation can be established among wild, cultivated, and promoted forms. In particular, traits such as plant size, leaf characteristics, and the size of terminal spines and teeth are key indicators of human management influence. Wild plants tend to be smaller, with fewer leaves and larger terminal spines and teeth, while cultivated plants exhibit intermediate characteristics. Those managed in situ display traits that are closer to those of cultivated forms. These patterns, which have been documented in other plant species [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e], reinforce the idea that domestication can occur in both cultivated plants and wild populations that have been incorporated into human-managed environments.\u003c/p\u003e \u003cp\u003eIntegrating morphological data with cultural and ecological knowledge is crucial for developing effective conservation strategies for \u003cem\u003eAgave\u003c/em\u003e species. The study emphasizes the importance of maintaining genetic diversity through both sexual and asexual reproduction, and highlights the need to incorporate seed production into management practices. \u003cem\u003eAgave\u003c/em\u003e populations that rely on clonal propagation, in particular may experience a reduction in genetic diversity, which could limit their ability to adapt to changing environmental conditions. Therefore, preserving sexual reproduction is essential for maintaining the long-term viability of these plants. Furthermore, research into the reproductive biology of \u003cem\u003eAgave\u003c/em\u003e species, particularly focusing on hybridization events and gene flow between wild and cultivated populations, would provide valuable insights into the evolutionary processes shaping these plants.\u003c/p\u003e \u003cp\u003eOverall, these results (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e and Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e) indicate that management intensity influences several vegetative and defensive traits. Cultivated plants generally develop larger vegetative structures and reduced spine dimensions compared with wild populations, reflecting the effects of human selection under cultivation.\u003c/p\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003eRecommendations for Future Research\u003c/h2\u003e \u003cp\u003eThe \u003cem\u003eAgave americana\u003c/em\u003e complex exhibits significant morphological diversity, driven by both natural and human-induced evolutionary forces. The multivariate analyses provide a deep understanding of how taxonomic factors, geographic conditions, and management regimes have shaped the vegetative, defensive and reproductive traits of these species. This knowledge is essential for sustainable management and conservation of \u003cem\u003eAgave\u003c/em\u003e species, offering new perspectives on how these plants adapt to selective pressures and how conservation efforts can support their survival under changing conditions [\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e, \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e70\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eTo advance our understanding of \u003cem\u003eAgave\u003c/em\u003e domestication, future studies should focus on expanding the geographic and ecological scope of research to include a broader range of taxa and populations. Sampling should be increased among fertile individuals and collaboration with local communities should be established to document traditional management practices. Genetic analyses, particularly the use of next-generation sequencing techniques and phylogeographic approaches, would be crucial for answering questions about the origin and diffusion of \u003cem\u003eAgave\u003c/em\u003e taxa. Such studies will also shed light on the evolutionary relationships between species and the diffusion routes of \u003cem\u003eAgave\u003c/em\u003e domestication throughout history, particularly in relation to \u003cem\u003eAgave cordillerensis\u003c/em\u003e and its connection to the \u003cem\u003eA. americana\u003c/em\u003e complex [\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e74\u003c/span\u003e, \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e75\u003c/span\u003e] .\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn summary, this study deepens our understanding of the complex processes underlying the domestication of \u003cem\u003eAgave\u003c/em\u003e. The interplay between morphological traits, management practices, and reproductive characteristics sheds light on how these plants have evolved under both natural and anthropogenic selection pressures. Integrating morphological, ecological, and genetic data, alongside cultural knowledge, is essential to understanding the full scope of \u003cem\u003eAgave\u003c/em\u003e domestication and its implications for conservation and sustainable management. Future research should continue to explore the genetic diversity, management practices, and reproductive biology of \u003cem\u003eAgave\u003c/em\u003e species paying particular attention to the role of floral traits in species delimitation and ecological adaptability.\u003c/p\u003e \u003cp\u003eOur findings complement and extend the analysis of \u003cem\u003eAgave\u003c/em\u003e domestication of the morphological diversity [\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e] demonstrating that reproductive characters, particularly floral traits, play a pivotal role in the organization of \u003cem\u003eAgave\u003c/em\u003e diversity. The persistence of floral variability under domestication, the phenotypic differentiation it enables among taxa, and its potential contribution to ecological adaptability highlight its importance as both a taxonomic tool but and a conservation priority. Together, the two datasets advocate for the continued integration of floral morphology in the study, management, and protection of \u003cem\u003eAgave\u003c/em\u003e genetic resources.\u003c/p\u003e \u003cp\u003eThe identified patterns are not conclusive; further details of management practices are required. The people of Mexico have managed \u003cem\u003eAgave\u003c/em\u003es for a long time for multiple purposes, so it is important to document these details to improve our understanding of domestication. Morphometric studies should be conducted at more sites, to cover the distribution ranges of the taxa included in this analysis. While it is difficult, it is not impossible to broaden the sampling of fertile individuals is indispensable, but it requires agreements with people who make use of these \u003cem\u003eAgave\u003c/em\u003es. Studies of reproductive biology would help to identify hybridisation events and how this process occurs. Finally, population genetics studies are needed to evaluate the state of genetic diversity and gene flow among wild and managed populations, as well as among the different analysed taxa.\u003c/p\u003e \u003cp\u003ePhylogeographic studies would be particularly useful for answering questions about the origin and spread of the taxa that make up \u003cem\u003eAgave americana\u003c/em\u003e. The question of whether the domesticated taxa have one or more ancestors remains unanswered. Phylogeographic approaches using next-generation genetic markers could help to identify the evolutionary relationships among the taxa, confirm or refute their identities, and establish the possible diffusion routes throughout history. Of particular interest is the South American taxon \u003cem\u003eAgave cordillerensis\u003c/em\u003e, which was long considered to be \u003cem\u003eA. americana\u003c/em\u003e. This \u003cem\u003eAgave\u003c/em\u003e species is widely distributed in the Andean region, from Colombia to Bolivia, but it does not appear to have been present for long time. The evolutionary relationship between this \u003cem\u003eAgave\u003c/em\u003e species and the \u003cem\u003eA. americana\u003c/em\u003e complex is an interesting question that could be answered that could be addressed through phylogeographic approaches providing valuable insights into the historical exchange of crops and domestication techniques between Mesoamerica and the Andean region.\u003c/p\u003e \u003cp\u003eThis is merely an initial study of the domestication of \u003cem\u003eAgave americana\u003c/em\u003e complex. While we are aware of the limitations of the information current collected, it enables us to identify the relevant taxa, regions, cultures, and research approaches to answering questions and improving our understanding of this group of plants, and other \u003cem\u003eAgave\u003c/em\u003e complexes in their main domestication setting: Mesoamerica.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eN.C.S. conceived and designed the study, conducted fieldwork and data collection, and performed the morphological measurements and statistical analyses. N.C.S. organized the datasets, interpreted the results, and wrote the initial draft of the manuscript, including the preparation of tables and figures.\u003c/p\u003e\n\u003cp\u003eA.C. supervised the research as the principal doctoral advisor and contributed to the conceptual framework and scientific direction of the study. A.C. provided guidance throughout the research process, contributed to the interpretation and discussion of the results, and participated in the writing process through critical revision, editing, and improvement of the manuscript.\u003c/p\u003e\n\u003cp\u003eBoth authors discussed the results, reviewed the manuscript critically, and approved the final version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was supported by the National Council of Science and Technology of Mexico (CONACYT) through a doctoral scholarship granted to N.C.S. Additional financial support was provided by CONACYT (research project A1-S-14306), the Programa de Apoyo a Proyectos de Investigaci\u0026oacute;n e Innovaci\u0026oacute;n Tecnol\u0026oacute;gica (PAPIIT) of the Direcci\u0026oacute;n General de Asuntos del Personal Acad\u0026eacute;mico (DGAPA, UNAM, Mexico; project IN224023), and by the National Commission for the Knowledge and Use of Biodiversity (CONABIO) together with the FAO\u0026ndash;GEF project Agrobiodiversity of Mexico (ID 9380) through project RG023. The funding bodies had no role in the design of the study, data collection, analysis, interpretation of the data, or in writing the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analyzed during the current study are available from the corresponding author upon reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors thank the National Council of Science and Technology of Mexico (CONACYT) for the doctoral scholarship granted to the first author and the Posgrado en Ciencias Biol\u0026oacute;gicas at the Instituto de Investigaciones en Ecosistemas y Sustentabilidad (IIES), Universidad Nacional Aut\u0026oacute;noma de M\u0026eacute;xico (UNAM), campus Morelia, for academic support during the development of this research.\u003c/p\u003e\n\u003cp\u003eWe also acknowledge the financial support provided by CONACYT (project A1-S-14306), the Programa de Apoyo a Proyectos de Investigaci\u0026oacute;n e Innovaci\u0026oacute;n Tecnol\u0026oacute;gica (PAPIIT\u0026ndash;DGAPA, UNAM; project IN224023), and the National Commission for the Knowledge and Use of Biodiversity (CONABIO) together with the FAO\u0026ndash;GEF project Agrobiodiversity of Mexico (ID 9380) through project RG023.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eDarwin C. On the origin of species by means of natural selection, or the preservation of favored races in the struggle for life. London: John Murray; 1859.\u003c/li\u003e\n \u003cli\u003eDarwin, C. The variation of animals and plants under domestication. London: John Murray; 1868.\u003c/li\u003e\n \u003cli\u003eHarlan JR. Crops and man. Madison: American Society of Agronomy; 1975.\u003c/li\u003e\n \u003cli\u003eHarlan JR. Agricultural origins: centers and non-centers. Science. 1971;174:468\u0026ndash;474.\u003c/li\u003e\n \u003cli\u003eVavilov NI. Origin and geography of cultivated plants. Cambridge: Cambridge University Press; 1992.\u003c/li\u003e\n \u003cli\u003eHammer K. Das domestikationssyndrom. Die Kulturpflanze. 1984;32(1):11\u0026ndash;34.\u003c/li\u003e\n \u003cli\u003eHawkes JG. The diversity of crop plants. Cambridge: Harvard University Press; 1983.\u003c/li\u003e\n \u003cli\u003eGepts P. Crop domestication as a long-term selection experiment. Plant Breed Rev. 2004;24:1\u0026ndash;44.\u003c/li\u003e\n \u003cli\u003ePickersgill B. Domestication of plants in the Americas. Ann Bot. 2007;100:925\u0026ndash;940.\u003c/li\u003e\n \u003cli\u003ePickersgill B. Parallel vs convergent evolution in domestication. Front Ecol Evol. 2018;6:56.\u003c/li\u003e\n \u003cli\u003eSmith BD. The initial domestication of Cucurbita pepo. Science. 1997;276:932\u0026ndash;934.\u003c/li\u003e\n \u003cli\u003eMacNeish RS. The origins of agriculture and settled life. Norman: University of Oklahoma Press; 1992.\u003c/li\u003e\n \u003cli\u003eFlannery K, editor. Guil\u0026aacute; Naquitz. New York: Academic Press; 1986.\u003c/li\u003e\n \u003cli\u003eCasas, A., Caballero, J., Mapes, C. \u0026amp; Z\u0026aacute;rate, S.. Manejo de la vegetaci\u0026oacute;n, domesticaci\u0026oacute;n de plantas y origen de la agricultura en Mesoam\u0026eacute;rica. Boletin De La Sociedad Botanica De Mexico. 2017.\u003c/li\u003e\n \u003cli\u003eCasas, A., Valiente-Banuet, A., Viveros, J. L., Caballero, J., Cort\u0026eacute;s, L., D\u0026aacute;vila, P., Lira, R. \u0026amp; Rodr\u0026iacute;guez, I.. Plant resources of the Tehuac\u0026aacute;n-Cuicatl\u0026aacute;n valley, Mexico. Economic Botany. 2000;55(1).\u003c/li\u003e\n \u003cli\u003eCasas, A., Otero-Arnaiz, A., P\u0026eacute;rez-Negr\u0026oacute;n, E. \u0026amp; Valiente-Banuet, A.. In situ Management and Domestication of Plants in Mesoamerica. Annals of Botany. 2007;100(5).\u003c/li\u003e\n \u003cli\u003eCasas A, Parra F, Aguirre-Dugua X, et al. Manejo y domesticaci\u0026oacute;n de plantas en Mesoam\u0026eacute;rica. In: Domesticaci\u0026oacute;n en el Continente Americano. Vol. 2. Morelia: UNAM; 2017. p. 69\u0026ndash;102.\u003c/li\u003e\n \u003cli\u003eCasas A, Torres I, Parra F, Torres-Guevara J. Centros de origen y diversificaci\u0026oacute;n de plantas cultivadas. De los cultivos nativos y el cambio del clima. Hallazgos (Hu\u0026aacute;nuco y Apur\u0026iacute;mac) (pp.23-56) Lima: UNALM; 2019. p. 23\u0026ndash;62.\u003c/li\u003e\n \u003cli\u003eBlancas, J., Casas, A., Rangel-Landa, S., Moreno-Calles, A. I., Torres, I., P\u0026eacute;rez-Negr\u0026oacute;n, E., Sol\u0026iacute;s, L., Delgado-Lemus, A., Parra, F., Arellanes, Y., Caballero, J., Cort\u0026eacute;s, L., Lira, R. \u0026amp; D\u0026aacute;vila, P.. Plant Management in the Tehuac\u0026aacute;n-Cuicatl\u0026aacute;n Valley, Mexico1. Economic Botany. 2010;64(4).\u003c/li\u003e\n \u003cli\u003eBoege E. El patrimonio biocultural de los pueblos ind\u0026iacute;genas de M\u0026eacute;xico. Mexico City: INAH; 2008.\u003c/li\u003e\n \u003cli\u003eBye RA, Burgess D, Mares-Trias A. Ethnobotany of the western Tarahumara. Bot Mus Leafl Harv Univ. 1975;24:85\u0026ndash;112.\u003c/li\u003e\n \u003cli\u003eGarc\u0026iacute;a-Mendoza AJ. Agav\u0026aacute;ceas. In: Biodiversidad de Oaxaca. M\u0026eacute;xico: UNAM; 2004. p. 159\u0026ndash;169.\u003c/li\u003e\n \u003cli\u003eGarc\u0026iacute;a-Mendoza AJ. Flora del Valle de Tehuac\u0026aacute;n-Cuicatl\u0026aacute;n. Agavaceae. M\u0026eacute;xico: UNAM; 2011.\u003c/li\u003e\n \u003cli\u003eGentry HS. Agaves of continental North America. Tucson: University of Arizona Press; 1982.\u003c/li\u003e\n \u003cli\u003ePalma-Cruz FJ. El g\u0026eacute;nero Agave y su distribuci\u0026oacute;n en Oaxaca. Tesis de licenciatura. UNAM; 1991.\u003c/li\u003e\n \u003cli\u003ePalma-Cruz FJ. Agaves productores de fibras duras en Oaxaca. Bol Soc Bot Mex. 2000;66:93\u0026ndash;102.\u003c/li\u003e\n \u003cli\u003eVargas-Ponce, O. and Zizumbo-Villarreal, D. In situ diversity and maintanenace of traditional Agave landraces used in spirits production in West-Central Mexico. Economic botany. 2006.\u003c/li\u003e\n \u003cli\u003eVargas-Ponce, O., Zizumbo-Villarreal, D., Mart\u0026iacute;nez-Castillo, J., Coello-Coello, J. \u0026amp; Colunga-Garc\u0026iacute;amar\u0026iacute;n, P.. Diversity and structure of landraces of \u003cem\u003eAgave\u003c/em\u003e grown for spirits under traditional agriculture: A comparison with wild populations of A. angustifolia (Agavaceae) and commercial plantations of \u003cem\u003eA. tequilana\u003c/em\u003e.. American-Eurasian journal of botany. 2009;96 2.\u003c/li\u003e\n \u003cli\u003eTorres, I., Blancas, J., Le\u0026oacute;n, A. \u0026amp; Casas, A. TEK, local perceptions of risk, and diversity of management practices of \u003cem\u003eAgave inaequidens\u003c/em\u003e in Michoac\u0026aacute;n, Mexico. Journal of Ethnobiology and Ethnomedicine. 2015;11(1).\u003c/li\u003e\n \u003cli\u003eTorres-Garc\u0026iacute;a, I., Rend\u0026oacute;n-Sandoval, F. J., Blancas, J., Casas, A. \u0026amp; Moreno-Calles, A.. The genus Agave in agroforestry systems of Mexico. Botan‪ical Sciences. 2019.\u003c/li\u003e\n \u003cli\u003eTorres-Garc\u0026iacute;a I, Delgado-Lemus AM, Casas A, \u0026Aacute;lvarez-R\u0026iacute;os GD, Rangel-Landa S, Mart\u0026iacute;nez-Jim\u0026eacute;nez R, et al. Agave americana L., Agave angustifolia Haw., Agave atrovirens Karw. ex Salm-Dyck, Agave asperrima Jacobi, Agave bovicornuta Gentry, Agave cupreata Trel. \u0026amp; A. Berger, Agave hookeri Jacobi, Agave inaequidens K. Koch, Agave karwinskii Zucc., Agave kerchovei Lem., Agave lechuguilla Torr., Agave mapisaga Trel., Agave marmorata Roezl, Agave maximiliana Baker, Agave montana Villareal, Agave potatorum Zucc., Agave rhodacantha Trel., Agave salmiana Otto ex Salm-Dyck, Agave scaposa Gentry, Agave tequilana F.A.C. Weber, Agave victoriae-reginae A. Berger (Asparagaceae). In: Casas A, Blancas V\u0026aacute;zquez JJ, editors. Ethnobotany of the Mountain Regions of Mexico. Cham: Springer International Publishing; 2022. p. 1\u0026ndash;52. doi: 10.1007/978-3-030-99357-3_26\u003c/li\u003e\n \u003cli\u003eClement, C., Casas, A., Parra-Rondinel, F., Levis, C., Peroni, N., Hanazaki, N., Cort\u0026eacute;s-Z\u0026aacute;rraga, L., Rangel-Landa, S., Alves, R. P., Ferreira, M. J., Cassino, M. F., Coelho, S. D., Cruz-Soriano, A., Pancorbo-Olivera, M., Blancas, J., Mart\u0026iacute;nez-Ballest\u0026eacute;, A., Lemes, G., Lotero-Vel\u0026aacute;squez, E., Bertin, V. \u0026amp; Mazzochini, G. G.. Disentangling Domestication from Food Production Systems in the Neotropics. Quaternary. 2021.\u003c/li\u003e\n \u003cli\u003eAlducin-Mart\u0026iacute;nez C, Ruiz Mondrag\u0026oacute;n KY, Jim\u0026eacute;nez-Barr\u0026oacute;n O, Aguirre-Planter E, Gasca-Pineda J, Eguiarte LE, Medellin RA. Uses, Knowledge and Extinction Risk Faced by Agave Species in Mexico. Plants. 2023; 12(1):124. https://doi.org/10.3390/plants12010124.\u003c/li\u003e\n \u003cli\u003e\u0026Aacute;lvarez-R\u0026iacute;os, G. D., Pacheco-Torres, F., Figueredo-Urbina, C. J. \u0026amp; Casas, A.. Management, morphological and genetic diversity of domesticated agaves in Michoac\u0026aacute;n, M\u0026eacute;xico.. Journal of Ethnobiology and Ethnomedicine. 2020;16.\u003c/li\u003e\n \u003cli\u003eCabrera-Toledo D, Vargas-Ponce O, Ascencio-Ram\u0026iacute;rez S, Valadez-Sandoval LM, P\u0026eacute;rez-Alquicira J, Morales-Saavedra J and Huerta-Galv\u0026aacute;n OF. Morphological and Genetic Variation in Monocultures, Forestry Systems and Wild Populations of Agave maximiliana of Western Mexico: Implications for Its Conservation. Front. Plant Sci. 2020;11:817. doi: 10.3389/fpls.2020.00817\u003c/li\u003e\n \u003cli\u003eFigueredo, C., Casas, A., Gonz\u0026aacute;lez‐Rodr\u0026iacute;guez, A., Nassar, J., Colunga-Garc\u0026iacute;amar\u0026iacute;n, P. \u0026amp; Rocha-Ram\u0026iacute;rez, V.. Genetic structure of coexisting wild and managed agave populations: implications for the evolution of plants under domestication. AoB PLANTS. 2015.\u003c/li\u003e\n \u003cli\u003eFigueredo-Urbina CJ, Casas A, Torres-Garc\u0026iacute;a I. Morphological and genetic divergence between Agave inaequidens, A. cupreata and the domesticated A. hookeri. Analysis of their evolutionary relationships. PLOS ONE 2017;12(11): e0187260. https://doi.org/10.1371/journal.pone.0187260\u003c/li\u003e\n \u003cli\u003eFigueredo-Urbina CJ, Casas A, Mart\u0026iacute;nez-D\u0026iacute;az Y, et al. Domestication and saponins contents in a gradient of management intensity of agaves: \u003cem\u003eAgave cupreata\u003c/em\u003e, \u003cem\u003eA. inaequidens\u003c/em\u003e and \u003cem\u003eA. hookeri\u003c/em\u003e in central Mexico. \u003cem\u003eGenet Resour Crop Evol\u003c/em\u003e. 2018;\u003cstrong\u003e65\u003c/strong\u003e, 1133\u0026ndash;1146 (2018). https://doi.org/10.1007/s10722-017-0601-6\u003c/li\u003e\n \u003cli\u003eFigueredo, C.J., Casas, A., Colunga-Garc\u0026iacute;aMar\u0026iacute;n, P. et al. Morphological variation, management and domestication of \u0026lsquo;maguey alto\u0026rsquo; (Agave inaequidens) and \u0026lsquo;maguey manso\u0026rsquo; (A. hookeri) in Michoac\u0026aacute;n, M\u0026eacute;xico. J Ethnobiology Ethnomedicine 2014;10:66.. https://doi.org/10.1186/1746-4269-10-66\u003c/li\u003e\n \u003cli\u003eGonz\u0026aacute;lez Elizondo, M., R. Galv\u0026aacute;n Villanueva, I. L. L\u0026oacute;pez Enriquez, L. Res\u0026eacute;ndiz Rojas y M. S. Gonz\u0026aacute;lez Elizondo. 2009. Agaves-magueyes, lechuguillas y noas-del Estado de Durango y sus alrededores. Centro Interdisciplinario de Investigaci\u0026oacute;n para el Desarrollo Integral Regional-Unidad Durango, Instituto Polit\u0026eacute;cnico Nacional-Comisi\u0026oacute;n Nacional para el Conocimiento y Uso de la Biodiversidad. Durango, Dgo. 163 pp.\u003c/li\u003e\n \u003cli\u003eHern\u0026aacute;ndez-Sandoval L, Gonz\u0026aacute;lez Romo C, Gonz\u0026aacute;lez Medrano F. Plantas \u0026uacute;tiles de Tamaulipas. An Inst Biol UNAM. 1991.\u003c/li\u003e\n \u003cli\u003eMacNeish RS. A summary of the subsistence. Prehistory of Tehuacan Valley. 1967;1:290\u0026ndash;309.\u003c/li\u003e\n \u003cli\u003eMiller, A. J. and Schaal, B.. Domestication of a Mesoamerican cultivated fruit tree, Spondias purpurea.. Proceedings of the National Academy of Sciences of the United States of America. 2005.\u003c/li\u003e\n \u003cli\u003eParra F, Blancas JJ, Casas A. Landscape management and domestication of Stenocereus pruinosus (Cactaceae) in the Tehuac\u0026aacute;n Valley: human guided selection and gene flow. J Ethnobiol Ethnomed. 2012;14;8:32. doi: 10.1186/1746-4269-8-32.\u003c/li\u003e\n \u003cli\u003eHancock JF. Plant evolution and the origin of crop species. Cambridge: CABI; 2006.\u003c/li\u003e\n \u003cli\u003eThiede, J. Agave Agavaceae. En U. Egli \u0026amp; R. Nyffeler (Eds.), Illustrated handbook of succulent plants: Monocotyledons. 2020 (2nd ed., pp. 21-311). Springer-Verlag.\u003c/li\u003e\n \u003cli\u003eVillase\u0026ntilde;or Jos\u0026eacute; Luis. Checklist of the native vascular plants of Mexico. Rev. Mex. Biodiv. 2016; 87( 3 ): 559-902. https://doi.org/10.1016/j.rmb.2016.06.017.\u003c/li\u003e\n \u003cli\u003eTrelease W. Agave in the West Indies. Whitefish: Kessinger; 1912.\u003c/li\u003e\n \u003cli\u003eValenzuela-Zapata, A. G.. Tequila: A Natural and Cultural History. . 2002.\u003c/li\u003e\n \u003cli\u003eUrrutia-Cruz SP. Etnobot\u0026aacute;nica de los agaves en Oaxaca. Tesis de licenciatura. UNAM; 1986.\u003c/li\u003e\n \u003cli\u003eBarbara Pickersgill, Domestication of Plants in the Americas: Insights from Mendelian and Molecular Genetics, Annals of Botany 2007;100:925\u0026ndash;940, https://doi.org/10.1093/aob/mcm193\u003c/li\u003e\n \u003cli\u003eSmith, B. D.. The Initial Domestication of Cucurbita pepo in the Americas 10,000 Years Ago. Science 276, no. 5314 (1997): 932\u0026ndash;34. http://www.jstor.org/stable/2893022.\u003c/li\u003e\n \u003cli\u003eRodr\u0026iacute;guez-Garay, B., Lomel\u0026iacute;-Senci\u0026oacute;n, J., Tapia-Campos, E., Guti\u0026eacute;rrez-Mora, A., Garc\u0026iacute;a-Galindo, J., Rodr\u0026iacute;guez-Dom\u0026iacute;nguez, J., Urbina-L\u0026oacute;pez, D. \u0026amp; Vicente-Ram\u0026iacute;rez, I.. Morphological and molecular diversity of Agave tequilana Weber var. Azul and Agave angustifolia Haw. var. Line\u0026ntilde;o.. Industrial Crops and Products. 2008;29(1).\u003c/li\u003e\n \u003cli\u003eOjeda-Linares C. , \u0026Aacute;lvarez-R\u0026iacute;os G. , Figueredo-Urbina C. , Islas L. , Lappe-Oliveras P. , Nabhan G. et al.. Traditional Fermented Beverages of Mexico: A Biocultural Unseen Foodscape. Foods 2021;10(10):2390. https://doi.org/10.3390/foods10102390\u003c/li\u003e\n \u003cli\u003eCasas, A., Otero-Arnaiz, A., P\u0026eacute;rez-Negr\u0026oacute;n, E. \u0026amp; Valiente-Banuet, A.. In situ Management and Domestication of Plants in Mesoamerica. Annals of Botany. 2007;100(5).\u003c/li\u003e\n \u003cli\u003eCl\u0026eacute;ment C. , Casas A. , Parra F. , Levis C. , Peroni N. , Hanazaki N. et al.. Disentangling Domestication from Food Production Systems in the Neotropics. Quaternary 2021;4(1):4. https://doi.org/10.3390/quat4010004\u003c/li\u003e\n \u003cli\u003eGarc\u0026iacute;a-Mendoza AJ. Agavaceae of Oaxaca. Biodiversidad de Oaxaca. 2004\u003c/li\u003e\n \u003cli\u003eBlancas J. , Casas A. , Rangel-Landa S. , Moreno-Calles A. , Torres-Garc\u0026iacute;a I. , P\u0026eacute;rez‐Negr\u0026oacute;n E. et al.. Plant Management in the Tehuac\u0026aacute;n-Cuicatl\u0026aacute;n Valley, Mexico1. Economic Botany 2010;64(4):287-302. https://doi.org/10.1007/s12231-010-9133-0\u003c/li\u003e\n \u003cli\u003eCasas A. , Ladio A. , \u0026amp; Cl\u0026eacute;ment C.. Editorial: Ecology and Evolution of Plants Under Domestication in the Neotropics. Frontiers in Ecology and Evolution 2019;7. https://doi.org/10.3389/fevo.2019.00231\u003c/li\u003e\n \u003cli\u003ePalma-Cruz FJ. Agave in Oaxaca. Bol Soc Bot Mex. 2000;66:93\u0026ndash;102.\u003c/li\u003e\n \u003cli\u003eTrejo L. , Soriano D. , Romano-Grande E. , S\u0026aacute;nchez-Carmona B. , \u0026amp; D\u0026aacute;vila-Navarro D.. Diversity of reproductive characters, seed set, and viability of Agave seeds used for pulque production and their wild relatives in Tlaxcala, Mexico. Genetic Resources and Crop Evolution 2023;71(6):2877-2903. https://doi.org/10.1007/s10722-023-01803-5\u003c/li\u003e\n \u003cli\u003eFigueredo-Urbina C. , Casas A. , Colunga‐Garc\u0026iacute;aMar\u0026iacute;n P. , Nassar J. , \u0026amp; Gonz\u0026aacute;lez‐Rodr\u0026iacute;guez A.. Morphological variation, management and domestication of \u0026lsquo;maguey alto\u0026rsquo; (Agave inaequidens) and \u0026lsquo;maguey manso\u0026rsquo; (A. hookeri) in Michoac\u0026aacute;n, M\u0026eacute;xico. Journal of Ethnobiology and Ethnomedicine 2014;10(1). https://doi.org/10.1186/1746-4269-10-66\u003c/li\u003e\n \u003cli\u003e\u0026Aacute;lvarez-R\u0026iacute;os G. , Pacheco-Torres F. , Figueredo-Urbina C. , \u0026amp; Casas A.. Management, morphological and genetic diversity of domesticated agaves in Michoac\u0026aacute;n, M\u0026eacute;xico. Journal of Ethnobiology and Ethnomedicine 2020;16(1). https://doi.org/10.1186/s13002-020-0353-9\u003c/li\u003e\n \u003cli\u003eKlimova, A.;Ruiz Mondrag\u0026oacute;n, K.Y.;Molina Freaner, F.;Aguirre-Planter, E.; Eguiarte, L.E. Genomic Analyses of Wild and Cultivated Bacanora Agave (\u003cem\u003eAgave angustifolia\u003c/em\u003e var. \u003cem\u003epaci\u003c/em\u003e\u003cem\u003efi\u003c/em\u003e\u003cem\u003eca\u003c/em\u003e) Reveal Inbreeding, Few Signs of Cultivation History and Shallow Population Structure. Plants 2022,11, 1426. https://doi.org/10.3390/plants11111426\u003c/li\u003e\n \u003cli\u003eCasas, A., S. Rangel-Landa, E. Torres-Garc\u0026iacute;a, E. P\u0026eacute;rez-Negr\u0026oacute;n, L. Sol\u0026iacute;s, F. Parra, A. Delgado, J. Blancas J, B. Farf\u0026aacute;n, and A. I. Moreno.. In situ management and conservation of plant resources. In: De Albuquerque, U. P., and M. Alves Ramos (eds). Current Topics in Ethnobotany. Research Signpost, Kerala, India. 2008 pp: 1-25.\u003c/li\u003e\n \u003cli\u003eParra, F., N. P\u0026eacute;rez-Nasser, R. Lira, D. P\u0026eacute;rez- Salicrup, and A. Casas.. Population genetics, and process of domestication of \u003cem\u003eStenocereus pruinosus\u003c/em\u003e (Cactaceae) in the Tehuacan Valley, Mexico. J. Arid Environ. 2008; 72: 1997-2010.\u003c/li\u003e\n \u003cli\u003eCasas A, Otero-Arnaiz A, P\u0026eacute;rez-Negr\u0026oacute;n E, Valiente-Banuet A. In situ management and domestication of plants in Mesoamerica. Ann Bot. 2007;100:1101\u0026ndash;1115.\u003c/li\u003e\n \u003cli\u003eClement CR, Casas A, Parra-Rondinel FA, et al. Disentangling domestication from food production systems in the Neotropics. Quaternary. 2021;4:4.\u003c/li\u003e\n \u003cli\u003eGepts P. Crop domestication as a long-term selection experiment. Plant Breed Rev. 2004;24:1\u0026ndash;44.\u003c/li\u003e\n \u003cli\u003eHancock JF. Plant evolution and the origin of crop species. 2nd ed. CABI; 2006.\u003c/li\u003e\n \u003cli\u003eParra F, Blancas J, Casas A. Landscape management and domestication of Stenocereus pruinosus. J Ethnobiol Ethnomed. 2012;8:32.\u003c/li\u003e\n \u003cli\u003eTrejo L, Reyes M, Cort\u0026eacute;s-Toto D, et al. Morphological diversity and genetic relationships in pulque production agaves. Front Plant Sci. 2020;11:524812.\u003c/li\u003e\n \u003cli\u003eRodr\u0026iacute;guez-Garay B, Lomel\u0026iacute;-Senci\u0026oacute;n JA, Tapia-Campos E, et al. Morphological and molecular diversity of Agave tequilana. Ind Crops Prod. 2009;29:220\u0026ndash;228.\u003c/li\u003e\n \u003cli\u003eOjeda-Linares C, \u0026Aacute;lvarez-R\u0026iacute;os GD, Figueredo-Urbina CJ, et al. Traditional fermented beverages of Mexico. Foods. 2021;10:2390.\u003c/li\u003e\n \u003cli\u003eTrejo L, Soriano D, Romano-Grande E, et al. Diversity of reproductive characters in Agave used for pulque. Genet Resour Crop Evol. 2024;71:2877\u0026ndash;2903.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table","content":"\u003cp\u003eTable 7. Means and standard error of 19 characters that presented significant differences when comparing the plants of\u0026nbsp;Agave americana\u0026nbsp;complex and associated species of\u0026nbsp;Americanae Group.\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"98%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 12px;\"\u003e\n \u003cp\u003eCharacter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" style=\"width: 21px;\"\u003e\n \u003cp\u003e\u003cem\u003evar americana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cem\u003evar expansa\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cem\u003eA abrupta\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8px;\"\u003e\n \u003cp\u003e\u003cem\u003evar oaxacensis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cem\u003evar subtilis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 14px;\"\u003e\n \u003cp\u003e\u003cem\u003esubsp protamericana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 14px;\"\u003e\n \u003cp\u003e\u003cem\u003eA. asperrima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e\u003cem\u003eA scaposa\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"31\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 12px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eP\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8px;\"\u003e\n \u003cp\u003eI\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eI\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eM\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003eW\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"20\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 12px;\"\u003e\n \u003cp\u003e\u003cem\u003eGROUP\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"20\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003ePlant length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.922 \u0026plusmn; 0.841d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.788\u0026plusmn; 0.566cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.296 \u0026plusmn; 0.584bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.936 \u0026plusmn; 0.215bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.158 \u0026plusmn; 0.163bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1.351\u0026plusmn; 0.613a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.674\u0026plusmn; 0.033ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.951\u0026plusmn; 0.441b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.055\u0026plusmn; 0.615bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.450 \u0026plusmn; 0.550ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.316\u0026plusmn; 0.386bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.875\u0026plusmn; 0.340 abcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eStem length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.620\u0026plusmn; 0.156d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.438\u0026plusmn; 0.082c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.406 \u0026plusmn; 0.069bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.658 \u0026plusmn; 0.079d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.650\u0026plusmn; 0.050d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.236 \u0026plusmn; 0.034a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.386 \u0026plusmn; 0.068bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.394 \u0026plusmn; 0.081bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.396 \u0026plusmn; 0.069bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.275\u0026plusmn; 0.086ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.383 \u0026plusmn; 0.040bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.141 \u0026plusmn; 0.014a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eDiameter***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.615 \u0026plusmn; 0.527g\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.966\u0026plusmn; 0.304def\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.583\u0026plusmn; 0.276cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.560\u0026plusmn; 0. 361fg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.334 \u0026plusmn; 0.189efg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1.725\u0026plusmn; 0.562bcd\u0026plusmn;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.456\u0026plusmn; 0.228efg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.242\u0026plusmn; 0.306ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.485\u0026plusmn; 0.570bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.850\u0026plusmn; 0.173a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.733\u0026plusmn; 0.103cde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.350\u0026plusmn; 0.264efg\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eLeaf length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.778 \u0026plusmn; 0.324f\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.506\u0026plusmn; 0.222ef\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.210\u0026plusmn; 0.170de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.356\u0026plusmn; 0.167de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.524 \u0026plusmn; 0.082ef\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.910\u0026plusmn; 0.202abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.274\u0026plusmn; 0.105cde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.893\u0026plusmn; 0.245ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.030\u0026plusmn; 0.334bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.627\u0026plusmn; 0.126a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.286\u0026plusmn; 0.080de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.10\u0026plusmn; 0.216bcde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eLeaf width***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.237 \u0026plusmn; 0.036\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.254\u0026plusmn; 0.052\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.182\u0026plusmn; 0.039\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.237\u0026plusmn; 0.012\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.276 \u0026plusmn; 0.020\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.092\u0026plusmn; 0.017\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.128\u0026plusmn; 0.004\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.212\u0026plusmn; 0.034\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.193\u0026plusmn; 0.042\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.192\u0026plusmn; 0.017\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.211\u0026plusmn; 0.022\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.285\u0026plusmn; 0.031\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eLL/LW***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e7.561 \u0026plusmn; 1.443de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.025\u0026plusmn; 0.928cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.985\u0026plusmn; 2.028cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e5.701\u0026plusmn; 0\u0026plusmn; .608bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e5.548 \u0026plusmn; 0.541bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e10.109\u0026plusmn; 2.473e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e9.947\u0026plusmn; 0.640e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e4.253\u0026plusmn; 1.177ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e5.310\u0026plusmn; 1.170bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.309\u0026plusmn; 0.904a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.129\u0026plusmn; 0.680cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.889\u0026plusmn; 0.871ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eLL/SL***\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.989 \u0026plusmn; 0.761abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.475\u0026plusmn; 0.447cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.033\u0026plusmn; 0.504bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.068\u0026plusmn; 0.172a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.353 \u0026plusmn; 0.192ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e5.853\u0026plusmn; 2.198e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.390\u0026plusmn; 0.701bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.305\u0026plusmn; 0.573a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.576\u0026plusmn; 0.560ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e2.352\u0026plusmn; 0.422ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.387\u0026plusmn; 0.397bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e28.108\u0026plusmn; 43.566de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eNumber of leaf***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e41.78 \u0026plusmn; 12.968bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e46.44\u0026plusmn; 7.55bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e38.846\u0026plusmn; 10.573bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e53.40\u0026plusmn; 6.618cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e79 \u0026plusmn; 1.73e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e62.10\u0026plusmn; 10.64de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e75.80\u0026plusmn; 6.61e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e33.21\u0026plusmn; 8.935ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e39.71\u0026plusmn; 13.09bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e20.25\u0026plusmn; 5.123a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e35.33\u0026plusmn; 5.60abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e34.0\u0026plusmn; 9.933abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eTerminal thorn length*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.029 \u0026plusmn; 0.006bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.056\u0026plusmn; 0.014e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.143\u0026plusmn; 0.207e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.026\u0026plusmn; 0.006ab\u0026plusmn;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.025 \u0026plusmn; 0.002ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.013\u0026plusmn; 0.006a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.027\u0026plusmn; 0.002bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.040\u0026plusmn; 0.006d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.044\u0026plusmn; 0.007de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.040\u0026plusmn; 0.008cde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.045\u0026plusmn; 0.013de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.060\u0026plusmn; 0.008e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eTerminal thorn width***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.005 \u0026plusmn; 0.002abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.014\u0026plusmn; 0.014c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.006\u0026plusmn; 0.003bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.006\u0026plusmn; 0.001bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.009 \u0026plusmn; 0.005bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.003\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.004\u0026plusmn; 0.0008abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.004\u0026plusmn; 0.001ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.005\u0026plusmn; 0.002ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.004\u0026plusmn; 0.0005abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.007\u0026plusmn; 0.002bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.005\u0026plusmn; 0.0005abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eTTL/TTW***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.524 \u0026plusmn; 3.776abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.027\u0026plusmn; 3.093abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e19.961\u0026plusmn; 19.409d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e4.110\u0026plusmn; 1.239ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e3.352 \u0026plusmn; 1.318a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e4.10\u0026plusmn; 1.235ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e5.890\u0026plusmn; 0.812abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e9.622\u0026plusmn; 2.583cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e9.613\u0026plusmn; 3.791cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e9.0\u0026plusmn; 2.0bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e6.916\u0026plusmn; 3.072abcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e11.416\u0026plusmn; 0.957cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eTTL/LL**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.017 \u0026plusmn; 0.006ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.037\u0026plusmn; 0.007d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.113\u0026plusmn; 0.152d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.019\u0026plusmn; 0.004ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.017 \u0026plusmn; 0.002ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.013\u0026plusmn; 0.004a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.021\u0026plusmn; 0.0005bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.047\u0026plusmn; 0.013d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.045\u0026plusmn; 0.011d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.067\u0026plusmn; 0.025d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.035\u0026plusmn; 0.010cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.056\u0026plusmn; 0.015d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 12px;\"\u003e\n \u003cp\u003eTeeth length***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.002c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.012\u0026plusmn; 0.002cde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.012\u0026plusmn; 0.001de\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.015\u0026plusmn; 0.001e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.004 \u0026plusmn; 0.0006ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8px;\"\u003e\n \u003cp\u003e0.004\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.002bcd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.011\u0026plusmn; 0.003cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.010\u0026plusmn; 0.003cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.012\u0026plusmn; 0.002cde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.009\u0026plusmn; \u0026nbsp; 0.001cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.010\u0026plusmn; 0.001cde\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"37\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eLTEE/LL***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.549\u0026plusmn; 0.523ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.005\u0026plusmn; 0.002a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.006\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.318\u0026plusmn; 0.082cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.606 \u0026plusmn; 0.101bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1.444\u0026plusmn; 0.287d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.710\u0026plusmn; 0.199d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.009\u0026plusmn; 0.003ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.007\u0026plusmn; 0.003a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.013\u0026plusmn; 0.002ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.005\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e1.871\u0026plusmn; 0.750d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eTeeth width***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.007\u0026plusmn; 0.001c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.003cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.007\u0026plusmn; 0.002bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.011\u0026plusmn; 0.0006d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.006 \u0026plusmn; 0.001bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.002\u0026plusmn; 0.0005a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.004\u0026plusmn; 0.001ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.002c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.007\u0026plusmn; 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0.083ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.591\u0026plusmn; 0.193ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eWTEE/LL***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.005\u0026plusmn; 0.001ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.002a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.010\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.001cd\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.004 \u0026plusmn; 0.0005bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.003\u0026plusmn; 0.0008d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.003\u0026plusmn; 0.0005d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.013\u0026plusmn; 0.005ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.0008\u0026plusmn; 0.002a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.020\u0026plusmn; 0.006ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.007\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.006\u0026plusmn; 0.003d\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 12px;\"\u003e\n \u003cp\u003eDistance between teeth**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.018\u0026plusmn; 0.008abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.015\u0026plusmn; 0.007ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.024\u0026plusmn; 0.011bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.024\u0026plusmn; 0.005bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.013 \u0026plusmn; 0.003ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0.018\u0026plusmn; 0.005abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.029\u0026plusmn; 0.005c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.021\u0026plusmn; 0.009bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.022\u0026plusmn; 0.007bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.017\u0026plusmn; 0.006abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.019\u0026plusmn; 0.009abc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 7px;\"\u003e\n \u003cp\u003e0.009\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd height=\"30\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 12px;\"\u003e\n \u003cp\u003eDTEE/LL ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.011\u0026plusmn; 0.007a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.010\u0026plusmn; 0.004a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.019\u0026plusmn; 0.008b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.018\u0026plusmn; 0.006ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008 \u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8px;\"\u003e\n \u003cp\u003e0.020\u0026plusmn; 0.006b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.023\u0026plusmn; 0.005b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.022\u0026plusmn; 0.005b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.024\u0026plusmn; 0.012b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.029\u0026plusmn; 0.012b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.015\u0026plusmn; 0.006ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 7px;\"\u003e\n \u003cp\u003e0.008\u0026plusmn; 0.001a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd height=\"38\" style=\"width: 0px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eDifferent letters indicate significant differences. Significant level 0.001***, 0.01** and 0.0\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-ethnobiology-and-ethnomedicine","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jeet","sideBox":"Learn more about [Journal of Ethnobiology and Ethnomedicine](http://ethnobiomed.biomedcentral.com/)","snPcode":"13002","submissionUrl":"https://submission.nature.com/new-submission/13002/3","title":"Journal of Ethnobiology and Ethnomedicine","twitterHandle":"@ethnobiomed","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Agave management, domestication, maguey, mescal, Mesoamerica, pulque","lastPublishedDoi":"10.21203/rs.3.rs-9078121/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9078121/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground\u003c/h2\u003e \u003cp\u003eDomestication is an evolutionary process guided by humans. It operates continuously and dynamically in multiple directions, with species undergoing adaption to diverse human-influenced environments and cultural and technological contexts. Mexico and the neighbouring Mesoamerican area are among the world\u0026rsquo;s main regions of domestication. The \u003cem\u003eAgave\u003c/em\u003e genus is widely used and managed, with at least eleven domesticated and semi-domesticated species. One species, \u003cem\u003eAgave americana\u003c/em\u003e, comprises two subspecies and four varieties, the diversification of which is hypothetically related to human management. This study explores this hypothesis.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eEthnobotanical studies were conducted on the use, management, and targets of human selection, as well as the patterns of morphological variation, in different ecological and cultural contexts. Semi-structured interviews were conducted with people managing these \u003cem\u003eAgave\u003c/em\u003es across their distribution range, from northern to southern Mexico. Morphometric studies were also conducted on populations distributed in contrasting social and ecological contexts to explore phenotypic variation and trends in relation to selection targets. Multivariate and univariate statistical analyses were performed to analyse patterns of variation in relation to environmental conditions, geographical distribution and forms of use and management.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003e \u003cem\u003eA. americana\u003c/em\u003e is used to extract sap for preparing the fermented beverage called pulque. Its stems have been used as food since prehistoric times, when they were cooked in underground ovens. This cooked matter forms the basis of current fermentation and the production of distilled mescal. The fibre of some varieties has been used to make cords and textiles. Both subspecies are used and managed, but the subspecies \u003cem\u003eprotamericana\u003c/em\u003e clearly has wild populations. Morphometric studies confirm the presence of traits indicative of domestication syndrome in most varieties of the subspecies \u003cem\u003eamericana\u003c/em\u003e.\u003c/p\u003e\u003ch2\u003eConclusions\u003c/h2\u003e \u003cp\u003ePhenotypic variation is clearly related to the taxonomic status and geographic distribution area of the analysed plants, but the influence of human selection favouring gigantism and reduced spines on the leaves is also evident. Further studies of population genetics and phylogeographic patterns would improve our understanding of these patterns.\u003c/p\u003e","manuscriptTitle":"Use, management and domestication of Agave americana in Mexico","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-19 12:21:51","doi":"10.21203/rs.3.rs-9078121/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"editorInvitedReview","content":"","date":"2026-05-04T14:56:36+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"14346040038671937453507752130198157549","date":"2026-04-23T15:31:37+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-08T16:19:26+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"266224142175631006367448501694733571463","date":"2026-03-19T16:27:42+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"170838595380363963642326896394307615457","date":"2026-03-18T13:11:33+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"204750945727466326708472855422863732561","date":"2026-03-16T07:43:09+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-16T07:35:38+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-03-16T05:27:16+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-03-16T05:26:33+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Ethnobiology and Ethnomedicine","date":"2026-03-10T02:17:46+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-ethnobiology-and-ethnomedicine","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jeet","sideBox":"Learn more about [Journal of Ethnobiology and Ethnomedicine](http://ethnobiomed.biomedcentral.com/)","snPcode":"13002","submissionUrl":"https://submission.nature.com/new-submission/13002/3","title":"Journal of Ethnobiology and Ethnomedicine","twitterHandle":"@ethnobiomed","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"d0e48931-5c15-40e1-86f7-a1f9b75ed6e7","owner":[],"postedDate":"March 19th, 2026","published":true,"recentEditorialEvents":[{"type":"editorInvitedReview","content":"","date":"2026-05-04T14:56:36+00:00","index":28,"fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-19T12:21:52+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-19 12:21:51","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9078121","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9078121","identity":"rs-9078121","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}