Use, management and domestication of Agave americana in Mexico

Nadia Campos-Salas; Alejandro Casas; Cuauhtémoc Jacques-Hernández

doi:10.1186/s13002-026-00924-2

. 2026 Jul 3;22:92. doi: 10.1186/s13002-026-00924-2

Use, management and domestication of Agave americana in Mexico

Nadia Campos-Salas ^1,^2,^✉, Alejandro Casas ^2,^✉, Cuauhtémoc Jacques-Hernández ³

PMCID: PMC13625399 PMID: 42399974

Abstract

Background

Domestication is an evolutionary process guided by humans. It operates continuously in multiple directions, with species undergoing adaption to diverse human-influenced environments and cultural and technological contexts. Mesoamerica is among the world’s main regions of domestication and the Agave genus is widely used and managed in this region, 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 by analysing morphological variation and its relation to environmental and management settings.

Methods

Ethnobotanical studies were conducted to document uses, management practices, and targets of human selection, together with patterns of morphological variation across different ecological and cultural contexts. Semi-structured interviews were carried out with agave managers throughout the species’ distribution range in Mexico. Morphometric analyses were performed on populations occurring under contrasting environmental and management conditions. Multivariate and univariate statistical analyses were used to evaluate morphological variation in relation to geographic distribution, environmental conditions, and management regimes.

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 managed, but the subspecies protamericana clearly has wild populations. Morphometric studies confirm the presence of traits indicative of domestication in most varieties of the subspecies americana.

Conclusions

Phenotypic variation within the Agave americana complex is associated with taxonomic identity, geographic distribution, and management intensity. The observed patterns are consistent with the effects of human selection, particularly in traits related to plant size and the reduction of defensive structures. However, further studies integrating common-garden experiments, population genetics, and phylogeographic analyses are required to clarify the evolutionary history and genetic basis of the observed variation.

Keywords: Agave management, domestication, maguey, mescal, Mesoamerica, pulque

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. Domesticated organisms have morphological and physiological attributes that are markedly (though not exclusively) aligned with human requirements [5, 6]. Although selection is the most studied evolutionary force in domestication, other forces—mutation, gene flow, genetic drift—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.) 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.

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 harbours one of the world’s richest reservoirs of agrobiodiversity [16, 21–23]. Documenting how this diversity originated and identifying the factors that maintain it remain key priorities for its conservation [24].

Indigenous peoples in Mesoamerica use nearly 6,500 native plant species [13, 25], 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. Other species, 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, but this time period is too short to identify signs of domestication [26]. The cultivation and management of most of the domesticated species, and the associated selective mechanisms, have been well documented [14, 27–32]. High levels of interspecific hybridization and genetic variation have also been reported [29–35].

Phenotypic variance results from the genotypic and environmental variances [36]. Therefore, researchers commonly assess domestication by examining divergence between wild and managed phenotypes [37, 38]. In Agave, phenotypes of managed populations reflect use type, often involving enlargement of useful parts—stems, leaves, fibres, or sap—and reduction of defensive or undesirable features such as spines, teeth, saponins, and other secondary metabolites [14, 28–30].

The interaction between natural and human selection makes Agave an excellent model for studying domestication. Mexico, with the greatest diversity and distribution of Agaves [39], offers ideal conditions for this research. Humans have managed and domesticated these plants since prehistoric times [10, 14, 22, 26], .

The case of Agave americana is of particular interest, given the extraordinary degree of diversification it has undergone during domestication. According to Gentry [22], 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. Examining these patterns helps reconstruct evolutionary pathways [9, 40].

Currently, Agave americana subsp. protamericana is the only taxon within the Agave americana complex for which clearly wild populations have been documented, although some populations may also be subject to low-intensity management. This taxon therefore provides an important reference for evaluating morphological divergence associated with domestication processes.

Human selection is expected to drive morphological and genetic differentiation among managed populations. However, a persistent challenge in evolutionary studies is determining whether observed phenotypic differences reflect genetic divergence, environmental influences, or the interaction of both factors. Because domestication is generally a gradual process, domesticated forms often retain traits that reveal relationships with their wild ancestors while exhibiting intermediate morphologies associated with different levels of management intensity.

We hypothesize that morphological variation within the Agave americana complex is associated with historical patterns of use and management. Because hybridization among Agave taxa is common, phenotypic variation is likely influenced by multiple interacting evolutionary processes, including selection, gene flow, and local environmental conditions. Consequently, the patterns documented in this study should be interpreted as a first approximation that will require complementary analyses based on population genetics, reproductive biology, and common-garden experiments.

This study aims to:1) document ethnobotanical and morphological information in order to analyse the Agave americana complex, its use and management, and the 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. We expected to observe morphological differentiation among the recognized varieties of Agave americana and trends consistent with domestication-related divergence relative to A. americana subsp. protamericana.

Materials and methods

Study species

The Agave americana complex comprises the following two subspecies and four varieties:

Agave americana subsp. protamericana, (Fig. 1G.) 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 [22] 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. 1A.) 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 [39] and Hernández-Sandoval et al. [41] reported putative wild populations near Ciudad Victoria, Tamaulipas. It is widespread in Mexico [42–44], though its wild status needs confirmation.

Agave americana subsp. americana var. expansa. (Fig. 1B.) 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 [22]. It has been reported in Chihuahua [45] and Jalisco [44].

Agave americana subsp. americana var. oaxacensis. (Fig. 1C.) It occurs in Oaxaca, where is commonly known as “maguey arroqueño”, “maguey coyote”, “maguey de Castilla”, or “maguey Sierra Negra” [23]. It has large, spreading, glaucous to whitish leaves with small teeth, producing large flowers and thick-walled capsules [22]. Because its stems accumulate high carbohydrate levels, it is used for high-quality mescal. Plants require 18–20 years to mature [44]. Historically, it was also used for pulque and fiber extraction [39, 46].

Agave americana subsp. americana var. subtilis (Fig. 1H.) 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. [47] 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. 1D.), closely related to var. expansa, this species was first described by Trelease (1912) and later regarded by Gentry [22] as part of var. expansa. It differs mainly in rosette size, leaf shape, and terminal spine.

Agave scaposa (Fig. 1F.), 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 [22]. According to García-Mendoza [48], A. scaposa is used for pulque production. Casas et al. [49] and Blancas et al. [50] reported its use in soil stabilization, fencing, and as a source of durable inflorescence stalks (or “quiote”) for construction.

Agave asperrima subsp. Asperrima (Fig. 1E.), the most widespread Agave in northern Mexico [22], 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 [51–53].

Study area

The field study sites for the analyzed taxa of the Agave americana complex are shown in Fig. 1 and Table 1.

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

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The study was conducted in three main regions of Mexico Fig. 2:

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.

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. Informed consent was obtained from the local authorities and the people interviewed prior to all interviews. In addition, some interviews were conducted in response to direct invitation from the agave managers. Information gathered during interviews was complemented with direct field observations, previously collected ethnobotanical records, and published literature. Direct observations were also conducted on wild populations of Agave asperrima, Agave scaposa and Agave americana subsp. protamericana and subsp. americana.

Ethnobotanical information was analyzed qualitatively to identify patterns of management intensity, domestication-related traits, and selection practices. Management systems were categorized as wild, managed in situ, promoted, cultivated, or intensively cultivated. These categories were subsequently compared with patterns of morphological variation observed in the studied 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, 28, 30, 54]. 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.

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

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Preliminary analyses indicated that reproductive characters did not contribute significantly to the discrimination among taxa. Consequently, subsequent analyses focused primarily on vegetative characters, which have been shown to be particularly informative in studies of domestication-related morphological variation in Agave [14, 28, 30, 54].

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) [55].

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 [56].

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.

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 [56]	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) [57,58]	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 [59] 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 [59] 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

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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.

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 flavour. 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, with a 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 Agaves 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 traits, 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.

Fig. 4 — Linear 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

Grouping by management category also revealed separation among wild, cultivated, and intensively managed populations. Cultivated populations formed a relatively compact cluster, whereas wild populations displayed greater dispersion within the discriminant space. 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.

Taxonomic groups showed varying degrees of separation. Some taxa formed distinct clusters, while others overlapped substantially. These results indicate that morphology reflects both taxonomic identity and management history, although the degree of differentiation varies among taxa.

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. Agaves 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 (Fig. 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).

Table 4.

Results of the multivariate analysis of variance (MANOVA) for the variables Area, Management, and Taxonomy

FUNCTION		WILKS´LAMBDA	CHI SQUARE	DF	P VALUE
AREA	H	3.5942e-05	86.98	32	0.09173
AREA	F	0.00101	615.38	76	2.2e-16
MANAGEMENT	H	1.8139e-05	92.79	32	0.06601
MANAGEMENT	F	0.0055	487.24	76	2.2e-16
TAXONOMY	H	---------------	-----------------	64	------------
TAXONOMY	F	0.00278	734.63	133	2.2e-16

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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

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%).

Table 5.

Characters evaluated in herbarium specimens and in the field and eigenvectors of the first and second principal components and discriminant functions

Character	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

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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 6).

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 ± 0.012 cd	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 ± 0.001 cd	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

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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.

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
Character	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 ± 0.566 cd	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 ± 0.527 g	1.966 ± 0.304def	1.583 ± 0.276 cd	2.560 ± 0. 361 fg	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 ± 0.928 cd	6.985 ± 2.028 cd	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 ± 0.680 cd	3.889 ± 0.871ab
LL/SL***	2.989 ± 0.761abc	3.475 ± 0.447 cd	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 ± 6.618 cd	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 ± 2.583 cd	9.613 ± 3.791 cd	9.0 ± 2.0bcd	6.916 ± 3.072abcd	11.416 ± 0.957 cd
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 ± 0.010 cd	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 ± 0.003 cd	0.010 ± 0.003 cd	0.012 ± 0.002cde	0.009± 0.001 cd	0.010 ± 0.001cde
LTEE/LL***	0.549 ± 0.523ab	0.005 ± 0.002a	0.006 ± 0.001a	1.318 ± 0.082 cd	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 ± 0.003 cd	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 ± 0.0005 cd	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 ± 0.001 cd	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

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Different letters indicate significant differences. Significant level 0.001***, 0.01** and 0.0

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 A. a. 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. a. var. 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 A. a. 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 A. a. 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 results of this study indicate that morphological variation within the Agave americana complex is associated with taxonomic identity, geographic distribution, and management intensity. The combination of ethnobotanical information and morphometric analyses revealed patterns that are consistent with the influence of human management on phenotypic variation. However, the observed variation is also likely influenced by environmental factors, historical population processes, and gene flow among taxa.

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 [60, 61].

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. a. 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.

Several morphological traits observed in cultivated and intensively managed populations are consistent with patterns previously documented in domesticated Agave species [62], 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 [60, 62]. This reproductive diversity supports long-term adaptability and evolutionary potential within the A. americana complex [61, 63].

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 [64, 65]. 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 [64, 65].

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 [66]. 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 [60, 61].

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 [67, 68].

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 [69, 70].

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 visualized among wild, cultivated, and promoted forms. In particular, traits such as plant size, leaf characteristics, and the size of terminal spines and teeth appear as 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 opposite 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 [64, 65], 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 (Tables 4 and 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 an overview of how, geographic conditions, and management regimes have shaped the vegetative, defensive and reproductive traits of these species. This knowledge offers new perspectives on how these plants adapt to selective pressures and how conservation efforts can support their survival under changing conditions [68, 71].

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 common garden experiments and next-generation sequencing techniques, 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 [72, 73] .

Conclusion

The Agave americana complex exhibits substantial morphological diversity associated with taxonomic identity, geographic distribution, and management intensity. Ethnobotanical and morphometric evidence indicates that human management has likely contributed to the differentiation of several cultivated forms, particularly those maintained under intensive production systems.

Although the observed patterns are consistent with domestication-related processes, morphology alone is insufficient to fully reconstruct evolutionary relationships or domestication pathways. Future studies integrating genomic, ecological, and experimental approaches will be necessary to clarify the origins and diversification of the Agave americana complex and to better understand the role of human management in shaping its diversity.

The identified patterns are not conclusive; further details of management practices are required. 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 Agaves. 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.

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.

Acknowledgements

The authors thank the grant provided to the first authors for doctoral studies by the National Council of Science and Technology (CONACYT, Mexico) and the postgraduate programme Posgrado en Ciencias Biológicas at the Instituto de Investigaciones en Ecosistemas y Sustentabilidad, UNAM, campus Morelia. Also, we thank financial support provided by CONACYT (research project A1-S-14306), the programme PAPIIT of the General Direction for Academic Support (DGAPA, UNAM, Mexico, research project IN224023), and the National Commission for the Knowledge and Use of Biodiversity (CONABIO) and FAO-GEF project Agrobiodiversity of Mexico (ID 9380) for the support to the project RG023.

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.C.J.H. participated in fieldwork particularly in northern Mexico, contributed to the acquisition and validation of morphological and ethnobotanical data, assisted in the interpretation of the results, and critically reviewed the manuscript.All authors contributed to the development of the final version of the manuscript, discussed the results and their implications, critically reviewed the content, approved the final manuscript for submission, and agree to be accountable for all aspects of the work.

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.

Data availability

The datasets generated and analyzed during the current study are available from the corresponding author upon reasonable request.

Declarations

Ethics approval and consent to participate

Information consent was obtained with local authorities and all people interviewed. The authors collaborate with a Mexican network of forest agave management; thus, some interviews were carried out by request of the agave managers.

Consent for publication

Information related to agave management was authorized for publication. Other data do not require specific consent.

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Contributor Information

Nadia Campos-Salas, Email: nadiacs@cieco.unam.mx.

Alejandro Casas, Email: acasas@cieco.unam.mx.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

The datasets generated and analyzed during the current study are available from the corresponding author upon reasonable request.

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PERMALINK

Use, management and domestication of Agave americana in Mexico

Nadia Campos-Salas

Alejandro Casas

Cuauhtémoc Jacques-Hernández

Abstract

Background

Methods

Results

Conclusions

Background

Materials and methods

Study species

Fig. 1.

Related taxa considered

Study area

Table 1.

Fig. 2.

Ethnobotanical studies

Studies of morphological variation

Table 2.

Statistical analyses of morphological characters

Results

Ethnobotanical uses

Table 3.

Management practices

Morphological patterns

Reproductive and vegetative characters analysed in herbarium specimens

Fig. 3.

Linear Discriminant Analysis (LDA)

Fig. 4.

Fig. 5.

Table 4.

Vegetative characters measured in the field

Fig. 6.

Table 5.

Fig. 7.

Fig. 8.

Table 6.

Morphological variation among taxa of the Agave americana complex

Table 7.

Discussion

Recommendations for future research

Conclusion

Acknowledgements

Author contributions

Funding

Data availability

Declarations

Ethics approval and consent to participate

Consent for publication

Competing interests

Footnotes

Contributor Information

References

Associated Data

Data Availability Statement

ACTIONS

PERMALINK

RESOURCES

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