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Photograph of Mammillaria pectinifera with fi ve whorls of fruits of different ages showing the possible fates of fl oral buds. (a) Flowers; (b) expelled fruit from the most recent whorl (only the gap between areoles is seen in the photo); (c) retained fruits from the previous whorl (over two years of age); (d) third whorl of fruits, three or more years of age, with eroded tips and most of the seeds dispersed; (e) the dry remains of a whorl of old empty fruits, clearly seen at the base of the photo but not drawn in the diagram. In this species, expelled fruits always belong to the most recent cohort. 

Photograph of Mammillaria pectinifera with fi ve whorls of fruits of different ages showing the possible fates of fl oral buds. (a) Flowers; (b) expelled fruit from the most recent whorl (only the gap between areoles is seen in the photo); (c) retained fruits from the previous whorl (over two years of age); (d) third whorl of fruits, three or more years of age, with eroded tips and most of the seeds dispersed; (e) the dry remains of a whorl of old empty fruits, clearly seen at the base of the photo but not drawn in the diagram. In this species, expelled fruits always belong to the most recent cohort. 

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Article
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Serotiny, or delayed seed dispersal, is common in fluctuating environments because it hedges the risks of establishment. Mammillaria pectinifera (Cactaceae) facultatively expels fruits in the year they are produced or retains them to disperse the seed over several years. We tested whether M. pectinifera increased fruit expulsion as a response to in...

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Context 1
... areoles possess white appressed spines that cover and shade the plant. The fl owers open in circular crowns around the apex. The fruit is a whitish berry that may be completely expelled or may remain inserted inside the stem of the plant, gradu- ally releasing seeds for 7 -8 years ( Boke, 1960 ; Bravo-Hollis and S á nchez- Mejorada, 1991 ) (Fig. 1). The species is endemic to the Tehuac á n Valley, a small semiarid area in the southeast of the State of Puebla, Mexico ( Bravo-Hollis and S á nchez-Mejorada, 1991 ), and is confi ned to deep alkaline soils with relatively high surface stoniness and high water retention capacity, receiving ca. 400 mm of annual precipitation ( ...
Context 2
... release from embedded fruits and viability -Twenty plants were col- lected near the study site, and the seeds that were retained inside 615 fruits of known age were extracted and counted. Age was easily estimated because an- nual rings of fruits are readily distinguishable in the stem (see Fig. 1 ). Curves were fi tted through a log-linear regression minimizing a χ 2 error function ( Crawley, 1993 ) to predict the number of seeds that remain in the fruits at dif- ferent ages. The seeds extracted from the fruits were placed in Petri dishes with agar gel and incubated at 25 ° C and a 12-h photoperiod to estimate viability loss in ...

Citations

... An alternative to soil seed banks is serotiny, which is an aerial seed bank (Lamont and Enright, 2000), and which has the additional advantage of protecting seeds in structures less accessible to soil-dwelling animals or that may reduce the attack of predators (Rodríguez-Ortega et al., 2006;Lamont et al., 2016;Lamont et al., 2020). Serotiny is considered a delayed dispersal mechanism due to the retention of mature seeds in mother plant structures for more than 1 year (Lamont, 1991;Peters et al., 2009;Lamont et al., 2020). This characteristic has been observed in approximately 1345 species of different fire-prone ecosystems from different continents (Lamont et al., 2020), including dry, Mediterranean and temperate ecosystems. ...
... Despite serotiny mostly occurring in fire-prone ecosystems (Lamont et al., 2020), it is also found in Cactaceae species inhabiting North American deserts such as the Mojave and Sonora (Martínez-Berdeja et al., 2015). Approximately 24 cactus species with mature seed retention have been reported so far (Barrios et al., 2020), although according to Peters et al. (2009), the work of Bravo-Hollis and Sánchez-Mejorada (1991) notes 25 species where some degree of seed retention is shown. Seed retention in cacti has been observed in tubercles axils in several Mammillaria species (Rodríguez-Ortega and Franco, 2001;Rodríguez-Ortega et al., 2006;Peters et al., 2009), in chained fruits in Cylindropuntia (Martínez-Berdeja et al., 2015), in the apex of stems, dry fruits and cephalia of several genera including Echinocactus (Chamberland, 1997), Melocactus (Bravo-Hollis andSánchez-Mejorada, 1991), Geohintonia, Pelecyphora, Ariocarpus (Anderson et al., 1994) and Sclerocactus (Earle, 1980). ...
... Approximately 24 cactus species with mature seed retention have been reported so far (Barrios et al., 2020), although according to Peters et al. (2009), the work of Bravo-Hollis and Sánchez-Mejorada (1991) notes 25 species where some degree of seed retention is shown. Seed retention in cacti has been observed in tubercles axils in several Mammillaria species (Rodríguez-Ortega and Franco, 2001;Rodríguez-Ortega et al., 2006;Peters et al., 2009), in chained fruits in Cylindropuntia (Martínez-Berdeja et al., 2015), in the apex of stems, dry fruits and cephalia of several genera including Echinocactus (Chamberland, 1997), Melocactus (Bravo-Hollis andSánchez-Mejorada, 1991), Geohintonia, Pelecyphora, Ariocarpus (Anderson et al., 1994) and Sclerocactus (Earle, 1980). However, to our knowledge there is little prior work on the role of seed retention in population dynamics of the cactus species where it has been observed (Barrios et al., 2020). ...
Article
Serotiny is a strategy in which the retention of mature seeds in parent structures allows plants to cope with environmental variability like heat, drought or fire. Although this phenomenon might be common in Cactaceae, and particularly in Melocactus, it has generally been scarcely addressed. The main goal of our work is to investigate if there are seeds hidden in the cepha-lium of Melocactus matanzanus and if there are, determine whether or not these seeds maintain their viability. We also discuss some advantages the cephalium may offer as diaspore after the death of individuals. Cephalia collected from dead individuals were divided into four slices and their seeds counted; we also assessed the viability and photoblastic response of the seeds by using growth chambers at 25/30°C, and by a cut test on the seeds that did not germinate. Our results showed retention of viable seeds of different ages in all slices of the cephalium. Seeds were photoblastic positive with germination between 11-22% and viability above 50% in the portion of the lots that did not germinate.
... Generally, the retention of seeds on the mother plant is associated with seed releases (dispersal) triggered by environmental factors such as fire, drought or wetting (Lamont, 1991). The duration of mature seeds retained on the mother plant varies with the species and ecosystems according to Cowling and Lamont (1987), and Bastida and Talavera (2002) in the Mediterranean area, Ma and Liu (2008) in Asia, and Peters et al. (2009) in America. ...
Article
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The ability of plants to safely retain seeds in the mother plant is an adaptive mechanism described in many desert plants. However, research about delayed seed dispersal species in the desert of the United Arab Emirates (UAE) is lacking. This study aims to identify these delayed seed dispersal species and assess the relationships of the presence of delayed seed dispersal with plant growth form, habit, spatial dispersal, antitelechoric mechanism, and seed release time. The relationships between the presence of delayed seed dispersal and the above studied traits were assessed by using the Pearson Chi-square test and Nonlinear Principal Components Analysis (NLPCA). Results showed that a total of 46 delayed seed dispersal species were recorded (15.0% of 307 studied species) and the highest incidence occurred in the Fabaceae family (17.4%). Delayed seed dispersal species were predominantly perennial plants (73.9%) with spatial restricted dispersal (67.4%), which released seed in the dry season (45.7%). The dominant groups of delayed seed dispersal species were persistent fruits species and synaptospermy (28.3%). All graminoids showed persistent lignified fruits, while prostrate annuals were basicarpic species with myxospermy. Sandy habitats had the highest number of delayed seed dispersal species (54.3%), whereas salt flats had the lowest (23.9%). In the desert of the UAE, delayed seed dispersal species spread seeds until the end of the dry and windy season, thus breaking seed dormancy at this time and ensuring seed germination in the next arrival of the rainy season. This morphological and ecological adaptation of delayed dispersal species is essential to the survival and sustainable development of vegetation in desert environments.
... Particularly in desert areas, the serotiny allows the species for: a. Synchronizing seed dispersal with moisture availability (Gutterman & Ginot, 1994;Hegazy et al., 2006) b. Regulating the frequency of seed dispersal and synchronize it with rainfall episode (Friedman et al., 1978;Ellner & Shmida, 1981;Peters et al., 2009). c. ...
... Santini and Martorell (2013) hypothesized that in dry areas, serotiny may be driven by different factor, namely seed priming within the mother plant which allow water to seep in seeds, thus, seeds experience repeated hydration and dehydration cycles throughout the rainy period. This was recorded for some species belonging to Agavaceae, Asteraceae, Cactaceae, Mesembryanthemaceae, Mimosaceae, Poaceae, and Ruscaceae (Rodríguez-Ortega et al., 2006;Peters et al., 2009Peters et al., , 2011 families. In consequence, the non-dormant seeds may start germinating when they imbibe (initial phase of germination, water uptake, or phase I), but they do not complete germination before they become dried up again. ...
... While whole-fruit release is triggered by a form of predictive cueing during favorable conditions, fruit retention followed by gradual seed release may act as a bethedging tool in a taxonomic group with exceptionally high seedling mortality (Godínez-Álvarez et al., 2003). Peters et al. (2009) worked with Mammillaria pectinifera, a rare and threatened plant of semi-arid area in the southeast of Mexico and reported two modes of seed release: (a) entire fruits may be dispersed with their complete seed content instantly after maturation, while (2) other fruits are retained in canopy, and they slowly release seeds via an apical aperture as they mature. The species may adapt one or other path in response of environmental variability, since the probability of their establishment may increase during wet period. ...
Chapter
Trait based studies can enhance our perceptive about the plant evolution, community assembly and ecosystem functioning. Serotiny (retention of aerial ripe seed in closed fruits or cones until an endogenous or exogenous trigger support dehiscence) is an important adaptive trait in fire-prone, and in desert ecosystems. This tactic has several advantages, and is exhibited by the angiosperm as well as by gymnosperm species. In this communication, I have discussed various features of serotiny habit, with light of current knowledge. An updated list of 113 serotinous plant species across the globe was prepared that consisted information on species and family names, habit and geographical locations of the species. Serotiny properties of many of these species were also discussed in detail. Other relevant information like hypotheses for serotiny, structural and external factors associated with this trait, aerial and soil seed bank properties, seed cone types, and degrees of their longevity and physiological aspects (maintenance costs) were also critically evaluated. Additionally, experimental works conducted on various factors like fire with relation to serotiny, their magnitude in desert ecosystems, as well as serotiny habit in cactus species were also discussed. Present effort revealed that, across the globe, most of the empirical studies on serotiny are focused on certain specific species, belonging to only a few genera such as Artemisia, Banksia, Callitris, Hakea, Hesperocyparis, Leucadendron, Mammillaria, Petrophile, Pinus, and Protea. However, this adaptive trait is still not fully explored for the Indian arid zone species. Study of serotiny habit for the Indian arid species would facilitate us to understand the factors (specifically related with site) associated with degradation of their population size
... Serotiny represents an alternative to the traditional seed bank and in many cases is known as an aerial form of seed bank, with the advantage of protecting the seeds in structures that are inaccessible to predators (Rodríguez-Ortega et al. 2006). Serotiny is known as a delayed dispersion mechanism because of the retention of mature seeds in structures of the mother plant for more than a year (Peters et al. 2009). In the Cactaceae family, serotiny probably is a common phenomenon in species that inhabit several North American deserts like the Mojave and the Sonora (Martínez-Berdeja et al. 2015). ...
... In the Cactaceae family, serotiny probably is a common phenomenon in species that inhabit several North American deserts like the Mojave and the Sonora (Martínez-Berdeja et al. 2015). So far 24 species of cacti with a retention of mature seeds have been listed (Table 1), although around 25 species showed some degree of retention in the work of Bravo-Hollis & Sánchez-Mejorada (1991) according to Peters et al. (2009) but the role that serotiny plays in the population dynamic of the species where it occurs has not been extensively studied. ...
... Seed retention in cacti has been observed in the axils of tubers (Rodríguez-Ortega et al. 2006, Peters et al. 2009), in chained fruits (Martínez-Berdeja et al. 2015), in the apex of the stems and in the cephalia (Bravo-Hollis & Sanchez-Mejorana 1991). The duration of the seeds in the stems has mainly been studied in the Mammillaria genus, where it has been demonstrated that the seeds can be retained from one (Santini & Martorell 2013) to eight years, forming an aerial seed bank (Boke 1960, Peters et al. 2009, Rodríguez-Ortega et al. 2006. ...
Article
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Background: Cactaceae is the fifth taxonomic group with the highest proportion of threatened species. One way to contribute to the preservation of this family is to understand the processes that promote seed germination. Questions: How common is dormancy and seed banks in Cactaceae? Are there general patterns in cacti germination response to temperature, light, water, salinity, phytohormones, hydration/dehydration cycles, mechanical or chemical scarification? Data description: A total of 333 studies on cactus germination with information on 409 taxa. Study site and dates: since 1939 to January 2020. Methods: A search of scientific articles in Google Scholar was performed with the words Cactaceae, cacti and cactus, in combination with various matters on germination in English, Spanish and Portuguese. Results: The main germination studies in cactus deal with photoblasticism (275 taxa), temperature (205 taxa) and seed longevity (142 taxa). Other lines of study in cactus germination (e.g., desiccation tolerance, vivipary, phytohormones, mechanical or chemical scarification, in vitro germination, hydration/dehydration cycles, water and saline stress, serotiny, storage in cold, high temperature tolerance and soil seed bank) include between 14 and 65 taxa. Cacti have only physiological dormancy and optimal germination for most species occur between 20 and 30 °C. Conclusions: Mexico, Brazil and Argentina are the three leading countries in the study of cactus germination.
... Delayed seed release can be adaptive by (1) spreading seed dispersal in time (Venable and Lawlor, 1980), (2) retaining seeds in a favourable microhabitat (Gutterman, 1994), (3) controlling the time of seed germination (Lamont, 1991), (4) changing the spatiotemporal pattern of soil seed banks (Günster, 1994) and (5) decreasing the opportunity for seed predation and death (Günster, 1994). The number of mature seeds retained on the mother plant and length of time vary with the species and ecosystem (Cowling and Lamont, 1987;Bastida and Talavera, 2002;Ma and Liu, 2008;Peters et al., 2009). Furthermore, the pattern of seed release and distribution away from mother plants, i.e. seed shadow, is an important part of plant fitness (Wenny, 2000). ...
Article
The pattern of seed dispersal in time and space can affect plant fitness and the soil seed bank, and thus information is needed on this aspect of the seed biology of a species before it is selected for use in habitat restoration projects. Zygophyllum xanthoxylon is a super-xerophilous shrub that is a potential pioneer species for use in revegetating highly disturbed areas of the cold deserts of northwest China. We studied fruit release and soil seed banks of Z. xanthoxylon for 3 years in two cold desert habitats characterized by different degrees of drought and wind velocity. In our study, fruit (a three-winged capsule) release began in summer (June 2010, August 2011, July 2012) and extended for 9–10 months, but plants can be found in the population with previous- and current-year fruits attached to them. More than 50% of the fruits were released in the first 3–4 months after maturity, while the others were released gradually over a 7–8 month period. The temporal pattern of fruit dispersal varied with habitat but not with amount of precipitation during summer. The pattern of fruit deposition on the soil surface was affected by neighbouring plants, wind velocity, wind direction and topography. In both habitats, >90% of the fruits were deposited beside large and small clusters of plants, mainly Ephedra przewalskii . To facilitate plant community development, we suggest that E. przewalskii should be planted (as a wind break) together with Z. xanthoxylon when native pioneer species are used for restoration of cold desert shrublands.
... This in combination with rain cues that trigger germination, could be advantageous in dry habitats indicating favourable conditions for the establishment. Actually, seed priming in combination with cued-dispersal increases the fitness of individuals of the cactus Mammillaria hernandezii (Santini and Martorell, 2013), and rain also promotes the release of fruits in M. pectinifera (Peters et al., 2009). This could be the case for our species, especially for F. hamatacanthus, in which fruit retention has been observed (MC Mandujano personal communication), and where HD-DH cycles increased the germination rate ( Table 1). ...
Article
In drylands, cacti seeds experience high variability amounts of precipitation throughout a year. Seeds undergo hydration and dehydration cycles (or priming) which can increase the proportion or the speed of germination. However, these advantages differ among cacti. In this study, we test the effect of priming on the germination percentage and the mean germination time of two cacti species. We show that priming does not increase the germination percentage for these cacti species, but does reduce the time to germination. We suggest that the response of cacti seeds could be an adaptation to their habitat and microhabitat conditions.
... Almost 80% of species in the genus Turbinicarpus are specialized to grow in extreme habitats, such as narrow cracks in low, sloping rocks and stones, often under full sun exposure, where the competition from other plants is limited (Donati & Zanovello, 2004). Seed dispersion in Turbinicarpus, which have some dormancy mechanisms (de la Rosa-Carrillo, Dom ınguez-Rosales, P erez-Reyes, & P erez-Molphe-Balch, 2012; Flores, Arredondo, & Jurado, 2005), is largely due to rainfall, as in other small cacti (Peters, Martorell, & Ezcurra, 2009;Rojas-Ar echiga & V azquez-Yanes, 2000), and myrmechory (Hofer, 2011). Consequently, Turbinicarpus populations are often isolated, frequently poor to very poor in individuals, with a high degree of morphological variability (Donati & Zanovello, 2004). ...
Article
The biogeography and the ecology of the genus Turbinicarpus are scarcely known and the taxonomic segregation of the genus Rapicactus from Turbinicarpus has been debated. The present study aims to (i) evaluate the distribution of all taxa of Turbinicarpus in the full distributional range of the genus (NE Mexico); (ii) investigate the potential ecological divergence between Rapicactus and Turbinicarpus; and (iii) analyse the influence of environmental variables on plant morphology. The distribution pattern of all taxa belonging to the genus Turbinicarpus s.l. (Turbinicarpus + Rapicactus) was mapped using information from 12 herbaria, as well as unpublished databases. The study area was divided into cells of 25 km × 25 km. The relationships between taxa richness, sampling effort and environmental variables were tested by using grid data, via spatially explicit models. The elevational distribution of populations and the effects of climate, latitude and elevation on the mean steam diameter of the populations were separately investigated for Turbinicarpus and Rapicactus. Turbinicarpus s.l. taxa richness is highest within 100°W and 101°W longitude, about 2500 km north of the equator, with maximum diversity in areas with a mean annual temperature of 17–19 °C and a mean annual rainfall of 550–650 mm. A significant relationship was observed between taxa richness and sampling effort, and between taxa richness and longitude, mean annual rainfall and calcium-rich substrate. Rapicactus taxa were found to grow at higher elevations compared with Turbinicarpus. Body size of the plants showed a negative relationship with latitude and elevation in Turbinicarpus, and a positive one with latitude in Rapicactus. The results showed that the genus Turbinicarpus s.l. presents diversity hotspots in the floristic regions of Chihuahuan Desert and Sierra Madre Oriental. The responses of Rapicactus and Turbinicarpus to latitudinal and elevational gradients were different, supporting the segregation of Rapicactus from Turbinicarpus.
... Está incluido en el Apéndice I de la CITES (CITES, 2013) y es considerado como una especie amenazada por la Nom-059 (Semarnat, 2010) y por la UICN (2013). Su estudio se ha centrado en la descripción básica de su morfología y taxonomía (Boke, 1960;Bravo-Hollis y Sánchez-Mejorada, 1991;Arias et al., 1997), y más recientemente en aspectos de su biología, ecología y filogeografía (Rodríguez-Ortega y Ezcurra, 2001; Zavala-Hurtado y Martorell y Peters, 2005;Rodríguez-Ortega et al., 2006;Valverde y Zavala-Hurtado, 2006;Peters et al., 2008;Peters et al., 2009;Valverde et al., 2009;Peters et al., 2011;Cornejo-Romero et al., 2014). Se trata de un cacto globoso de tamaño pequeño, simple y semigeofítico de 1-5 cm de diámetro y de 1-2 cm de altura; las areolas tienen espinas radiales blancas, pectinadas y adpresas que cubren totalmente a la planta (Arias et al., 1997). ...
Article
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Mammillaria pectinifera es un cacto amenazado y endémico del valle de Tehuacán. A mediados de la década de 1990 sólo se conocían 6 localidades con un número reducido de individuos, información que fue clave para proteger a la especie con instrumentos legales nacionales e internacionales. Para ampliar el conocimiento de la distribución geográfica y estado de conservación de esta especie, realizamos recorridos exploratorios en zonas ecológicas similares al de las poblaciones conocidas y, posteriormente, mediante un modelo predictivo de distribución geográfica. Los resultados indican que la especie está conformada por al menos 31 localidades y un número mucho mayor de individuos al registrado en estudios previos, lo que sugiere que para evaluar la distribución, abundancia y presión sobre especies poco conspicuas, es necesario dar un fuerte énfasis al trabajo de exploración, utilizando las mejores herramientas y métodos disponibles. No obstante, al incremento sustancial de localidades e individuos, y que el 55% de éstas se encuentran dentro de la Reserva de la Biosfera Tehuacán-Cuicatlán, este cacto presenta un alto grado de presión antrópica, lo que sugiere que la especie es vulnerable a desaparecer en algunas localidades si las tasas de deterioro de su hábitat no disminuyen en el corto plazo.
... In general, the retention of seeds in the plant canopy for one to 30 years or more is termed as serotiny (Lamont et al. 1991). It is common in fluctuating environments such as dry, fire-prone scrubs and forests (Peters et al. 2009). Plants are either non-serotinous, weakly serotinous (most seeds released within a few years) or strongly serotinous (most seeds still retained after a few years). ...
Article
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Canopy seed bank is an important adaptive evolutionary trait that provides various types of protection to the seeds. However, costing of such evolutionary trait on plant survival is largely unknown. Present investigation provided a new insight on the serotonious habit of Blepharis sindica associated with its endangerment status. Extinction probabilities of two available population of B. sindica were quantified using two types of census data, i.e., fruiting body number and actual population size. Population Viability Analysis (PVA) revealed that delayed seed release tendency (higher fruiting body number) was not synchronized with actual ground conditions (lower population size). PVA analysis based on actual population size indicated that both the available populations would vanish within 20 years. The mean time of extinction calculated from both type census data indicated its extinction within 48 years. For assessing the conservation criteria, a glass house experiment was carried out with different soil types and compositions. Pure sand and higher proportions of sand -silt were more suitable compared to clay; further, gravelly surface was the most unsuitable habitat for this species. Collection of the seeds from mature fruits/capsule and their sowing with moderate moisture availability with sandy soil could be recommended.
... Populations are distributed in the northern sector of the TVr, and separated by 3.66 to 62.68 km (Valverde and Zavala-Hurtado, 2006). Pollination is carried out by small insects (Valverde PL, unpublished results), and seeds can be retained several years within mature fruits limiting dispersal in space and time (Peters et al., 2009). To date, there have been no attempts to explain the processes responsible for the diversification of mammillarias in the TVr. ...
... Mature fruits and seeds can be retained within the maternal plant for several years, a phenomenon known as serotiny. Seeds are only released during episodes of high precipitation, increasing the chances of germination and establishment (Peters et al., 2009). Pollination by small insects, serotiny, and gravity dispersal can strongly limit pollen and seed mobility among populations, constraining genetic interchange. ...
Article
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The endemic Mexican cactus, Mammillaria pectinifera, shows low dispersal capabilities and isolated populations within the highly dissected landscape of Tehuacán Valley. These characteristics can restrict gene flow and act upon the genetic divergence and speciation in arid plants. We conducted a phylogeographic study to determine if the origin, current distribution, and genetic structure of M. pectinifera were driven by Quaternary geomorphic processes. Sequences of the plastids psbA-trnH and trnT-trnL obtained from 66 individuals from seven populations were used to estimate genetic diversity. Population differentiation was assessed by an analysis of molecular variance. We applied a stepwise phylogenetic calibration test to determine whether species origin and genetic divergence among haplotypes were temporally concordant with recognizable episodes of geomorphic evolution. The combination of plastid markers yielded six haplotypes, with high levels of haplotype diversity (h = 0.622) and low nucleotide diversity (π = 0.00085). The populations were found to be genetically structured (FST = 0.682; P < 0.00001), indicating that geographic isolation and limited dispersal were the primary causes of genetic population differentiation. The estimated origin and divergence time among haplotypes were 0.017-2.39 and 0.019-1.237 mya, respectively, which correlates with Pleistocene tectonics and erosion events, supporting a hypothesis of geomorphically-driven geographical isolation. Based on a Bayesian skyline plot, these populations showed long term demographic stability, indicating that persistence in confined habitats has been the main response of this species to landscape changes. We conclude that the origin and haplotype divergence of M. pectinifera were a response to local Quaternary geomorphic evolution.