Impact of climate change on the distribution of populations of an endemic Mexican columnar cactus in the Tehuacán-Cuicatlán Valley,Mexico

In this paper, we present a bioclimatic approach to (1) differentiate populations of the endemic Mexican columnar cactus Neobuxbaumia tetetzo within the Tehuacán-Cuicatlán Valley and (2) evaluate, under two possible future scenarios (years 2050 and 2080), the effects of climate change on the total species distribution in this area, as well as on groups of populations defined by their bioclimatic models. Four population groups were identified, and principal component analysis showed that the variables that explained more than 40% of the climatic variation were precipitation of the wettest quarter and temperature seasonality. Bioclimatic models under the different scenarios indicated that when the overall species distribution was analyzed, this area will probably have contracted by 19.5% by the year 2050 and 47.05% by the year 2080, whereas the separate analysis of population groups projected area contractions of 18.4% by the year 2050 and 51.95% by the year 2080. These results demonstrate the importance of exploring new approaches for evaluating and predicting current and future distribution of plant species.     

Demography of the columnar cactus Neobuxbaumia macrocephala: a comparative approach using population projection matrices

Neobuxbaumia macrocephala is a long-lived columnar cactus endemic to the Tehuacán-Cuicatlán Valley in south-central Mexico. This plant has a very restricted distribution and few recruitment events have been detected in its populations. In this study, we analyze the N. macrocephala demographic pattern using a projection matrix in order to determine the main limiting factors of this species. To accomplish this goal, we compare our results with those obtained for another species of the same genus, N. tetetzo. Considering that both species inhabit the same valley, we believe that this comparative study will offer insights into the main demographic limitations of N. macrocephala. Results showed that these species of columnar cacti have similar demographic patterns in which survival is the process with the highest relative contribution to λ, followed by growth and reproduction. Of all the life cycle stages, seeds and seedlings have the lowest survival probabilities due to a high mortality caused by seed predation and effects of direct solar radiation on germinated seeds. The estimated growth rates indicate that populations of these species of Neobuxbaumia are in a numerical equilibrium. With respect to reproduction, N. macrocephala produce a lower number of seeds per plant than N. tetetzo. This low level of sexual reproduction may decrease the probability of establishment of new individuals in N. macrocephala populations. It is suggested that pollen limitation and pre-dispersal seed predation could be some factors that limit the distribution and abundance of this columnar cactus. This revised version was published online in June 2006 with corrections to the Cover Date.     

Genetic diversity and population genetic structure of three endemic species of Mammillaria (Cactaceae) from the Tehuacán Valley in central México

Biodiversity and Conservation - Mammillaria hernandezii, M. dixanthocentron and M. lanata are endemic and threatened cactus species that inhabit the Tehuacán Valley, an arid zone in southern...     

Pollination biology of Oreocereus celsianus (Cactaceae), a columnar cactus inhabiting the high subtropical Andes

The geographical dichotomy hypothesis suggests that columnar cacti in the tropics depend primarily on bats for pollination. This dependence may to be less in the outer tropics where many columnar cactus species (or their populations) show a relatively generalized pollination system with both nocturnal (moths and bats) and diurnal pollinators (bees and hummingbirds) (geographical dichotomy hypothesis). This hypothesis has been mostly tested in the northern tropics; nonetheless, our knowledge of columnar cactus species inhabiting the southern tropics is still scarce. The aim of this project was to evaluate the pollination biology of Oreocereus celsianus, a columnar cactus with restricted distribution in the subtropical Andes, to determine if the pollination system of this cactus tends to be more generalized than specialized because of the geographical position where it occurs. Observations of frequency of visit showed that Patagona gigas (Giant Hummingbird) is the main pollinator of the flowers, visiting them when they are opening (afternoon of the first day). Bees, wasps and moths were occasional visitors of the flowers. None of them seem to act as pollinator. Autogamy, geitonogamy and xenogamy treatments produced high fruit set, showing that O. celsianus has an unusual mixed mating system. The results suggest that this Andean columnar cactus is partially specialized on hummingbirds, with most pollination service performed by a single species, and it has the capacity of selfing (??fail-safe?? pollination system). This mixed mating system (both outcrossing and selfing) may be a response to the unpredictable environment of the Prepuna in the subtropical Andes.     

Isolation and characterization of microsatellites in the columnar cactus: Polaskia chichipe and cross‐species amplification within the Tribe Pachycereeae (Cactaceae)

Microsatellite markers were developed for the columnar cactus Polaskia chichipe from central Mexico. After an enrichment procedure and three screening steps 87% of colonies contained microsatellites. A pair of primers for 10 loci (seven polymorphic) were developed, tested and used to estimate variation in samples of 18–45 individuals from the Tehuacan Valley, Mexico. Alleles per locus ranged from two to eight (mean 5.28; SD 2.5). Range of expected heterozygosity values was 0.188–0.797 (mean 0.502; SD 0.25). These loci are particularly useful for more precise evolutionary studies, such as gene flow and breeding systems, for this cactus species.     

Stem tilting,pseudocephalium orientation,and stem allometry in <Emphasis Type="Italic">Cephalocereus columna-trajani</Emphasis> along a short latitudinal gradient

Cephalocereus columna-trajani is a giant columnar cactus endemic of the Tehuacán-Cuicatlán Valley in Central Mexico. Stem tilting and northward pseudocephalium azimuth in C.␣columna-trajani have functional advantages in terms of interception of direct solar radiation at the northernmost portions of its range. Since the success of both characters strongly depends on the apparent position of the sun during the growing season, in this paper we test the hypothesis that the occurrence of such columnar morphology is restricted geographically and imposes mechanical restrictions that limit column height. Following a latitudinal gradient along the Tehuacán-Cuicatlán Valley, we selected five populations, recorded tilting angle and pseudocephalium azimuth, and carried out allometric and biomechanical analyses of height–diameter relationships. Northern populations showed higher tilting angles. Pseudocephalium azimuth significantly differed among populations, and pseudocephalium orientation was consistently North-Northwestern. Stem allometry showed that the stems of the southern populations increased in height at a far greater rate with respect to diameter than the northern populations. The southernmost population showed the lowest safety factor. These results support the hypothesis that stem tilting in C.␣columna-trajani is functionally advantageous in a restricted geographical range, and imposes mechanical restrictions to column height.     

Pollination ecology of Stenocereus queretaroensis (Cactaceae), a chiropterophilous columnar cactus, in a tropical dry forest of Mexico

Flowers of columnar cacti are animal-pollinated, often displaying a chiropterophylic syndrome. This study examined if the columnar cactus Stenocereus queretaroensis, a tropical species endemic to western Mexico, is bat-pollinated, by studying its pollination biology and the foraging behavior of potential pollinators. Flowers were produced in winter through spring, peaking in April. Anthesis was nocturnal, and stigma and anther turgidity began around 2200 hours. Production of nectar secretion and highest sugar concentration and energy supply were nocturnal, peaking between 2200 and 2400 hours. Manual auto-pollination and exclusion experiments showed that self-pollination yielded no fruits, while nocturnal pollinators resulted in high fruit set and seed set compared to diurnal pollination treatments. The nectar-feeding bat Leptonycteris curasoae (Phyllostomidae) was the main nocturnal pollinator with the highest effective pollination. Peak bat visitation coincided with peaks in nectar production. The high abundance of L. curasoae throughout the 4-yr study, suggests that it is a seasonally reliable pollinator for this columnar cactus. While pollination syndromes have been increasingly called into question in recent years, this study suggests that at least for this system, there is a fairly close fit between pollinator and pollination syndrome.     

Traditional Management and Morphological Patterns of <Emphasis Type="Italic">Myrtillocactus schenckii</Emphasis> (Cactaceae) in the Tehuacán Valley,Central Mexico<Superscript>1</Superscript>

Traditional Management and Morphological Patterns of Myrtillocactus schenckii (Cactaceae) in the Tehuacán Valley, Central Mexico Myrtillocactus schenckii is a columnar cactus endemic to central México and appreciated for its edible fruit. In the Tehuacán Valley it occurs wild in thorn-scrub forests, but it is also under silviculture management—tolerate in agroforestry systems and cultivated in home gardens. Ethnobotanical and morphometric studies were conducted to document its use and management forms, as well as consequences of artificial selection on morphological patterns in managed populations. In silviculture populations artificial selection occurs through selective tolerance of the phenotypes producing more, larger, and sweeter fruits, whereas in home gardens people plant branches of the preferred phenotypes. Morphological differences were found especially in fruit size and production, the main targets of artificial selection. The highest average values of fruit size and production were found in cultivated populations (0.993 cm³ and 204.45 fruits per branch, respectively), intermediate in silviculture populations (0.819 cm³ and 70.63 fruits per branch), and the lowest in the wild (0.68 cm³ and 59.75 fruits per branch). Multivariate statistical analyses differentiated populations according to their management type. Morphological diversity was higher in managed populations (0.703 ± 0.029 and 0.679 ± 0.019 in silviculture and cultivated populations, respectively) than in the wild (0.652 ± 0.016). Managed plants of M. schenckii do not show signs of depending on humans for survival and reproduction and, therefore, domestication should be considered incipient.     

Facilitation of the non-native annual plant Mesembryanthemum crystallinum (Aizoaceae) by the endemic cactus Eulychnia acida (Cactaceae) in the Atacama Desert

The Atacama Desert is one of the most stressful environments worldwide and represents a strong barrier for the establishment of native and non-native plants. In this study, we report the establishment of a non-native annual plant through facilitation by a native endemic cactus in a relatively undisturbed coastal area in north-central Chile. Soil collected under Eulychnia acida contained more available nutrients (N, P and K), water, and soluble salts than soils collected away from E. acida. Co-occurrence analyses showed a strong positive spatial association (facilitation) between the native cactus E. acida and the non-native annual, Mesembryanthemum crystallinum. The aboveground biomass of M. crystallinum individuals was 4-fold higher under the influence of E. acida. Native halophytes occasionally shared the cactus understory with the non-native species, but dominant native shrubs and perennial herbs did not co-occur with the cactus at scales of 1 and 4 m². All these results support facilitation of the native cactus on the non-native herb. The combination of direct and indirect positive effects could explain the assembly of the non-native annual plant in these undisturbed areas of the Atacama Desert and have major implications on M. crystallinum capacity to colonize new areas.     

Global Diversity and Conservation Priorities in the Cactaceae

The purpose of this paper was to analyze the diversity patterns of Cactaceae at a global scale, to identify those countries where conservation actions should be performed. In order to do this, the species richness and the number of endemic species for 34 American countries were determined. With these data, the relationship between the total number of species or the number of endemic species and the area of the countries were analyzed. In addition, a complementarity analysis was conducted to determine the most important countries for cactus conservation. Results showed that Mexico had the highest number of total and endemic species followed by Argentina, Bolivia, Brazil, and Peru, among others. There was a significant positive relationship between both, the total and endemic species, and the area of the countries. Despite this fact, the cactus diversity in Mexico, Argentina, Peru, Bolivia, Chile, and Costa Rica was higher than expected according to their area. Further, these countries also presented the highest proportions of endemic species. The complementarity analysis indicated that 24 countries are necessary to preserve all cactus species. However, 94% of all species could be preserved with only 10 countries. Considering the diversity patterns and the complementarity analysis, three important groups for cactus conservation were identified: (1) Mexico, Argentina, Peru, Bolivia, Chile, and Costa Rica, (2) Paraguay and Cuba, and (3) Brazil and USA. Conservation efforts should be focused on these countries in order to preserve cactus diversity.     

Experimental Evolution of Alkaloid Tolerance in Sibling <Emphasis Type="Italic">Drosophila</Emphasis> Species with Different Degrees of Specialization

Drosophila buzzatii and Drosophila koepferae are sibling species with marked ecological differences related to their patterns of host exploitation. D. buzzatii is a polyphagous species with a sub-cosmopolitan distribution, while D. koepferae is endemic to the mountain plateaus of the Andes, where it exploits alkaloidiferous columnar cacti as primary hosts. We use experimental evolution to study the phenotypic response of these cactophilic Drosophila when confronting directional selection to cactus chemical defenses for 20 generations. Flies adapted to cactus diets also experienced higher viability on alkaloid-enriched media, suggesting the selection of adaptive genetic variation for chemical-stress tolerance. The more generalist species D. buzzatii showed a rapid adaptive response to moderate levels of secondary metabolites, whereas the columnar cacti specialist D. koepferae tended to maximize fitness under harder conditions. The evolutionary dynamic of fitness-related traits suggested the implication of metabolic efficiency as a key mediator in the adaptive response to chemical stress. Although we found no evidence of adaptation costs accompanying specialization, our results suggest the involvement of compensatory evolution. Overall, our study proposes that differential adaptation to secondary metabolites may contribute to varying degrees of host specialization, favoring niche partitioning among these closely related species.     

Reproductive biology of Ferocactus recurvus (Mill.) Borg subsp. recurvus (Cactaceae) in the Tehuacán‐Cuicatlán Valley,Mexico

• Mexico has one of the highest diversities of barrel cacti species worldwide; however, all are threatened and require conservation policies. Information on their reproductive biology is crucial, but few studies are available. Ferocactus recurvus subsp. recurvus is a barrel cactus endemic to the Tehuacán‐Cuicatlán Valley. Our research aimed to characterise its floral and pollination biology. We hypothesised bee pollination, as suggested by its floral morphology and behaviour, and self‐incompatibility, like most barrel cacti studied.
• Three study sites were selected in the semiarid Zapotitlán Valley, Mexico. We examined 190 flowers from 180 plants to determine: morphometry and behaviour of flowers, flower visitors and probable pollinators, and breeding system.
• Flowers showed diurnal anthesis, lasting 2–5 days, the stigma being receptive on day 2 or 3 after the start of anthesis. Flowers produced scarce/no nectar and main visitors were bees (Apidae), followed by flies (Muscidae), ants (Formicidae), thrips (Thripidae) and hummingbirds (Throchilidae); however, only native bees and occasionally wasps contacted the stigma and anthers. Pollination experiments revealed that this species is self‐incompatible and xenogamous. In natural conditions, fruit set was 60% and cross‐pollination fruit set was 100%. Percentage seed germination resulting from cross‐pollination was higher than in the control treatment.
• Our results provide ecological information for conservation programmes to ensure a high probability of breeding and seed production in natural populations of F. recurvus.

     

Diversity, spatial arrangement, and endemism of Cactaceae in the Huizache area, a hot-spot in the Chihuahuan Desert

We studied several aspects of the cactus diversity in the Huizache, an area located in the southern extreme of the Chihuahuan Desert Region, in San Luis Potosí, Mexico. Fieldwork was conducted in a square-shaped polygon (22°30–23°00 N, 100°00–100°30 W), where a total of 80 systematically-chosen localities were sampled for cactus species. Results showed that the Huizache is an important focal point for the conservation of the Cactaceae, due to the remarkably high diversity of members of this plant family in the area. With 75 species recorded, this area has the highest concentration of cactus species in the American Continent, in comparison with other comparably-sized regions. The outstanding diversity of cactus species in the Huizache area may be explained by the relatively favorable climate of this desert area, its environmental heterogeneity, and its location in a transitional position between three regions with distinct cactus floras: Chihuahuan Desert Region, Queretaroan-Hidalgoan Arid Zone, and Tula-Jaumave Valleys. Species richness is not distributed evenly in the area; the zones of highest species concentration are primarily located in the lowland valleys, in the northwestern portion of the grid square, where typical Chihuahuan Desert conditions prevail. Of the 75 cactus species found in the area, 63% are endemic to the Chihuahuan Desert. Among these, ten species are outstanding for being endemic or nearly endemic to the Huizache area. The Huizache grid square is part of a recently declared natural reserve, The Real de Guadalcázar Natural Reserve.     

Mycorrhizal perennials of the "matorral xerófilo" and the "selva baja caducifolia" communities in the semiarid Tehuacán-Cuicatlán Valley,Mexico

We investigated the mycorrhizal status of perennial xeric plant species occurring in the "matorral xerófilo" (arid tropical scrub) and the ecotone of the "selva baja caducifolia" (tropical deciduous forest) communities in the semiarid valley of Tehuacán-Cuicatlán, south-central Mexico. The perennial species examined are dominant/codominant elements within the "matorral xerófilo" and the "selva baja caducifolia", both endangered communities in the Biosphere Reserve Tehuacán-Cuicatlán Valley. Of the 50 sampled species, 45 were mycorrhizal. To our knowledge, we report arbuscular mycorrhizae (AM) for the first time in 37 species, of which 21 are endemic to Mexico and nine are endemic to the Valley. We also report AM for the first time in three genera, Buddleja, Hechtia and Zornia, and in one plant family, Buddlejaceae. Beaucarnea gracilis, a threatened species, and Mimosa purpusii, a potentially rare species, are both mycorrhizal. This is the first study of the mycorrhizal status of plant species within the Valley.     

Floral biology of Echinopsis chiloensis ssp. chiloensis (Cactaceae): Evidence for a mixed pollination syndrome

In this study the flower biology of Echinopsis chiloensis ssp. chiloensis, a columnar cactus occurring in Central Chile, is investigated, in particular its pollination syndrome, its visitors, their frequencies and behaviors and their pollination efficiencies. As statements on floral anthesis of this species are contradictory, this study also intended to elucidate both its beginning and duration. A pollinator exclusion study of a total of 162 flowers from 12 plants was conducted at one of the two study sites. Fruit and seed production as well as seed viability were documented to evaluate pollinator efficiencies.     

Genetic differentiation of island populations: geographical barrier or a host switch?

In the Sonoran desert, there exists a diverse community of cactophilic drosophilids that exploit toxic, rotting cactus tissue as a food resource. The chemistry of the necrotic cactus tissue varies among species, and several drosphilid species have evolved specialized detoxification mechanisms and a preference for certain cactus types. In the present study, we compared the genetic structure of two columnar cactus species, Drosophila mettleri and Drosophila mojavensis, and two prickly pear species, Drosophila mainlandi and Drosophila hamatofila, which have all recently colonized Catalina Island off the coast of southern California. Because there are no columnar cactus species on Catalina Island, the two columnar specialists underwent a host switch to prickly pear cactus, the only cactus present on the island. Previous genetic studies of D. mettleri and D. mojavensis showed significant genetic differentiation between mainland and island populations, which could result from restricted gene flow as a result of the San Pedro Channel, or because of a host switch to prickly pear. To distinguish between these possibilities, we analyzed the genetic structure of the prickly pear species aiming to isolate the effects of geography versus host switching. The results obtained show little to no genetic differentiation for the prickly pear species, supporting the hypothesis that the genetic differentiation of the two columnar species is a result of a host switch from columnar cacti to prickly pear. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Effectiveness of bats as pollinators of Stenocereus stellatus (Cactaceae) in wild, managed in situ, and cultivated populations in La Mixteca Baja, central Mexico

Stenocereus stellatus is an endemic, self-incompatible, columnar cactus found in central Mexico where many of its wild populations have been fragmented. As an economically important species of fruit-producing cactus, S. stellatus occurs in wild, managed in situ, and cultivated populations. The objectives of this study were to determine the effective pollinators of S. stellatus, to compare pollinator visits and reproductive parameters among the three types of populations, and to determine if nectar feeding-bats are moving among populations. Effective pollinators were the nectarivorous bats Choeronycteris mexicana, Leptonycteris curasoae, and L. nivalis. Fewer total visits per flower per night and fewer visits by Choeronycteris were observed in cultivated populations, while the opposite pattern was observed for Leptonycteris. One aggressive interaction was filmed in which Choeronycteris was physically displaced by Leptonycteris, and Choeronycteris visits were significantly affected by Leptonycteris visits. Cultivated populations received more pollen grains and had more fruit set. Variation in pollinator visits between different populations and the consequent effects on reproductive success were likely a result of competition between bat species, and differences in foraging and in sensitivity of bat species to human populations. Three marked L. curasoae traveled 15 km from their roosting site to their foraging area, and one visited cultivated and managed populations, suggesting that this species may be particularly important in moving pollen among populations.     

Host-plant shifts and adult survival in the cactus breeding Drosophila mojavensis

Abstract. 1. In order to gain insight into how Drosophila mojavensis Patterson and Crow (which lives on the rotting tissue of cactus) successfully shifted host plants, from an Opuntia species to columnar cactus species, over evolutionary time, specific components of columnar cactus and Optunia tussue were examined for their effects on adult longevity.
2. The availability of free sugars affects the survival of adult D.mojavensis . The addition of a sugar to columnar cactus tissue significantly increased the longevity of adults that fed on those tissues. This increase in adult longevity was equal to the longevity observed on unsupplemented Opuntia tissue, which has free sugars present.
3. Live microorganisms added to Opuntia tissue had a slight detrimental effect on adult longevity (longevity was decreased by approximately 1 week).
4. Low concentrations of atmospheric ethanol (independent of diet) increased the longevity of adult D.mojavensis to levels comparable in magnitude to that provided by the addition of free sugars to the diet.
5. This information and the apparent lack of a year-round supply of free sugars for D.mojavensis in nature suggests that the ethanol (produced by fermentation) rather than free sugars per se is the most important component of rotting cactus tissue for the survival of this species and may explain its successful host-plant shift.     

Bat and hummingbird pollination of an AUTOTETRAPLOID COLUMNAR CACTUS,Weberbauerocereus weberbaueri (Cactaceae)

I present data on flower morphology, pollination, breeding system, and genetic diversity of the columnar cactus Weber-bauerocereus weberbaueri at 2500 m elevation in southwestern Peru. Weberbauerocereus weberbaueri is a self-compatible columnar cactus that is visited and pollinated by one species of rare endemic bat, Platalina genovensium, and two species of hummingbirds, Patagona gigas and Rhodopis vesper. W. weberbaueri exhibits pronounced interplant variation in floral color and size, and flowers exhibit traits corresponding to both bat and hummingbird pollination syndromes. Starch-gel electrophoresis of flower bud tissue indicated that W. weberbaueri is an autotetraploid and that genetic diversity (Hep = 0.257) of the study population was high relative to diploid plants but similar to other tetraploid species. Initial fruit set from pollinator exclusion experiments conducted in 1991 and 1993, at the onset of and after a drought associated with the El Niño event of 1991–1992, revealed that bats were the most important pollinators in 1991, but that hummingbirds and diurnal insects were most important in 1993. In both years, however, autogamy and lepidopteran larval infestation of fruits reduced differences in mature fruit production among pollinator exclusion treatments so that differences in mature fruit set were not statistically significant. Reduced bat pollination in 1993 is attributed to the reduced abundance of bats at the study site during a drought caused by El Niño. I hypothesize that interaction among several factors, including tetraploidy, autogamy, larval infestation of developing fruits, and variation in pollinator abundance, may not result in strong selection for a bat vs. hummingbird floral morph, thus allowing the persistence of floral variation in this cactus.