Comparative pollination biology of Venezuelan columnar cacti and the role of nectar-feeding bats in their sexual reproduction

The floral biology, reproductive system, and visitation behavior of pollinators of four species of columnar cacti, Stenocereus griseus, Pilosocereus moritzianus, Subpilocereus repandus, and Subpilocereus horrispinus, were studied in two arid zones in the north of Venezuela. Our results support the hypothesis that Venezuelan species of columnar cacti have evolved toward specialization on bat pollination. Additional information on the floral biology of a fifth species, Pilosocereus lanuginosus, was also included. All species showed the typical traits that characterize the pollination syndrome of chiropterophily. All species but Pilosocereus moritzianus were obligate outcrossers. Nectar and pollen were restricted to nocturnal floral visitors. Two species of nectar-feeding bats, Leptonycteris curasoae Miller and Glossophaga longirostris Miller, were responsible for practically all the fruit set in these cacti. Frequency of bat visitation per flower per night was highly variable within and between species of cactus, with average frequencies varying between 27 and 78 visits/flower/night. In general terms, the pattern of floral visitation through the night was significantly correlated with the pattern of nectar production and nectar sugar concentration for all species of cactus. Under natural pollination, fruit:flower ratios varied from 0.46 in Subpilocereus repandus to 0.76 in Stenocereus griseus. The efficiency of bat pollination in terms of seed:ovule ratio was high in all species, varying between 0.70 and 0.94.     

Stem Anatomy of Uebelmannia (Cactaceae) — with Special Reference to Uebelmannia gummifera

The three species of Uebelmannia (Cactaceae: Cactoideae; endemic to Minas Cerais, Brazil) are noteworthy for their tough, bumpy stem surface and the presence of conspicuous mucilage cavities restricted to the outer part of the stems. The anatomy of the mature structures and their ontogeny was investigated from microtome sections. The uncommon surface relief is caused by groups of unequally elongated hypodermis cells. The mucilage cavities consist of solitary and considerably enlarged mucilage cells (U.buiningii and U. pectinifera). In U.gummifera groups of mucilage cells disintegrate and form large cavities which are finally united into longitudinal ducts. A comparison of these stem features with species of the superficially similar genera Islaya and Wigginsia (representing tribe Notocacteae), does not indicate a close phylogenetic relationship between Uebelmannia and members of tribe Notocacteae. Finally, some short comments touch on the adaptational aspect of the characteristic stem features of Uebelmannia.     

Characterization of microsatellite loci for the critically endangered cactus Ariocarpus bravoanus

Ariocarpus bravoanus is common in trade but critically endangered in its natural habitat. With the ultimate aim of developing a certification scheme to aid in the conservation of this species, we have isolated A. bravoanus microsatellites from a nonenriched library. Fifty-four sequences contained a microsatellite array, of which eight were polymorphic among 23 individuals, 20 from one population and three plants from trade.     

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.     

Organogenesis of vegetative shoots from in vitro cultured flower buds of Mammillaria albicoma (Cactaceae)

We tested the organogenetic capacity of floral buds of Mammillaria albicoma Böed. (Cactaceae). Buds were incubated on solid MS medium supplemented with 0.1 mg l^?1 α-naphthaleneacetic acid (NAA) and 5.0 mg l^?1 6-benzylaminopurine (BA). Callus growth was observed from the cut explant base and from within the perianth. These calli during subsequent subcultures to the same medium gave rise to adventitious shoots. Shoots formed also directly from the perianth, as confirmed by observations in the light microscope and scanning electron microscope (SEM). On transfer to a fresh medium, the shoots produced proliferating cultures. This is the first report of regeneration of cactus shoots from floral explants.     

Population Genetic Structure of Two Columnar Cacti with a Patchy Distribution in Eastern Brazil

The genetic variability and population genetic structure of six populations of Praecereus euchlorus and Pilosocereus machrisii were investigated. The genetic variability in single populations of Pilosocereus vilaboensis, Pilosocereus aureispinus, and Facheiroa squamosa was also examined. All of these cacti species have a patchy geographic distribution in which they are restricted to small areas of xeric habitats in eastern Brazil. An analysis of genetic structure was used to gain insights into the historical mechanisms responsible for the patchy distribution of P. euchlorus and P. machrisii. High genetic variability was found at the populational level in all species (P=58.9–92.8%, A_p=2.34–3.33, H_e=0.266–0.401), and did not support our expectations of low variability based on the small population size. Substantial inbreeding was detected within populations (F_IS=0.370–0.623). In agreement with their insular distribution patterns, P. euchlorus and P. machrisii had a high genetic differentiation (F_ST=0.484 and F_ST=0.281, respectively), with no evidence of isolation by distance. Accordingly, estimates of gene flow (N_m) calculated from F_ST and private alleles were below the level of N_m=1 in P. machrisii and P. euchlorus. These results favored historical fragmentation as the mechanism responsible for the patchy distribution of these two species. The genetic distance between P. machrisii and P. vilaboensis was not compatible with their taxonomic distinction, indicating a possible local speciation event in this genus, or the occurrence of introgression events.     

Allozyme diversity and morphometrics of Melocactus paucispinus (Cactaceae) and evidence for hybridization with M. concinnus in the Chapada Diamantina, North-eastern Brazil

BACKGROUND AND AIMS: Melocacatus paucispinus (Cactaceae) is endemic to the state of Bahia, Brazil, and due to its rarity and desirability to collectors it has been considered threatened with extinction. This species is usually sympatric and inter-fertile with M. concinnus, and morphological evidence for hybridization between them is present in some populations. Levels of genetic and morphological variation and sub-structuring in populations of these species were assessed and an attempt was made to verify the occurrence of natural hybridization between them. METHODS: Genetic variability was surveyed using allozymes (12 loci) and morphological variability using multivariate morphometric analyses (17 vegetative characters) in ten populations of M. paucispinus and three of M. concinnus occurring in the Chapada Diamantina, Bahia. KEY RESULTS: Genetic variability was low in both species (P = 0.0-33.3, A = 1.0-1.6, H(e) = 0.000-0.123 in M. paucispinus; P = 0.0-25.0, A = 1.0-1.4, H(e) = 0.000-0.104 in M. concinnus). Deficit of heterozygotes within the populations was detected in both species, with high values of F(IS) (0.732 and 0.901 in M. paucispinus and M. concinnus, respectively). Evidence of hybridization was detected by the relative allele frequency in the two diaphorase loci. High levels of genetic (F(ST) = 0.504 in M. paucispinus and 0.349 in M. concinnus) and morphological (A = 0.20 in M. paucispinus and 0.17 in M. concinnus) structuring among populations were found. CONCLUSIONS: The Melocactus spp. displayed levels of genetic variability lower than the values reported for other cactus species. The evidence indicates the occurrence of introgression in both species at two sites. The high F(ST) values cannot be explained by geographical substructuring, but are consistent with hybridization. Conversely, morphological differentiation in M. paucispinus, but not in M. concinnus, is probably due to isolation by distance.     

The impact of early Quaternary climate change on the diversification and population dynamics of a South American cactus species

Aim

Climatic oscillations have been suggested to promote speciation and changes in species distributions, mostly in connection with the Last Glacial Maximum (LGM). However, the LGM is just the most recent of the 20+ glacial‐interglacial periods that characterise the Quaternary. Here, we investigate the role of climatic changes and geomorphological features in shaping the evolution, distribution and population dynamics of the South American cactus Cereus hildmannianus.

Location

South‐eastern South America.

Methods

We built a large fossil‐calibrated phylogeny for cacti (family Cactaceae), comprising 128 species distributed in all subfamilies, using a Bayesian relaxed clock. We used the results to derive a secondary calibration for a population‐level phylogeny in C. hildmannianus. We amplified two plastid (trnQ‐5′rps16 and psbJ‐petA) and one nuclear marker (PhyC) for 24 populations. We estimated population dynamics, ancestral areas, and species distribution models to infer the clade's evolutionary history in time and space.

Results

Our results show a major population divergence of C. hildmannianus at c. 2.60 Ma, which is strikingly coincident with the transition of the Pliocene–Pleistocene and onset of Quaternary glaciations. This was followed by a complex phylogeographic scenario involving population expansions across ecologically diverse regions.

Main conclusions

Contrary to the dominant research focus on the LGM, our study indicates a major impact of the first Quaternary glaciation on the distribution and population divergence of a South American plant species. Further intraspecific events seem related to successive climatic changes and geomorphology, including the development of the coastal plain and its peculiar diversity. We propose that the first Quaternary glaciation acted as a major evolutionary bottleneck, whereby many warm‐adapted lineages succumbed, while those that survived could diversify and better cope with subsequent climatic oscillations.     

Isoenzymes of peroxidase and esterase related to morphogenesis in Mammillaria gracillis Pfeiff. tissue culture

In vitro propagated plants of the cactus Mammillaria gracillis Pfeiff. (Cactaceae) spontaneously produced callus. The habituated callus regenerated normal and hyperhydric shoots without the addition of grown regulators. Tumours were obtained by infecting cactus explants with Agrobacterium tumefaciens; the wild strain B6S3 (tumour TW) or with the rooty mutant GV3101 (tumour TR). Both tumour lines grew vigorously, never expressing any morphogenic potential. In this study, cactus shoots, callus, normal and hyperhydric regenerants and TW and TR tumours were compared with regard to peroxidase (EC 1.11.1.7) and esterase activity, and isoenzyme patterns. Guaiacol peroxidase activity was the lowest in the cactus shoots and in the normal regenerants. Callus, hyperhydric regenerants and tumours had peroxidase activity of 6 to 7 times higher. Esterase activity was measured with 1- and 2-naphthylacetate as broad-spectrum substrates. The highest esterase activity was determined in tumours with both substrates. All tissues, except the TR tumour, had higher esterase activity for 2-compared to 1-naphtylacetate. Peroxidase and esterase isoenzyme patterns were not completely identical among the investigated tissues.     

Senita cactus: a plant with interrupted sterol biosynthetic pathways

Locereol (4α-methylcholesta-8,14-dien-3β-ol) and 5α-cholesta-8,14-dien-3β-ol, not previously isolated from plants, 24-methylenelophenol, lathosterol, 5α-campest -7-en-3β-ol and spinasterol are present in senita cactusin addition to the lophenol and schottenol described previously.