A unique cactus with scented and possibly bat-dispersed fruits: Rhipsalis juengeri

Rhipsalis juengeri was described in 1995 as an unusual representative of epiphytic cacti, forming more than 3 m long curtains, hanging from the canopy of the Atlantic Rainforest in eastern Brazil. At the apex of thin, pendant shoots, green-brownish berries are formed. We report here as a novelty for the Cactaceae that these berries are strongly scented, and present an odour analysis along with an olfactoric survey of fruits of about 50 species and varieties of the cactus tribe Rhipsalideae. The volatile blend of berries of R. juengeri is dominated by ketones, some of which are responsible for the characteristic blackcurrant-like scent, as is shown by GC-olfactometry. The odour and inconspicuous colour stand out among fruits of other epiphytic cacti that are thought to be consumed by birds. Fruit characters of R. juengeri and the flagellicarpic presentation indicate adaptation to chiropterochory.     

Programmed cell death promotes male sterility in the functional dioecious Opuntia stenopetala (Cactaceae)

Background and Aims

The sexual separation in dioecious species has interested biologists for decades; however, the cellular mechanism leading to unisexuality has been poorly understood. In this study, the cellular changes that lead to male sterility in the functionally dioecious cactus, Opuntia stenopetala, are described.

Methods

The spatial and temporal patterns of programmed cell death (PCD) were determined in the anthers of male and female flowers using scanning electron microscopy analysis and histological observations, focusing attention on the transition from bisexual to unisexual development. In addition, terminal deoxynucleotidyl transferase-mediated dUTP nick-end labelling assays were used as an indicator of DNA fragmentation to corroborate PCD.

Key results

PCD was detected in anthers of both female and male flowers, but their patterns differed in time and space. Functionally male individuals developed viable pollen, and normal development involved PCD on each layer of the anther wall, which occurred progressively from the inner (tapetum) to the outer layer (epidermis). Conversely, functional female individuals aborted anthers by premature and displaced PCD. In anthers of female flowers, the first signs of PCD, such as a nucleus with irregular shape, fragmented and condensed chromatin, high vacuolization and condensed cytoplasm, occurred at the microspore mother cell stage. Later these features were observed simultaneously in all anther wall layers, connective tissue and filament. Neither pollen formation nor anther dehiscence was detected in female flowers of O. stenopetala due to total anther disruption.

Conclusions

Temporal and spatial changes in the patterns of PCD are responsible for male sterility of female flowers in O. stenopetala. Male fertility requires the co-ordination of different events, which, when altered, can lead to male sterility and to functionally unisexual individuals. PCD could be a widespread mechanism in the determination of functionally dioecious species.     

Spider communities in two plant architectures of epiphytic cacti in the Brazilian Atlantic forest

The vegetative architecture of the Cactaceae provides sites used by spiders as hideout, foraging, mating and oviposition. We found more web weavers in plants with cylindrical cladodia and more hunters in plants with flat cladodia. Between February 2014 and January 2015 1551 spiders distributed in 20 families were collected: one of these families, Palpimanidae, had only juvenile individuals collected; of the other 19 families, 80 species/morphospecies were identified. This work aims to contribute to the knowledge of biodiversity in the canopy of the Atlantic Forest, especially the relationship between cacti and spiders.     

Revisiting Bergmann's rule for saguaros (Carnegiea gigantea (Engelm.) Britt. and Rose): stem diameter patterns over space

Aim The influence of winter temperatures and other climate variables are explored to determine which variables are associated with saguaro stem diameter and to determine if Bergmann's rule is applicable to saguaros. Location The northern Sonoran Desert in Arizona, USA. Methods Thirty saguaro populations were sampled (height, diameter, number of branches), and after adjustment for population height structure, mean relative thickness of saguaros was calculated for each plot (population). Fifty‐seven climate variables were calculated for the thirty populations. Regression was run to determine which variables (cold winter, hot summer and precipitation) best predict relative thickness. Previous studies have demonstrated a significant positive relationship between winter precipitation and saguaro branching ( Drezner, 2003 , Ecography, in press). To determine if relative thickness may be an artefact of branching (branches), partial correlation analysis was employed. Results Mean March precipitation best predicts relative thickness. When only winter temperature variables are considered, none are significantly related to relative thickness. Relative thickness is not an artefact of branches. Main conclusions (1) Rainfall, not temperature, best predicts saguaro stem thickness. In addition, despite the focus on summer rains in the literature, winter precipitation is the best predictor of thickness. (2) Bergmann's rule is not applicable to saguaro populations as has been previously suggested [e.g. Niering et al. (1963) Science 142 , 15], as thickness does not increase significantly with latitude. In addition, the suggested mechanism for Bergmann's rule, cold winter temperature, does not significantly predict saguaro stem diameters over the area studied. In some populations that experience high winter rainfall as well as cold temperatures, individuals likely derive thermal benefits from a larger stem diameter; however, the trend is not observed over the area studied and it does not appear to be adaptive.     

Minor betalains in fruits of Hylocereus species

Betacyanins in peel and flesh of fruits of different Hylocereus species were identified by means of GC/MS, electrospray MS/MS, HPLC as well as (1)H and (13)C NMR techniques. As hitherto unknown pigments: betanidin 5-O-(2'-O-beta-D-apiofuranosyl)-beta-D-glucopyranoside, betanidin 5-O-(4'-O-malonyl)-beta-D-glucopyranoside and betanidin 5-O-[(5'-O-E-sinapoyl)-2'-O-beta-D-apiofuranosyl]-beta-D-glucopyranoside were elucidated. The sinapoyl moiety attachment position in the structure of betacyanins was established for the first time. The peel contained a more complex pattern of betacyanins with apiofuranosyl moiety. Other recently identified pigments were also present in the samples and their (1)H or (13)C NMR spectra were recorded. In the case of phyllocactin and its 4'-isomer the migration of the malonyl group was noticed.     

Pollination biology of Harrisia portoricensis (Cactaceae), an endangered Caribbean species

Traits associated with self-pollination are common features of island plant communities. In this work, we studied the pollination biology and the breeding system of Harrisia portoricensis, an island columnar cactus, to test for the presence of inbreeding and inbreeding depression. For H. portoricensis, which bears flowers with typical outcrossing morphology, the results from 322 h of direct observations and videotaping showed that visits to flowers by animals were uncommon. Controlled pollinations demonstrated that H. portoricensis has a partially self-compatible breeding system that it is not autogamous and thus requires an external mechanism for the movement of pollen to set fruit. We detected differences in seed size, seed mass, germination success, and multiplicative fitness estimates between self- and cross-pollination treatments. We found that progeny resulting from natural and self-pollination treatments showed signs of inbreeding depression compared with progeny resulting from cross-pollination; however, the magnitude of the inbreeding depression was less than 50%. Our combined results suggest that for this species an endogamous breeding system should be favored by natural selection.     

Spiny invaders – Patterns and determinants of cacti invasion in Europe

In this paper, we analyse the patterns and determinants of cacti invasion in 22 European countries. We compiled a checklist for each country. Cacti were classified for each country according to their invasion status as casuals, locally established (1–5 localities of small population size) and widely established (>5 localities of considerable population size).We used generalised linear models (GLM) from the Poisson family with a log-link function and a set of seven country-specific explanatory variables to account for geographical, climatic, habitat-related and economic determinants to test which features of the recipient area determine invasion success and if distribution patterns of species at different invasion stages are governed by the same interplay of explanatory variables. Separate models were fitted with the same predictor variables for casual, locally established, widely established and all cacti. Further, we analysed the temporal invasion trend, and tested if niche breadth (expressed as the number of habitat types colonized) is influenced by the range size (measured as the number of countries invaded). Finally, we reviewed the consequences of cacti invasion for nature conservation.In total, 26 cacti species have been recorded in Europe. Ten species are more widespread and occur in at least three countries, Opuntia humifusa (six countries) being the most widespread species. The country with most cacti is Spain (21 species), whereas in 13 countries no cactus species have been recorded. By far the most important genus is Opuntia with 20 alien species. The temporal invasion pattern shows an exponential increase of the cumulative number of invasion events, increasing from three (1801–50) to nine (1951–2000) invasion events over a 50-year period.Regardless of the invasion stage, the factor explaining most of the variance in the models is the presence of the Mediterranean biogeographic region, and a significant positive effect of the country size on species numbers was identified.Considering the invasion stage, some interesting deviations in the models can be observed. Invasion of casual cacti is only influenced by the presence of the Mediterranean biogeographic region. For locally established cacti, precipitation is negatively correlated with the invasion rate, and the presence of the Alpine biogeographic region is positively correlated; the latter is due the local occurrence of few hardy cacti (Opuntia phaeacantha, O. humifusa) in low-lying valleys of the Alps. As all widely established cactus species are restricted to the Mediterranean region, only this factor was included in the model.All cacti are confined to dry, open habitats on acid siliceous bedrock. Thus, the predominant habitats invaded are rock vegetation, dry grassland, open Mediterranean scrub and dry ruderal habitats. The niche breadth of cacti increases with the numbers of countries colonized. Further, the niche breadth of cacti exhibits a geographic gradient towards the Mediterranean region.Until the 19th century, the dominant pathway of invasion was agriculture, as some cactus species had been introduced for the production of forage and fruits. However, in the last decades horticulture and deliberate planting in the wild have become the dominant pathways.The invasion of cacti in natural and semi-natural habitats in the Mediterranean region changes habitat structure and species composition. However, dense and extensive stands of cacti are restricted to few species (e.g. Opuntia ficus-indica).     

Epidermal and hypodermal characteristics in North American Cactoideae (Cactaceae)

Dermal and hypodermal anatomical features of 70 species representing 21 genera of North American Cactoideae were studied. Results show that all species examined have parallelocytic stomata and anticlinal wall surface varies from straight to undulate. Cuticle thickness is mostly narrow (1–10 μm) contrary to the general opinion that cuticle is thick in most cacti; however, few species such as Ariocarpus fissuratus and several species of Pachycereus show a distinctive thick cuticle. More than 80% of the species studied have a single-layered epidermis. Papillae occur in eight species belonging to four genera. Notable papillae are a feature shared by all members of Peniocereus subg. Peniocereus. Other species show a bullate surface produced by irregular patches of secondary epidermal cell divisions. Commonly, the hypodermis is composed of more than two cell layers with distinctive collenchymatous walls as reported in many South American species. Silica bodies, prismatic crystals, druses, sphaerocrystals, and tannins are the most common cellular inclusions that distinguish several genera and appear to have taxonomic value. However, a more thorough search in species of Cephalocereus, Coryphantha, Echinocereus, Mammillaria, Neobuxbaumia, Pilosocereus, and Turbinicarpus is needed to support the previous assertion. Electronic Publication     

Identification of self-incompatibility groups in<Emphasis Type="Italic"> Hatiora</Emphasis> and<Emphasis Type="Italic"> Schlumbergera</Emphasis> (Cactaceae)

The Cactaceae, a family of about 1,800 species of succulent perennials, contains numerous species that exhibit self-incompatibility (SI). The objective of the current study was to determine the number of incompatibility groups present among diploid (2n=2x=22) cultivars of the genera Schlumbergera Lem. (Christmas cacti) and Hatiora Britton & Rose (Easter cacti). Two partial diallel crosses were performed, one with 19 cultivars of Christmas cacti [= S. truncata (Haworth) Moran and S. × buckleyi (Buckley) Tjaden] and the other with 10 cultivars of Easter cacti [= H. gaertneri (Regel) Barthlott, H. rosea (Lagerheim) Barthlott, and H. × graeseri Barthlott ex D. Hunt]. The compatibility/incompatibility status of crosses was determined by percent fruit set and presence of seed in mature fruit. None of the cultivars set fruit when selfed or crossed with a cultivar in the same incompatibility group, but fruit set ranged from 35% to 100% following compatible crosses. For the Christmas cacti, eight intra-incompatible but reciprocally compatible groups were identified, with 13 of the 19 cultivars assigned to three incompatibility groups (68%). The ten cultivars of Easter cacti yielded nine intra-incompatible but reciprocally compatible groups, with two cultivars in one incompatibility group and the other eight cultivars each assigned to a unique group. One cultivar of Christmas cactus ('Abendroth 6') was incompatible when crossed as a male with cultivars in incompatibility group 2 but was compatible in reciprocal crosses, results that suggest that this cultivar is an S-allele homozygote. The crossing relationships are consistent with a one-locus, gametophytic SI system with multiple alleles. Allozyme locus Lap-1, shown previously to be linked with the S locus (recombination frequency 7%) in Schlumbergera, exhibited insufficient allelic diversity for determining the S genotypes of the 19 cultivars of Christmas cacti. Based on the number of incompatibility groups in each diallel, at least five S-alleles occur in the 19 Christmas cacti and the 10 Easter cacti.Publication 3337 of the Massachusetts Agricultural Experiment Station. This material is based on work supported in part by the Cooperative State Research, Extension, Education Service, United States Department of Agriculture, Massachusetts Agricultural Experiment Station, under Project No. 746