Latitudinal gradients in biotic interactions: Are cacti pollination systems more specialized in the tropics?

Biotic interactions are said to be more specialized in the tropics, and this was also proposed for the pollination systems of columnar cacti from North America. However, this has not yet been tested for a wider set of cactus species. Here, we use the available information about pollination in the Cactaceae to explore the geographic patterns of this mutualistic interaction, and test if there is a latitudinal gradient in its degree of specialization. We performed a bibliographic search of all publications on the pollination of cacti species and summarized the information to build a database. We used generalized linear models to evaluate if the degree of specialization in cacti pollination systems is affected by latitude, using two different measures: the number of pollinator guilds (functional specialization) and the number of pollinator species (ecological specialization). Our database contained information about the pollination of 148 species. The most frequent pollinator guilds were bees, birds, moths and bats. There was no apparent effect of latitude on the number of guilds that pollinate a cactus species. However, latitude had a small but significant effect on the number of pollinator species that service a given cactus species. Bees are found as pollinators of most cactus species, along a wide latitudinal gradient. Bat and bird pollination is more common in the tropics than in the extra-tropics. The available information suggests that cacti pollination systems are slightly more ecologically specialized in the tropics, but it does not support any trend with regard to functional specialization.

Do cacti form soil seed banks? An evaluation using species from the Southern Central Andes.

There is controversy over whether cactus species form soil seed banks. Although it is commonly assumed that cacti do not form seed banks, very few studies have evaluated them. In this work, we analysed whether cactus species form soil seed banks, studying seed distribution, seed density and seed longevity in the Southern Central Andes. Soil samples were collected in two microhabitats (under nurse plants and in bare areas) at 12 selected sites. We determined seed presence-absence, density and distribution for 32 native cactus species. Seed longevity for six of these species was determined through a burial experiment. We recorded viable seeds for 62.5% of the 32 evaluated species, finding variation in seed density between microenvironments and among populations. In some species, the highest seed density was found under potential nurse plants. Seed germination and seed viability decreased with burial time, with seed longevity always being <24 months after burial. Our results show strong evidence that cactus species do form seed banks. Seed density can vary between microenvironments and among populations, suggesting that cactus-nurse plant associations can also be explained by differential seed dispersal and not only by differential establishment. We found that Echinopsis and Gymnocalycium species form short-term seed banks. Our results will help to better understand the population dynamics of cactus species, a focal species group for conservation actions because many of them are threatened by human activities.

Alternating temperature combined with darkness resets base temperature for germination (Tb ) in photoblastic seeds of Lippia and Aloysia (Verbenaceae).

Thermal time models for seed germination assume a continuum of rate responses in the sub-optimal temperature range. Generally, the models describe germination performance in non-dormant seeds at constant temperatures, yet alternating temperature (AT) is a feature of many natural environments. We studied the possible interacting effects of AT on germination progress in photoblastic seeds of three aromatic-medicinal Verbenaceae species in the genera Lippia and Aloysia. For Lippia turbinata f. turbinata and L. turbinata f. magnifolia seed, germination only occurred in light conditions, while for L. integrifolia and Aloysia citriodora it was significantly higher in the light than in darkness. Although relative light germination (RLG) was not different between constant and AT in the sub-optimal range, AT raised the base temperature for germination progress (Tb ) from ca. 3-6 °C in constant temperature to 7-12 °C in AT. Among the species, thermal time for 50% seed germination [θT(50) ] was 55-100 °Cd at constant temperature. Although AT resulted in slight modifications to θT(50) , the germination rate at comparable average temperatures in the sub-optimal range was slower than under constant temperatures. For all species, the proportion of germinated seeds was similar for constant and AT. Our results suggest that an interaction between cool temperature and darkness during AT treatment limits the temperature range permissive for germination in these positively photoblastic seed, reflecting both close adaptation to the natural ecology and niche requirements of the species.

How specialised is bird pollination in the Cactaceae?

Many cactus species produce 'bird' flowers; however, the reproductive biology of the majority of these species has not been studied. Here, we report on a study of the pollination of two species from the Cleistocactus genus, cited as an ornithophilous genus, in the context of the different ways in which they are specialised to bird pollination. In addition, we re-evaluate the level of specialisation of previous studies of cacti with bird pollination and evaluate how common phenotypic specialisation to birds is in this family. Both Cleistocactus species exhibited ornithophilous floral traits. Cleistocactus baumannii was pollinated by hummingbirds, whereas Cleistocactus smaragdiflorus was pollinated by hummingbirds and bees. Pollination by birds has been recorded in 27 cactus species, many of which exhibit ornithophilous traits; however, they show generalised pollination systems with bees, bats or moths in addition to birds being their floral visitors. Of all cactus species, 27% have reddish flowers. This trait is associated with diurnal anthesis and a tubular shape. Phenotypic specialisation to bird pollination is recognised in many cactus species; however, it is not predictive of functional and ecological specialisation in this family.

[Seeds germination of Caesalpinia paraguariensis (Fabaceae): scarificator agents and cattle effects]. / Germinación de semillas de Caesalpinia paraguariensis (Fabaceae): agentes escarificadores y efecto del ganado.

The tree Caesalpinia paraguariensis grows in the Chaco region, Argentina. Fruits are indehiscent with many seeds. This species is an important source of wood and the fruits are consumed by cattle in Salta province. We studied seed germination under chemical, mechanical and biological scarification. Seeds from controls (without scarification) and those with biological scarification had a smaller (and similar) germination rate. The non-germinated seeds from biological treatments were mechanically scarified and their germination rate was similar to others under the mechanical treatment. Passage by digestive tracts would not enhance germination because viable seeds are still dormant due to their hard coats.

[Floral biology and pollinators of Trichocereus pasacana (Cactaceae) in Parque Nacional Los Cardones, Argentina]. / Biología floral y polinizadores de Trichocereus pasacana (Cactaceae) en el Parque Nacional Los Cardones, Argentina.

Many columnar cacti are bat pollinated. It has been suggested that this kind of pollination would be more important in tropical than in temperate regions where flowers are open only one night. Thrichocereus pasacana produces big and resistant white flowers. We analyzed flower characteristics, floral cycle, stigmatic receptivity, nectar production, pollen presence and floral visitors in a T. pasacana population at National Park Los Cardones (Salta, Argentina) in November 1997. Flower features were constant between individuals of the population. Flowers start opening at evening and anthesis time is from 18 to 40 hs. The estigma was receptive throughout the floral cycle. Anther dehiscence occurs with flower opening. Nectar production was highest between 18 to 24 hs. Although T. pasacana are open during the night, floral visitors are diurnal. The most frequent was Xylocopa sp. In the study area, nectarivorous bats were not detected. The morphological features of T. pasacana flowers were similar but bigger compared to other columnar cacti. Anthesis time was also longer while nectar production was lower. T. pasacana pollination at National Park Los Cardones is done by bees.

Pollen:ovule ratio and its relationship with other floral traits in Papilionoideae (Leguminosae): an evaluation with Argentine species.

The pollen:ovule ratio (P/O) has traditionally been used as a rough estimator of plant breeding systems. It has been shown that plant breeding systems are associated with particular floral traits. In this study, we determined the P/O in 21 Leguminosae species from Argentina and explored relationships between P/O and taxonomic position, flower size, floral rewards, pollen presentation and pollination mechanisms. According to the results, 15 out of the 21 species classified were obligate xenogamous, although some of them have been recorded as facultative xenogamous in previous studies. There was a significant effect of taxonomic position (genus), reward type and pollination mechanism on P/O. Species offering only nectar as a floral reward (which were species with a brush mechanism) had a significantly lower P/O than species offering pollen or pollen and nectar. Species with the brush pollination mechanism had the lowest P/O, while species with valvular and pump mechanism had the highest P/O. However, pollen presentation (primary and secondary) and flower size did not have a significant effect on P/O. Our results demonstrate that P/O variability is determined by taxonomic position and pollination mechanism in this plant group.

Reproductive biology of Echinopsis terscheckii (Cactaceae): the role of nocturnal and diurnal pollinators.

The aim of this study was to analyse the reproductive biology of Echinopsis terscheckii, a species endemic to northwest Argentina that has nocturnal flowers. We expected that this species had a generalised pollination system, with moths and diurnal visitors as the primary pollinators. To test this, we studied the floral biology, breeding system and floral visitors of this species and the effectiveness of nocturnal and diurnal visitors. Floral biology was defined based on floral morphology, floral cycle and nectar production of the flowers. The breeding system and relative contributions of diurnal and nocturnal visitors to fruit and seed set were analysed through field experiments. E. terscheckii flowers opened at sunset and closed the following day. The peak of nectar production occurred at midnight. Flowers were determined to be self-incompatible. Moths, bees and birds were identified as floral visitors. Moths were the most frequent visitors at night, whereas bees were the most frequent visitors during the day. Fruit production by diurnal pollinators was less than that by nocturnal pollinators; among all floral visitors, moths were the most effective pollinators. We have demonstrated for the first time that moths are the primary pollinators of columnar cacti of the genus Echinopsis. Our results suggest that moths might be important pollinators of columnar cactus species with nocturnal flowers in the extra-tropical deserts of South America.