Mating system of Helianthemum squamatum (Cistaceae), a gypsophile specialist of semi-arid Mediterranean environments

Aragán C. F. and Escudero A. 2008. Mating system of Helianthemum squamatum (Cistaceae), a gypsophile specialist of semi-arid Mediterranean environments. Bot. Helv. 118: 129–137 Plants living on gypsum have received remarkably little study, despite being one of the most important groups of endemic and rare plants in arid and semi-arid regions. This work is among the first to study the mating system of a gypsophyte, Helianthemum squamatum (Cistaceae), which is a short-lived perennial endemic to gypsum habitats of the Iberian Peninsula. We carried out a hand-pollination experiment, considering the following pollination treatments: autogamy, geitonogamy, xenogamy and control (open-pollination). Generalized linear mixed models were fitted to evaluate the effect of the treatment on the fruit set and the number of viable seeds yielded per fruit. Both response variables were significantly higher in the xenogamy treatment than in the autogamy and geitonogamy treatments. No significant differences were detected between the xenogamy treatment and the control group, indicating that H. squamatum is mainly an outcrosser although partially self-compatible. Our results also suggest that pollen limitation may only slightly affect the reproductive success of H. squamatum. In contrast with the general tendency towards self-compatibility previously described in short-lived species of the Cistaceae, this outcrossing mating system resembles the one of long-lived perennials Submitted 10 April 2008; Accepted 25 August 2008 Subject editor: Christian Parisod     

Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation

 The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide distribution in Europe. It is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in E. helleborine, we sampled 13 populations from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden. In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations. Received November 6, 2001; accepted April 27, 2002 Published online: December 3, 2002     

Entomophily of the cloudberry (Rubus chamaemorus)

Cloudberry (Rubus chamaemorus L.), a dioecious perennial plant of boreal circumpolar distribution, is greatly prized for its berries. We crossed two treatments, pollinator exclusion and supplementary hand-pollination, to determine i) the relative importance of insects as pollinators, ii) if pollinator activity was a limiting factor for the sexual reproduction of the plant, and iii) the relative contribution of diurnal vs. nocturnal visitors to pollination. The activity of natural pollinators resulted in 97.5 and 88.5% fruit set, along with 76.7 and 62.5% seed set in 1998 and 1999, respectively. When insects were excluded, fruit-set dropped significantly to 18.4 (1998) and 12.8% (1999) and seed-set to 5.4 (1998) and 5.0% (1999) showing the importance of mid- and large-sized insects as pollinators. Natural levels of insect activity were sufficient to ensure complete pollination in both years as supplementary hand-pollination did not significantly increase either parameter in plots where pollinators had free access. Nocturnal insects may serve as pollinators (fruit-set = 41%), although they were less effective than diurnal pollinators (fruit-set = 93%).     

Reproductive biology of the invasive exotic shrub, Rhododendron ponticum L. (Ericaceae)

The reproductive biology of an exotic species will affect its ability to become naturalized and invasive in non-native habitats. Rhododendron ponticum is an ecologically damaging exotic weed in the British Isles, which spreads predominantly by seed. I investigated how inbreeding and outcrossing affect seed production and germination in a wild population of this species in Ireland. Experimental manipulations revealed low fruit and seed set when insects were excluded from flowers, suggesting that this species has limited capability for spontaneous autogamy. Hand-pollination treatments showed that, although flowers are self-compatible (with self and same plant pollen), higher levels of seed set occur following outcrossing (xenogamy). There was no significant difference in rate of germination of seeds from inbred or outcrossed treatments. The addition of xenogamous pollen to open flowers did not increase fruit or seed set, suggesting that flowers in this population are not pollen limited: native generalist pollinators, mainly bumblebees ( Bombus spp.), are providing an adequate pollinator service. This work demonstrates that outcrossing increases seed set and pollinators are required to facilitate this. Hence, generalist native pollinators can promote invasion by exotic plants. Native pollinators can clearly play an important part in alien species invasion.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 373–381.     

Differential costs of reproduction in females and hermaphrodites in a gynodioecious plant

Background and Aims

Plants exhibit a variety of reproductive systems where unisexual (females or males) morphs coexist with hermaphrodites. The maintenance of dimorphic and polymorphic reproductive systems may be problematic. For example, to coexist with hermaphrodites the females of gynodioecious species have to compensate for the lack of male function. In our study species, Geranium sylvaticum, a perennial gynodioecious herb, the relative seed fitness advantage of females varies significantly between years within populations as well as among populations. Differences in reproductive investment between females and hermaphrodites may lead to differences in future survival, growth and reproductive success, i.e. to differential costs of reproduction. Since females of this species produce more seeds, higher costs of reproduction in females than in hermaphrodites were expected. Due to the higher costs of reproduction, the yearly variation in reproductive output of females might be more pronounced than that of hermaphrodites.

Methods

Using supplemental hand-pollination of females and hermaphrodites of G. sylvaticum we examined if increased reproductive output leads to differential costs of reproduction in terms of survival, probability of flowering, and seed production in the following year.

Key Results

Experimentally increased reproductive output had differential effects on the reproduction of females and hermaphrodites. In hermaphrodites, the probability of flowering decreased significantly in the following year, whereas in females the costs were expressed in terms of decreased future seed production.

Conclusions

When combining the probability of flowering and seed production per plant to estimate the multiplicative change in fitness, female plants showed a 56 % and hermaphrodites showed a 39 % decrease in fitness due to experimentally increased reproduction. Therefore, in total, female plants seem to be more sensitive to the cost of reproduction in terms of seed fitness than hermaphrodites.     

Pollen and resource limitations to lifetime seed production in a wild population of the endangered plant Disanthus cercidifolius var. longipes H. T. Chang (Hamamelidaceae)

Disanthus cercidifolius Maxim. var. longipes H. T. Chang, a plant species that only occurs in a few counties in Hunan, Jiangxi and Zhejiang Provinces and with a relatively small number of individuals, is ranked as a second Class endangered species for conservation in China. We have studied the effect of pollen and resources available to female reproduction, and the reproductive mechanism of “excess flowers with low fruit set” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang was discussed. Results are as follows Pollen from different sources has significant effects on fruit set and seed set of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. The pollen source rather than pollen numbers significantly affected reproduction of this species. In wild populations, producing one fruit needs about 54.8 flowers, and one satiation seed needs about 6.60 flowers or 83.19 ovules. After fertilizing, which was propitious to flower development, the abortion rate of flower buds was decreasing, but the flowering rate was increasing. The fruit set and seed set was also significantly increasing, while abortion rate of fruit was significantly decreasing. With the increasing percentages of cutting leaves, the fruit set decreased, but the abortion rate of fruit shows no significant differentiation among treatments. After cutting branches that were puny, broken and insectin-fested branches, the flower number seemed to be decreasing, but the fruit set and seed set all increased significantly. After removing some flowers, the fruit set was calculated with respect to the number of flowers remaining after the treatment increased with increasing of percentages of flower removal, whereas fruit set calculated with respect to the initial number of flowers remained constant, and the mean weights of per fruit and per seed all decreased significantly. Sufficient spatial or temporal heterogeneities in nutrient levels might allow limitation of seed set by resources and pollen in a natural population, while supplying resources may indirectly affect pollination by increasing attraction of the flowers to pollinators. There were very low fruit and seed sets in natural populations of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. Different factors may have interacted to effect a low fruit set. A joint adoption of the “selection abortion hypothesis”, “ovary reserve hypothesis” and “male function hypothesis” seems to be the most likely explanation for the reproductive strategy of “excess flowers with few fruit sets” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang. __________ Translated from Acta Ecologica Sinica, 2006, 26(2): 496–502 [译自: 生态学报]