Reproductive biology and the process of domestication of the columnar cactus Stenocereus Stellatus in Central Mexico

Pollination biology, breeding system, and floral phenology of the columnar cactus Stenocereus stellatus were studied in wild, wild managed in situ and cultivated populations of central Mexico, in order to examine whether these aspects have been modified under domestication and whether they determine reproductive barriers between wild and manipulated individuals. Individuals of both wild and manipulated populations are self-incompatible, indicating that artificial selection has not modified the breeding system. Their pollination biology is also similar. Anthesis is mainly nocturnal, with a peak of nectar production between 0200 and 0400 when the stigma presents maximum turgidity. Nocturnal visitors are the effective pollinators. Nearly 75% of flowers exposed for nocturnal pollination set fruit, while none of the flowers exposed for diurnal pollination produced fruits. The bats Leptonycteris curasoae, L. nivalis, and Choeronycteris mexicana (Glossophaginae) are the most likely pollinators, and their time of foraging is synchronized with the time of nectar production and stigma receptivity in S. stellatus. Bats potentially move pollen over a considerable distance, so there is apparently no spatial isolation to prevent pollen exchange between wild and cultivated populations. Phenological studies showed that there are also no apparent temporal barriers. However, manual cross pollination failed between some domesticated and wild phenotypes, suggesting that gene flow between wild and cultivated populations might be limited by pollen incompatibility.     

Pollination system of the Pilosocereus leucocephalus columnar cactus (tribe Cereeae) in eastern Mexico

It has been suggested that there is a geographic dichotomy in the pollination systems of chiropterophilous columnar cacti: in intra‐tropical areas they are pollinated almost exclusively by bats, whereas in extratropical areas they are pollinated by bats, birds and bees. However, currently the studies are clumped both taxonomically (mainly Pachycereeae species) and geographically (mainly in the Tehuacan Valley and the Sonoran Desert). This clumping limits the possibility of generalising the pattern to other regions or cactus tribes. Only four of the 36 chiropterophilous cacti in Pilosocereus have been studied. Despite the tropical distribution of two Pilosocereus species, bees account for 40–100% of their fruit set. We examined how specialised is the pollination system of P. leucocephalus in eastern Mexico. As we studied tropical populations, we expected a bat‐specialised pollination system. However, previous studies of Pilosocereus suggest that a generalised pollination system is also possible. We found that this cactus is mainly bat‐pollinated (bats account for 33–65% of fruit set); although to a lesser degree, diurnal visitors also caused some fruit set (7–15%). Diurnal visitors were more effective in populations containing honeybee hives. P. leucocephalus is partially self‐compatible (14–18% of fructification) but unable to set fruit without visitors. Despite the variation in pollination system, P. leucocephalus shows more affinity with other columnar cacti from tropical regions than with those from extratropical regions. Although we report here that a new species of tropical Pilosocereus is relatively bat‐specialised, this Cereeae genus is more flexible in its pollination system than the Pachycereeae genera.     

Effect of habitat disturbance on pollination biology of the columnar cactus Stenocereus quevedonis at landscape‐level in central Mexico

Stenocereus quevedonis (‘pitire’) is a columnar cactus endemic to central Mexico, grown for its edible fruit. Phenology, pollination biology and behaviour of flower visitors of this species were compared in six conserved and disturbed sites, hypothesising that: (i) pitire pollination is self‐incompatible, requiring animal vectors; (ii) higher incidence of radiation on plants in cleared forest may lead to a higher number of flowers per pitire plant and longer blooming season, and disturbing and differential spatial availability of flower resources may determine differential attraction of pollinators to conserved and disturbed areas; (iii) if pitire pollination system is specialised, reproductive success would decrease with pollinator scarcity, or other species may substitute for main pollinators. In all sites, pitire reproduction started in January, flowering peak occurring in April, anthesis duration was 15 h and predominantly nocturnal (9 h), pollen was released at 23:00 h, nectar was produced throughout anthesis, and breeding system was self‐incompatible. Flower production per plant was similar in disturbed and conserved sites, but flower availability was higher (because of higher tree density) and longer in disturbed sites. Pollination is nocturnal, the most frequent legitimate pollinator being the bat Leptonycteris yerbabuenae; diurnal pollination is rare but possible, carried out by bee species. Fruit and seed set in control and nocturnal pollination treatments at disturbed sites were higher than in conserved sites. Frequency of L. yerbabuenae visits was similar among site types, but more visits of complementary nocturnal and diurnal pollinators were recorded in disturbed sites, which could explain differences in reproductive success.     

Morphological variation and the process of domestication of Stenocereus stellatus (Cactaceae) in Central Mexico

Morphological variation was analyzed in wild, managed in situ, and cultivated populations of the columnar cactus Stenocereus stellatus in central Mexico. The purpose was to evaluate whether morphological divergence between manipulated and wild populations has resulted from domestication processes. Variation of 23 morphological characters was analyzed among 324 individuals from 19 populations of the Tehuacán Valley and La Mixteca Baja. Multivariate statistical analyses were used to group individuals and populations according to their morphological similarity. Individuals grouped according to the way of management and fruit characteristics were the most relevant for grouping. Within each region, sweet fruits with pulp colors other than red were more frequent in cultivated populations, where fruits were also larger, contained more and bigger seeds, and had thinner peel and fewer spines than fruits from wild individuals. Phenotypes common in managed in situ and cultivated populations generally occur in the wild but in lower frequencies. Artificial selection has thus operated by enhancing and maintaining desirable rare phenotypes in managed in situ and cultivated populations, causing divergent patterns of morphological variation from wild populations. Cultivation has caused the strongest level of divergence, but divergence has also been significant with management of wild populations in situ.     

Diurnal and nocturnal pollination of Marginatocereus marginatus (Pachycereeae: Cactaceae) in Central Mexico

BACKGROUND AND AIMS: Chiropterophillous and ornithophillous characteristics can form part of a single reproductive strategy in plants that have flowers with diurnal and nocturnal anthesis. This broader pollination strategy can ensure seed set when pollinators are scarce or unpredictable. This appears to be true of hummingbirds, which presumably pollinate Marginatocereus marginatus, a columnar cactus with red nocturnal and diurnal flowers growing as part of dense bat-pollinated columnar cacti forests in arid regions of central Mexico. The aim of this study was to study the floral biology of M. marginatus, and evaluate the effectiveness of nocturnal vs. diurnal pollinators and the contribution of each pollinator group to overall plant fitness. METHODS: Individual flower buds were marked and followed to evaluate flower phenology and anthesis time. Flowers and nectar production were measured. An exclusion experiment was conducted to measure the relative contribution of nocturnal and diurnal pollinators to seed set. KEY RESULTS: Marginatocereus marginatus has red hermaphroditic flowers with nocturnal and diurnal anthesis. The plant cannot produce seeds by selfing and was pollinated during the day by hummingbirds and during the night by bats, demonstrating that both pollinator groups were important for plant reproduction. Strong pollen limitation was found in the absence of one of the pollinator guilds. CONCLUSIONS: Marginatocereus marginatus has an open pollination system in which both diurnal and nocturnal pollinators are needed to set seeds. This represents a fail-safe pollination system that can ensure both pollination, in a situation of low abundance of one of the pollinator groups (hummingbirds), and high competition for nocturnal pollinators with other columnar cacti that bloom synchronously with M. marginatus in the Tehuacan Valley, Mexico.     

Evolution of Polaskia chichipe (Cactaceae) under domestication in the Tehuacan Valley, central Mexico: reproductive biology

Polaskia chichipe, a columnar cactus, is cultivated for its edible fruits in central Mexico. This study analyzed whether artificial selection has modified its reproduction patterns and caused barriers to pollen exchange between wild, managed in situ, and cultivated populations. Anthesis was diurnal (～16 h in winter, ～10 h in spring) as well as partly nocturnal (～12 h in winter, ～3 h in spring), and flowers were pollinated by bees, hummingbirds, and hawk moths. Manual cross-pollination was ～37-49% effective in all populations. Self-pollination was ～12% successful in the wild, but twice as successful (～22-27%) in managed and cultivated populations. Diurnal pollination was ～35-55% effective in winter and 100% in spring. Nocturnal pollination was successful only in winter (15%). Crosses among individuals were more effective within populations than among populations, including populations under a similar management regimen. The least successful crosses were between wild and cultivated populations. Flowers were produced in all populations from January to March, but flowering peaks differed by 1 mo among wild, managed, and cultivated populations and by 2 mo between wild and cultivated populations. The latter interrupted pollen exchange in May. Seeds from managed and cultivated populations germinated faster than those from wild individuals. Domestication has seemingly favored self-compatible P. chichipe plants with higher fruit yield, a longer period of fruit production, and faster seed germination, attributes that have resulted in partial reproductive barriers between wild and manipulated populations.     

Pollination and Breeding System of two Co-Occurring Hirtella Species (Chrysobalanaceae) in Central Brazil

The phenology, major floral characteristics, breeding systems and fruiting success of two co-occurring species of Hirtella: H. glandulosa and H.gracilipes (Chrysobalanaceae) were studied in Central Brazil. The two species occur as trees in mesophyllous forests but H. glandulosa is frequent also in dense savanna areas. Both species flower at the end of the dry season (September) and have flowers with one-day longevity. In both species flowers produce nectar and are pollinated exclusively by butterflies. The number of visits recorded and pollination rate in each species were significantly different and indicate that pollinators prefer H. glandulosa flowers. Natural and controlled fruit sets were low in both species. The index of self-incompatibility (ISI) was 0.17 in H. glandulosa and 0.86 in H. gracilipes. Although ISI in H. glandulosa could denote an incompatibility system, the presence of fruits with aborted embryos at different degrees of development point to an inbreeding depression situation. Low pollen viability and fruit set in H. gracilipes suggested reproductive problems which may be linked to hybridization events.     

广东地区外来种五爪金龙的传粉生物学研究（英文）

五爪金龙(Ipomoea cairica)是一种匍匐或攀援生长的入侵杂草。2006-2007年作者在广东地区开展了对该入侵种的传粉生物学研究。五爪金龙在广东全年均可开花,花序为聚伞花序,花冠为紫色或蓝紫色喇叭状。开花时间在早晨4:30-5:20之间,下午17:40闭合,持续大约12h。每朵花平均分泌花蜜量为5.82±2.47μL(n=10),花蜜平均含糖量为41.03±6.52%,氨基酸含量为0.042mg/mL。开花过程中花粉活性和柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为4,778.18±264.58(n=10)。木蜂(Xylocopasp.)是其有效的传粉昆虫,蝇类和蝶类为随机访问者。开花过程中柱头与花药相接触,有利于自花授粉。然而,繁育系统的研究表明五爪金龙是自交不亲和植物,其种子仅能通过异花授粉获得。     

Reproductive biology and mating system estimates of two Andean melocacti, Melocactus schatzlii and M. andinus (Cactaceae)

BACKGROUND AND AIMS: The genus Melocactus comprises 36 species of globose cacti with the most derived traits in the Cereeae tribe. It is the proper study system to examine what are the most derived reproductive strategies within that tribe. This study aims to characterize the reproductive biology and to estimate the mating system parameters of two Andean melocacti, Melocactus schatzlii and M. andinus. METHODS: The reproductive attributes of the two species were described, including floral morphology, anthesis patterns, floral rewards, floral visitors and visitation patterns. Levels of self-compatibility and autonomous self-pollination were estimated by hand-pollination experiments. Mating system estimates were obtained by conducting progeny array analyses using isozymes. KEY RESULTS: The flowers of the two species present the typical hummingbird-pollination syndrome. Despite their morphological resemblance, the two species differ in flower size, pollen and ovule production and anthesis pattern. Their main pollinator agents are hummingbirds, four species in M. schatzlii and one species in M. andinus. Both cacti are self-compatible and capable of self-pollination without the aid of pollen vectors. Population-level outcrossing rate was higher for M. schatzlii (t(m)=0.9) than for M. andinus (t(m)=0.4). At the family level, outcrossing rates for most mothers of M. schatzlii were higher (t(m)>0.8) than for M. andinus (t(m)<0.5). CONCLUSIONS: Although the two cacti are capable of selfing, M. schatzlii is a predominantly outcrossing species, while M. andinus behaves as a mixed-mating cactus. Hummingbirds are the only pollinators responsible for outcrossing and gene flow events in these species. In their absence, both melocacti set seeds by selfing. Based on its low population size, restricted distribution in Venezuela, low rates of floral visits, and high levels of inbreeding, M. andinus is considered to be an endangered species deserving further study to define its conservation status.     

Pollination in Brazilian Syngonanthus (Eriocaulaceae) species: evidence for entomophily instead of anemophily

BACKGROUND AND AIMS: The reproductive biology of Syngonanthus mucugensis and S. curralensis (Eriocaulaceae) was studied in areas of 'campo rupestre' vegetation in the Chapada Diamantina, north-eastern Brazil. These species are herbaceous and the individuals have a grouped distribution. Their leaves are united in a rosette, and their inflorescence is monoecious, of the capitulum type. The staminate and pistillate rings mature in a centripetal manner on the capitulum. METHODS: A field study was conducted, including observations concerning the morphology and biology of the flowers, fruit development, insect visits and anemophily, in both S. mucugensis and S. curralensis. Experimental pollinations were also carried out to study the mating systems of S. mucugensis. KEY RESULTS: Both species flower from June to August. The staminate cycle lasts approx. 7 d, and the pistillate cycle from 3 to 4 d, with no temporal overlap between them on the same capitulum. The pollen viability of S. mucugensis was 88.6%, and 92.5% for S. curralensis. The inflorescences of both species demonstrated ultraviolet absorbance, and a sweet odour was detected during both the staminate and pistillate phases. No nectar production was ever noted, although nectaries are present. Both species were visited by numerous groups of insects, with the Diptera being the principal pollinators, especially the species of Syrphidae and Bombyliidae. There were secondary pollinators among species of Coleoptera and Hymenoptera. There was no evidence of wind pollination. Syngonanthus mucugensis is a self-compatible species, and forms fruits by agamospermy at low frequencies. CONCLUSIONS: This is apparently the first report for pollination biology and mating systems of Eriocaulaceae. Conversely to that stated by some authors, entomophily, mainly effected by species of Diptera but also by species of Coleoptera and Hymenoptera, is probably the only pollination system in these species. In spite of the monoecious inflorescences without overlap of the staminate and pistillate phases, geitonogamy may occur in S. mucugensis, as the species is self-compatible and different capitula in the same plant at different phases is common.     

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.     

Relative fitness of two hermaphroditic mating types in the androdioecious clam shrimp,Eulimnadia texana

Androdioecy (populations of males and hermaphrodites) is a rare reproductive form, being described from only a handful of plants and animals. One of these is the shrimp Eulimnadia texana, which has populations comprised of three mating types: two hermaphroditic types (monogenics and amphigenics) and males. In a recent study, the amphigenic hermaphrodites were found to be in greater abundance than that predicted from a model of this mating system. Herein, we compare the relative fitness of offspring from amphigenic and monogenic siblings, attempting to understand the greater relative abundance of the former. Populations started with offspring from selfed monogenic hermaphrodites had a net reproductive rate (R) 87% that of offspring from their amphigenic siblings. Additionally, within populations of amphigenic offspring (which included males, monogenics and amphigenics), amphigenics survived longer than monogenics. These differences help to explain the increased relative abundance of amphigenics in natural populations, but amphigenics continue to be more abundant than expected.     

Inbreeding depression in two species of Mimulus (Scrophulariaceae) with contrasting mating systems

We examined the effect of self- and cross-pollination on germination success, flowering probability, pollen and ovule production, survivorship, and adult aboveground biomass in two species of Mimulus with contrasting mating systems: the highly seifing M. micranthus and an outcrossing population of M. guttatus. Cross-pollinations were performed both within and between populations in order to examine the scale at which the genetic load is distributed. We found significant inbreeding depression in M. guttatus in four of the six traits, with the highest inbreeding depression observed in biomass (68% and 69% based on within- and between-population crosses, respectively) and lowest in ovule production (21% based on between-population crosses only). M. micranthus displayed significant inbreeding depression in only two of the six traits examined. Again, we observed the highest inbreeding depression in biomass (47–60% based on within- and between-population crosses, respectively), but both traits showing significant differences between self and outcross progeny expressed lower inbreeding depression than in M. guttatus. We detected no significant inbreeding depression for either pollen or ovule production in M. micranthus. An estimate of total inbreeding depression based on the multiplicative effects of all traits was also lower in M. micanthus than∗∗∗ in M. guttatus. Our results are consistent with the expected purging of genetic load in populations with high selfing rates. The absence of inbreeding depression in M. micranthus pollen and ovule production, two traits with strong links to fitness in a selfing annual, further suggests the important role of directional selection in determining the population's genetic load. Comparison of cross-pollinations made within and between populations revealed little evidence of divergence of genetic load among the M. micranthus and M. guttatus populations examined.     

Pollination biology and sexual differentiation ofOsyris alba (Santalaceae) in the Mediterranean region

Osyris alba L. is a widespread dioecious hemiparasitic shrub of S Europe, N Africa, and SW Asia. Male inflorescences are multiflowered whereas each female inflorescence is reduced to a single flower with persistent enlarged bracts. Pollination is a prerequisite for fruit and seed development and wind is unlikely to be an effective means of pollen spread. In southern Italy pollen is transported by small unspecialized flies and beetles. Both male and female flowers produce an indistinguishable sweet odour. Male flowers are produced in large numbers and over a larger period than the females and provide pollen, nectar, and staminal hairs as rewards for pollinators. The presence and function of staminal hairs with tip cells inOsyris alba has been reported for the first time. Female flowers are rewardless, producing neither mature pollen, nectar nor staminal hairs, but possess three modified yellow indehiscent anthers containing no viable pollen which may provide a strong visual feeding stimulus for pollinators. It is suggested that pollinators are attracted by deceit to female flowers by mimicry of the males and the floral mimicry is, therefore, intraspecific and intersexual. The floral characteristics and flowering phenology of male and female plants are consistent with this kind of mimicry. The female flower possesses a tricarpellary ovary with three ovules of which only one develops. The single seed, containing a small embryo and a large, rich endosperm, is borne in a red fleshy bird-dispersed fruit. The reduction in seed number per flower to one highly nutrient-invested seed, together with a reduction of the multiflowered inflorescence to a solitary flower and the sequential production of ripe fruits over an extended fruiting season, suggest that the female function is markedly resource-limited. It is suggested that, although all the reproductive characteristics present inOsyris alba, as well as hemiparasitism, had probably evolved before the end of the tropical Tertiary, they are of adaptive advantage in the nutrient and water-limited environment of the Mediterranean maquis.     

Crane flies and microlepidoptera also function as pollinators in Epidendrum (Orchidaceae: Laeliinae): the reproductive biology of E. avicula

Crane flies and microlepidoptera have been recorded as pollinators in unrelated orchid groups, but these insects have never been recorded in Epidendroideae, the most species‐rich orchid subfamily, which includes one of the most diverse genera among Orchidaceae, Epidendrum. Based on data on phenology, floral morpho‐anatomy, pollinators, pollination mechanisms and breeding system, the reproductive biology of E. avicula was studied in south‐eastern Brazil. Epidendrum avicula possess osmophores that produce a citric fragrance at night. The flowers attract Tipulidae flies and several families of microlepidoptera that drink the nectar produced in a tube formed by the adnation of the labellum and column. As is common in Epidendrum, after removing the pollinarium, both crane flies and micro‐moths get trapped by the proboscis, which frightens the insects and inhibits any possible intent to immediately visit another flower. The behavior of the pollinators on flowers, plus the retention of the anther cap by the pollinarium, results in a reduction in the occurrence of geitonogamy. Because E. avicula is self‐incompatible, the consequence of pollinator behavior and the floral mechanisms tend to reduce the pollen loss. As far as we know, this is the first study to report the reproductive biology of a species of Epidendroideae pollinated by crane flies and microlepidoptera. Based on more recent concepts of plant–pollinator interactions, although E. avicula is pollinated by several species belonging to two distinct orders, suggesting an unspecialized pollination system is involved, nectar‐seeking microlepidoptera and Tipulidae flies can be recognized as a single functional group.     

Comparative study of pollination biology of two closely related alpine Primula species,namely Primula beesiana and P.bulleyana (Primulaceae)

The comparative pollination biology of a population of Primula beesiana,a population of P.bulleyana,and an overlapping population of these two species in Yulong Shan,Yunnan Province,China,was studied in 2004 and 2005.The results indicate that both P.beesiana and P.bulleyana are typical heterostylous and obligate outcrossing species;the main pollinators of the two species were bees and butterflies.At the sites of all three populations,the main pollinating visitors of the two species showed preference for one...     

Reproductive biology of the melon cactus, Melocactus curvispinus (Cactaceae)

We examined the reproductive biology of Melocactus curvispinus Pfeiffer (Cereeae, Cactoideae) in xeric areas of northern Venezuela. Floral traits correspond to a classic hummingbird-pollination syndrome; however, pollination is shared between hummingbirds, Leucippus fallax (Bourcier 1843), and anthophorid bees, Ceratina sp. Reproduction occurs during most of the year. Anthesis and nectar secretion occur between noon and sunset. Average daily nectar production per flower was 163.1 l, nectar sugar concentration between 29.1 and 30.2% (w/w). Hummingbirds promoted inter-plant pollen movement and were relatively more reliable floral visitors than anthophorid bees, but these insects had a higher frequency of floral visits (28.75/day) than hummingbirds (4.96/day). M. curvispinus is self-compatible and autogamous. By combining extended reproductive activity, frequent animal-mediated pollination, and selfing capabilities, this cactus possesses a mating strategy that guarantees reproduction under variable environmental conditions. We argue that based on its reproductive biology, M. curvispinus should be considered an example of evolutionary transition towards selfing within tribe Cereeae.     

Geographic variation in the breeding system and the evolutionary stability of trioecy in Pachycereus pringlei (Cactaceae)

The Sonoran Desert columnar cactus Pachycereus pringlei has a geographically variable, non-hermaphroditic breeding system. It is trioecious (separate males, females and hermaphrodites) in the northern two-thirds of its range in Sonora, Mexico, and in the southern three-quarters of its range in Baja California, Mexico, and is gynodioecious (separate females and hermaphrodites) elsewhere. Trioecy occurs near known maternity roosts of its major pollinator, the nectar-feeding bat Leptonycteris curasoae; gynodioecy occurs>50km from known bat roosts. The observed geographic patterns cannot be explained by limited gene flow or by the geographic distributions of diurnal avian pollinators. Our field observations plus a theoretical analysis suggest that the abundance of chiropteran pollinators plays an important role in the maintenance of trioecy in this plant. Under pollinator limitation, trioecy can be a stable breeding system in this species. This revised version was published online in July 2006 with corrections to the Cover Date.