Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

Orchid conservation in China from 2000 to 2020: Achievements and perspectives

We review achievements in the conservation of orchid diversity in China over the last 21 years. We provide updated information on orchid biodiversity and suggestions for orchid conservation in China. We outline national policies of biodiversity conservation, especially of orchid conservation, which provide general guidelines for orchid conservation in China. There are now approximately 1708 known species of Orchidaceae in 181 genera in China, including five new genera and 365 new species described over the last 21 years. The assessment of risk of extinction of all 1502 known native orchid species in China in 2013 indicated that 653 species were identified as threatened, 132 species were treated as data-deficient, and four species endemic to China were classified as extinct. Approximately 1100 species (ca. 65%) are protected in national nature reserves, and another ~66 species in provincial nature reserves. About 800 native orchid species have living collections in major botanical gardens. The pollination biology of 74 native orchid species and the genetic diversity and spatial genetic structure of 29 orchid species have been investigated at a local scale and/or across species distributions. The mycorrhizal fungal community composition has been investigated in many genera, such as Bletilla, Coelogyne, Cymbidium, Cypripedium, and Dendrobium. Approximately 292 species will be included in the list of national key protected wild plants this year. Two major tasks for near future include in situ conservation and monitoring population dynamics of endangered species.     

四川黄龙沟草本层植物种类组成与数量特征

四川黄龙沟草本植物群落在所调查样方中(N=662)共出现维管植物124种,分属于37科91属,有54个中国特有种:其中兰科植物多达21属33种,中国特有种有12个,新种一个。如此众多的地生兰科植物聚集在面积不到1公里的沟内,而且部分兰科植物种类在沟内形成优势草本群落,这在中国地生兰的分布区域中是不多见的。大多数物种,包括兰科植物在内,出现的频率都较小,并且分布不均匀。黄龙沟兰科植物主要分布在两种生境中,即钙化滩流地和森林中。在这两种生境中微环境条件以及兰科植物的种类组成、数量特征和分布格局的差异都很大。钙化滩流地的兰科植物种类数目和每个样方中包含的兰科植物种类数目都比森林生境中的高。钙化滩流地中发现有30种兰科植物,最为常见的是无苞杓兰、黄花杓兰、西藏杓兰、广布小碟兰、二叶根茎兰和少花鹤顶兰。森林生境中有21种兰科植物分布,其中筒距兰和斑叶兰出现最多,光照强度可能对兰科植物的分布起到重要的决定作用。钙化滩流地中的溪流对建立和维持兰科植物生存所必须的稳定环境条件起到至关重要的作用,如果随意改变水流的方向或减少水流的流量,将给某些兰科植物带来灾难性的后果。     

Effects of floral display and plant abundance on fruit production of Ryncholaelia glauca (Orchidaceae)

Flowering plant density can increase number of visits and fruit set in multi-flowering plants, however this aspect has not been studied on few flower species. We studied the effects of individual floral display and plant density on the fruit production of the epiphytic, moth-pollinated orchid, Ryncholaelia glauca, in an oak forest of Chavarrillo, Veracruz, Mexico. Species is non-autogamous, and produced one flower per flowering shoot each flowering season. We hypothesized that orchids with more flowering shoots and those on trees with clumps of conspecific should develop more fruits than isolated ones. R. glauca population flowers synchronouly, and individual flowers last up to 18 days, with flowers closing rapidly after pollination. Individuals produced few flowers per year, although some plants developed flowers in both seasons and fewer of them developed fruits both years. There was no relationship between flower number per orchid, or per host tree, with the number of fruits developed per plant. Host trees with flowering and fruiting orchids were randomly dispersed and the pattern of distribution of flowering and fruiting plants was not related. Apparently, pollinators visit the flowers randomly, with no evidence of density dependence. The fruit set of R. glauca was as low as fruit set of multi-flowered orchids moth pollinated, suggesting that fruit set on moth pollinated orchids could be independent of the number of flowers displayed.     

Observations of putative pollinators of Hemipilia flabellata Bur.et Franch.(Orchidaceae) in north-west Yunnan Province, China

Interactions between Hemipilia flabellata and anthophilous insects were studied in northwestern Yunnan, China during June-July of 1996 and 1997. Twenty-seven species of insects were seen visiting the flowers of this species, although only male and female individuals of the bee Anthophora mangkamensis Wu were observed carrying the pollinaria. The frequency of visits by A.mangkamensis to this orchid was rather low. Pollinating bees normally sampled only the lowermost flowers in an inflorescence, staying for 1–3 seconds. This species is also characterized by low pollination, low fruit set (1–22%) and a very striking decrease in fruit production from bottom to top of the inflorescence. All of these indicate that the orchid may rely on deception to attract visitors. Ajuga forrestii Diels (Labiatae) appears to be the main or exclusive subsidiary nectar source for A.mangkamensis during the flowering period of H.flabellata . It appears that H.flabellata utilizes a food deceit pollination strategy for pollination. The orchid benefits from the great variation or flowers in colour and shape. Phenological, functional-morphometric and distribution data indicate that A.mangkamensis is an optimal pollinator of H.flabellala and thus the anthecological adaptation of the orchid to the pollen vector may be unique.     

Reproductive Biology of the Monoecious Understory Palm Prestoea schultzeana in Amazonian Ecuador1

Prestoea schultzeana is a monoecious, protandrous palm in the forest understory of Amazonian Ecuador. We studied its leaf production, population density, sexual expression, phenology, pollination, and the specificity of the floral visitors. On average, 1.4 leaves and 0.9 inflorescences are produced per individual per year. The number of staminate flowers per inflorescence is relatively constant compared with the number of pistillate flowers which varies greatly. Flowering occurs in staminate and pistillate phases of approximately 19 and 0–7 days duration, respectively. Flowers open in the morning, and staminate flowers abscise in the afternoon of the same day whereas pistillate flowers last for two days. Flowers are whitish-yellow with a sweet odor and produce nectar. They were visited by Coleoptera (Chrysomelidae, Curculionidae, Nitidulidae, Ptiliidae, Staphylinidae), Hemiptera, Diptera (Drosophilidae, Syrphidae, Ceratopogonidae), Lepidoptera (Nymphalidae), and Hymenoptera (Formicidae, Halictidae). All examined individuals of the syrphid fly Copestylum sp. visiting pistillate flowers carried 100–500 grains of P. schultzeana pollen. Pollen occurred on all body parts, but especially on the legs, and this makes Copestylum sp. the most important pollinator. Most floral visitors were also frequent on the flowers of co-occurring plant species; notably the palm Hyospathe elegans shared most visitor species with P. schultzeana.     

Functional Significance of Labellum Pattern Variation in a Sexually Deceptive Orchid (Ophrys heldreichii): Evidence of Individual Signature Learning Effects

Mimicking female insects to attract male pollinators is an important strategy in sexually deceptive orchids of the genus Ophrys, and some species possess flowers with conspicuous labellum patterns. The function of the variation of the patterns remains unresolved, with suggestions that these enhance pollinator communication. We investigated the possible function of the labellum pattern in Ophrys heldreichii, an orchid species in which the conspicuous and complex labellum pattern contrasts with a dark background. The orchid is pollinated exclusively by males of the solitary bee, Eucera berlandi. Comparisons of labellum patterns revealed that patterns within inflorescences are more similar than those of other conspecific plants. Field observations showed that the males approach at a great speed and directly land on flowers, but after an unsuccessful copulation attempt, bees hover close and visually scan the labellum pattern for up to a minute. Learning experiments conducted with honeybees as an accessible model of bee vision demonstrated that labellum patterns of different plants can be reliably learnt; in contrast, patterns of flowers from the same inflorescence could not be discriminated. These results support the hypothesis that variable labellum patterns in O. heldreichii are involved in flower-pollinator communication which would likely help these plants to avoid geitonogamy.     

Pollination and floral ecology of Arundina graminifolia (Orchidaceae) at the northern border of the species’ natural distribution

Arundina graminifolia is an early successional plant on Iriomote Island, the Ryukyus, Japan, where it is endangered. Populations flower for more than half a year, and many inflorescences bloom for one to several months. The nectarless gullet flowers, which open for up to six days, are self-compatible but cannot self-pollinate spontaneously; thus they rely on pollinating agents for capsule production. Field observations at two habitats identified at least six species of bees and wasps, primarily mate-seeking males of Megachile yaeyamaensis and Thyreus takaonis, as legitimate pollinators. Thus, this orchid is a pollinator generalist, probably owing to its long blooming period and simple flower morphology. Carpenter bees, which were previously reported to pollinate this orchid, frequently visited flowers but were too large to crawl into the labellum chamber and never pollinated the flowers. Extrafloral nectaries on inflorescences attracted approximately 40 insect taxa but were not involved with pollination. Fruit-set ratios at the population level varied spatiotemporally but were generally low (5.2–12.4 %), presumably owing to infrequent flower visits by mate-seeking pollinators and the lack of food rewards to pollinators.     

Geitonogamy in rewarding and unrewarding inflorescences: modelling pollen transfer on actual foraging sequences

Many orchid species are unusual in that they provide no nectar or pollen rewards for their pollinators. Absence of reward is expected to have a fundamental effect on pollinator visitation patterns. In particular the number of flowers visited per inflorescence is expected to be affected in both unrewarding and co-flowering rewarding species. We used arrays of artificial inflorescences, which could be either rewarding or unrewarding and were differentiated by their colour, to test how many flowers bumblebees visit in each type of inflorescence. The frequency of the two colours was varied, thus modelling the case where different frequencies of both an unrewarding and rewarding species were present in a patch. We found that bumblebees visited more flowers per rewarding inflorescence after they have experienced unrewarding or partially emptied rewarding inflorescences. We used these results to simulate pollen transfer and thus predict selfing rates on rewarding inflorescences. We found these increased when nectar depleted or when there was a greater proportion of unrewarding inflorescences in the patch. Conversely, we found that the number of flowers bumblebees visited on each unrewarding inflorescence did not significantly change through experiments. Selfing rates for unrewarding inflorescences were predicted to depend principally on the number of these inflorescences bumblebees visited rather than on the number of flowers they visit per inflorescence. This was because most visitors to orchids are supposed to be naive, and pollinators that commence foraging carrying no pollen will necessarily self any flower they pollinate on the first inflorescence they visit. Thus the average selfing rate is expected to increase as the sequence of inflorescences visited decreases in length.     

Diversity of inflorescences in the Boutelouinae subtribe (Poaceae: Chloridoideae: Cynodonteae)

The Boutelouinae subtribe is comprised of one monophyletic genus, Bouteloua, with 57 species inhabiting the semi-arid regions of the New World. The inflorescences show significant structural variations, which provides an interesting system to examine their morphological evolution and identify characters and processes that may help to understand the group systematics. The structure of inflorescences was studied in 25 species of Bouteloua. All the species covered under this study have truncated polytelic inflorescences. Structural variations in the inflorescence unit among species may be accounted for by: (1) symmetry of the inflorescence unit, (2) total number of long primary branches, (3) total number of spikelets per branch, (4) number of perfect flowers per spikelet, (5) number of rudimentary flowers, and (6) reproductive system. Homogenization and truncation processes account for the diversity of mature inflorescences that exists in Bouteloua. In this work, we discuss the systematic and taxonomic value of the inflorescence in the Boutelouinae subtribe.     

Flowering and fruiting phenology of Normanbya normanbyi (W. Hill) L. H. Bailey (Arecaceae), a palm endemic to the lowland tropical rainforest of north‐eastern Australia

Abstract Normanbya normanbyi (W. Hill) L. H. Bailey (Arecaceae) is a monoecious, arborescent palm with a very small distribution area within the Daintree rainforest in north‐eastern Australia. Our 2‐year study was focused on the reproductive phenology at the individual and population level. At the population level flowering peaked in the dry season, whereas fruiting was confined to the wet season. Each palm can bear up to three inflorescences/infructescences at the same time. Flowering of each inflorescence is separated from each other by a couple of weeks. A single inflorescence consists of about 1900 staminate and 800 pistillate flowers. The flowering of N. normanbyi is protandrous with a staminate phase lasting 40 days and a pistillate phase of approximately 2 weeks. Between both phases is a non‐flowering phase of about 9 days. Fruit ripening takes 21 weeks, with an average of about 280 ripe fruit per tree. Comparison of three study plots revealed a moderate synchrony of flowering and fruiting initiation in this species of palm. The male phase of flowering shows a higher degree of synchrony than the female phase at the population level. Seasonal regularity of flowering and fruiting peaks appears to be predictable. The general flowering and fruiting phenology of N. normanbyi follows a subannual pattern with a strong tendency towards a continual pattern.     

The pollination ecology of Grevillea beadleana McGillivray,an endangered shrub from northern New South Wales,Australia

Grevillea beadleana (Proteaceae) is an endangered species known from five populations in northern New South Wales, Australia. The reproductive ecology of G. beadleana was compared in two populations with a ten-fold difference in the number of plants. Grevillea beadleana was found to be self-compatible in both populations and an examination of pollen viability and stigma maturation revealed that the species is protandrous. Flowering within inflorescences is acropetallous. In the first season plants in the largest population produced approx. ten-fold more inflorescences than those in the smaller population and, although the number of flowers per inflorescence did not vary significantly between populations the first season, the larger population produced more fruit per inflorescence than the smaller population. However, fruit to flower ratios were less than 0.2 in both seasons and populations. In both populations the number of fruit was significantly greater at the proximal end of the inflorescence, where flowers open first, compared with medial and distal positions. Several bird species were observed visiting flowers, although few birds were recorded foraging at plants in the smaller population. Within both populations, birds tended to make more within- than between-plant visits. Self-compatibility, acropetally and proximal fruit-set, combined with the predominantly within-plant movement of honeyeaters, suggests inbreeding may be common within both populations of G. beadleana. Pollination and fruiting success are discussed for G. beadleana and breeding systems among rare and common taxa in Grevillea are reviewed.     

Bee pollination and evidence of substitutive nectary in <Emphasis Type="Italic">Anadenanthera colubrina</Emphasis> (Leguminosae-Mimosoideae)

Anadenanthera colubrina (Vell.) Brenan (Leguminosae-Mimosoideae) is a widely-distributed tree in seasonally dry tropical forests of South America that was classified previously as lacking nectaries. However, some studies have stated that its flowers produce nectar, while others analyzed the composition of unifloral honey produced from A. colubrina flowers, raising the question about nectar production in the species. We studied the pollination and reproductive biology of A. colubrina var. cebil (Griseb.) Altschul in a natural population in the Caatinga, northeastern Brazil. Reproductive phenology, sexual system, floral biology, resource, and pollinators were investigated. We analyzed the breeding system through controlled pollinations for addressing its dependence on pollen vectors for reproduction. Anadenanthera colubrina flowered in the dry season, flower heads are heteromorphic, with staminate flowers at the base and perfect flowers at the apex of the inflorescence, characterizing andromonoecy. Anthesis is diurnal. We observed small drops of nectar at the apex of the petals of some flowers per inflorescence. Together with observations on flower visitor behavior and histochemical tests, we propose that A. colubrina produces floral nectar at the apex of the corolla, characterizing a substitutive nectary (sensu Vogel). This is the first record of substitutive nectary in the Mimosoideae and the first record of andromonoecy in the genus. Bees were the main pollinators (higher frequency), although other insects such as wasps, butterflies, and small beetles were also observed collecting nectar and/or pollen. The species is self-incompatible, thus depending on insect pollen vectors, mainly bees, for reproduction.     

Inflorescence architecture affects pollinator behaviour and mating success in Spiranthes sinensis (Orchidaceae)

? Despite the wide inflorescence diversity among angiosperms, the effects of inflorescence architecture (three-dimensional flower arrangement) on pollinator behaviour and mating success have not been sufficiently studied in natural plant populations. ? Here, we investigated how inflorescence architecture affected inter- and intra-plant pollinator movements and consequent mating success in a field population of Spiranthes sinensis var. amoena (S. sinensis). In this species, the flowers are helically arranged around the stem, and the degree of twisting varies greatly among individuals. The large variation in inflorescence architecture in S. sinensis results from variation in a single structural parameter, the helical angle (the angular distance between neighbour-flower directions). ? The numbers of visits per inflorescence and successive probes per visit by leaf-cutting bees decreased with helical angle, indicating that individual flowers of tightly twisted inflorescences received less visitations. As expected from pollinator behaviour, pollinia removal and fruit set of individual flowers decreased with helical angle. Meanwhile, geitonogamy decreased in tightly twisted inflorescences. ? Our novel findings demonstrate that natural variation in inflorescence architecture significantly affects pollinator behaviour and reproductive success, suggesting that inflorescence architecture can evolve under pollinator-mediated natural selection in plant populations. We also discuss how diverse inflorescence architectures may have been maintained in S. sinensis populations.     

A new species of Govenia (Orchidaceae) from Chiapas, Mexico

Govenia rubellilabia, a new terrestrial orchid from Sierra Madre de Chiapas, Mexico, is described and illustrated. Diagnostic characters of this new species are compared with other species ofGovenia. Govenia rubellilabia is similar in appearance toG. matudae, but it can be distinguished from this species by a shorter inflorescence with fewer flowers, green-yellowish tepals, a reddish lip lacking spots in the margin of the apex, and by a shorter anther beak.     

Direct and indirect selection on floral pigmentation by pollinators and seed predators in a color polymorphic South African shrub

The coexistence of different color morphs is often attributed to variable selection pressures across space, time, morph frequencies, or selection agents, but the routes by which each morph is favored are rarely identified. In this study we investigated factors that influence floral color polymorphisms on a local scale in Protea, within which approximately 40% of species are polymorphic. Previous work shows that seed predators and reproductive differences likely contribute to maintaining polymorphism in four Protea species. We explored whether selection acts directly or indirectly on floral color in two populations of Protea aurea, using path analysis of pollinator behavior, nectar production, seed predation, color, morphology, and maternal fecundity fitness components. We found that avian pollinators spent more time on white morphs, likely due to nectar differences, but that this had no apparent consequences for fecundity. Instead, the number of flowers per inflorescence underpinned many of the reproductively important differences between color morphs. White morphs had more flowers per inflorescence, which itself was positively correlated with nectar production, seed predator occurrence, and total long-term seed production. The number of seeds per plant to survive predation, in contrast, was not directly associated with color or any other floral trait. Thus, although color differences may be associated with conflicting selection pressures, the selection appears to be associated with the number of flowers per inflorescence and its unmeasured correlates, rather than with inflorescence color itself.     

Floral display size in comfrey, Symphytum officinale L. (Boraginaceae): relationships with visitation by three bumblebee species and subsequent seed set

The fecundity of insect-pollinated plants may not be linearly related to the number of flowers produced, since floral display will influence pollinator foraging patterns. We may expect more visits to plants with more flowers, but do these large plants receive more or fewer visits per flower than small plants? Do all pollinator species respond in the same way? We would also expect foragers to move less between plants when the number of flowers per plant are large, which may reduce cross-pollination compared to plants with few flowers. We examine the relationships between numbers of inflorescence per plant, bumblebee foraging behaviour and seed set in comfrey, Symphytum officinale, a self-incompatible perennial herb. Bumblebee species differed in their response to the size of floral display. More individuals of Bombus pratorum and the nectar-robbing B.?terrestris were attracted to plants with larger floral displays, but B. pascuorum exhibited no increase in recruitment according to display size. Once attracted, all bee species visited more inflorescences per plant on plants with more inflorescences. Overall the visitation rate per inflorescence and seed set per flower was independent of the number of inflorescences per plant. Variation in seed set was not explained by the numbers of bumblebees attracted or by the number of inflorescences they visited for any bee species. However, the mean seed set per flower (1.18) was far below the maximum possible (4 per flower). We suggest that in this system seed set is not limited by pollination but by other factors, possibly nutritional resources.     

Surprisingly high orchid diversity in travertine and forest areas in the Huanglong valley, China, and implications for conservation

The presence of such a large number of terrestrial orchid species in a small area (ca. 1 km²) of the Huanglong valley in southwestern China is uncommon for this country. Studying the relationship between the distribution patterns of these orchid species and their microenvironments may help us understand this uncommon phenomenon. We established 662 1 m × 1 m plots, measured the cover of each species and found that there were 33 orchid species distributed mainly in two different habitats, i.e. travertine areas and forest. In the travertine areas, 30 orchid species were found; the six most common ones being Cypripedium bardolphianum, Cypripedium flavum, Cypripedium tibeticum, Galearis diantha, Ponerorchis chusua and Phaius delavayi. However in the forested habitat, we found 21 orchid species; the most common ones being Tipularia szechuanica and Goodyera repens. Travertine areas had a higher number of orchid species as well as higher numbers of orchid species per plot as compared to forest. Light availability seems critical to the performance and distribution of orchid species. Stream flow through the travertine area during the orchids growing season appears to be an important factor in shaping and maintaining stable microenvironments favorable to the growth and reproduction of orchids. The results presented in this study suggest that some orchid species in the travertine area might be threatened if the travertine stream flows were to change or be disrupted.     

Density-dependent variation in reproductive success in a terrestrial orchid

 Within population variation in plant density can affect reproductive success and breeding systems. We examined such effects in Listera cordata R. Br., a north temperate terrestrial orchid, in a subalpine population in Colorado, USA. Hand pollinations showed that L. cordata was self-compatible as all pollinations produced fruits. Cross-pollinations differed significantly from self-pollinations, and had a higher number of seeds per capsule and higher % of fertilized ovules. Together they could compensate for the transmission advantage of self-pollinations. Average number of flowers per inflorescence was 4.2–4.4 and average fruit set was 20.7–23.7%. By establishing 36 plots with varying plant densities (1, 4, 8 and 16) and emasculating all flowers in target plants of a set of 20 plots, we estimated density effects on fruit and seed production. Plant density had no effect on the amount of reproduction except in reduced seed production of highly dispersed plants. Our results suggest that inbreeding depression may be an important selective factor in L. cordata. Further studies are needed to define its role in the evolution of mating systems in this species. Received February 21, 2000 Accepted December 26, 2000