More than symbioses : orchid ecology ; with examples from the Sydney Region

The Orchidaceae are one of the largest and most diverse families of flowering plants. Orchids grow as terrestrial, lithophytic, epiphytic or climbing herbs but most orchids native to the Sydney Region can be placed in one of two categories. The first consists of terrestrial, deciduous plants that live in fire-prone environments, die back seasonally to dormant underground root tubers, possess exclusively subterranean roots, which die off as the plants become dormant, and belong to the subfamily Orchidoideae. The second consists of epiphytic or lithophytic, evergreen plants that live in fire-free environments, either lack specialised storage structures or possess succulent stems or leaves that are unprotected from fire, possess aerial roots that grow over the surface of, or free of, the substrate, and which do not die off seasonally, and belong to the subfamily Epidendroideae. Orchid seeds are numerous and tiny, lacking cotyledons and endosperm and containing minimal nutrient reserves. Although the seeds of some species can commence germination on their own, all rely on infection by mycorrhizal fungi, which may be species-specific, to grow beyond the earliest stages of development. Many epidendroid orchids are viable from an early stage without their mycorrhizal fungi but most orchidoid orchids rely, at least to some extent, on their mycorrhizal fungi throughout their lives. Some are completely parasitic on their fungi and have lost the ability to photosynthesize. Some orchids parasitize highly pathogenic mycorrhizal fungi and are thus indirectly parasitic on other plants. Most orchids have specialised relationships with pollinating animals, with many species each pollinated by only one species of insect. Deceptive pollination systems, in which the plants provide no tangible reward to their pollinators, are common in the Orchidaceae. The most common form of deceit is food mimicry, while at least a few taxa mimic insect brood sites. At least six lineages of Australian orchids have independently evolved sexual deception. In this syndrome, a flower mimics the female of the pollinating insect species. Male insects are attracted to the flower and attempt to mate with it, and pollinate it in the process. Little is known of most aspects of the population ecology of orchids native to the Sydney Region, especially their responses to fire. Such knowledge would be very useful in informing decisions in wildlife management

Further advances in orchid mycorrhizal research

Orchid mycorrhizas are mutualistic interactions between fungi and members of the Orchidaceae, the world’s largest plant family. The majority of the world’s orchids are photosynthetic, a small number of species are myco-heterotrophic throughout their lifetime, and recent research indicates a third mode (mixotrophy) whereby green orchids supplement their photosynthetically fixed carbon with carbon derived from their mycorrhizal fungus. Molecular identification studies of orchid-associated fungi indicate a wide range of fungi might be orchid mycobionts, show common fungal taxa across the globe and support the view that some orchids have specific fungal interactions. Confirmation of mycorrhizal status requires isolation of the fungi and restoration of functional mycorrhizas. New methods may now be used to store orchid-associated fungi and store and germinate seed, leading to more efficient culture of orchid species. However, many orchid mycorrhizas must be synthesised before conservation of these associations can be attempted in the field. Further gene expression studies of orchid mycorrhizas are needed to better understand the establishment and maintenance of the interaction. These data will add to efforts to conserve this diverse and valuable association

Pemanfaatan Anggrek Sebagai Bahan Obat Tradisional Pada Etnis Batak Sumatera Utara [Utilitation of Orchids as Medicinal Plants by Ethnic Batak of North Sumatra]

Sumatra has rich diversity of orchids. The local communities in Sumatra have been used orchids as a ornamental plant, food, and medicine. Research on utilitation of orchids as medicinal plants by ethnic Batak of North Sumatra was conducted using ethnobotanical methods. The objectives of the research was to know species of orchids that were used as medicinal plants by Batak ethnic in North Sumatra. Respond-ents consisted of traditional medicine plants traders in the traditional markets and traditional healers. We found as many as seven species of 6 genera of orchids have been used as traditional medicine. Those orchids used as medicine for fever, aphrodisiac, maintain stamina, respira-tory disorders, and gastrointestinal disorders

The effect of population structure, plant size, herbivory and reproductive potential on effective population size in the temperate epiphytic orchid, Sarcochilus australis

Distribution of plant size and reproductive success is investigated in the temperate epiphytic orchid Sarcochilus australis (Lindl.) Rchb. f. at Kinglake National Park, Victoria, in south-eastern Australia, and applied to estimating the effective population size. Plant size distribution (leaf number, length of longest leaf and number of flowers) was not normally distributed. Most individuals were vegetative and it is estimated that more than half of all individuals are too small to flower, however exceptionally large individuals even though rare are able to have more than one active inflorescence. Flowering probability is plant size dependent and follows a sigmoid curve. The minimum observed leaf size of a flowering individual was 26 mm, however these small individuals have a low probability of flowering ( 80 mm) have a much higher probability of flowering (90%). The effective population size (Ne) of the Kinglake population of Sarcochilus australis was estimated from the distribution of flower production, and shown to be small (Ne = 10–19%) and comparatively similar to some of the other published estimates of effective populations size in orchids. From this basic survey of size distribution in Sarcochilus australis it is predicted that genetic diversity is low

Sam Van Aken: New Edens

Hybridized fruit trees, grafted orchids on shiny, reflective aluminum pedestals, fluorescent lights placed vertically on stands, and sheets of silver Mylar create a lush and somewhat disorienting space in contemporary artist Sam Van Aken’s most recent body of work New Edens. Van Aken makes Gettysburg College’s Schmucker Art Gallery into a kind of fantastical and futuristic winter garden. Without daylight and despite the cool fall weather of the Northeast, the dozen trees in the gallery are leafy and green, some even bearing fruit. Peach, plum, cherry, nectarine and apricot branches emerge from a single trunk and grow productively alongside their sister fruits. These surprising new plants, carefully designed and created by the artist, are titled Trees of 40 Fruits, and as time passes the artist will continue to graft more branches of various kinds of fruits onto each “parent” rootstock until he has reached forty. The saplings on display are relatively small, but eventually these trees will reach an approximate height of twenty feet. Van Aken created a nursery as part of his studio in Syracuse, New York. As an artist-cum-horticulturalist, he, like a nurturing parent, cares for his grafted fruit trees with a steadfast devotion. In his studio Van Aken carefully concocts the best fertilizers, waters carefully and diligently, removes hoards of Japanese beetles from the leaves one-by-one, and provides adequate warmth and protection for the young trees (with huge mounds of mulch and careful wrappings) during harsh New York winters. [excerpt]https://cupola.gettysburg.edu/artcatalogs/1007/thumbnail.jp

The role of pollinator attracting scent in the sexually deceptive orchids Ophrys chestermanii, O. normanii and O. tenthredinifera

Sexual deception of male bees is one of the most remarkable mechanisms of pollination (Ackermann 1986, Proctor & al. 1996). Flowers of the orchid genus Ophrys mimic females of their pollinator species, usually bees and wasps, to attract males, which try to copulate with the flowers. During this so-called “pseudocopulation” the male removes the pollinia and transfers them to another flower to ensure pollination. Apart from visual and tactile cues, floral scent was shown to be most important for eliciting mating behaviour in males (Kullenberg 1961, Schiestl & al. 1999, Ayasse & al. 2003). Pollination in Ophrys is highly specific and usually each Ophrys species attracts only one pollinator species (Paulus & Gack 1990). The high degree of specialization provides the means of reproductive isolation between the intercrossable Ophrys-species (Ehrendorfer 1980). The complex odour-bouquets released by the flowers are species-specific and often consist of more than 100 different chemical compounds (Borg-Karlson & al. 1985, Ayasse 2006). Speciation in Ophrys-orchids may be brought about by changes in the pollinator attracting floral scent. The attraction of a new pollinator may act as a pre-zygotic isolation barrier (Stebbins 1970, Paulus & Gack 1990, Soliva & al. 2001). We investigated three sympatrically occuring Ophrys-species on Sardinia. O. chestermanii and O. normanii are endemic and are both pollinated by males of the bumblebee B. vestalis. O. tenthredinifera is pollinated by Eucera nigrilabris. There are different opinions concerning the taxonomic status of O. normanii. It has been described as an actual hybrid between O. chestermanii and O. tenthredinifera (Wood 1983). Paulus & Gack (1995) suggested that it is an own species, that either has developed from a hybrid between O. chestermanii and O. normanii or that has evolved by radiation from O. tenthredinifera. By conducting behavioural-tests with B. vestalis males, performing gas chromatographic analyses and electrophysiological studies we wanted to identify pollinator attracting scent and to clarify the taxonomic status of O. normanii.Sexualtäuschorchideen der Gattung Ophrys (Orchidaceae) imitieren die Weibchen ihrer Bestäuber in Duft, Form und Farbe. Insektenmännchen versuchen mit dem Labellum der Blüte zu kopulieren und transportieren den Pollen von Blüte zu Blüte, wodurch die Orchidee bestäubt wird. In dieser Arbeit untersuchten wir die Bestäuber anlockenden Duftstoffe der beiden endemisch auf Sardinien vorkommenden Arten O. normanii und O. chestermanii, die beide von Bombus vestalis Männchen (Hymenoptera: Apidae) bestäubt werden und von O. tenthredinifera, die Eucera nigrilabris (Hymenoptera: Apidae) zur Bestäubung anlockt. O. normanii wurde von Wood (1983) als Primärhybride beschrieben. Nach Paulus und Gack (1995) handelt es sich um eine hybridogene Art oder um eine Art die durch Abspaltung von O. tenthredinifera entstanden ist. Das Ziel der Untersuchungen war die Identifizierung Männchen-anlockender Verbindungen. Die Attraktivität der drei Arten für B. vestalis Männchen sollte Hinweise auf den Artstatus von O. normanii geben. In Biotests mit B. vestalis-Männchen lösten Blütenextrakte von O. normanii und O. chestermanii ebenso wie B. vestalis-Weibchen Kopulationsverhalten der Männchen aus, nicht jedoch Extrakte von O. tenthredinifera. Folglich handelt es sich bei O. normanii nicht um einen aktuellen Hybriden zwischen O. chestermanii und O. tenthredinifera. Ein Vergleich der GC-EAD-aktiven Duftbouquets mittels Diskriminanzanalyse ergab große Ähnlichkeiten zwischen O. normanii und O. chestermanii für die Substanzklassen der Ester, Alkohole und Fettsäuren, die daher vermutlich eine Schlüsselfunktion bei der Bestäuberanlockung haben

Nectar and oleiferous trichomes as floral attractants in Bulbophyllum saltatorium Lindl. (Orchidaceae)

Although many Orchidaceae have deceit flowers that produce no reward, the most common reward, when present, is nectar. Bulbophyllum, however, is unusual in that the labellar secretions of most species investigated to date lack sugars, and, therefore, cannot be considered true nectar. The African species Bulbophyllum saltatorium is an exception in that it produces not only nectar but also possesses specialized, capitate oleiferous trichomes. The nectary of B. saltatorium is borne on the labellum and is represented by a deep, narrow, median longitudinal groove, having a small aperture, and flanked by trichomes. Isodiametric epidermal cells lining this groove secrete nectar which collects both in the groove and on the surface of the labellum. As well as a nectary, the labellum of B. saltatorium also bears three types of unicellular trichomes: the longest trichomes are borne distally and abaxially; the marginal ones form a rim around the entire labellum, and finally, massive, capitate trichomes occur proximally and adaxially. These are oleiferous, containing large quantities of oil which might function as precursors of volatile components of fragrance or provide a food-reward. To the best of our knowledge, this is the first time for such oleiferous trichomes to be described for Bulbophyllum. Therefore, apart from their color and markings, flowers of this species are able to attract pollinators in at least two, possibly three ways: food-reward in the form of nectar; fragrance; and possibly food-rewards in the form of food-hairs

Molecular and morphological phylogenetics of the digitate-tubered clade within subtribe Orchidinae s.s. (Orchidaceae: Orchideae)

The digitate-tubered clade (Dactylorhiza s.l. plus Gymnadenia s.l.) within subtribe Orchidinae is an important element of the North-temperate orchid flora and has become a model system for studying the genetic and epigenetic consequences of organism-wide ploidy change. Here, we integrate morphological phylogenetics with Sanger sequencing of nrITS and the plastid region trnL-F in order to explore phylogenetic relationships and phenotypic character evolution within the clade. The resulting morphological phylogenies are strongly incongruent with the molecular phylogenies, instead reconstructing through parsimony the genus-level boundaries recognised by traditional 20th Century taxonomy. They raise fresh doubts concerning whether Pseudorchis is sister to Platanthera or to Dactylorhiza plus Gymnadenia. Constraining the morphological matrix to the topology derived from ITS sequences increased tree length by 20%, adding considerably to the already exceptional level of phenotypic homoplasy. Both molecular and morphological trees agree that D. viridis and D. iberica are the earliest- diverging species within Dactylorhiza (emphasising the redundancy of the former genus Coeloglossum). Morphology and ITS both suggest that the former genus Nigritella is nested within (and thus part of) Gymnadenia, the Pyrenean endemic 'N.' gabasiana apparently forming a molecular bridge between the two radically contrasting core phenotypes. Comparatively short subtending molecular branches plus widespread (though sporadic) hybridisation indicate that Dactylorhiza and Gymnadenia approximate the minimum level of molecular divergence acceptable in sister genera. They share similar tuber morphologies and base chromosome numbers, and both genera are unusually prone to polyploid speciation. Another prominent feature of multiple speciation events within Gymnadenia is floral paedomorphosis. The 'traditional' morphological and candidate-gene approaches to phylogeny reconstruction are critically appraised.Peer reviewedFinal Published versio

Pollination ecology of New Zealand orchids : a thesis presented in partial fulfilment of the requirements for the degree of Master of Science in Ecology at Massey University

The New Zealand orchid flora comprises twenty-five genera and at least 100 species occurring throughout the country. Although the number of endemic species is high (69%) only four genera are endemic to New Zealand. The main physical threats to orchid survival in New Zealand are habitat destruction, modification and fragmentation. The effect of the disruption of interactions with their pollinators has never been considered. This study concentrates on this mutualistic interaction, by assessing the breeding system, pollination syndromes and pollinator-dependence of four widespread terrestrial (Gastrodia cunninghamii, Thelymitra longifolia, Pterostylis alobula and P. patens) and four widespread epiphytic orchids (Earina autumnalis, E. aestivalis, E. mucronata and Winika cunninghamii) occurring in the southern portion of the North Island. In order to determine the breeding system and the presence of self-incompatibility, hand-pollination treatments were conducted in all eight orchid species during the flowering seasons of 2001 and 2002. Pollen grains and ovules numbers, pollen:ovule ratio and presence of floral scent glands were assessed. In those nectariferous species (E. autumnalis, E. aestivalis, E. mucronata and W. cunninghamii), the nectar standing crop was determined using the anthrone colorimetric assay for total carbohydrates. The activity of pollinator was observed both in the field and in captivity. Insects observed foraging in these orchids were identified and ranked according to their likely pollination effectiveness. Finally, measurements of pollination success and pollinia removal and deposition were used to assess whether fruit-set is pollen limited in these species and explore the effect contrasting rewarding strategies (nectar v/s deception) has on the pollination success of these orchids. Pollination treatments in three terrestrial (T. longifolia, P. alobula and P. patens) and two epiphytic (E. autumnalis and E. mucronata) orchids confirmed the absence of genetic incompatibility. Despite these five orchids being self-compatible, their reproduction relies on contrasting reproductive strategies. T. longifolia is predominantly self-pollinated, whereas Pterostylis and Earina species are incapable of autonomous selfing and completely dependent on pollinators. The epiphytic species E. aestivalis and W. cunninghamii are partially self-incompatible and also completely dependent on pollinators. Agamospermy is likely to occur in G. cunninghamii but not involved in seed-production in any of the remaining seven orchids. Both terrestrial and epiphytic species showed a positive reaction to neutral red except E. autumnalis. This indicated the presence of scent glands, mainly located around the column, lip and sepal tips. Pollen:ovule ratios calculated for these species ranged from 20:1 in E. mucronata and E. aestivalis to 320: 1 in P. alobula. Of the four terrestrial orchids studied, insect visitation was observed only in P. alobula. This orchid is pollinated by male fungus gnats of the genus Zygomyia (Diptera: Mycetophilidae). Pollination by sexual deception is likely to occur in species of this genus. Numerous insects were recorded visiting the nectariferous epiphytic orchids (3 orders, 13 families). Insects considered as "probable pollinator" were Eristalis tenax (Diptera: Syrphidae) for Earina autumnalis, Dilophus nigrostigmus (Diptera: Bibionidae) for E. mucronata, and Melangyna novaezealandiae (Diptera: Syrphidae), Calliphora quadrimaculata (Diptera: Calliphoridae), the Ichneumonid wasp Aucklandella sp. (Hymenoptera: Ichneumonidae), Hylaeus sp. (Hymenoptera: Colletidae) and an unidentified weevil (Coleoptera: Curculionidae) for E. aestivalis. In W. cunninghamii the species Apis mellifera and the native syrphid flies Helophilus antipodus and M. novaezealandiae were considered as "probable pollinators". Levels of natural fruit-set were similarly low in rewarding and non-rewarding species fluctuating from 4.3% (P. alobula) to 40% (P. patens). Fruiting in these orchids is pollen limited, as supplementary hand-pollinations increased fruit set above 40% in all species except P. patens. The degree of pollen limitation varied from 0.32 (P. patens) and 0.94 (P. alobula and E. mucronata). Pollen limitation in these orchids may be caused by the simplicity of their flowers, the poor efficiency of their pollinators in depositing pollinia and the use of species-specific pollination systems (e.g. Pterostylis). The survival capability and conservation requirements of these orchids are discussed in the light of the specific reproductive requirements revealed by this study