Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems

Scientific approaches to Australian temperate terrestrial orchid conservation

Mark C. Brundrett

Terrestrial Ecosystems Branch, Policy and Coordination Division, EPA Service Unit, Department of Environment, PO Box K822, Perth, WA 6000, Australia and Faculty of Natural and Agricultural Sciences, School of Plant Biology, The University of Western Australia, Crawley, WA 6009, Australia. Email:

Australian Journal of Botany 55(3) 293-307
Submitted: 21 June 2006  Accepted: 23 October 2006   Published: 18 May 2007


This review summarises scientific knowledge concerning the mycorrhizal associations, pollination, demographics, genetics and evolution of Australian terrestrial orchids relevant to conservation. The orchid family is highly diverse in Western Australia (WA), with over 400 recognised taxa of which 76 are Declared Rare or Priority Flora. Major threats to rare orchids in WA include habitat loss, salinity, feral animals and drought. These threats require science-based recovery actions resulting from collaborations between universities, government agencies and community groups.

Fungal identification by DNA-based methods in combination with compatibility testing by germination assays has revealed a complex picture of orchid–fungus diversity and specificity. The majority of rare and common WA orchids studied have highly specific mycorrhizal associations with fungi in the Rhizoctonia alliance, but some associate with a wider diversity of fungi. These fungi may be a key factor influencing the distribution of orchids and their presence can be tested by orchid seed bait bioassays. These bioassays show that mycorrhizal fungi are concentrated in coarse organic matter that may be depleted in some habitats (e.g. by frequent fire). Mycorrhizal fungi also allow efficient propagation of terrestrial orchids for reintroduction into natural habitats and for bioassays to test habitat quality.

Four categories of WA orchids are defined by the following pollination strategies: (i) nectar-producing flowers with diverse pollinators, (ii) non-rewarding flowers that mimic other plants, (iii) winter-flowering orchids that attract fungus-feeding insects and (iv) sexually deceptive orchids with relatively specific pollinators. An exceptionally high proportion of WA orchids have specific insect pollinators. Bioassays testing orchid-pollinator specificity can define habitats and separate closely related species. Other research has revealed the chemical basis for insect attraction to orchids and the ecological consequences of deceptive pollination. Genetic studies have revealed that the structure of orchid populations is influenced by pollination, seed dispersal, reproductive isolation and hybridisation. Long-term demographic studies determine the viability of orchid populations, estimate rates of transition between seedling, flowering, non-flowering and dormant states and reveal factors, such as grazing and competition, that result in declining populations.

It is difficult to define potential new habitats for rare orchids because of their specific relationships with fungi and insects. An understanding of all three dimensions of orchid habitat requirements can be provided by bioassays with seed baits for fungi, flowers for insects and transplanted seedlings for orchid demography. The majority of both rare and common WA orchids have highly specific associations with pollinating insects and mycorrhizal fungi, suggesting that evolution has favoured increasing specificity in these relationships in the ancient landscapes of WA.


Adams PB Lawson SD 1993 Pollination of Australian orchids: a critical assessment of the literature 1882–1992. Australian Journal of Botany 41 553 575 doi:10.1071/BT9930553

Arditti J Ghani AKA 2000 Numerical and physical properties of orchid seeds and their biological implications. New Phytologist 145 367 421 doi:10.1046/j.1469-8137.2000.00587.x

Batty AL Dixon KW Brundrett MC Sivasithamparam K 2001 a Constraints to symbiotic germination of terrestrial orchid seeds in mediterranean woodland. New Phytologist 152 511 520 doi:10.1046/j.0028-646X.2001.00277.x

Batty AL Dixon KW Brundrett MC Sivasithamparam K 2001 b The long-term storage of mycorrhizal fungi and seed as a tool for the conservation of endangered terrestrial orchids. Australian Journal of Botany 49 619 628 doi:10.1071/BT01029

Batty AL , Dixon KW , Brundrett MC , Sivasithamparam K (2002) Orchid conservation and mycorrhizal associations. In ‘Microorganisms in plant conservation and biodiversity’. (Eds K Sivasithamparam, KW Dixon, RL Barrett) pp. 195–226. (Kluwer Academic Publishers: Dordrecht, The Netherlands)

Batty AL Brundrett MC Dixon KW Sivasithamparam K 2006 a In situ symbiotic seed germination and propagation of terrestrial orchid seedlings for establishment at field sites. Australian Journal of Botany 54 375 381 doi:10.1071/BT04024

Batty AL Brundrett MC Dixon KW Sivasithamparam K 2006 b New methods to improve symbiotic propagation of temperate terrestrial orchids from axenic culture to soil. Australian Journal of Botany 54 367 374 doi:10.1071/BT04023

Bayerman P , Otero T (2006) Microbial endophytes of orchids. In ‘Microbial root endophytes’. (Eds B Schulz, C Boyle, T Sieber) pp. 153–177. (Springer-Verlag: Berlin)

Beardsell DV Clements MA Hutchinson JF Williams EG 1986 Pollination of Diuris maculata R.Br. (Orchidaceae) by floral mimicry of the native peas Daviesia spp. and Pultanaea scabra R.Br. Australian Journal of Botany 34 165 173 doi:10.1071/BT9860165

Bell DT 2001 Ecological response syndromes in the flora of southwestern Western Australia: fire resprouters versus reseeders. Botanical Review 67 417 440

Benzing DH Atwood JT Jr 1984 Orchidaceae: ancestral habitats and current status in forest canopies. Systematic Botany 9 155 165

Bernhardt P (1990) ‘Wily violets and underground orchids: revelations of a botanist.’ (Allen and Unwin: Sydney)

Bidartondo MI Burghardt B Gebauer G Bruns TD Read DJ 2004 Changing partners in the dark: isotopic and molecular evidence of ectomycorrhizal liaisons between forest orchids and trees. Proceedings of the Royal Society of London. Series B. Biological Sciences 271 1799 1806 doi:10.1098/rspb.2004.2807

Bonnardeaux Y Brundrett M Batty A Dixon K Koch J Sivasithamparam K 2007 Diversity of mycorrhizal fungi of Western Australian terrestrial orchids: compatibility webs, brief encounters, lasting relationships and alien invasions. Mycological Research 111 51 61

Bougoure JJ Bougoure DS Cairney JWG Dearnaley JDW 2005 ITS–RFLP and sequence of endophytes from Acianthus, Caladenia and Pterostyis (Orchidaceae) in southeastern Queensland. Mycological Research 109 452 460

Bower CC 1996 Demonstration of pollinator-mediated reproductive isolation in sexually deceptive species of Chiloglottis (Orchidaceae). Australian Journal of Botany 44 15 33 doi:10.1071/BT9960015

Bowles M Zettler L Bell T Kelsey P 2005 Relationship between soil characteristics, distribution and restoration potential of the federal threatened eastern prairie fringed orchid Platanthera leucophaea (Nutt.) Lindl. American Midland Naturalist 154 273 286 doi:10.1674/0003-0031(2005)154[0273:RBSCDA]2.0.CO;2

Brown A (1991) Western Australian native orchids: the masters of deceit. In ‘Proceedings of the 12th Australian orchid conference’. pp. 24–29. (Orchid Society of Western Australia: Perth)

Brown EM , Burbidge AH , Dell J , Edinger D , Hopper SD , Wills RT (1997) ‘Pollination in Western Australia: a database of animals visiting flowers. Handbook no. 15.’ (Western Australian Naturalists’ Club: Perth)

Brown A , Thomson-Dans C , Marchant N (1998) ‘Western Australia’s threatened flora.’ (Department of Conservation and Land Management: Perth)

Brzosko E 2002 Dynamics of island populations of Cypripedium calceolus in the Biebrza river valley (north-east Poland). 2002. Botanical Journal of the Linnean Society 139 67 77 doi:10.1046/j.1095-8339.2002.00049.x

Brundrett MC 1991 Mycorrhizas in natural ecosystems. Advances in Ecological Research 21 171 313

Brundrett M 2002 Coevolution of roots and mycorrhizas of land plants. New Phytologist 154 275 304

Brundrett M 2004 Diversity and classification of mycorrhizal associations. Biological Reviews 79 473 495 doi:10.1017/S1464793103006316

Brundrett MC (2006) Understanding the roles of multifunctional mycorrhizal and endophytic fungi. In ‘Microbial root endophytes’. (Eds B Schulz, C Boyle, T Sieber) pp. 281–297. (Springer-Verlag: Berlin)

Brundrett MC Abbott LK 1991 Roots of jarrah forest plants. I. Mycorrhizal associations of shrubs and herbaceous plants. Australian Journal of Botany 39 445 457 doi:10.1071/BT9910445

Brundrett MC Scade A Batty AL Dixon KW Sivasithamparam K 2003 Development of in situ and ex situ seed baiting techniques to detect mycorrhizal fungi from terrestrial orchid habitats. Mycological Research 107 1210 1220 doi:10.1017/S0953756203008463

Burgman MA , Lindenmayer DB (1998) ‘Conservation biology for the Australian environment.’ (Surrey Beatty: Sydney)

Calvo RN 1993 Evolutionary demography of orchids: intensity and frequency of pollination and cost of fruiting. Ecology 74 1033 1042 doi:10.2307/1940473

Cameron DD Leake JR Read DJ 2006 Mutualistic mycorrhiza in orchids: evidence from plant–fungus carbon and nitrogen transfers in the green-leaved terrestrial orchid Goodyera repens. New Phytologist 171 405 416 doi:10.1111/j.1469-8137.2006.01767.x

Chase MW , Cameron KM , Barrett RL , Freudenstein JV (2003) DNA data and Orchidaceae systematics: a new phylogenetic classification. In ‘Orchid conservation’. (Eds KW Dixon, SP Kell, RL Barrett, PJ Cribb) pp. 69–89. (Natural History Publications: Kota Kinabalu, Sabah, Malaysia)

Chung MY Nason JD Chung MG 2004 Spatial genetic structure in populations of the terrestrial orchid Cephalanthera longibracteata (Orchidaceae). American Journal of Botany 91 52 57

Clark S deLacey C Chamberlain S 2004 Using environmental variables and multivariate analysis to delineate preferred habitat for Cryptostylis hunteriana, the leafless tongue orchid, in the Shoalhaven Local Government Area, NSW. Cunninghamia 8 467 476

Coates DJ Atkins KA 2001 Priority setting and the conservation of Western Australia’s diverse and highly endemic flora. Biological Conservation 97 251 263

Coates F Lunt ID Tremblay RL 2006 Effects of disturbance on population dynamics of the threatened orchid Prasophyllum correctum D.L.Jones and implications for grassland management in south-eastern Australia. Biological Conservation 129 59 69 doi:10.1016/j.biocon.2005.06.037

Collins M Koch J Brundrett M Sivasithamparam K 2005 Recovery of terrestrial orchids in the post-mining landscape. Selbyana 25 255 264

Commonwealth of Australia (2002) ‘Australian terrestrial biodiversity assessment 2002.’ (National Land and Water Resources Audit: Canberra)

Cozzolino S Widmer A 2005 Orchid diversity: an evolutionary consequence of deception? Trends in Ecology & Evolution 20 487 494

Cozzolino S Noce ME Musacchio A Widmer A 2003 Variation at a chloroplast minisatellite locus reveals the signature of habitat fragmentation and genetic bottlenecks in the rare orchid Anacamptis palustris (Orchidaceae). American Journal of Botany 90 1681 1687

Cozzolino S Scheistl FP Müller A De Castro O Nardella AM Widmer A 2005 Evidence for pollinator sharing in Mediterranean nectar-mimic orchids: absence of premating barriers? Proceedings of the Royal Society of London. Series B. Biological Sciences 272 1271 1278

Cozzolino S Nardella AM Impagliazzo S Widmer A Lexer C 2006 Hybridization and conservation of Mediterranean orchids: should we protect the orchid hybrids or the orchid hybrid zones? Biological Conservation 129 14 23 doi:10.1016/j.biocon.2005.09.043

Currah RS , Zelmer CD , Hambleton S , Richardson KA (1996) Fungi from orchid mycorrhizas. In ‘Orchid biology: reviews and perspectives. VII’. (Eds J Arditti, A Pridgeon) pp. 117–170. (Kluwer Academic Publishers: Dordrecht, The Netherlands)

Dafni A 1984 Mimicry and deception in pollination. Annual Review of Ecology and Systematics 15 259 278 doi:10.1146/

Dafni A Calder DM 1987 Pollination by deceit and floral mimesis in Thelymitra atennifera (Orchidaceae). Plant Systematics and Evolution 158 11 22 doi:10.1007/BF00936140

Darwin C (1904) ‘The various contrivances by which orchids are fertilized by insects.’ 2nd edn. (John Murray: London)

Dixon K (1991) Seeder/clonal concepts in Western Australian orchids. In ‘Population ecology of terrestrial orchid’. (Eds TCE Wells, JH Willems) pp. 111–123. (SPB Academic Publishing: The Hague, The Netherlands)

Edwards W Westoby M 2000 Families with highest proportions of rare species are not consistent between floras. Journal of Biogeography 27 733 740 doi:10.1046/j.1365-2699.2000.00424.x

Elliott CP Ladd PG 2002 Pollen limitation of fruit set in Western Australian terrestrial orchids. Journal of the Royal Society of Western Australia 85 165 168

Erickson R (1965) ‘Orchids of the west.’ (Paterson Brokensha: Perth)

Esitken A Ercisli S Eken C 2005 Effects of mycorrhiza isolates on symbiotic germination of terrestrial orchids (Orchis palustris Jacq. and Serapias vomeracea subsp. vomeracea (Burm.f.) Briq.) in Turkey. Symbiosis 38 59 68

Feuerherdt L Petit S Jusaitis M 2005 Distribution of mycorrhizal fungus associated with the endangered pink-lipped spider orchid (Arachnorchis (syn. Caladenia) behrii) at Warren Conservation Park in South Australia. New Zealand Journal of Botany 43 367 371

George AS 1980 Rhizanthella gardneri R.S.Rogers – the underground orchid of Western Australia. American Orchid Society Bulletin 49 631 644

Gonzales D Carling DE Kuninaga S Vilgalys R Cubeta MA 2001 Ribosomal DNA systematics of Ceratobasidium and Thanatephorus with Rhizoctonia anamorphs. Mycologia 93 1138 1150

Grant CD Koch J 2003 Orchid species succession in rehabilitated bauxite mines in Western Australia. Australian Journal of Botany 51 453 457

Hoffman N , Brown A (1998) ‘Orchids of South-west Australia.’ 2nd edn. (University of Western Australia Press: Perth)

Hollick P (2004) Mycorrhizal specificity in endemic Western Australian terrestrial orchids (tribe Diurideae): implications for conservation. PhD Thesis, Murdoch University, Perth.

Hopper SD Brown AP 2004 Robert Brown’s Caladenia revisited, including a revision of its sister genera Cyanicula, Ericksonella and Pheladenia (Caladeniinae: Orchidaceae). Australian Systematic Botany 17 171 240 doi:10.1071/SB03002

Hopper SD Gioia P 2004 The Southwest Australian Floristic Region: evolution and conservation of a global hotspot of biodiversity. Annual Review of Evolution and Systematics 35 623 650 doi:10.1146/annurev.ecolsys.35.112202.130201

Honnay O Bossuyt B 2005 Prolonged clonal growth: escape route or route to extinction? Oikos 108 427 432 doi:10.1111/j.0030-1299.2005.13569.x

Hutchings MJ 1987 The population biology of the early spider orchid, Ophrys sphegodes Mill. I. A demographic study from 1975 to 1984. Journal of Ecology 75 711 727 doi:10.2307/2260201

Huynh TT McLean CB Coates F Lawrie AC 2004 Effect of developmental stage and peloton morphology on success in isolation of mycorrhizal fungi in Caladenia formosa (Orchidaceae). Australian Journal of Botany 52 231 241 doi:10.1071/BT03099

Jacquemyn H Brys R Hermy M Willems JH 2005 a Does nectar reward affect rarity and extinction probabilities of orchid species? An assessment using historical records from Belgium and the Netherlands. Biological Conservation 121 257 263 doi:10.1016/j.biocon.2004.05.002

Jacquemyn H Micheneau C Roberts DL Pailler T 2005 b Elevational gradients of species diversity, breeding system and floral traits of orchid species on Reunion Island. Journal of Biogeography 32 1751 1761 doi:10.1111/j.1365-2699.2005.01307.x

Johnson SD Neal PR Harder LD 2005 Pollen fates and the limits on male reproductive success in an orchid population. Biological Journal of the Linnean Society 86 175 190 doi:10.1111/j.1095-8312.2005.00541.x

Jones DL (1988) ‘Native orchids of Australia.’ (Reed Books Pty Ltd: Sydney)

Julou T Burghardt B Gebauer G Berveiller D Damesin C Selosse M-A 2005 Mixotrophy in orchids: insights from a comparative study of green individuals and nonphotosynthetic individuals of Cephalanthera damasonium. New Phytologist 166 639 653 doi:10.1111/j.1469-8137.2005.01364.x

Kelly CL Pickering CM Buckley RC 2003 Impacts of tourism on threatened plant taxa and communities in Australia. Ecological Management & Restoration 4 37 44 doi:10.1046/j.1442-8903.2003.00136.x

Kéry M Gregg KB 2004 Demographic analysis of dormancy and survival in the terrestrial orchid Cypripedium reginae. Journal of Ecology 92 686 695 doi:10.1111/j.0022-0477.2004.00885.x

Kéry M Gregg KB Schaub M 2005 Demographic estimation methods for plants with unobservable life-states. Oikos 108 307 320 doi:10.1111/j.0030-1299.2005.13589.x

Kristiansen KA Freudenstein JV Rasmussen FN Rasmussen HN 2004 Molecular identification of mycorrhizal fungi in Newiedia veratrifolia (Orchidaceae). Molecular Phylogenetics and Evolution 33 251 258 doi:10.1016/j.ympev.2004.05.015

Kull T Hutchings MJ 2006 A comparative analysis of decline in the distribution ranges of orchid species in Estonia and the United Kingdom. Biological Conservation 129 31 39 doi:10.1016/j.biocon.2005.09.046

Leake JR 1994 The biology of myco-heterotrophic (‘saprophytic’) plants. New Phytologist 127 171 216 doi:10.1111/j.1469-8137.1994.tb04272.x

Light MHS MacConaill M 2005 Long-term studies: a case for orchid species survival. Selybyana 26 174 188

Linder HP 1995 Setting conservation priorities: the importance of endemism and phylogeny in the southern African orchid genus Herschelia. Conservation Biology 9 585 595

Machon N Bardin P Mazer SJ Moret J Godelle B Austerlitz F 2003 Relationship between genetic structure and seed and pollen dispersal in the endangered orchid Spiranthes spiralis. New Phytologist 157 677 687 doi:10.1046/j.1469-8137.2003.00694.x

Mant J Peakall R Weston PH 2005 a Specific pollinator attraction and the diversification of sexually deceptive Chiloglottis (Orchidaceae). Plant Systematics and Evolution 253 185 200 doi:10.1007/s00606-005-0308-6

Mant J Bower CC Weston PH Peakall R 2005 b Phylogeography of pollinator-specific sexually deceptive Chiloglottis taxa (Orchidaceae): evidence for sympatric divergence? Molecular Ecology 14 3067 3076 doi:10.1111/j.1365-294X.2005.02659.x

Masuhara G Katsuya K 1994 In situ and in vitro specificity between Rhizoctonia spp. and Spiranthes sinensis (Persoon.) Ames. var. amoena (M.Beiberstein) Hara (Orchidaceae). New Phytologist 127 711 718 doi:10.1111/j.1469-8137.1994.tb02974.x

McCormick MK Whigham DF O’Neill J 2004 Mycorrhizal diversity in photosynthetic terrestrial orchids. New Phytologist 163 425 438 doi:10.1111/j.1469-8137.2004.01114.x

McKendrick SL 1995 The effects of herbivory and vegetation on laboratory-raised Dactylorhiza praetermissa (Orchidaceae) planted into grassland in southern England. Biological Conservation 73 215 220 doi:10.1016/0006-3207(94)00111-3

Milligan MJ Williams PG 1988 The mycorrhizal relationship of multinucleate rhizoctonias from non-orchids with Microtis (Orchidaceae). New Phytologist 108 205 210 doi:10.1111/j.1469-8137.1988.tb03697.x

Molvray M , Kores PJ , Chase MW (2000) Polyphyly of mycoheterotrophous orchids and functional influences on floral and molecular characters. In ‘Monocots: systematics and evolution’. (Eds KL Wilson, DA Mossison) pp. 441–448. (CSIRO Publishing: Melbourne)

Murren CJ 2002 Effects of habitat fragmentation on pollination: pollinia viability and reproductive success. Journal of Ecology 90 100 107 doi:10.1046/j.0022-0477.2001.00638.x

Murren CJ Ellison AM 1998 Seed dispersal characteristics of Brassavola nodosa (Orchidaceae). American Journal of Botany 85 675 680 doi:10.2307/2446537

Myers N Mittermeier RA Mittermeier CG da Fonseca GAB Kent J 2000 Biodiversity hotspots for conservation priorities. Nature 403 853 858 doi:10.1038/35002501

Myklestad A Saetersdal M 2005 Effects of fertilization and afforestation on community structure of traditionally managed hay meadows in western Norway. Nordic Journal of Botany 23 593 606

Nicolè F Brzosko E Till-Bottraud I 2005 Population viability analysis of Cypripedium calceolus in a protected area: longevity, stability and persistence. Journal of Ecology 93 716 726

Neiland MRM Wilcox CC 1998 Fruit set, nectar reward, and rarity in the Orchidaceae. American Journal of Botany 85 1657 1671 doi:10.2307/2446499

Nilsson LA 1992 Orchid pollination biology. Trends in Ecology & Evolution 7 255 259 doi:10.1016/0169-5347(92)90170-G

Otero JT Flanagan NS 2006 Orchid diversity – beyond deception. Trends in Ecology & Evolution 21 64 65 doi:10.1016/j.tree.2005.11.016

Pacini E Hesse M 2002 Types of pollen dispersal units in orchids, and their consequences for germination and fertilization. Annals of Botany 89 653 664 doi:10.1093/aob/mcf138

Pate JS , Hopper SD (1994) Rare and common plants in ecosystems, with special reference to the south-west Australian flora. In ‘Biodiversity and ecosystem function’. (Eds E-D Schulze, HA Mooney) pp. 293–325. (Springer-Verlag: Berlin)

Peakall R 1990 Responses of male Zaspilothynnus trilobatus Turner wasps to females and the sexually deceptive orchid it pollinates. Functional Ecology 4 159 167 doi:10.2307/2389335

Peakall R Beattie AJ 1991 The genetic consequences of worker ant pollination in a self-compatible clonal orchid. Evolution 45 1837 1848 doi:10.2307/2409835

Peakall R Beattie AJ 1996 Ecological and genetic consequences of pollination by sexual deception in the orchid Caladenia tentaculata. Evolution 50 2207 2220 doi:10.2307/2410692

Perkins AJ McGee PA 1995 Distribution of the orchid mycorrhizal fungus, Rhizoctonia solani, in relation to its host, Pterostylis acuminata, in the field. Australian Journal of Botany 43 565 575 doi:10.1071/BT9950565

Perkins AJ Masuhara G McGee PA 1995 Specificity of the associations between Microtis parvifolia (Orchidaceae) and its mycorrhizal fungi. Australian Journal of Botany 43 85 91 doi:10.1071/BT9950085

Peterson RL Uetake Y Zelmer C 1998 Fungal symbioses with orchid protocorms. Symbiosis 25 29 55

Petit S Dickson CR 2005 Grass-tree (Xanthorrhoea semiplana, Liliaceae) facilitation of the endangered pink-lipped spider orchid (Caladenia syn. Arachnorchis behrii, Orchidaceae) varies in South Australia. Australian Journal of Botany 53 455 464

Pfeifer M Wiegand K Heinrich W Jetchke G 2006 Long-term demographic fluctuations in an orchid species driven by weather implications for conservation planning. Journal of Applied Ecology 43 313 324 doi:10.1111/j.1365-2664.2006.01148.x

Pope EJ Carter DA 2001 Phylogenetic placement and host specificity of mycorrhizal isolates belonging to AG-6 and AG-12 in the Rhizoctonia solani species complex. Mycologia 93 712 719

Primack R Stacy E 1998 Cost of reproduction in the pink lady’s slipper orchid (Cypripedium acule, Orchidaceae): an eleven-year experimental study of three populations. American Journal of Botany 85 1672 1679

Ramsay MM , Dixon KW (2003) Propagation science, recovery and translocation of terrestrial orchids. In ‘Orchid conservation’. (Eds KW Dixon, SP Kell, RL Barrett, PJ Cribb) pp. 259–288. (Natural History Publications: Kota Kinabalu, Sabah, Malaysia)

Ramsay RR Sivasithamparam K Dixon KW 1987 Anastomosis groups among Rhizoctonia-like endophytic fungi in southwestern Australian Pterostylis species (Orchidaceae). Lindleyana 2 161 166

Rasmussen HN (1995) ‘Terrestrial orchids, from seed to mycotrophic plant.’ (Cambridge University Press: Cambridge)

Rasmussen HN 2002 Recent developments in the study of orchid mycorrhiza. Plant and Soil 244 149 163

Rasmussen HN Whigham D 1993 Seed ecology of dust seeds in situ: a new study technique and its application in terrestrial orchids. American Journal of Botany 80 1374 1378 doi:10.2307/2445665

Reinhammar L-G Olsson EGA Sørmeland E 2002 Conservation biology of an endangered grassland plant species, Pseudorchis albida, with some references to the closely related alpine P. straminea (Orchidaceae). Botanical Journal of the Linnean Society 139 47 66 doi:10.1046/j.1095-8339.2002.00041.x

Risdall-Smith JJ 1970 The behaviour of Hemithynnus hyalinatus (Hymenoptera: Tiphiidae), with notes on some other Thynninae. Journal of the Australian Entomological Society 9 196 208

Roberts P (1999) ‘Rhizoctonia-forming fungi.’ (Whitstable Litho Printers Ltd: Kent, UK)

Scade A Brundrett MC Batty AL Dixon KW Sivasithamparam K 2006 Survival of transplanted terrestrial orchid seedlings in urban bushland habitats with high or low weed cover. Australian Journal of Botany 54 383 389

Schiestl FP 2005 On the success of a swindle: pollination by deception in orchids. Die Naturwissenschaften 92 255 264 doi:10.1007/s00114-005-0636-y

Schiestl FP Peakall R Mant J 2004 Chemical communication in the sexually deceptive orchid genus Cryptostylis. Botanical Journal of the Linnean Society 144 199 205 doi:10.1111/j.1095-8339.2003.00249.x

Schwartz MW Simberloff D 2001 Taxon size predicts rates of rarity in vascular plants. Ecology Letters 4 464 469 doi:10.1046/j.1461-0248.2001.00241.x

Selosse M-A Faccio A Scappaticci G Bonfante P 2004 Chlorophyllous and achlorophyllous specimens of Epipactis microphylla (Neottieae, Orchidaceae) are associated with ectomycorrhizal septomycetes, including truffles. Microbial Ecology 47 416 426 doi:10.1007/s00248-003-2034-3

Sharma IK Jones DL French CJ 2003 Unusually high genetic variability revealed through allozymic polymorphism of an endemic and endangered Australian orchid, Pterostylis aff. picta (Orchidaceae). Biochemical Systematics and Ecology 31 513 526 doi:10.1016/S0305-1978(02)00186-2

Shefferson RP Kull T Tali K 2005 a Adult whole-plant dormancy induced by stress in long-lived orchids. Ecology 86 3099 3104

Shefferson RP Weiss M Kull T Taylor DL 2005 b High specificity generally characterizes mycorrhizal association in rare lady’s slipper orchids, genus Cypripedium. Molecular Ecology 14 613 626

Sivasithamparam K 1993 Ecology of root infecting pathogenic fungi in mediterranean environments. Advances in Plant Pathology 10 245 279

Soliva M Widmer A 2003 Gene flow across species boundaries in sympatric, sexually deceptive Ophrys (Orchidaceae) species. Evolution 57 2252 2261

Stewart SL Zettler LW 2002 Symbiotic germination of three semi-aquatic rein orchids (Habenaria repens, H. quinquiseta, H. macroceratitius) from Florida. Aquatic Botany 72 25 35 doi:10.1016/S0304-3770(01)00214-5

Stoutamire WP 1983 Wasp-pollinated species of Caladenia (Orchidaceae) in south-western Australia. Australian Journal of Botany 31 383 394 doi:10.1071/BT9830383

Taylor DL , Bruns TD , Leake JR , Read DJ (2002) Mycorrhizal specificity and function in myco-heterotrophic plants. In ‘Mycorrhizal ecology. Ecological studies vol. 157’. (Eds MGA van der Heijden, I Sanders) pp. 375–413 (Springer-Verlag: Berlin)

Taylor DL Bruns TD Szaro TM Hodges SA 2003 Divergence in mycorrhizal specialization within Hexalectris spicata (Orchidaceae), a nonphotosynthetic desert orchid. American Journal of Botany 90 1168 1179

Tremblay RL Ackerman JD Zimmerman JK Calvo RN 2005 Variation in sexual reproduction in orchids and its evolutionary consequences: a spasmodic journey to diversification. Biological Journal of the Linnean Society 84 1 54

Vallius E 2001 Factors affecting fruit set and seed production in Dactylorhiza maculata (Orchidaceae). Botanical Journal of the Linnean Society 135 89 95 doi:10.1006/bojl.2000.0355

Wallace LE 2002 Examining the effects of fragmentation on genetic variation in Platanthera leucophaea (Orchidaceae): inferences from allozyme and random amplified polymorphic DNA markers. Plant Species Biology 17 37 49 doi:10.1046/j.1442-1984.2002.00072.x

Wallace LE 2003 The cost of inbreeding in Platanthera leucophaea (Orchidaceae). American Journal of Botany 90 235 242

Warcup JH 1973 Symbiotic germination of some Australian terrestrial orchids. New Phytologist 72 387 392

Warcup JH 1988 Mycorrhizal associations of isolates of Sebacina vermifera. New Phytologist 110 227 231 doi:10.1111/j.1469-8137.1988.tb00256.x

Weiss M Selosse M-A Rexer K-H Urban A Oberwinkler F 2004 Sebacinales: a hitherto overlooked cosm of heterobasidiomycetes with a broad mycorrhizal potential. Mycological Research 108 1003 1010 doi:10.1017/S0953756204000772

Whigham DF O’Neill JP Rasmussen HN Caldwell BA McCormick MK 2006 Seed longevity in terrestrial orchids – potential for persistence in situ seed banks. Biological Conservation 129 24 30 doi:10.1016/j.biocon.2005.10.029

Willems JH Dorland E 2000 Flowering frequency and plant performance and their relation to age in the perennial orchid Spiranthes spiralis (L.) Chevall. Plant Biology 2 344 349 doi:10.1055/s-2000-3707

Willems JH Melser C 1998 Population dynamics and life-history of Coeloglossum virde (L.) Harm.: an endangered orchid species in The Netherlands. Botanical Journal of the Linnean Society 126 83 89 doi:10.1006/bojl.1997.0146

Willems JH Balounova Z Kindlmann P 2001 The effect of experimental shading on seed production and plant survival of Spiranthes spiralis (Orchidaceae). Lindleyana 16 31 37

Wotavová K Balounová Z Kindlmann P 2004 Factors affecting persistence of terrestrial orchids in wet meadows and implications for their conservation in a changing agricultural landscape. Biological Conservation 118 271 279

Zelmer CD Currah RS 1997 Symbiotic germination of Spiranthes lacera (Orchidaceae) with naturally occurring endophyte. Lindleyana 12 142 148

Zelmer CD Cuthbertson L Currah RS 1996 Fungi associated with terrestrial orchid mycorrhizas, seeds and protocorms. Mycoscience 37 439 448

Export Citation Cited By (33)