Table of Contents
The protea stands as one of the most remarkable and distinctive flowering plants on Earth, representing not only the natural heritage of South Africa but also a fascinating example of botanical evolution and adaptation. The genus Protea is one of the most well-known and charismatic of the Cape Floristic Region’s (CFR) Fynbos Biome, and the King Protea (Protea cynaroides) is South Africa’s national flower. This comprehensive guide explores the protea’s distribution, evolutionary history, ecological adaptations, and cultural significance across the Southern Hemisphere.
The Ancient Origins of the Proteaceae Family
The story of the protea begins hundreds of millions of years ago, making it one of the oldest flowering plant families on our planet. The family Proteaceae to which Protea species belong is an ancient one among angiosperms, with evidence from pollen fossils suggesting Proteaceae ancestors grew in Gondwana, in the Upper Cretaceous, 75–80 million years ago. Some sources suggest an even more ancient lineage, with proteas considered to be among the oldest families of flowering plants on the planet, dating back approximately 300 million years.
This extraordinary age places the Proteaceae family among the most ancient flowering plant lineages, having evolved alongside and outlived the dinosaurs. Evidence from pollen fossils suggests Proteaceae ancestors grew in Gondwana, in the Upper Cretaceous, 75–80 million years ago, during a time when the supercontinent was still intact. The family’s distribution across multiple continents today serves as living evidence of continental drift and the breakup of Gondwana.
Gondwanan Distribution and Continental Drift
The Proteaceae is a good example of a Gondwanan family, with taxa occurring on virtually every land mass considered a remnant of the ancient supercontinent Gondwana, except Antarctica, and the family and subfamilies are thought to have diversified well before the fragmentation of Gondwana, implying all of them are well over 90 million years old. This ancient distribution pattern has resulted in the family’s presence across the Southern Hemisphere today.
The Proteaceae are divided into two subfamilies: the Proteoideae, best represented in southern Africa, and the Grevilleoideae, concentrated in Australia and South America and the other smaller segments of Gondwana that are now part of eastern Asia. This division reflects the evolutionary history of the family and the geographic separation that occurred as Gondwana fragmented.
Interestingly, Africa shares only one genus with Madagascar, whereas South America and Australia share many common genera – this indicates they separated from Africa before they separated from each other. This biogeographic pattern provides crucial insights into the timing and sequence of continental separation during the Mesozoic era.
Global Distribution of the Proteaceae Family
Today, the Proteaceae family exhibits a predominantly Southern Hemisphere distribution that reflects its Gondwanan origins. Proteaceae are mainly a Southern Hemisphere family, with its main centres of diversity in Australia and South Africa, and it also occurs in Central Africa, South and Central America, India, eastern and south eastern Asia, and Oceania.
The family comprises 83 genera with about 1,660 known species, making it a substantial and diverse plant family. Australia and South Africa have the greatest concentrations of diversity, with these two regions serving as the primary centers of speciation and endemism for the family.
Distribution in New Zealand and Oceania
While the Proteaceae family is widespread across former Gondwanan landmasses, its representation varies considerably by region. Only two species are known from New Zealand, although fossil pollen evidence suggests there were more previously. This reduced diversity in New Zealand compared to Australia and South Africa suggests that environmental changes and extinctions have significantly reduced the family’s presence in some regions over geological time.
The fossil record of some areas, such as New Zealand and Tasmania, show a greater biodiversity for Proteaceae than currently exists, which supports the fact that the distribution of many taxa has changed drastically with the passage of time and that the family has suffered a general decline, including high levels of extinction during the Cenozoic.
Dispersal Mechanisms: Continental Drift and Oceanic Crossing
For many years, scientists believed that the Proteaceae family’s distribution was solely the result of continental drift, with ancestral populations simply riding along as Gondwana fragmented. However, recent molecular studies have revealed a more complex story. Using DNA sequence data, botanists have shown that the large southern hemisphere plant family Proteaceae lived on the super-continent Gondwanaland almost 120 million years ago, and as Gondwanaland broke up, it was originally thought that these plants merely moved with the newly formed continents, but a new study has shown that, while this is the case for some of these plants, others are far too recent to have lived at the time when the super-continent broke up, and they must therefore have dispersed across oceans to reach their current distribution ranges.
This discovery has important implications for understanding plant biogeography. Ancestors of some of the modern Proteaceae must have crossed the Atlantic and Indian Oceans, and in Africa, for example, the spectacular genus Protea is truly Gondwanan, but 250 species from other genera that occur in the ‘fynbos’ vegetation of the highly diverse south-western Cape biodiversity hotspot are much younger, and have Australian relatives.
The Protea Genus: Diversity and Distribution in Africa
Within the Proteaceae family, the genus Protea itself represents a remarkable concentration of diversity, particularly in southern Africa. There are around 112 Protea species, 89 of which are restricted to southern Africa and most of which only occur within South Africa’s CFR. This extraordinary level of endemism makes the Cape Floristic Region one of the world’s most important centers of plant diversity.
Concentration in the Cape Floristic Region
About 92% of the species occur only in the Cape Floristic Region, a narrow belt of mountainous coastal land from Clanwilliam to Grahamstown, South Africa. This remarkable concentration of species in such a relatively small geographic area represents one of the highest levels of plant endemism anywhere on Earth.
The Cape Floristic Region itself is an extraordinary biodiversity hotspot. The Cape Floristic Region is home to the greatest non-tropical concentration of higher plant species in the world, with 9,000 species crammed into its small extent, and incredibly, more than 6,200 (69 percent) of these species are found nowhere else in the world. It occupies less than 0.5 percent of the area of Africa but supports approximately 20 percent of its plant life, and about two-thirds of the estimated 9,000 species that grow there are endemic.
Protea Species Beyond the Cape
While the vast majority of protea species are concentrated in the Cape Floristic Region, some species have much wider distributions across Africa. Most protea species are found south of the Limpopo River, but there are notable exceptions that extend into tropical Africa.
Protea madiensis grows in Afromontane enclaves across tropical Africa, from Guinea to Sudan, Mozambique, and Angola. Additionally, Protea afra ranges from the Cape region to Uganda and Kenya, including in the chaparral zone of Mount Kenya National Park. These wide-ranging species demonstrate the genus’s ability to adapt to diverse environmental conditions across the African continent.
Evolutionary Origins and Diversification
Recent phylogenetic research has revealed surprising insights into the evolutionary history of the genus Protea. Contrary to previous views, the Cape is identified as the ancestral area for the radiation of the extant lineages: most species in subtropical and tropical Africa are derived from a single invasion of that region. This finding challenges earlier assumptions about the direction of species migration and diversification within the genus.
Diversification rates have been similar within and outside the Cape region, but migration out of the Cape has opened up vast areas, though those lineages have not diversified as extensively at fine spatial scales as lineages in the Cape. This pattern suggests that the unique environmental conditions of the Cape Floristic Region have promoted exceptional levels of speciation and species persistence at small spatial scales.
The Proteaceae Family in Southern Africa
The diversity of the Proteaceae family in southern Africa extends well beyond the genus Protea itself. Fourteen genera of the Proteaceae occur in southern Africa, namely Aulax (featherbushes – 3 taxa), Brabejum (Wild Almond being the single taxon), Diastella (silkypuffs – 9 taxa), Faurea (beechwoods – 5 taxa), Leucadendron (conebushes – 98 taxa), Leucospermum (pincushions – 54 taxa), Mimetes (pagodas – 13 taxa), Orothamnus (Marsh Rose being the single taxon), Paranomus (sceptres – 19 taxa), Protea (sugarbushes – 87 taxa), Serruria (spiderheads – 59 taxa), Sorocephalus (clusterheads – 11 taxa), Spatalla (spoons – 20 taxa), and Vexatorella (vexators – 5 taxa).
Naming and Taxonomy
The family Proteaceae was named in 1767 by Carl Linnaeus after Proteus, one of the gods in Greek mythology who had the ability to constantly change his form, and the name Proteaceae and Protea thus echo the variability of many different plant forms found within the large family. This naming reflects the extraordinary morphological diversity found within the family, with species ranging from prostrate shrubs to tall trees, and flowers exhibiting an astonishing array of forms, colors, and sizes.
The first record of any plant ever described in southern Africa was Protea neriifolia, described by Clusius, 50 years before Jan van Riebeeck landed at the Cape of Good Hope, highlighting the early recognition of these distinctive plants by European botanists.
The Fynbos Biome: Home of the Proteas
The fynbos biome of South Africa provides the primary habitat for the majority of protea species and represents one of the world’s most unique vegetation types. Most of the region is treeless and covered in a type of evergreen shrubland called fynbos, and the fynbos teems with reeds, heathers, and succulents, as well as colorful orchids, lilies, and proteas.
The king protea (Protea cynaroides), whose flower heads can reach 30 centimeters (12 inches) in diameter, is South Africa’s national flower and present throughout the fynbos. This iconic species serves as the flagship for the entire ecosystem and has become synonymous with South African natural heritage.
Environmental Conditions of the Fynbos
The environmental conditions in the fynbos biome might appear inhospitable to many plants, but proteas have evolved remarkable adaptations to thrive in this challenging environment. Fynbos plants are adapted to the Mediterranean climate of hot, dry summers and wet winters, and they thrive in the sandy nutrient-poor soil, pummeled by wind and sun.
The soil conditions are particularly challenging. Fynbos plants are adapted to the Mediterranean climate of hot, dry summers and wet winters, and they thrive in the sandy nutrient-poor soil, pummeled by wind and sun. These nutrient-poor, acidic soils would be unsuitable for most plant species, but proteas have evolved specialized root systems to extract nutrients efficiently from these challenging substrates.
Fire as an Ecological Driver
Fire plays a crucial and essential role in the fynbos ecosystem, and proteas have evolved remarkable adaptations to survive and even benefit from periodic wildfires. The health of the ecosystem is reliant upon the periodic fires that burn there, both for seed dispersal and germination and for killing off invasive species.
Like many other Protea species, P. cynaroides is adapted to an environment in which bushfires are essential for reproduction and regeneration, and most Protea species can be placed in one of two broad groups according to their response to fire: reseeders are killed by fire, but fire also triggers the release of their canopy seed bank, thus promoting recruitment of the next generation; resprouters survive fire, resprouting from a lignotuber or, more rarely, epicormic buds protected by thick bark, and P. cynaroides is a resprouter as it shoots up new stems from buds in its thick underground stem after a fire.
The relationship between proteas and fire is so intimate that some species are described as “pyrophylic” or fire-loving. They do so by being either a re-sprouter (growing back from a protected root-stock or resilient stump) or by being a re-seeder. This dual strategy ensures that protea populations can persist through the frequent fires that characterize their native habitats.
Remarkable Adaptations of Proteas
Proteas have evolved an impressive suite of adaptations that allow them to thrive in the challenging conditions of their native habitats. These adaptations encompass root systems, leaf structures, water management strategies, and reproductive mechanisms.
Specialized Root Systems
One of the most distinctive features of proteas is their specialized root system. Like most other Proteaceae, P. cynaroides has proteoid roots, roots with dense clusters of short lateral rootlets that form a mat in the soil just below the leaf litter, and these enhance solubilisation of nutrients, thus allowing nutrient uptake in the low-nutrient, phosphorus-deficient soils of its native fynbos habitat.
These proteoid roots, also called cluster roots, represent a remarkable adaptation to nutrient-poor soils. Their roots grow mostly horizontally, just below the surface of the soil, allowing them to efficiently capture nutrients from the shallow soil layers where organic matter accumulates.
Drought Tolerance and Water Management
Proteas have evolved multiple strategies to cope with the hot, dry summers characteristic of Mediterranean-type climates. Protea cynaroides grows in a harsh environment with dry, hot summers and wet, cold winters, and several adaptions include tough, leathery leaves, which helps to prevent excessive loss of moisture, and a large taproot which penetrates deep into the soil to reach underground moisture.
The combination of a deep taproot for accessing groundwater and shallow proteoid roots for nutrient acquisition represents an elegant solution to the dual challenges of nutrient scarcity and water stress. The tough, leathery leaves with their waxy cuticles minimize water loss through transpiration during the hot, dry summer months.
Fire Survival Strategies
Proteas have evolved diverse strategies for surviving and reproducing in fire-prone environments. Protea cynaroides is adapted to survive wildfires by its thick underground stem, which contains many dormant buds; these will produce the new growth after the fire. This resprouting strategy allows individual plants to survive multiple fire cycles, potentially living for decades.
Other protea species employ a reseeding strategy. Some re-seeding proteas – Leucospermums and Serrurias – drop their seeds before a fire, and these are at risk from predators – mice, canaries and insects – but each seed comes with an edible attachment – an elaiosome – which attracts foraging ants, and the obliging ants carry the seed down into their underground nest, eat the elaiosome but leave the seed intact, and now the seed has a nice warm, dark and safe place to lie dormant until the heat from a fire propels it into germination.
The technique is called myrmecochory and about 30% of fynbos plants use it for seed dispersal. This mutualistic relationship between plants and ants represents a sophisticated adaptation to fire-prone environments, ensuring that seeds are protected underground until fire creates favorable conditions for germination.
Pollination Ecology of Proteas
Proteas exhibit diverse pollination strategies, with different species adapted to different pollinators. The flowers are fed at by a range of nectarivorous birds, mainly sunbirds and sugarbirds, including the orange-breasted sunbird (Anthobaphes violacea), southern double-collared sunbird (Cinnyris chalybeus), malachite sunbird (Nectarinia famosa), and the Cape sugarbird (Promerops cafer).
The relationship between proteas and their bird pollinators is so close that specialized bird species have evolved to exploit protea nectar. The area is home to a number of true fynbos species such as the Cape sugarbird (Promerops cafer), the orange-breasted sunbird (Nectarinia violacea), the Protea canary (Serinus leucopterus) and the Cape siskin (Serinus totta).
Beyond bird pollination, proteas also utilize other pollination vectors. Some species of Proteaceae exude a ‘yeasty’ scent, which is a typical trait of plants pollinated by small mammals, and visits by small mammals do not however preclude visits by other pollinators, such as sunbirds and insects, and the mammal-pollinated flowers are more open-shaped and with a diameter wider than the length of the inflorescence, e.g., Protea canaliculata, P. sulphurea, and P. humiflora.
Habitat Requirements and Growing Conditions
Understanding the natural habitat requirements of proteas is essential for successful cultivation, whether in their native range or in suitable climates elsewhere in the world.
Soil Requirements
Soil drainage is the single most critical factor for growing proteas successfully. The single most critical factor in growing Proteas is to provide adequate water drainage; if the soil drains well, good results will usually be achieved. Proteas rarely succeed in heavy clay soils, notorious for their poor drainage.
Proteas require acidic soils with low nutrient levels. Proteas thrive in slightly acidic soil (pH 5-7) and generally do not need fertilizers. In fact, Protea plants thrive in acidic, nutrient-poor soil, and an abundance of phosphorus, in particular, will kill them. This sensitivity to phosphorus is a direct consequence of their adaptation to the phosphorus-deficient soils of their native habitats.
Light and Temperature Requirements
Proteas are sun-loving plants that require full exposure to thrive and flower abundantly. For growing, Proteas must have a full sun location with good air movement around the plants. Insufficient sunlight typically results in poor flowering or no flowers at all.
Proteas can stand a range of temperatures, from 23 F. (-5 C.) to 100 F. (38 C.), though they may not survive long exposure beyond that. While proteas are generally hardy, Protea plants are generally hardy but should be protected from frost, and in winter they can usually handle frosts around 25-30°F.
Water Requirements
Despite their drought tolerance once established, proteas require careful water management, especially during establishment. Despite loving the sun, protea evolved in arid conditions, and they struggle in high humidity and waterlogged soil, and a drip system providing moderate watering, especially during the first three years, supports their horizontal root growth without overwatering, particularly in winter when they require little additional water.
Water your established plants every two to three weeks, and if your plants are just starting out, water them more frequently. The key is to provide deep, infrequent watering rather than frequent shallow watering, which encourages the development of the deep taproot system.
Cultivation and Commercial Production
Proteas have become increasingly popular as ornamental plants and cut flowers, leading to commercial cultivation in suitable climates around the world.
Global Cultivation
Members of the genus are exported as cut flowers all over the world, prized for their beauty, diversity and longevity. The long vase life and striking appearance of protea flowers make them highly valued in the floriculture industry. The flower has a long vase life in flower arrangements, and makes for an excellent dried flower.
California has emerged as a significant center for protea cultivation outside of South Africa. The plant requires full sun, acidic soil, good drainage and air circulation, as well as mild winters, so proteas flourish just as well in a place like San Diego County as if they were in their native environment. Soil and weather conditions are the same as what you would find on a South African protea farm, and the soil is pure decomposed granite with a pH that ranges between 5 and 7.
Propagation Methods
Proteas can be propagated from both seeds and cuttings, though each method presents its own challenges. While seeds often need a smoke primer to germinate and might not thrive in all conditions, cuttings are easier but still delicate, risking mold, and it’s a labor of love, requiring patience, especially since protea, including the king and pink ice varieties, bloom after 5-6 years.
For seed propagation, The best time to sow is in autumn or spring, when the difference between day and night temperatures is about 12oC (54oF), and choose the season that gives the protea seedlings the most time to grow under favourable conditions, for example, if your summer is very hot and dry, but your winter is moderate and wet – sowing in autumn will give the protea seedlings a whole winter and spring to become strong before the harsh summer, and in colder climates, it is best to sow in spring so that protea plants can become hardy before the frosts of winter.
Fertilization Considerations
One of the most important aspects of protea cultivation is avoiding excessive fertilization, particularly phosphorus. Proteas require very little fertilizer—too much produces fast-growing plants with crooked stems and heavier foliage. Are phosphorus intolerant and should not be fertilized.
Since the proteaceae are adapted to nutrient-poor conditions, chemical fertilizer or manure will burn their sensitive root system, and use an organic plant food such as a fish or seaweed emulsion, and avoid any fertilizers containing Phosphates, and generally, no additions are needed to Proteaceae, as their roots systems are so well adapted at extracting whatever they need, even from nutrient poor soil.
Cultural Significance and Symbolism
Beyond their botanical interest, proteas hold deep cultural significance, particularly in South Africa where they serve as powerful national symbols.
National Symbol of South Africa
The king protea is the national flower of South Africa and as such lends its name to the national cricket team, whose nickname is the Proteas. The flower appears on South African birth certificates and passports, as well as on the South African 5-Rand coin, and the Proteas, South Africa’s cricket team, also took its name from the genus.
The choice of the king protea as South Africa’s national flower is deeply symbolic. South Africa’s motto is Diverse People Unite, so the king protea flower truly symbolizes their national spirit. The flower’s diversity of forms and its ability to thrive in challenging conditions mirror the nation’s own diversity and resilience.
Symbolic Meanings
In South African culture, proteas have long been regarded as symbols of hope, resilience, and strength, and the flower is often used in traditional healing practices and is believed to bring good luck and fortune. Proteas represents change and hope in the African tradition, and in Europe’s flower language they stand for diversity and culture.
The protea flower symbolizes change and transformation across cultures. This symbolism is particularly apt given the flower’s ability to regenerate after fire, emerging renewed from the ashes of destruction. The protea’s resilience in the face of adversity makes it a powerful metaphor for personal transformation and national renewal.
Conservation Challenges and Efforts
Despite their hardiness and adaptability, many protea species face significant conservation challenges in their native habitats.
Threatened Species
There are other species of Protea teetering at the brink of extinction and needing all efforts possible to conserve them, and Protea odorata, also known as the Swartland Sugarbush, is listed as Critically Endangered on the Red List of South African Plants, and only around 30 plants of this species now survive in the wild where it is distributed in lowland vegetation from Klapmuts to Riverlands.
Although proteas are well adapted to their natural habitat, a number of varieties are considered at risk of extinction, and of the 360 African species, about 120 of them are listed as endangered, due to loss of habitat and over-collection. This represents approximately one-third of all African protea species, highlighting the urgent need for conservation action.
Threats to Protea Habitats
It is thought that the Cape Floristic Region is experiencing one of the most rapid rates of extinction in the world due to habitat loss, land degradation, and invasive alien plants. The introduction of non-native species poses a particular threat to the fynbos ecosystem.
Thirstier non-native trees such as acacia, pine, and eucalyptus have spread into the floristic region, threatening endemic species and using more water than native plants. These invasive species not only compete with native proteas for resources but also alter fire regimes and water availability, fundamentally changing the ecosystem dynamics that proteas depend upon.
Conservation Initiatives
Recognizing the importance of proteas and the broader fynbos ecosystem, various conservation initiatives are underway. Because of its distinct, diverse, and abundant plant communities, the area is not only recognized as its own major floral kingdom, but a number of properties within it are also protected as UNESCO World Heritage sites, national parks, and nature reserves.
The Protea Atlas Project represents an important citizen science initiative for monitoring and conserving protea populations. The king protea is also the flagship of the Protea Atlas Project, run by the South African National Botanical Institute. Such projects engage the public in conservation efforts while gathering valuable data on species distributions and population trends.
Proteas in Australia and Other Regions
While South Africa is the primary center of diversity for the genus Protea, the broader Proteaceae family shows remarkable diversity in Australia and other Southern Hemisphere regions.
Australian Proteaceae
Australia hosts an extraordinary diversity of Proteaceae species, though these belong primarily to different genera than the African proteas. World wide around 79 genera and 1700 species, and Australia around 46 genera and 1100 species. This means that Australia contains more than half of all Proteaceae species globally, though distributed across different genera than those found in Africa.
Well-known Australian genera include Banksia, Grevillea, Hakea, and the spectacular Waratah (Telopea). Well-known Proteaceae genera include Protea, Banksia, Embothrium, Grevillea, Hakea, and Macadamia, and species such as the New South Wales waratah (Telopea speciosissima), king protea (Protea cynaroides), and various species of Banksia, Grevillea, and Leucadendron are popular cut flowers.
Economic Importance
Beyond their ornamental value, some Proteaceae species have significant economic importance. The nuts of Macadamia integrifolia are widely grown commercially and consumed, as are those of Gevuina avellana on a smaller scale. The macadamia nut industry represents a major agricultural enterprise, particularly in Hawaii, Australia, and South Africa.
The Future of Proteas
As climate change and human activities continue to impact natural ecosystems, the future of proteas depends on effective conservation strategies and sustainable cultivation practices.
Climate Change Impacts
Climate change poses significant challenges for protea populations in their native habitats. Changes in temperature, rainfall patterns, and fire regimes could all impact protea survival and distribution. Research is ongoing to understand how different species might respond to changing conditions and to identify populations that may be particularly vulnerable or resilient.
Sustainable Cultivation
Commercial cultivation of proteas for the cut flower industry can help reduce pressure on wild populations while providing economic benefits. However, it’s essential that cultivation practices are sustainable and that genetic diversity is maintained in cultivated populations. The development of new cultivars should not come at the expense of wild populations or lead to genetic pollution through hybridization with wild plants.
Education and Awareness
Increasing public awareness about the importance of proteas and the fynbos ecosystem is crucial for long-term conservation success. Educational programs, botanical gardens, and ecotourism initiatives all play important roles in connecting people with these remarkable plants and fostering appreciation for their conservation.
The economical worth of fynbos biodiversity, based on harvests of fynbos products (e.g. wildflowers) and eco-tourism, is estimated to be in the region of R77 million (~US$5 million) a year. This economic value provides additional incentive for conservation while supporting local communities.
Conclusion
The protea stands as a testament to the power of evolution and adaptation, representing one of the oldest and most distinctive flowering plant lineages on Earth. From its ancient origins on the supercontinent Gondwana to its current distribution across the Southern Hemisphere, the protea’s story is one of survival, diversification, and remarkable adaptation to challenging environments.
The extraordinary concentration of protea diversity in South Africa’s Cape Floristic Region makes this small area one of the world’s most important centers of plant biodiversity. The unique adaptations that allow proteas to thrive in nutrient-poor soils, survive periodic fires, and flourish in Mediterranean climates demonstrate the remarkable evolutionary innovations that have occurred in this ancient plant family.
As both a national symbol and a botanical treasure, the protea embodies themes of resilience, transformation, and diversity that resonate far beyond the realm of botany. Whether admired in its native fynbos habitat, cultivated in gardens around the world, or featured in floral arrangements, the protea continues to captivate and inspire.
The conservation challenges facing proteas remind us of the fragility of even the most resilient species in the face of habitat loss, invasive species, and climate change. Protecting these remarkable plants and their ecosystems requires ongoing commitment, scientific research, and public engagement. By understanding and appreciating the unique biology and cultural significance of proteas, we can work toward ensuring that these ancient flowers continue to thrive for generations to come.
For those interested in learning more about proteas and supporting conservation efforts, consider visiting botanical gardens that feature Proteaceae collections, supporting organizations working to protect the Cape Floristic Region, or exploring opportunities for sustainable ecotourism in South Africa’s fynbos regions. Additional information can be found through the South African National Biodiversity Institute, the Critical Ecosystem Partnership Fund, and various botanical research institutions worldwide.