A Closer Look at Hyacinths

Photo by Radu Chibzii licensed under CC BY-SA 2.0

They say that our sense of smell is very closely tied with the formation of memories. It is around this time of year that I am strongly reminded of the power of that link. All I have to do is catch a whiff of a blooming hyacinth and I am immediately transported back to childhood where spring time gatherings with the family were always accompanied by mass quantities of these colorful bulbs. Indeed, the smell of hyacinths in bloom will forever hold a special place in my mind (and heart).

Because it is spring in my neck of the woods and because my partner recently came home with a wonderful potted hyacinth to add some springy joy our apartment, I decided to take a dive into the origins of these plants. Where do they come from and how do they live in the wild? Certainly they didn’t originate in our gardens.

To start with, there are surprisingly few true hyacinths in this world these days. Whereas many more spring flowering bulbs were once considered members, today the genus Hyacinthus is comprised of only three species, H. litwinovii, H. transcaspicus, and the most famous of them all, H. orientalis. All other “hyacinths” are hyacinths in name only. These plants were once considered members of the lily family (Liliaceae) but more recent genetic work places them in the asparagus family (Asparagaceae).

All three species of hyacinth are native to the eastern Mediterranean region, throughout the Middle East, and well into southwestern Asia. As you might imagine, there is a fair amount of geographical variation across populations of these plants. For instance, H. orientalis itself contains many putative subspecies and varieties. However, their long history of human cultivation has seen them introduced and naturalized over a much wider area of the globe. Generally speaking, these plants tend to prefer cool, higher elevation habitats and loose soils.

As many of you already know, hyacinths are bulbous plants. Throughout most of the year, they lie dormant beneath the soil waiting for warming spring weather to signal that it’s growing time. And grow they do! Because their leaves and inflorescence are already developed within the bulb, hyacinths can rapidly emerge, flower, and leaf out once snow thaws and releases water into the soil. And flower they do! Though selective breeding has resulted in myriad floral colors and strong, pleasant odors, the wild species are nonetheless put on quite a display.

The flowers of wild hyacinths are generally fewer in number and can range in color from almost white or light blue to nearly purple. Their wonderful floral scent is not a human-bred characteristic either, though we have certainly capitalized on it in the horticulture trade. In the wild, these scent compounds call in pollinators who are rewarded with tiny amounts of nectar. It is thought that bees are the primary pollinators of hyacinths both in their native and introduced habitats.

Of course, all of their floral beauty comes down to seed production. Upon ripening, each fruit (capsule) opens to reveal numerous seeds, each with a fleshy attachment called an elaiosome. The elaiosome is very attractive to resident ants that quickly go to work collecting seeds and bringing them back to their colony. However, it isn’t the seed itself the ants are interested in, but rather the elaiosome. Once it is removed and consumed, the seed is discarded, usually in a waste chamber within the colony where it is free to germinate far away from potential seed predators.

Once growth and reproduction are over, hyacinths once again retreat back underground into their bulb phase. Amazingly, these plants have a special adaptation to make sure that their bulbs are tucked safely underground, away from freezing winter temperatures. Throughout the growing season, hyacinths produce specialized roots that are able to contract. As they contract, they literally pull the base of the plant deeper into the soil. This is very advantageous for plants that enjoy growing in loose soils that are prone to freezing. Once underground and away from frost and snow, they lie dormant until spring returns.

I don’t know about you but getting to know how common garden plants like hyacinths make a living in the wild only makes me appreciate them more. I hope this brief introduction will have you looking at the hyacinths in your neighborhood in a whole new light.

Further Reading: [1] [2]

Of Bluebells and Fungi

Photo by Christophe Couckuyt licensed under CC BY 2.0

Whether in your garden or in the woods, common bluebells (Hyacinthoides non-scripta) are a delightful respite from the dreary months of winter. It should come as no surprise that these spring geophytes are a staple in temperate gardens the world over. And, as amazing as they are in the garden, bluebells are downright fascinating in the wild.

Bluebells can be found growing naturally from the northwestern corner of Spain north into the British Isles. They are largely a woodland species, though finding them in meadows isn't uncommon. They are especially common in sites that have not experienced much soil disturbance. In fact, large bluebell populations are used as indicators of ancient wood lots.

Photo by RX-Guru licensed under CC BY-SA 3.0

Being geophytes, bluebells cram growth and reproduction into a few short weeks in spring. We tend to think of plants like this as denizens of shade, however, most geophytes get going long before the canopy trees have leafed out. As such, these plants are more accurately sun bathers. On warm days, various bees can be seen visiting the pendulous flowers, with the champion pollinator being the humble bumble bees.

The above ground beauty of bluebells tends to distract us from learning much about their ecology. That hasn't stopped determined scientists though. Plenty of work has been done looking at how bluebells make their living and get on with their botanical neighbors. In fact, research is turning up some incredible data regarding bluebells and mycorrhizal fungi.

Photo by Mick Garratt licensed under CC BY-SA 2.0

Bluebell seeds tend not to travel very far, most often germinating near the base of the parent. Germination occurs in the fall when temperatures begin to drop and the rains pick up. Interestingly, bluebell seeds actually germinate within the leaf litter and begin putting down their initial root before the first frosts. Often this root is contractile, pulling the tiny seedling down into the soil where it is less likely to freeze. During their first year, phosphorus levels are high. Not only does the nutrient-rich endosperm supply the seedling with much of its initial needs, abundant phosphorus near the soil surface supplies more than enough for young plants. This changes as the plants age and change their position within the soil.

Photo by MichaelMaggs licensed under CC BY-SA 3.0

Over the next 4 to 5 years, the bluebell's contractile roots pull it deeper down into the soil, taking it out of the reach of predators and frost. This also takes them farther away from the nutrient-rich surface layers. What's more, the roots of older bluebells are rather simple structures. They do not branch much, if at all, and they certainly do not have enough surface area for proper nutrient uptake. This is where mycorrhizae come in.

Bluebells partner with a group of fungi called arbuscular mycorrhiza, which penetrate the root cells, thus greatly expanding the effective rooting zone of the plant. Plants pay these fungi in carbohydrates produced during photosynthesis and in return, the fungi provide the plants with access to far more nutrients than they would be able to get without them. One of the main nutrients plants gain from these symbiotic fungi is phosphorus.

Photo by Oast House Archive licensed under CC BY-SA 2.0

For bluebells, with age comes new habitat, and with new habitat comes an increased need for nutrients. This is why bluebells become more dependent on arbuscular mycorrhiza as they age. In fact, plants grown without these fungi do not come close to breaking even on the nutrients needed for growth and maintenance and thus live a shortened life of diminishing returns. This is an opposite pattern from what we tend to expect out of mycorrhizal-dependent plants. Normally its the seedlings that cannot live without mycorrhizal symbionts. It just goes to show you that even familiar species like the bluebell can offer us novel insights into the myriad ways in which plants eke out a living.

Photo Credits: [1] [2] [3] [4] [5]

Further Reading: [1] [2]

Daffodil Insights

Photo by Amanda Slater licensed under CC BY-SA 2.0

Daffodils seem to be everywhere. Their horticultural popularity means that, for many of us, these plants are among the first flowers we see each spring. Daffodils are so commonplace that it's as if they evolved to live in our gardens and nowhere else. Indeed, daffodils have had a long, long history with human civilization, so much so that it is hard to say when our species first started to cohabitate. Our familiarity with these plants belies an intriguing natural history. What follows is a brief overview of the world of daffodils. 

If you are like me, then you may have gone through most of your life not noticing much difference between garden variety daffodils. Though many of us will be familiar with only a handful of daffodil species and cultivars, these introductions barely scratch the surface. One may be surprised to learn that as of 2008, more than 28,000 daffodil varieties have been named and that number continues to grow each and every year. Even outside of the garden, there is some serious debate over the number of daffodil species, much of this having to do with what constitutes a species in this group.

Narcissus poeticus

As I write this, all daffodils fall under the genus Narcissus. Estimates as to the number of species within Narcissus range from as few as 50 to as many as 80. The genus itself sits within the family Amaryllidaceae and is believed to have originated somewhere between the late Oligocene and early Miocene, some 18 to 30 million years ago. Despite its current global distribution, Narcissus are largely Mediterranean plants, with peak diversity occurring on the Iberian Peninsula. However, thanks to the aforementioned long and complicated history in cultivation, it has become quite difficult to understand the full range of diversity in form and habitat of many species. To understand this, we first need to understand a bit about their reproductive habits.

Much of the evolution of Narcissus seems to center around floral morphology and geographic isolation. More specifically, the length of the floral tube or "corona" and the position of the sexual organs within, dictates just who can effectively pollinate these plants. The corona itself is not made up of petals or sepals but instead, its tube-like appearance is due to a fusion of the stamens into the famous trumpet-like tube we know and love.

Variation in corona shape and size has led to the evolution of three major pollination strategies within this genus. The first form is the daffodil form, whose stigma is situated at the mouth of the corolla, well beyond the 6 anthers. This form is largely pollinated by larger bees. The second form is the paperwhite form, whose stigma is situated more closely to or completely below the anthers at the mouth of the corona. This form is largely pollinated by various Lepidoptera as well as long tongued bees and flies. The third form is the triandrus form, which exhibits three distinct variations on stigma and anther length, all of which are situated deep within the long, narrow corona. The pendant presentation of the flowers in this group is thought to restrict various butterflies and moths from entering the flower in favor of bees.

Narcissus tazetta. Photo by Fanghong licensed under CC BY-SA 3.0

The variations on these themes has led to much reproductive isolation among various Narcissus populations. Plants that enable one type of pollinator usually do so at the exclusion of others. Reproductive isolation plus geographic isolation brought on by differences in soil types, habitat types, and altitudinal preferences is thought to have led to a rapid radiation of these plants across the Mediterranean. All of this has gotten extremely complicated ever since humans first took a fancy to these bulbs.

Narcissus cyclamineus. Photo by Francine Riez licensed under CC BY-SA 3.0

Reproductive isolation is not perfect in these plants and natural hybrid zones do exist where the ranges of two species overlap. However, hybridization is made much easier with the helping hand of humans. Whether via landscape disturbance or direct intervention, human activity has caused an uptick in Narcissus hybridization. For centuries, we have been mixing these plants and moving them around with little to no record as to where they originated. What's more, populations frequently thought of as native are actually nothing more than naturalized individuals from ancient, long-forgotten introductions. For instance, Narcissus populations in places like China, Japan, and even Great Britain originated in this manner.

All of this mixing, matching, and hybridizing lends to some serious difficulty in delineating species boundaries. It would totally be within the bounds of reason to ask if some of the what we think of as species represent true species or simply geographic varieties on the path to further speciation. This, however, is largely speculative and will require much deeper dives into Narcissus phylogenetics.

Narcissus triandrus. Photo by Dave Gough licensed under CC BY 2.0

Despite all of the confusion surrounding accurate Narcissus taxonomy, there are in fact plenty of true species worth getting to know. These range in form and habit far more than one would expect from horticulture. There are large Narcissus and small Narcissus. There are Narcissus with yellow flowers and Narcissus with white flowers. Some species produce upright flowers and some produce pendant flowers. There are even a handful of fall-blooming Narcissus. The variety of this genus is staggering if you are not prepared for it.

Narcissus viridiflorus - a green, fall-blooming daffodil. Photo by A. Barra licensed under CC BY 3.0

After pollination, the various Narcissus employ a seed dispersal strategy that doesn't get talked about enough in reference to this group. Attached to each hard, black seed are fatty structures known as eliasomes. Eliasomes attract ants. Like many spring flowering plant species around the globe, Narcissus utilize ants as seed dispersers. Ants pick up the seeds and bring them back to their nests. They go about removing the eliasomes and then discard the seed. The seed, safely tucked away in a nutrient-rich ant midden, has a much higher chance of germination and survival than if things were left up to simple chance. It remains to be seen whether or not Narcissus obtain similar seed dispersal benefits from ants outside of their native range. Certainly Narcissus populations persist and naturalize readily, however, I am not aware if ants have any part in the matter.

The endangered Narcissus alcaracensis. Photo by José Luis López González licensed under CC BY-SA 4.0

Despite their popularity in the garden, many Narcissus are having a hard go of it in the wild. Habitat destruction, climate change, and rampant collecting of wild bulbs are having serious impacts on Narcissus numbers. The IUCN considered at least 5 species to be endangered and a handful of some of the smaller species already thought to be extinct in the wild. In response to some of these issues, protected areas have been established that encompass at least some of the healthy populations that remain for some of these species.

If you are anything like me, you have ignored Narcissus for far too long. Sure, they aren't native to the continent on which I live, and sure, they are one of the most commonly used plants in a garden setting, but every species has a story to tell. I hope that, armed with this new knowledge, you at least take a second look at the Narcissus popping up around your neighborhood. More importantly, I hope this introduction makes you appreciate their wild origins and the fact that we still have much to learn about these plants. I have barely scratched the surface of this genus and there is more more information out there worth perusing. Finally, I hope we can do better for the wild progenitors of our favorite garden plants. They need all the help they can get and unless we start speaking up and working to preserve wild spaces, all that will remain are what we have in our gardens and that is not a future I want to be a part of.

Photo Credits: [1] [2] [3] [4] [5] [6] [7]

Further Reading: [1] [2] [3] [4] [5] [6] [7] [8] [9]

A Green Daffodil From Spain

Photo by A. Barra licensed under CC BY 3.0

There are some plants that are so ubiquitous in horticulture that I almost forget that they have wild constituents. Every plant in our gardens can trace its lineage back to the wild. As is often the case, I find the wild congeners of our most beloved horticultural curiosities to be far more fascinating. Take, for instance, the genus Narcissus. Who doesn't recognize a daffodil? The same cannot be said for their wild cousins. In fact, there exists some pretty fantastic species within this genus including a small handful of species that flower in autumn. 

A unique fall flowering daffodil is a species known scientifically as Narcissus viridiflorus. This lovely little plant is quite restricted in its range. You will only find it growing naturally in a small region around Gibraltar where it is restricted to rich, clay and/or rocky soils. During years when it is not in flower, N. viridiflorus produces spindly, rush-like leaves. As such, it can be hard to find. 
 

When Narcissus viridiflorus does decide to flower, it forgoes leaf production. From the bulb arises a single green scape. From that scape emerges the flower. The flowers of this bizarre daffodil are decidedly not very daffodil-like. They are rather reduced in form, with long, slender green petals and a nearly nonexistent daffodil cone. Also, they are green. Though I have not seen this investigated directly, it has been suggested that the green scape and flowers contain enough chlorophyll that they plant can recoup at least some of the energy involved in producing flowers and eventually seed. 

The flowers themselves open at night and are said to be very fragrant. Again, no data exists on who exactly pollinates this species but the timing, color, and smell all suggest nocturnal insects like moths. Like the other daffodils of this region, Narcissus viridiflorus is poorly understood. Taken in combination with its limited distribution one can easily see how such a species may be quite vulnerable to human disturbance. As it stands now, this species and many of its cousins are no more than horticultural curiosities for more niche bulb societies. In other words, Narcissus viridiflorus is in need of some real attention. 

Photo Credit: [1] [2]

Further Reading: [1]

The Evolution of Bulbs

Photo by Ewan Bellamy licensed under CC BY-NC-ND 2.0

Spring time is bulb time. As the winter gives way to warmer, longer days, bulbs are among the first of our beloved botanical neighbors to begin their race for the sun. Functionally speaking, bulbs are storage organs. They are made up of a short stem surrounded by layers of fleshy leaves, which contain plenty of energy to fuel rapid growth. Their ability to maintain dormancy is something most of us will be familiar with.

As you might expect, bulbs are an adaptation for short growing seasons. Their ability to rapidly grow shoots gives them an advantage during short periods of time when favorable growing conditions arrive. Despite the energetic costs associated with supplying and maintaining such a relatively large storage organ, the ability to rapidly deploy leaves when conditions become favorable is very advantageous.

Contrast this with rhizomatous species, which are often associated with a life in the understory (though not exclusively) or in crowded habitats like grasslands where competition for light and space can be fierce. Their ambling subterranean habit allows them to vegetatively "explore" for light and nutrients. What's more, the connected rhizomes allow the parent plant to provide nutrients to the developing clones until they grow large enough to support themselves. Under such conditions, bulbs would be at a disadvantage.

Bulbs have evolved independently throughout the angiosperm tree. Many instances of a switch from rhizomatous to bulbous growth habit occurred during the Miocene (23.03 to 5.332 million years ago) and has been associated with a global decrease in temperature and an increase in seasonality at higher latitudes. The decrease in growing season may have favored the evolution of bulbous plants such as those in the lily family. Today, we take advantage of this hardy habit, making bulbous species some of the most common plants used in gardens.


Further Reading: [1]