plant taxonomy

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Tea Time: Linden Flower Tea

Lindens make great trees for urban areas. A few species and hybrids in particular are commonly planted in parks, yards, and along the streets of cities across the northern hemisphere and have been for decades – centuries even. They cast dense shade, are tolerant of a variety of climates and soil conditions, and are generally easy to maintain. For much of the year as you move throughout the city you live in, you likely pass by dozens of lindens without thinking twice about them. They are ubiquitous, conventional, ordinary, common. Unless they’re in bloom. For a few weeks in early to mid-summer, flowering lindens produce an impossibly sweet fragrance that can’t be ignored. Along with the scent comes the sound of hundreds of buzzing bees collecting pollen and nectar from the pendulous blooms.

Lindens are trees and shrubs in the family Malvaceae and genus Tilia. Around 30 or so species are found in temperate regions across the northern hemisphere, mostly in Europe and Asia. Depending on who you ask, there are between one and three species native to North America. Tilia caroliniana and Tilia heterophylla are considered by some to be varieties of Tilia americana, or American basswood, which is distributed across central and eastern United States and north into parts of Canada. Another common name for linden is lime because words used to refer to the tree in older languages were similar to the word lime. The name basswood comes from the tree’s fibrous inner bark, known as bast.

Linden leaves are generally heart-shaped and asymmetrical with serrate margins. Small clusters of little yellow to white flowers form at the end of a slender stem attached to a narrow, ribbon-like, yellow-green bract. The bract aids in seed dispersal by helping the fruits float on the wind away from the parent tree in a manner similar to the samaras of maple trees. The fruits are small, round, hardened drupes that resemble little peas. The fragrant, nectar-rich flowers are not only favored by beekeepers for honey production, but also have a long history of being harvested for making tea (i.e. tisane). Linden flower tea is said to have a number of medicinal uses and health benefits, all of which I take with a grain of salt. This series of posts isn’t meant to be an investigation into the health claims of plants, but instead an opportunity for me – out of sheer curiosity – to try making tea out of a variety of different plants . If medicinal uses interest you, I encourage you to seek out credible, peer-reviewed sources.

I made linden flower tea from flowers I collected from Tilia cordata, commonly known as littleleaf linden. It was an easy one to find due to its popularity as an urban tree. The natural distribution of littleleaf linden extends from Britain across Europe and into western Asia. Its triangular-ovate shaped leaves are 4-10 centimeters long, glossy green on top, and pale green on the bottom with tufts of orange hairs along the leaf veins, concentrated at the base of the leaf where the leaf blade meets the petiole. The tree can reach up to 21 meters tall and has an oval or rounded-pyramidal shape, though many trees in urban areas are cultivars and can be smaller and more compact.

I harvested the flowers – bracts and all – in late June. It’s advised that they not be harvested directly after a rain (or after being hit by sprinklers), and that they are harvested when the flowers are newly opened. I presume this is because the flowers are at their freshest at this point and will be the best for making tea. I layed the flowers out to dry on a clean kitchen towel on top of a metal cake rack. It only takes 2 or 3 days for them to dry. After drying I removed and saved all the flowers and threw out the bracts and stems, but apparently you can use the entire inflorescence if you’d like.

There are several linden flower tea recipes online. I went with 3 cups of boiling water poured over 1 tablespoon dried linden flowers, covered and steeped for 15 minutes. The resulting tea was an appealing pastel yellow color. I tried it plain as well as sweetened with a little bit of honey. I preferred it sweetened, but unsweetened wasn’t too bad, just a little bitter. It has a floral taste and pleasant smell. Sierra said it tasted earthy, like something she wasn’t supposed to be drinking. Despite that odd review, she said she liked it. Since several sources discussed the calming, sleep-inducing effects of the tea, I made sure to drink it in the evening when it would be normal for me to be feeling sleepy. I suggest you do the same.

More Tea Time Posts on Awkward Botany

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Seed Dispersal by Way of Tree Climbing Goats

Goats are surprisingly good climbers. Given the opportunity, they’ll climb just about anything, including each other. So what’s stopping them from climbing a tree, especially if there is something up there they can eat? And so they do. Tree climbing goats are such a fascinating sight, they even have their own calendar. But the story doesn’t end there. The goats find food in the trees, entertaining humans as they go; meanwhile, the trees have a reliable partner in the goats, who inadvertently help disperse the tree’s seeds.

In general, goats don’t need to climb trees to find food. Goats aren’t known to be picky eaters, and there is usually plenty for them to eat at ground level. However, in arid climates where food can become limited, ascending trees to eat foliage and fruits is a matter of survival. This is the case in southwestern Morocco, where goats can be found in the tops of argan trees every autumn gorging on the fruits of this desert tree.

goats in Argania spinosa via wikimedia commons

Argan (Argania spinosa) is a relatively short tree with a sprawling canopy and thorny branches. It is the only species in its genus and is endemic to parts of Morocco and neighboring Algeria. The tree is economically important to the area due to the oil-rich seeds found within its bitter fruits. Argan oil has a variety of culinary uses and is also used medicinally and in cosmetics. To get to the oil, goats are often employed in harvesting the fruits. The goats retrieve the fruits from the tops of the trees and consume their fleshy outer layer. The hard, seed-containing pits are expelled, collected, and cracked open to get to the seeds.

This is where a team of researchers from Europe come in. There has been some confusion as to how the pits are expelled, with some reports claiming that they pass through the goats digestive track and are deposited in their manure. This is a common way for the seeds of many other plant species to be dispersed, and is carried out not only by goats and other ruminants, but also by a wide variety of mammals, as well as birds and even reptiles. However, considering the average size of the pits (22 mm long x 15 mm wide), the researchers thought this to be unlikely.

fruits of Argania spinosa via wikimedia commons

Others reported that the seeds were spat out in the goats’ cud while they ruminated. Goats, like other ruminants, have stomachs composed of multiple compartments, the first of which being the rumen. Partially digested food, known as cud, is sent back into the mouth from the rumen for further chewing and may be spat out or swallowed again. Goats are known to ruminate in the same location that they defecate, which results in confusion as to when and how certain seeds, like those of the argan tree, are deposited.

By feeding various fruits to a group of goats, the researchers were able to test the hypothesis that seeds could be regurgitated and spat from the cud and that this is a viable method of seed dispersal. The researchers reported that larger seeds were more commonly spat out than smaller seeds, but that “almost any seed could be ejected during, mastication, spat from the cud, digested, or defecated.” The viability of spat out seeds was tested, and over 70% of them were found to be viable.

pits and seeds of Argania spinosa via wikimedia commons

This discovery suggests that seed dispersal via spitting by ruminants could be a common occurrence – possibly far more common than previously considered. The researchers postulate that studies that have only considered seeds dispersed in manure “may have underestimated an important fraction of the total number of dispersed seeds” and that the seeds spat from the cud likely represent different species from those commonly dispersed in dung. In addition, the seeds of some species don’t survive the digestive tract of ruminants, so “spitting from the cud may represent their only, or at least their main, dispersal mechanism.”

This study surrounding the argan trees was followed up by the same group of researchers with a literature review that was published last month. The review looked into all available studies that mentioned seed dispersal via regurgitation by ruminants. While they considered over 1000 papers, only 40 published studies were found to be relevant for the review. From these studies, they determined that the seeds of 48 plant species (representing 21 different families) are dispersed by being spat from a ruminant’s cud, and that most of these plant species are trees and shrubs whose fruits contain large seeds. Also of note is that ruminants across the globe are doing this – representatives from 18 different genera were mentioned in the studies.

ruminating goat via wikimedia commons

The researchers conclude that this is a “neglected” mechanism of seed dispersal. It’s difficult to observe, and in many cases it hasn’t even been considered. Like so many other animals, ruminants can disperse seeds in a variety of ways. Seeds can attach to their fur and be transported wherever they go. They can pass through their digestive track and end up in their dung, potentially far from where they were first consumed. And, as presented here, they can be spat out during rumination. Investigations involving all of these mechanisms and the different plant species involved will allow us to see, in a much clearer way, the role that ruminants play in the dispersal of seeds.

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Spring Weeds in the Mustard Family

Is there a plant family that consists of more weedy species than the mustard family? Asteraceae and Poaceae, for sure. Fabaceae or Lamiaceae, perhaps. Regardless, Brassicaceae is replete with dozens of species – mostly annual – that are skilled at taking advantage of the disturbed environments that humans are in the habit of creating.

It helps that the mustard family is so large: 372 genera and over 4,000 species distributed across the globe. Around 55 genera are said to occur in North America. Most of the plants in this family are herbaceous; few are shrubs. Foliage is aromatic, especially when crushed. Flowers are particularly distinctive. Each flower has four petals – in some species petals are divided, giving the impression that there are more than four – arranged in the shape of a cross or “X.” Flowers are often small, have 4 tall stamens and 2 short stamens, and commonly come in white, yellow, pink, or purple. They are arranged on a raceme, which is typically either tall and straight or compact and flat-topped.

Fruits in the mustard family are capsules with two compartments separated by a clear membrane. The capsules may be at least three times longer than they are wide, in which case they are referred to as a silique; or they may be less than three times longer than they are wide and referred to as a silicle. This is a curious distinction, and it doesn’t tell you all that much. It’s more important to understand that the capsules of mustards can come in various sizes and shapes, and that some can be long and narrow while others are short and either round or angular.

mustard seeds via wikimedia commons

Despite the size or shape of the capsule, enclosed are numerous seeds – sometimes dozens. Surely one of the reasons why plants in the mustard family are so successful at proliferating is their ability to produce thousands, even tens of thousands, of seeds per plant. The seeds are typically tiny; and while they may not make it very far from the parent plant, they are numerous. Depending on the species, they can also remain viable for years, affording them the opportunity to sprout whenever conditions are right. You may have heard the biblical verse about faith the size of a mustard seed giving one the ability to move mountains. Size seems irrelevant here, so how about faith as tough, resilient, opportunistic, and resourceful as a mustard seed? If a mountain can be moved, mustards might be the one to do it.

While it isn’t the scope of this post, it’s worth mentioning the chemical compounds present in mustards that give them the flavors and health benefits we enjoy as well as the toxicity that can harm us and any other organisms that dare consume them. Glucosinolates, which are present in various concentrations depending on the species, are a defining characteristic of plants in the mustard family. They contribute to the spicy-ness of things like horseradish, radish, and condiment mustard while also acting as a natural insecticide, deterring herbivory.

And now on to the cast of characters:

Whitetop (Lepidium spp.)

Garlic mustard (Alliaria petiolata) – a noxious weed in many parts of North America – is fortunately not an issue in southwestern Idaho, otherwise it would be first on the list. Instead, we deal with whitetop – a noxious weed in Idaho and several other states. As the common name suggests, individual plants – up to two feet tall – are topped with a dense cluster of tiny, white flowers. Seed production in this group isn’t as abundant as other mustards; instead, the tour de force are their rhizomes. Whitetop is a perennial plant that spreads aggressively via underground stems as well as root fragments and can easily form expansive, dense patches, outcompeting other plants in the area.

Another common name for this group is hoary cress on account of their gray-green, fuzzy foliage. They are further distinguished by the shape of their seed pods: lens-podded hoary cress (L. chalepense), heart-podded hoary cress (L. draba), and globe-podded hoary cress (L. applelianum).

white top (Lepidium sp.)

white top (Lepidium sp.)

Tansymustard (Descurainia spp.)

There are two species of tansymustard (also known as flixweed) that occur in my part of the world, one is native and the other is introduced from Europe. They are indistinguishable to my untrained eye. If I have seen them side by side, I wouldn’t have known it. They are both annuals and can be as short as a few inches to over two feet tall. They have highly dissected, fern-like leaves and tiny, pale yellow or green-yellow flowers. The seed pods are very skinny and around an inch long. Each pod can hold 40 seeds, and a large plant can produce over 75,000 seeds. They are quick to take advantage of disturbed soil and come up in abundance after a fire. I’m not sure what it is about this year, but they have been particularly prolific this spring.

tansymustard (Descurainia sp.)

tansymustard (Descurainia sp.)

Blue Mustard (Chorispora tenella)

Also known as musk mustard or crossflower, this sticky, stinky, annual plant apparently makes cow’s milk taste funny; however some people still enjoy eating it. It can get to about a foot and a half tall, and is adorned with pretty, little, blue-purple flowers. The pointy seed pods split crosswise rather than lengthwise, an uncommon trait in mustards.

blue mustard (Chorispora tenella)

blue mustard (Chorispora tenella)

Desert Madwort (Alyssum desertorum)

Like tansymustard, this species is very similar in appearance to another closely related species, Alyssum alyssoides (commonly known as pale madwort or yellow alyssum). Both are annuals under a foot tall, covered in tiny hairs, with minuscule yellow flowers, and numerous round seed pods. They are adapted to dry, neglected sites.

yellow alyssum (Alyssum dessertorum)

Annual Honesty (Lunaria annua)

If you don’t recognize this plant when it’s flowering, you will when its seed pods ripen. They are thin, round discs up to three inches across. Eventually, the outer layers of the seed pods fall away, and translucent membranes remain, sometimes with seeds still attached. This trait has earned this species common names like money plant and silver dollar. The plants are attractive, reach up to three feet tall, and produce showy, purple flowers, which explains why they are popular ornamentals. However, like other mustards, the proficiency with which they reproduce in abundance via seeds, means they also easily migrate into natural areas and neglected sites.

annual honesty (Lunaria annua)

Hairy Bittercress (Cardamine hirsuta)

This little, quick-growing, fast-spreading annual is a common nuisance in greenhouses and nurseries. On stalks above compact rosettes are borne clusters of white flowers that, as Ken Thompson writes in The Book of Weeds, “are so tiny they are almost invisible.” The slender seed pods burst open at maturity, sending minuscule seeds flying. Brush your hand over a patch of mature hairy bittercress and you will be bombarded.

hairy bittercress (Cardamine hirsuta)

And the list goes on…

I’ve observed several other weedy species in this family recently, but to keep the length of this post reasonable I will just list them here: shepherd’s purse (Capsella bursa-pastoris), clasping pepperweed (Lepidium perfoliatum), spring draba (Draba verna), tumble mustard (Sisymbrium altissimum), and pennycress (Thlaspi arvense). This list only scratches the surface; there are many other weeds in the mustard family. All deserve to be mentioned, so perhaps another time.

See Also: In Defense of Plants – One Mustard, Many Flavors

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Charles Darwin and the Phylogeny of State Flowers and State Trees

This is a guest post by Rachel Rodman. Photos by Daniel Murphy.

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Every U.S. state has its own set of symbols: an official flower, an official tree, and an official bird. Collectively, these organisms form the stuff of trivia and are traditionally presented in the form of a list.

But, lists…well. As charming as lists can sometimes be, lists are rarely very satisfying.

So I decided to try something different.

I am not, of course, the first person to be unhappy with the eclectic, disordered nature of many biological assemblages. In the 18th century, Linnaeus developed a classification system in order to make sense of that untidiness. Kingdom, Phylum, Class, and so on.

In the 19th century, Darwin set biodiversity into an even more satisfying intellectual framework, outlining a model that linked organisms via descent from a series of common ancestors. And, as early as 1837, he experimented with a tree-like structure, in order to diagram these relationships.

Following Darwin’s lead, I’ve worked to reframe the state flowers and state trees in terms of their evolutionary history (*see the methods section below). And today, in honor of Darwin’s 209th birthday, I am delighted to present the results to you.

Let’s start with the state flowers.

In this tree, Maine’s “white pine cone and tassel” forms the outgroup. Among all the state “flowers,” it is the only gymnosperm—and therefore, in fact, not actually a flower.

Notice, also, that the number of branches in this tree is 39—not 50. Most of this stems from the untidy fact that there is no requirement for each state to select a unique flower. Nebraska and Kentucky, for example, share the goldenrod; North Carolina and Virginia share the dogwood.

With the branch labeled “Rose,” I’ve compressed the tree further. The state flowers of Georgia, Iowa, North Dakota, New York, and Oklahoma are all roses of various sorts; with my data set (*see methods below), however, I was unable to disentangle them. So I kept all five grouped.

This is a rich tree with many intriguing juxtapositions. Several clades, in particular, link geographical regions that are not normally regarded as having a connection. Texas’ bluebonnet, for example, forms a clade with Vermont’s red clover. So, similarly, do New Hampshire’s purple lilac and Wyoming’s Indian paintbrush.

Texas bluebonnet (Lupinus texensis) – the state flower of Texas

The second tree—the tree of state trees—is similarly rewarding. This tree is evenly divided between angiosperms (19 species) and gymnosperms (17 species).

Iowa’s state tree is simply the “oak”—no particular species was singled out. To indicate Iowa’s selection, I set “IA” next to the node representing the common ancestor of the three particular oak species: white oak, red oak, and live oak, which were selected as symbols by other states.

Arkansas’ and North Carolina’s state tree, similarly, is the “pine,”—no particular species specified. I’ve indicated their choice in just the same way, setting “AR” and “NC” next to the node representing the common ancestor of the eight particular pine species chosen to represent other states.

In this tree of trees, as with the tree of flowers, several clades link geographical regions that are not usually linked—at least not politically. Consider, for example, the pairing of New Hampshire’s white birch with Texas’ tree, the pecan.

Another phylogenetic pairing also intrigued me: Pennsylvania’s eastern hemlock and Washington’s western hemlock. It evokes, I think, a pleasing coast-to-coast symmetry: two states, linked via an east-west cross-country bridge, over a distance of 2,500 miles

The corky bark of bur oak (Quercus macrocarpa). Oak is the state tree of Iowa.

In this post, I’ve presented the U.S. state flowers and U.S. state trees in evolutionary framework. The point in doing that was not to denigrate any of the small, human stories that lie behind these symbols—all of the various economic, historical, and legislative vagaries, which led each state to select these particular plants to represent them. (Even more importantly, I have no wish to downplay the interesting nature of any of the environmental factors that led particular plants to flourish and predominate in some states and not others.)

The point, instead, was to suggest that these stories can coexist and be simultaneously appreciated alongside a much larger one: the many million year story of plant evolution.

With Darwin’s big idea—descent with modification—the eclectic gains depth and meaning. And trivia become a story—a grand story, which can be traced back, divergence point by divergence point: rosids from asterids (~120 mya); eudicots from monocots (~160 mya); angiosperms from gymnosperms (~300 mya), and so on and so on.

So today, on Darwin’s 209th, here, I hope, is one of the takeaways:

An evolutionary framework really does make everything—absolutely everything: U.S. state symbols included—more fun, more colorful, more momentous, and more intellectually satisfying.

Thanks, Darwin.

*Methods:

To build these two trees, I relied on a data set from TimeTree.org, a website maintained by a team at Temple University. At the “Load a List of Species” option at the bottom of the page, I uploaded two lists of species in .txt format; each time, TimeTree generated a phylogenetic tree, which served as a preliminary outline.

Later, once I’d refined my outlines, I used the “Get Divergence Time For a Pair of Taxa” feature at the top of the page in order to search for divergence time estimates. As I reconstructed my trees in LibreOffice, I used these estimates to make my branch lengths proportional.

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Rachel Rodman has a Ph.D. in Arabidopsis genetics and presently aspires to recontextualize all of history, literature, and popular culture in the form of a phylogenetic tree. Won’t you help her?

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On the Genus Euphorbia

This is a guest post. Words and photos by Jeremiah Sandler.

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Suspicion

I collect cacti and succulents. The more I collect plants, the more and more I become interested in taxonomic and phylogenetic relationships between them. Not just my own plants – all of them. Most recently, the genus Euphorbia has been on my mind. My favorite species are E. meloformis var. valida and E. horrida.

I’m mostly familiar with the succulent and cacti-looking euphorbia (they are not true cacti) and a few ornamental annuals. Sometimes I would come across a species that I could determine was a euphorbia; but in trying to identify exactly which species, I found countless possibilities within the genus. It seemed odd to me that a single genus could contain so many different forms.

Turns out, Euphorbia consists of over 1800 separate species. What?! That is an insanely high number! Only about 20 genera of plants contain over 1000 separate species. Euphorbia is the fourth most populated genus among all genera of plants.

That staggering number got me thinking: how can a single genus have so many different species? How has the classification worked that out? Has the genus been phylogenetically examined? There’s no way a genus can be so huge. You know what breeders and collectors can do with that much genetic material in a single genus? The man-made hybrids seem endless.

Euphorbia globosa in bloom

Taxonomy

In older taxonomic practices, morphological similarities were the primary method of grouping individuals together. While that is still a common practice today, phylogenetic testing is now an accessible tool for organizing species into related groups.

Organizations such as the Angiosperm Phylogeny Group (APG) have been doing this advanced scientific research – analyzing DNA, doing detailed dissection, etc. Ultimately, they organize plant taxonomy and systematics with greater detail, and examine plant relationships genetically – phylogenetics.

Analyzing genomes is much more expensive and time consuming than observing morphologies. Now, a mix of methods is used, but DNA sequencing has definitely changed the systematics game in a big way. As a result of the APG’s incorporation of widespread phylogenetic DNA analyses, their taxonomical classifications are quickly becoming the generally accepted classifications among plant taxonomists.

Since the inclusion of genetic testing, many plant orders, families, and genera have been reorganized, renamed, expanded, or shrunk.

Euphorbia

One of the identifying features of euphorbias are their very unique flowers. All species in the genus have a cyathium, an inflorescence exclusively produced by euphorbias. Lacking in true petals, sepals, or nectaries, monoecious euphorbia flowers possess only the most essential parts of reproduction. However, bracts, extra-floral nectaries, and other structures surrounding the reproductive parts of the flowers make them appear superficially different.

It would be very time consuming to sequence the DNA of every member of this genus to see where they all fit. Approximately 10% of the euphorbias have been phylogenetically examined, and they confirm the traditional morphological placement. How about that?

Interestingly, of the species genetically analyzed, some were subsequently placed into the genus Euphorbia after historically being considered members of other genera.

Euphorbia horrida and Euphorbia obesa

So? What’s that mean?

Species within the same genus when crossed can (but not always) produce viable offspring. Sometimes they don’t because of differences in pollinators, flowering times, or geographic location, which prevents hybridization. Clades within plant genera also can affect intra-genus reproduction. For example, hard maples won’t naturally hybridize with soft maples, despite both being in the genus Acer. Perhaps the case is similar between the groups within Euphorbia.

As a plant collector and cacti and succulent enthusiast, imagining the endless amounts of hybrids within a massive genus is a fancy idea to me. The APG’s confirming of the initial classifications of Euphorbia into a massive genus makes the idea of endless hybrids all the more real.

Additional guest posts by Jeremiah Sandler:

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Jeremiah Sandler lives in southeast Michigan, has a degree in horticultural sciences, and is an ISA certified arborist. Follow him on Instagram: @j.deepsea