Flowers in the aster family have one of the most recognizable shapes in botany – a circle with a series of petals surrounding it. If you were asked to draw a flower, there is a good chance your drawing would look something like a sunflower, a daisy, a cosmos, or an aster. It’s one of the most basic flower shapes, and yet it isn’t a single flower; it’s a pseudanthium – a false flower. This is because what might appear as a single flower is actually a collection of tens, hundreds, or even thousands of tiny flowers. This aggregation of flowers into a single compact unit is the reason the family was once given the name Compositae, and even now is often informally referred to as the composites.
Another reason why a flower in the aster family – or Asteraceae – might be the first thing you would draw is because it is the largest family of flowering plants, numbering up to 33,000 species worldwide. Chances are you’ve seen a few of them around. In the contiguous U.S. alone, there are more than 2400 species, and that doesn’t include the plethora of species brought in from regions across the world either intentionally (to be grown in our gardens) or unintentionally (as weeds). Of course, not all of the species in this family are going to have a typical sunflower-like flower head, but they do all have a specific type of inflorescence called a capitulum. Capitula are made up of densely packed, miniature flowers called florets, which are stalkless (or sessile) and arranged on a flattened central stem (or axis). There are at least four different types of florets in the aster family, but we’ll leave that discussion for another time.
In this post, we’re specifically interested in what is happening at the base or underside of the capitulum. All of the florets in a capitulum are held within a cup or bowl-shaped series of bracts called an involucre. Bracts are modified leaves, and this whorl of tightly held or loosely arranged bracts are initially found surrounding a developing flower bud. As the inflorescence opens, the involucre opens as well and its bracts persist at the base of the flower head. The bracts that make up the involucre are called phyllaries, and they vary in shape, number, and size depending on plant species. In fact, the features of phyllaries are so unique they are often relied on to help identify a plant in the aster family to genus, species, and infraspecies (variety, subspecies, etc.).
phyllaries of blanketflower (Gaillardia aristata)
When it comes to flowers in the aster family, there is more than meets the eye. After you take some time to appreciate the intricate beauty of its collection of florets, turn the flower head over and take in its phyllaries. They come in various colors, they can be hairy or smooth, their margins can be entire or adorned with hairs, teeth, etc., they can be flat and straight or they can curve outwards in interesting ways, their tips can be pointed, spine-tipped, rounded, or keel shaped. Phyllaries can be laid out very evenly, tightly overlapping each other like shingles on a roof (i.e. imbricate) or their arrangement can be slightly uneven and irregular (i.e. subimbricate). Use a hand lens to get a closer look at all of these features. As you get in the habit of observing the details of the involucre and its phyllaries, chances are each time you come across a flower in the aster family, you’ll find yourself flipping it over to get a look at its undercarriage. What will you find?
phyllaries of dandelion (Taraxacum officinale)
phyllaries of Mexican sunflower (Tithonia rotundifolia)
phyllaries of stemless four-nerve daisy (Tetraneuris acaulis)
phyllaries of hoary tansyaster (Dieteria canescens)
phyllaries of aromatic aster (Symphyotrichum oblongifolium ‘October Skies’)
phyllaries of curlycup gumweed(Grindelia squarrosa)
If phyllaries have piqued your interest and you’d like to learn more about plants in the family Asteraceae, I highly recommend getting your hands on the book, The Sunflower Family by Richard Spellenberg and Naida Zucker. It has a North American focus, but it’s a great place to start learning more about this massive plant family.
While we’re on the subject of pollen-gluing mechanisms, there is another material apart from pollenkitt that a limited number of flowering plant families use to link their pollen grains together. It functions, much like pollenkitt, by aiding in the attachment of pollen to visiting animals. However, unlike pollenkitt, it isn’t sticky, oily, or viscous, and is instead more like a series of threads. Viscin threads to be exact.
One of the major differences between pollenkitt and viscin threads is their composition. The lipid-rich coating that surrounds pollen grains, which we call pollenkitt, is derived from breakdown materials of an inner layer of the anther. It is added to pollen grains after they are formed and before the anther dehisces. Viscin threads are made up of sporopollenin, the same biopolymer that exine (the outer wall of a pollen grain) is composed of. Viscin threads have points of attachment on an outer layer of the exine called the ektexine. Unlike pollenkitt, viscin threads don’t add new color to pollen grains, nor do they contain scent compounds. Their thickness, length, abundance, and texture are dependent on the species of plant they are found on, much like pollenkitt varies in form and composition depending on species.
pollen strands of tufted evening primrose (Oenothera caespitosa)
Viscin threads evolved independently in three distantly related plant families. These include Onagraceae (the evening primrose family), Ericaceae (the heath family), and a subfamily in the pea family known as Caesalpinioideae (the peacock flower subfamily). Viscin threads are found in many, but not all, of the species in these three families. Some species in other plant families have what appear to be viscin threads but are actually ropy strands of pollenkitt, as they are composed of pollenkitt and not sporopollenin. Because they are made up of the same durable material as exine, viscin threads can be preserved in the fossil record. A paper published in Grana (1996) looked at the morphology of pollen grains with viscin threads from the Tertiary Period and concluded that “this advanced pollination syndrome using viscin threads as a pollen connecting agent” dates back to at least the Eocene and perhaps much earlier.
While pollenkitt’s stickiness adheres pollen grains together, viscin threads are more of a tangling device. Single pollen grains or pollen grain groupings called tetrads become tangled up together and then become entangled with a visiting insect, bird, or bat and carried away to a nearby flower. Disentanglement from the pollinator ideally happens when the threads are brushed against the sticky surface of a stigma. The viscin threads themselves vary by species and family. Micheal Hesse, in a paper published in Grana (1981), describes the threads in Onagraceae as “long, numerous, thin, and sculptured” with “knobs, furrows, etc.,” while those in Ericaceae are thin and smooth and those in Caesalpinioideae are thick and smooth.
smooth azalea, pink form (Rhododendron arborescens)
The length and size of tangled pollen masses also differ by species and can offer clues as to which pollinators visit which flowers. Research published in New Phytologist (2019) looked at the size of pollen thread tangles (PTT) in 13 different species of Rhododendron. They also noted which pollinators visited each species and how often they visited. The researchers found that species presenting pollen in small but abundant PTT were visited by bees, and those with large but few PTT were visited by birds and Lepidoptera (butterflies and moths). Bees also visited the flowers more frequently than birds and Lepidoptera. Bees collect and consume pollen. Between visits to anthers, they spend time grooming themselves, removing pollen clusters from their bodies and packing them into corbiculae (i.e. pollen baskets) for later*. Birds and Lepidoptera don’t groom pollen from their bodies and don’t collect it. In the authors terms, this “suggests pollinator-mediated selection on pollen packaging strategies.” Since flowers pollinated by bees lose much of their pollen in the process, they present it in smaller packages, and since flowers pollinated by birds and Lepidoptera are visited less frequently, their pollen packages are larger.
This is an example of the pollen presentation theory, and is something we will revisit as the Year of Pollination continues.
*This applies specifically to bee species that have corbiculae, and many bee species do not.
Pollination in flowering plants is the process of moving pollen grains, which carry sperm cells, from the anthers to the stigmas of either the same flower or a separate flower. If things go well from there, sperm cells will be transported via pollen tubes into the ovaries where fertilization with egg cells can take place and seeds can form. Pollen grain development occurs within the anthers, and by the time the anthers dehisce – or split open – they are ready for transport.
In order to protect the enclosed sperm cells and aid in their movement, pollen grains consist of a series of layers that, among other things, help ensure safe travel. Two major layers are an internal layer called intine, composed largely of cellulose, and an external layer called exine, composed mainly of sporopollenin (a highly durable and complex biopolymer). In many flowering plants, especially those that rely on animals to help carry their pollen, an additional outer layer called pollenkitt is added to the pollen grains before anthers dehisce.
three different pollen grains (image credit: wikimedia commons/Asja Radja)
Pollenkitt is an oily, viscous, hydrophobic layer composed of lipids, carotenoids, flavonoids, proteins, and carbohydrates derived from the breakdown of an internal layer of the anther called the tapetum. Pollenkitt forms a sticky layer around the pollen grains and can add color to the pollen other than the typical yellow. The thickness of the pollenkitt and its composition is species specific. In fact, the look, size, and shape of pollen grains themselves are unique to each species and can even be used to help identify plants. Pollenkitt is found in almost all families of flowering plants and is particularly prevalent in species that are animal-pollinated. One exception is the mustard family (Brassicaceae), whose pollen grains are coated in a substance known as tryphine, which functions similar to pollenkitt but whose formation and composition differ enough to be considered separately.
The sticky nature of pollenkitt has numerous functions. For one, it helps pollen grains remain on anthers until an animal comes along to remove them. It also holds pollen grains together in clumps, helps pollen grains stick to insect (and other animal) pollinators during transport, and helps adhere them to stigmas when deposited. A paper published in Flora (2005) lists twenty possible functions for pollenkitt, many of which have been confirmed in certain species and some of which are hypothetical. In addition to functions having to do with pollen movement and placement, pollenkitt may also provide protection from water loss, UV radiation, and fungal and bacterial invasions. In species where pollen is offered as food to pollinating insects, pollenkitt is a more easily digestible food source than the pollen grain itself. Thanks to carotenoids, pollenkitt can make pollen more colorful, which may help attract pollinating insects, or, depending on the color, can also hide pollen from insect visitors.
Another important function of pollenkitt is to give pollen a scent. Odors can help encourage insect visitors or deter them, so depending on the situation, scented pollenkitt may be attracting pollinators or discouraging pollen consumers. In a study published in American Journal of Botany (1988), Heidi Dobson analyzed the chemical composition of 69 different species of flowering plants. She isolated numerous scent compounds in pollenkitt and suggested that “some of the chemicals in pollenkitt may … serve as identification cues to pollen-foraging bees.” Most of the species she analyzed were pollinated by bees (which consume pollen), but the few that were mainly pollinated by hummingbirds and butterflies tended to have fewer scent compounds. Since birds and butterflies are there for the nectar and not the pollen, it would make sense that the pollen of these plant species wouldn’t need to carry a scent.
In flowers that are wind-pollinated, the pollenkitt layer is either very thin or absent altogether. In this case, pollen grains need to be easily released from the anther and are better off when they aren’t sticking to other pollen grains. That way, they are free to be carried off in the breeze to nearby flowers. Some plant species are amphiphilous, meaning they can be both animal-pollinated and wind-pollinated, and according to the authors of the paper published in Flora (2005), pollenkitt layers in these species exhibit intermediate characteristics of both types of pollen grains, generally with thinner, less-sticky pollenkitt and more pollenkitt found within the cavities of the exine.
It’s clear that this unique pollen-glueing substance plays a critical role in the pollination process for many plant species. Considering that each species of plant has its own story to tell, there is still more to learn about the forms and functions that pollenkitt takes.
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This is the first in a series of posts in 2024 in which, once again, I am exploring the world of pollinators and pollination. You can read more about this effort in last month’s Year in Review post.
Sweetgum is a distinctive tree at any time of year. It stands out among most other deciduous trees with its excurrent growth habit, which gives it a narrowly conical or pyramidal shape. Only in its advanced age does it begin to develop a more rounded and irregular form. Its leaves are star-shaped with either five or seven lobes and span between four and seven inches. Their glossy green color gives way to a wide range of colors in the fall, including yellow, orange, red, and maroon, remaining on the tree for several weeks depending on weather conditions. The fruits are particularly distinct, especially in the winter after the leaves have dropped. Woody balls made up of a series of capsules with pointed beaks hang from long stalks throughout the winter, windborne seeds having been released from small openings in the fall. Without even bothering to look at the twigs, seeing these durable, spiky balls hanging from branches (or feeling them under your feet) is a quick indication that you are looking at a sweetgum.
one of manyfall colors found on sweetgum (Liquidambar styraciflua)
Liquidamabar styraciflua (one of the funnest botanical names to say) was previously placed in the family Hamamelidaceae but is now one of the few members of Altingiaceae. Its natural distribution is broad, covering a large portion of the eastern United States and west into Missouri then down into Texas, Mexico, and much of Central America. Outside of its natural range it has been widely planted as an ornamental, and there are several popular sweetgum cultivars currently in cultivation. Both the common and botanical names for this tree refer to the resin found in its bark, which historically has had many uses.
The winter twigs of sweetgum are stout, round, smooth, and yellow-brown to green or olive-green to brown-purple in color. They can also be glossy and feature a few scattered lenticels. Older twigs (or branchlets) are brown at first and then light grey with dark grey lenticels. They are occasionally adorned with corky wings similar to those of bur oak. Because the wings aren’t always present, it can be a fun thing to encounter when you are out looking at twigs.
corky wings on the branchlets of sweetgum
The winter buds of sweetgum twigs are egg-shaped and made up of bud scales with acute tips and ciliate margins. They are green to orange-brown or reddish in color and occasionally sticky. Lateral buds are alternately arranged, are much smaller than terminal buds, and are stalkless and flattened against the twig. They sit above a slightly raised leaf scar that is half-elliptical to triangular in shape and has three distinct vascular bundle traces. The pith of sweetgum twigs is solid, continuous, brownish, and irregularly shaped.
winter twig of sweetgum tree
Sweetgum bark is light to dark grey and is made up of a series of rough, vertically arranged, scaly ridges that become deeply furrowed with age. The mace-like fruits of sweetgum are about one inch wide and, at a glance, are similar in appearance to the seed balls of sycamore trees; however, sycamore seed balls easily break apart when compressed, while the pointed, woody capsules that make up a sweetgum ball are held firmly together and can hold their shape for long periods of time. When these “gumballs” collect on the ground below, they can become a hazard, especially where there is lots of foot traffic. Speaking from experience, they are also obnoxious when operating a mower. This polarizing feature has resulted in bad opinions of the sweetgum tree. Luckily, some people are out there defending it.
Things were pretty quiet on the blog in 2023, and I apologize for that. I have no excuses really. It’s just life. Fewer posts doesn’t mean I’m any less committed to writing and sharing about plants since the day I started this project, it’s more about quality over quantity. I would never want this to become a half-hearted affair, so even if months go by without hearing from me, just know that there are great things in the works, which I hope will be worth the wait.
Recently, while writing an article for Wildflowermagazine, I came across this giant tome, Pollination and Floral Ecology by Pat Willmer. The previous year I had read Jeff Ollerton’s book, Pollinators and Pollination, and really got a lot out of it. These incredible resources on the science of pollination reminded me of a time early in Awkward Botany’s history in which I spent a year posting about pollination. I called it Year of Pollination, and by the time the year came to a close, I was struck by how much I still wanted to share about this topic. So now, armed with these new resources, I think it’s time for Another Year of Pollination.
In 2024, I plan on posting another series of pollinator and pollination themed posts. I may not be able to match what I accomplished in 2015, but I will aim for at least one a month. Just something to look forward to in the coming year.
If this entices you enough to continue to follow Awkward Botany (or to start), please do. Relevant links are here on my linktree. Awkward Botany can be found on a number of different social media platforms, but there are a few that I am more active on than others. With the fall of Twitter, I have moved on to other things. This is where you can find me most often at this point in time:
In part one of this two part series, I introduced you to the Boise State University campus, located in the heart of Boise, Idaho. I’ve been spending the past year walking the campus and cataloging the weeds that I find there. Boise has a fairly mild climate compared to the rest of Idaho, so weeds are generally easy to find just about any time of year. What weeds are present depends on what time of year it is. To get a complete picture of the suite of weeds that can be found on a site, it’s important to make observations throughout the year. Weeds can also come and go, with certain species becoming more abundant in some years than others, so making observations over multiple years also helps. This is why I try to update posts that are part of the Weeds of Boise series as I make return visits and encounter additional weed species.
What follows is the second half of the list of weeds I’ve documented so far at Boise State University. I’m including a photograph for each month of the year (July – December), as well as a list of what I’ve encountered up to this point. I’m also including a list of weeds that I didn’t come across but that are documented on iNaturalist.
birdsfoot trefoil (Lotus corniculatus) at BSU in July 2023
yellow nutsedge (Cyperus esculentus) at BSU in August 2023
velvetleaf (Abutilon theophrasti) at BSU in September 2023
chicory (Cichorium intybus) at BSU on October 2023
puncturevine (Tribulus terrestris) at BSU in November 2023
bull thistle (Cirsium vulgare) at BSU in December 2023
Additional weeds found on the BSU campus from July – December 2023:
Abutilon theophrasti (velvetleaf)
Cichorium intybus (chicory)
Cirsium vulgare (bull thistle)
Cyperus esculentus (yellow nutsedge)
Eragrostis cilianensis (stinking lovegrass)
Lotus corniculatus (birdsfoot trefoil)
Medicago sativa (alfalfa)
Melilotus alba (white sweetclover)
Solanum nigrum (black nightshade)
Sonchus asper (prickly sowthistle)
Tribulus terrestris (puncturevine)
Additional weeds observed on the BSU campus by iNaturalist users as of December 2023:
lf you’ve seen one fireweed, you’ve probably seen several. As an early successional species, growing in large numbers across a vast amount of space is kind of its thing. Any disturbance that leaves bare ground in its wake, such as a wildfire or a windstorm, gives fireweed the opportunity to colonize. It grows quickly and spreads via rhizomes, producing thousands of airborne seeds in the process, sending them off to continue colonization or contribute to soil seed banks in preparation for future disturbances. The role of plants like fireweed is vital – promptly covering bare ground to stave off erosion and acting as a nurse plant to new saplings destined to become the future forest. In a garden setting or in locations where aggressively spreading plants are discouraged, fireweed and its weedy behavior may be unwelcome, but in other contexts, its services are essential.
fireweed (Chamerion angustifolium)
Fireweed (Chamerion angustifolium) is a species in Onagraceae, commonly known as the evening primrose family. It has an impressive distribution, widespread across much of North America and Eurasia. This is owed largely to its adaptability. Deep shade and overly dry soil are two conditions that it does not tolerate well, otherwise it seems to grow in a wide variety of soil types, moisture levels, and sun exposures, particularly in areas where there is regular disturbance. Swaths of towering plants topped with rose-pink flower spikes make fireweed impossible to ignore and a favorite of wildflower enthusiasts.
Fireweed stems reach from three to nine feet tall and are rarely branched. Long, narrow, lance-shaped leaves are arranged alternately along the lengths of stems and give the plant a willow-like appearance, which explains another common name, rosebay willowherb. The undersides of leaves have a distinct venation pattern, in which the veins don’t reach the leaf margins, a feature that can help with identification.
distinct leaf veins of fireweed(Chamerion angustifolium)
A series of rose-pink to purple flowers top the stems of fireweed. Each flower has four sepals and four petals with eight stamens and a four-lobed stigma extending prominently from the center. Its long, narrow ovary can be confused for a flower stalk. Rich with nectar, fireweed flowers are a favorite of honeybee keepers. They are also edible, like much of the rest of the plant. Narrow, four-chambered capsules form in place of fertilized flowers and later split open to release abundant, small seeds with a tuft of fluff attached to each one to aid in wind dispersal.
Fireweed has a long history of being used as food and medicine. Stem fibers are also useful for making cord, and seed fluff is useful for weaving and padding. Certainly, fireweed’s abundance and ubiquity contribute to its utility. Having never eaten fireweed before, I decided that a good way to introduce it to my diet would be to make a tea. Fireweed leaves are commonly collected for tea and are said to make an excellent non-caffeinated replacement for black tea.
fireweed tealeaves
Making fireweed tea starts by stripping young leaves from fireweed stems. Recipes I encountered all called for fermenting the leaves before drying them. I did this by squeezing handfuls of leaves in my fists just enough to break and bruise them a bit and then packing them into a quart size Mason jar. I closed the lid tight and kept them in the jar for about five days, shaking it up a couple times a day supposedly to help prevent mold issues. After that, I dried the leaves on a baking sheet in the hot sun. From there, they are ready for making tea the same way you would make any other loose leaf tea, chopping the leaves up a bit before immersing them in hot water.
I found the taste of fireweed tea to be mild and pleasant. Despite several sources comparing it to black tea, I thought it was more similar to green tea. Sierra liked the smell more than the taste and wished it had honey in it. Compared to other teas I’ve tried in this short series of posts, this is definitely one of the better ones, and a tea I could see myself making again sometime.
If you live in a major city (or even a minor one), there is a good chance it is home to a college or university (perhaps several). Universities tend to take up a lot of space, which means there is often a plethora of landscaping accompanying their buildings, hardscaping, and other impervious surfaces. Among all the turf, flower beds, tree wells, and other greenspaces, there is bound to be a fair share of weeds. In spite of how hard the groundskeepers may work, the campus is not likely to ever be completely weed-free. Lucky for us, this means that institutions of higher learning are excellent places to familiarize ourselves with many of the weed species that occur in our cities, particularly weeds that are common in garden beds and turfgrass.
Near downtown Boise, on the southside of the Boise River, you will find the ever-expanding campus of Boise State University, home of the Broncos and their famous blue turf. According to the internet’s favorite encyclopedia, the campus is 285 acres in size, plenty of space for weeds to grow and abudant opportunities to hunt them out. Tallying the number of weed species in a place like this takes time. The benefit of botanizing for weeds is that you can find them at just about any time of year. While some species only show up in certain seasons, others can be seen practically year-round.
In order to document the weeds of Boise State University, I’m spending the entire year walking the campus listing and photographing the weeds I find. What follows is the first half of what’s been documented so far. I’m including a photograph for each month of the year, as well as a list of what I’ve encountered. In part two, I’ll share a list of any additional weeds found throughout the remainder of the year. While you’re waiting for that, check out the other posts in the Weeds of Boise series.
common groundsel (Seneciovulgaris) at BSU in January 2023
chickweed (Stellaria media) at BSU in February 2023
hairy bittercress (Cardamine hirsuta) at BSU in March 2023
ivyleaf speedwell (Veronica hederifolia) at BSU in April 2023
black medic (Medicago lupulina) at BSU in May 2023
creeping thistle (Cirsium arvense) at BSU in June 2023
List of weeds found on the campus of Boise State University as of June 2023:
Ailanthus altissima (tree of heaven)
Anthriscus caucalis (bur chervil)
Amaranthus retroflexus (redroot pigweed)
Arctium minus (lesser burdock)
Bassia scoparia (kochia)
Bromus tectorum (cheatgrass)
Capsella bursa-pastoris (shepherd’s purse)
Cardamine hirsuta (hairy bittercress)
Ceratocephala testiculata (bur buttercup)
Chenopodium album (lamb’s quarters)
Chondrilla juncea (rush skeletonweed)
Cirsium arvense (creeping thistle)
Claytonia perfoliata (miner’s lettuce)
Convolvulus arvensis (field bindweed)
Conyza canadensis (horseweed)
Descurainia sophia (flixweed)
Digitaria sanguinalis (crabgrass)
Draba verna (spring draba)
Epilobium ciliatum (willowherb)
Erodium cicutarium (redstem filare)
Euphorbia maculata (spotted spurge)
Galium aparine (cleavers)
Geum urbanum (herb Bennet)
Holosteum umbellatum (jagged chickweed)
Hordeum jubatum (foxtail barley)
Lactuca serriola (prickly lettuce)
Lamium purpureum (purple deadnettle)
Lepidium sp. (whitetop)
Malva neglecta (common mallow)
Medicago lupulina (black medic)
Oxalis corniculata (creeping woodsorrel)
Parthenocissus quinquefolia (Virginia creeper)
Plantago lanceolata (narrowleaf plantain)
Plantago major (broadleaf plantain)
Poa annua (annua bluegrass)
Poa bulbosa (bulbous bluegrass)
Polygonum aviculare (prostrate knotweed)
Portulaca oleracea (purslane)
Prunella vulgaris (self-heal)
Ranunculus repens (creeping buttercup)
Senecio vulgaris (common groundsel)
Sonchus sp. (sow thistle)
Stellaria media (chickweed)
Taraxacum officinale (dandelion)
Tragopogon dubius (salsify)
Trifolium repens (white clover)
Ulmus pumila (Siberian elm)
Veronica hederifolia (ivyleaf speedwell)
Vulpia myuros (rat’s tail fescue)
Do you frequent the BSU campus? Have you seen anything not on my list? Comment below or send me a message and let me know what you’ve seen and where.
Have you ever tried to identify a grass? Most of us who like to look at plants and learn their names will probably admit that we often give up on grasses pretty quickly, or just ignore them entirely. They aren’t the easiest plants to identify to species, and there are so many of them. Without close inspection, they can all look pretty similar. Their flowers aren’t particularly showy, and their fruits are fairly forgettable. They are strands or clumps of green that create a backdrop for more intriguing forms of vegetation. Yet, they are among the most ecologically and economically important groups of plants on the planet. And actually, if you can ascend the hurdles that come with getting to know them, they are beautiful organisms and really quite amazing.
Kōura in the Grass
The grass family – Poaceae – consists of nearly 8oo genera and about 12,000 species. Grasses occur in a wide range of habitats across the globe. Wherever you are on land, a grass is likely nearby. Grasses play vital roles in their ecosystems and, from a human perspective, are critical to life as we know it. We grow them for food, use them for building materials and fuel, plant them as ornamentals, and rely on them for erosion control, storm water management, and other ecosystem services. We may not acknowledge their presence most of the time, but we very likely wouldn’t be here without them.
The sheer number of grass species is one thing that makes them so difficult to identify. Key identifying features of grasses and grass-like plants (also known as graminoids) tend to be very small and highly modified compared to similar features on other flowering plants. This requires using a hand lens and learning a whole new vocabulary in order to begin to understand a grass’s anatomy. It’s a time commitment that goes beyond a lot of other basic plant identification, and it’s a learning curve that few dare to follow. However, once you learn the basic features, it becomes clear that grasses are relatively simple organisms, and once you start identifying them, it can actually be an exciting and rewarding experience.
Quackgrass (Elymus repens) and Its Rhizome
Depending on the species, grasses can be annuals – completing their life cycle within a single year – or perennials – coming back year after year for two or more years. Most grasses have a fibrous root system; some are quite shallow and simple while others are extremely deep and extensive. Some species of perennial grasses spread by either rhizomes (underground stems), stolons (horizontal, above ground stems), or both. Some grasses also produce tillers, which are essentially daughter plants that form at the base of the plant. The area where roots, rhizomes, stolons, and tillers meet the shoots and leaves of a grass plant is called the crown. This is an important region of the plant, because it allows for regrowth even after the plant has been browsed by a grazing animal or mown down by a lawn mower.
The stem or shoot of a grass is called a culm. Leaves are formed along the lengths of culms, and culms terminate in inflorescences. Leaves originate at swollen sections of the culm called nodes. They start by wrapping around the culm and forming what is called a leaf sheath. Leaves of grasses are generally long and narrow with parallel venation – a trait typical of monocotyledons. The part of the leaf that extends away from the culm is called the leaf blade or lamina. Leaves are alternatively arranged along the length of the stem and are two-ranked, meaning they form two distinct rows opposite of each other along the stem.
The area where the leaf blade meets the leaf sheath on the culm is called the collar. This collar region is important for identifying grasses. With the help of a hand lens, a closer look reveals the way in which the leaf wraps around the culm (is it open or closed?), whether or not there are hairs present and what they are like, if there are auricles (small flaps of leaf tissue at the top of the collar), and what the ligule is like. The ligule is a thin membrane (sometimes a row of hairs) that forms around the culm where the leaf blade and leaf sheath intersect. The size of the ligule and what its margin is like can be very helpful in identifying grasses.
The last leaf on the culm before the inflorescence is called the flag leaf, and the section of the culm between the flag leaf and the inflorescence is called a peduncle. Like the collar, the flower head of a grass is where you’ll find some of the most important features for identification. Grass flowers are tiny and arranged in small groupings called spikelets. In general, several dozen or hundreds of spikelets make up an inflorescence. They can be non-branching and grouped tightly together at the top of the culm, an inflorescence referred to as a spike, or they can extend from the tip of the culm (or rachis) on small branches called pedicels, an inflorescence referred to as a raceme. They can also be multi-branched, which is the most common form of grass inflorescence and is called a panicle.
Either way, you will want to take an even closer look at the individual spikelets. Two small bracts, called glumes, form the base of the spikelet. Above the glumes are a series of florets, which are enclosed in even smaller bracts – the outer bract being the lemma and the inner bract being the palea. Certain features of the glumes, lemmas, and paleas are specific to a species of grass. This includes the way they are shaped, the presence of hairs, their venation, whether or not awns are present and what the awns are like, etc. If the grass species is cleistogamous – like cheatgrass – and the florets never open, you will not get a look at the grass’s sex parts. However, a close inspection of an open floret is always a delight. A group of stamens protrude from their surrounding bracts bearing pollen, while feathery stigmas reach out to collect the pollen that is carried on the wind. Depending on the species, an individual grass floret can have either only stamens, only pistils (the stigma bearing organs), or both. Fertilized florets form fruits. The fruit of a grass is called a caryopsis (with a few exceptions) and is indistinguishable from the seed. This is because the seed coat is fused to the wall of the ovary, unlikeother fruit typesin which the two are separate and distinct.
If all this doesn’t make you want to run outside and take a close look at some grasses, I don’t know what will. What grasses can you identify in your part of the world? Let me know in the comment section below or check out the linktree and get in touch by the means that suits you best.
To understand the current state of rangeland wildfires in the Intermountain West, you must first familiarize yourself with a plant commonly referred to as cheatgrass. This annual grass moved into the region over a century ago, and its spread has had a massive impact on the environment, as well as the economy and our way of life. Just the very mention of cheatgrass in the West will get some people’s blood boiling. It’s a menace, a scourge, a pest, and yet it’s here to stay. It’s a result of us being here, yet somehow it’s the invader. Its success is largely due to the way we’ve chosen to operate in this region, yet it’s the one to blame for our troubles. When you really start to learn about this plant, it’s hard not to develop an appreciation for it, despite the tragic ways in which it has shaped our region for the worse. It’s not a plant that is showy or grandiose in any significant way. Everything about its appearance screams for it to be dismissed and overlooked, yet it’s story – at least here in the American West – is larger than life.
cheatgrass (Bromus tectorum)– illustration credit:Selected Weeds of the United States, Agriculture Handbook No. 366 (ARS/USDA)
Bromus tectorum goes by more than a dozen common names, but the ones you tend to hear most often are downy brome and cheatgrass. Downy because of how fuzzy its leaf blades can be and cheat because its presence on wheat farms cheats farmers of their yield. It is distributed widely across Europe, eastern Asia, and northern Africa where it originates, and was introduced to North America in the mid-19th century. How and why it got here isn’t totally clear. It likely had multiple introductions, both as a contaminant in seeds and attached to fur, clothing, packaging materials, etc., as well as intentionally as a forage crop for livestock. Regardless, it managed to establish readily in the east and then quickly spread across the country, spanning the continent by the early 20th century. It found the Great Basin particularly habitable due to its hot, dry summers and cold, wet winters and largely treeless landscape.
Apart from the climate, a significant factor behind cheatgrass’s establishment in the Intermountain West are all the cows. For a number of reasons, the Great Basin isn’t really suitable for largescale farming operations, but livestock grazing is another story. Many of the animals native to the region are grazing animals after all, so why not graze cattle and sheep? But there is a limit. Too many animals stuck in one spot for too long leads to overgrazing, and overgrazed sites take time for the native vegetation to recover. Cheatgrass exploits this opportunity by establishing itself quickly in disturbed and overgrazed locations and begins the process of outcompeting nearby plants for limited water and nutrients. Once it begins to dominate these sites, it has another trick up its sleeve.
Cheatgrass actually makes good forage for livestock early in the spring when it’s green and tender, but that quickly changes as the plants start to dry out and go to seed. By early summer, cheatgrass has completed its lifecycle and what’s left is a dried-up plant that, due to the silica in its cells, does not break down readily. Where cheatgrass is abundant, this means large swaths of standing brown grass as far as the eye can see. What’s more, this dead vegetation is highly flammable, and the slightest spark can set off a roaring blaze that moves quickly across the landscape, igniting everything in its path. In a region where fires once occurred decades apart, they now occur on a nearly annual basis. And because fire had been historically infrequent, the native vegetation is not adapted to regular fire and can take years to recover, whereas cheatgrass bounces right back, again exploiting the void left by the decimation of native plants and is flowering again the following spring. It’s a self-perpetuating cycle, and cheatgrass excels at it.
cheatgrass on fire
Cheatgrass is a winter annual, meaning that it germinates in the fall as soon as moisture becomes available. It then lies mostly dormant, its shallow, fibrous roots still growing as long as the ground isn’t frozen. Employing this strategy means cheatgrass is ready to resume growth at a quick pace as soon as the weather warms in the spring. Its roots spread horizontally in the soil and essentially rob water from nearby, more deeply rooted native vegetation. Its deep green, hairy leaves form a little tuft or rosette and provide early spring forage for livestock, gamebirds, and other grazing animals. As the spring progresses flower stalks form and the plants reach heights of around 2 feet (60 centimeters). Their inflorescence is a prominently drooping, open panicle and each spikelet has between 4-8 florets, each with a single, straight awn. The flowers of cheatgrass are cleistogamous, which means they don’t ever open. Self-pollination occurs inside the closed floret, and viable seeds soon develop. As the plant matures, it takes on a purple-reddish hue, after which it turns crispy and light brown as the seeds disperse.
The stiff awns remain on the seeds and aid in dispersal. They also cause injury to animals that dare consume them, poking into the soft tissues of their mouths. Passing animals are also injured when the awns work their way into their feet, ears, and other vulnerable body parts. The ability of the awns to attach so easily to fur and clothing is one of the reasons why cheatgrass spreads so readily. Wind also helps distribute the seed. A single plant can produce hundreds, if not thousands, of seeds, which are ready to germinate upon dispersal. They remain viable in the soil for only a few short years, but since they germinate so easily and are produced so abundantly, their short lifespan isn’t much of a downside.
dried inflorescence of cheatgrass (Bromus tectorum)
In many ways, cheatgrass is the perfect weed. It is able to grow under a broad range of conditions. Its seeds germinate readily, and the plant grows during a time when most other plants have gone dormant. It excels at capturing water and nutrients. It self-pollinates and produces abundant viable seed, which are reliably and readily dispersed thanks to persistent awns. Disturbed areas are ripe for a plant like cheatgrass, but even nearby undisturbed areas can be invaded as seeds are dispersed there. With the help of fire, cheatgrass also creates its own disturbance, which it capitalizes on by then growing even thicker, more abundant stands with now even less competition from native vegetation. And because it is available so early in the season and is readily consumed by livestock and gamebirds, what motivation is there for humans to totally replace it with something else? As James Young and Charlie Clements ask in their book, Cheatgrass, “How can we come to grips with the ecological and economic consequences of this invasive alien species that can adapt to such a vast range of environmental conditions?” In another section they lament, “cheatgrass represents a stage in transition toward an environment dominated by exotic weeds growing on eroded landscapes.”
The topic of cheatgrass and other introduced annual grasses, as well as the even broader topic of rangeland wildfires, is monstrous, but it is one that I hope to continue to cover in a series of posts over the coming months and years. It’s not an easy (or necessarily fun) thing to tackle, but it’s an important one, especially for those of us who call the cheatgrass sea our home.
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