The Dispersal of Ancient and Modern Apples by Humans and Other Megafauna

Crop domestication often involves selection for larger fruits. In some crops, humans took plant species with relatively small fruits and, over many generations of artificial selection, developed a plant with much larger fruits. Consider giant pumpkins as an extreme example. Yet in the case of apples, relatively large fruits already existed in the wild. Producing larger apples happened quickly and, perhaps even, unconsciously. Apples were practically primed for domestication, and as Robert Spengler explains in a paper published last year in Frontiers in Plant Science, looking back in time at the origins of the apple genus, Malus, can help us understand how the apple we know and love today came to be.

Apples are members of the rose family (Rosaceae), a plant family that today consists of nearly 5000 species. According to the fossil record, plants in the rose family were found in large numbers across North America as early as the Eocene (56 – 33.9 million years ago). They were present in Eurasia at this time as well, but Spengler notes, “there is a much clearer fossil record for Rosaceae fruits and seeds in Europe and Asia during the Miocene and Pliocene (20 – 2.6 million years ago).” Around 14 million years ago, larger fruits and tree-form growth habits evolved in Rosaceae subfamilies, giving rise to the genera Malus and Pyrus (apples and pears). Small, Rosaceae fruits were typically dispersed by birds, but as Sprengler writes, “it seems likely that the large fruits [in Malus and Pyrus] were a response to faunal dispersers of the late Miocene through the Pliocene of Eurasia.” Larger animals were being recruited for seed dispersal in a changing landscape.

Glacial advances and retreats during the Pleistocene (2.6 million – 11,700 years ago) brought even more changes. Plants with effective, long distance seed dispersal were favored because they were able to move into glacial refugium during glacial advances. Even today, these glacial refugium are considered genetic hot spots for Malus, and could be useful for future apple breeding. As the Pleistocene came to a close, many megafauna were going extinct. This continued into the Holocene. Large-fruited apple species lost their primary seed dispersers, and their ranges became even more contracted.

Humans have had an extensive relationship with apples, which began long before domestication. Foraging for apples was common, and seeds were certainly spread that way (perhaps even intentionally). Favorable growing conditions were also created when forests were cleared and old fields were left fallow. Apple trees are early successional species that easily colonize open landscapes, gaps in forests, and forest edges, so human activity that would have created such conditions “could have greatly promoted the spread and success of wild Malus spp. trees during the Holocene.”

The earliest evidence we have of apple domestication (in which “people were intentionally breeding and directing reproduction”) occurred around 3000 years ago in the Tian Shan Mountains of Kazakhstan, where Malus sieversii – a species that is now facing extinction – was being cultivated. This species was later brought into contact with other apple species, a few of which were also being cultivated, including M. orientalis, M. sylvestris, and M. baccata. These species easily hybridized, giving us the modern, domesticated apple, M. domestica. As Spengler writes, “the driving force of apple domestication appears to have been the trans-Eurasian crop exchange, or the movement of plants along the Silk Road.” Continued cultivation and further hybridization among M. domestica cultivars over the past 2000 years has resulted in thousands of different apple varieties.

The unique thing about domesticated apples is that their traits are not fixed in the same way that traits of other domesticated crops are. Growing an apple from seed will result in a very different apple than the apple from which the seed came. Apple traits instead have to be maintained through cloning, which is accomplished mainly through cuttings and grafting. Apples hybridize with other apple species so readily that most apple trees found in the wild are hybrids between wild and cultivated populations.

Spengler considers the study of apple domestication to be “an important critique of plant domestication studies broadly, illustrating that there is not a one-size fits-all model for plant domestication.” The “key” for understanding apple domestication “rests in figuring out the evolutionary driver for large fruits in the wild – seed dispersal through megafaunal mammals – and the process of evolution for these large fruits – hybridization.” He notes that “domestication studies often ignore evolutionary processes leading up to human cultivation,” which, in the case of apples, involves “hybridization events in the wild” that led to the evolution of large fruits “selected for through the success in recruiting large megafaunal mammals as seed disperses.” Many of those mammals went extinct, but humans eventually assumed the role, selecting and propagating “large-fruiting hybrids through cloning and grafting – creating our modern apple.”

Excerpt from Fruit from the Sands by Robert N. Spengler:

Indeed, the relationship between apples and people is close and complex, spanning at least five millennia. The story of the apple begins along the Silk Road… In recent years genetic studies have resolved much of the debate over these origins. Nevertheless, the ancestry of the apple is highly complex. Cloning, inbreeding, and reproduction between species have created a genealogy that looks more like a spider’s web than a family tree. To growers, the beauty of the apple lies not in its rosy skin but in its genetic variability and plasticity, its ability to cross with other species of Malus and other distant lines of M. domestica, and the ease with which it can be grafted onto different rootstocks and cloned.

See Also: Science Daily – Exploring the Origins of the Apple

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Interested in learning more about how plants get around. Check out the first issue of our new zine Dispersal Stories.

Book Review: Fruit from the Sands

“By dispensing plants and animals all around the world, humans have shaped global cuisines and agricultural practices. One of the most fascinating and least-discussed episodes in this process took place along the Silk Road.” — Fruit from the Sands by Robert N. Spengler III

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My understanding of the origins of agriculture and the early years of crop domestication are cursory at best. The education I received was mostly concerned with the Fertile Crescent, as well as crops domesticated by early Americans. The Silk Road, as I understood it, was the route or routes used to move goods across Asia and into eastern Europe well after the domestication of many of the crops we know today. Other than the fact that several important crops originate there, little was ever taught to me about Central Asia and its deep connection to agriculture and crop domestication. I suppose that’s why when I picked up Fruit from the Sands by Robert N. Spengler III, published last year by University of California Press, I wasn’t entirely prepared for what I was about to read.

It wasn’t until I read a few academic reviews of Fruit from the Sands that I really started to understand. Spengler’s book is groundbreaking, and much of the research he presents is relatively new. The people of Central Asia played a monumental role in discovering and developing much of the food we grow today and some of the techniques we use to grow them, and a more complete story is finally coming to light thanks to the work of archaeologists like Spengler, as well as advances in technology that help us make sense of their findings.

This long history with agricultural development can still be seen today in the markets of Central Asia, which are loaded with countless varieties of fruits, grains, and nuts, many of which are unique to the area. Yet this abundance is also at risk. Crop varieties are being lost at an alarming rate with the expansion of industrial agriculture and the reliance on a small selection of cultivars. With that comes the loss of local agricultural knowledge. Yet, with climate change looming, diversity in agriculture is increasingly important and one of the tools necessary for maintaining an abundant and reliable food supply. Unveiling a thorough history of our species’ agricultural roots will not only give us an understanding as to how we got here, but will also help us learn from past successes and failures. Hence, the work that Spengler and others in the field of archaeobotany are doing is crucial.

To set the stage for a discussion of “the Silk Road origins of the foods we eat,” Spengler offers his definition of the Silk Road. The term is misleading because there isn’t (and never was) a single road, and the goods, which were transported in all directions across Asia, included much more than silk. In fact, some of what was transported wasn’t a good at all, but knowledge, culture, and religion. In Spengler’s words, “The Silk Road … is better thought of as a dynamic cultural phenomenon, marked by increased mobility and interconnectivity in Eurasia, which linked far-flung cultures….This network of exchange, which placed Central Asia at the center of the ancient world, looked more like the spokes of a wheel than a straight road.” Spengler also sees the origins of the Silk Road going back at least five thousand years, much earlier than many might expect.

Most of Spengler’s book is organized into chapters discussing a single crop or group of crops, beginning with grains (millet, rice, barley, wheat) then moving on to fruits, nuts, and vegetables before ending with spices, oils, and teas. Each chapter compiles massive amounts of research that can be a bit overwhelming to take in all at once. Luckily each section includes a short summary, which nicely distills the information down into something more digestible.

Spengler’s chapter on broomcorn millet (Panicum miliaceum) is particularly powerful. While in today’s world millet has largely “been reduced to a children’s breakfast food in Russia and bird food in Western Europe and America,” it was “arguably the most influential crop of the ancient world.” Originally domesticated in East Asia, “it passed along the mountain foothills of Central Asia and into Europe by the second millennium BC.” It is a high-yielding crop adapted to hot, dry conditions that, with the development of summer irrigation, could be grown year-round. These and other appealing qualities have led to an increase in the popularity of millet, so much so that 2023 will be the International Year of Millets.

The “poster child” for Spengler’s book may very well be the apple. Popular the world over, the modern apple began its journey in Central Asia. As Spengler writes, “the true ancestor of the modern apple is Malus sieversii,” and “remnant populations of wild apple trees survive in southeastern Kazakhstan today.” As the trees were brought westward, they hybridized with other wild apple species, bringing rise to the incredible diversity of apple cultivars we know today. Sadly though, most of us are only familiar with the small handful of common varieties found at our local supermarkets.

Of course, as Spengler says, “No discussion of plants on the Silk Road would be complete without the inclusion of tea (Camellia sinensis),” a topic that could produce volumes on its own. Despite the brevity of the section on tea, Spengler has some interesting things to say. One in particular involves the transport of tea to Tibet in the seventh century, where “an unquenchable thirst for tea” had developed. But the journey there was long and difficult. Fermented and oxidized tea leaves traveled best. Along the way, “the leaves were exposed to extreme cold as well as hot and humid temperatures in the lowlands, and all the time they were jostled on the backs of sweaty horses and mules.” This, however, only improved the tea, as teas exposed to such conditions “became a highly sought-after commodity among the elites.”

“In Central Asia, Mongolia, and Tibet, tea leaves were oxidized, dried, and compressed into hard bricks from which chunks could be broken off and immersed in water.” – Robert N. Spengler III in Fruit from the Sands (photo via wikimedia commons)

As dense as this book is, it’s also quite approachable. The information presented in each of the chapters is thorough enough to be textbook material, but Spengler does such a nice job summing up the main points, that there are plenty of great takeaways for the casual reader. For those wanting a deeper dive into the history of our food (which in many ways is the history of us), Spengler’s book is an excellent starting point.

More Reviews of Fruit from the Sands:

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Ground Beetles as Weed Seed Predators

As diurnal animals, we are generally unaware of the slew of animal activity that occurs during the night. Even if we were to venture out in the dark, we still wouldn’t be able to detect much. Our eyes don’t see well in the dark, and shining a bright light to see what’s going on results in chasing away those creatures that prefer darkness. We just have to trust that their out there, and in the case of ground beetles, if they’re present in our gardens we should consider ourselves lucky.

Ground beetles are in the family Carabidae and are one of the largest groups of beetles in the world with species numbering in the tens of thousands. They are largely nocturnal, so even though they are diverse and relatively abundant, we rarely get to see them. Look under a rock or log during the day, and you might see a few scurry away. Or, if you have outdoor container plants, there may be a few of them hiding out under your pots with the pillbugs. At night, they leave the comfort of their hiding places and go out on the hunt, chasing down grasshoppers, caterpillars, beetle grubs, and other arthropods, as well as slugs and snails. Much of their prey consists of common garden pests, making them an excellent form of biological control. And, as if that weren’t enough, some ground beetles also eat the seeds of common weeds.

Harpalus affinis via wikimedia commons

Depending on the species, a single ground beetle can consume around a dozen seeds per night. In general, they prefer the seeds of grasses, lambsquarters (Chenopodium album), pigweeds (Amaranthus spp.), and various plants in the mustard family (Brassicaceae). The seeds of these species are small with seed coats that are easily crushed by a beetle’s mandibles. Providing suitable habitat, avoiding insecticides, and minimizing soil disturbance (i.e. reducing or eliminating tillage) are ways that healthy ground beetle populations can be encouraged and maintained. Ground beetles prefer dense vegetation where they can hide during the daytime. Strips of bunchgrasses and herbaceous perennials planted on slightly raised bed (referred to as beetle banks) are ideal because they provide good cover and keep water from puddling up in the beetles’ hiding spots.

The freshness of weed seeds and the time of year they are available may be determining factors in whether or not ground beetles will help control weed populations. A study published in Weed Science (2014), looked at the seed preferences of Harpalus pensylvanicus, a common species of ground beetle that occurs across North America. When given the choice between year old seeds and freshly fallen seeds of giant foxtail (Setaria faberi), the beetles preferred the fresh ones. The study also found that when giant foxtail was shedding the majority of its seeds, the density of beetles was on the decline, meaning that, at least in this particular study, most of the seeds would go uneaten since fewer beetles were around when the majority of the seeds were made available. Creating habitat that extends the ground beetles’ stay is important if the goal is to maximize the number of weed seeds consumed.

Harpalus pensylvanica via wikimedia commons

Of course, the seeds of all weed species are not considered equal when it comes to ground beetle predation. Several studies have sought to determine which species ground beetles prefer, offering seeds of a variety of weeds in both laboratory and field settings and seeing what the beetles go for. Pinning this down is difficult though because there are numerous species of ground beetles, all varying in size and activity. Their abundances vary from year to year and throughout the year, as do their food sources. Since most of them are generalists, they will feed on what is available at the time. A study published in European Journal of Entomology (2003) found a correlation between seed size and body mass – small beetles were consuming small seeds and large beetles were consuming large seeds, relatively speaking.

Another study published in European Journal of Entomology (2014) compared the preferences of ground beetles in the laboratory to those in the field and found that, in both instances, the seeds of field pansy (Viola arvensis) and shepherd’s purse (Capsella bursa-pastoris) were the preferred choice. The authors note that both species have lipid-rich seeds (or high “energy content”). Might that be a reason for their preference? Or maybe it’s simply a matter of availability and “the history of individual predators and [their] previous encounters with weed seed.” After all, V. arvensis was “the most abundant seed available on the soil surface” in this particular study.

Pterostichus melanarius via wikimedia commons

A study published in PLOS One (2017), looked at the role that scent might play in seed selection by ground beetles. Three species of beetles were offered the seeds of three different weed species in the mustard family. The seeds of Brassica napus were preferred over the other two by all three beetle species. The beetles were also offered both imbibed and non-imbibed seeds of all three plants. Imbibed simply means that the seeds have taken in water, which “can result in the release of volatile compounds such as ethanol and acetaldehyde.” The researchers wondered if the odors emitted from the imbibed seeds would “affect seed discovery and ultimately, seed consumption.” This seemed to be the case as all three beetle species exhibited a preference for the imbibed seeds.

Clearly, ground beetles are fascinating study subjects, and there is still so much to learn about them and their eating habits. If indeed their presence is limiting the spread of weeds and reducing weed populations, they should be happily invited into our farms and gardens and efforts should be made to provide them with quality habitat. For a bit more about ground beetles, check out this episode of Boise Biophilia.

Further Reading:

From Cut Flower to Noxious Weed – The Story of Baby’s Breath

One of the most ubiquitous plants in cut flower arrangements hails from the steppes of Turkey and neighboring countries in Europe and Asia. It’s a perennial plant with a deep taproot and a globe-shaped, multi-branched inflorescence loaded with tiny white flowers. In full bloom it looks like a small cloud hovering above the ground. It’s airy appearance earns it the common name baby’s breath, and the attractive and durable nature of its flowers and flower stalks, both fresh and dried, have made it a staple in the floral industry. Sadly, additional traits have led to it becoming a troublesome weed outside of its native range.

baby’s breath (Gypsophila paniculata) via wikimedia commons

Gypsophila paniculata is in the family Caryophyllaceae – sharing this distinction with other cut flowers like carnations and pinks, as well as other weeds like chickweed and soapwort. At maturity and in full bloom, baby’s breath might reach three to four feet tall; however, its thick taproot extends deep into the ground as much as four times its height. Its leaves are unremarkable and sparse, found mostly towards the base of the plant and sometimes with a blue or purplish hue. The flowers are numerous and small, have a sweet scent to them (though not appreciated by everyone), and are pure white (sometimes light purple or pink).

Each flower produces just a few seeds that are black, kidney-shaped, and minuscule. Many of them drop from their fruits and land near their parent plant, but some are retained within their little capsules as the flower stalk dries and becomes brittle. Eventually a stiff breeze knocks the entire inflorescence loose and sends it tumbling across the ground. Its rounded shape makes it an effective tumbleweed, as the remaining seeds are shaken free and scattered far and wide.

baby’s breath flowers close up (via wikimedia commons)

Being a tumbleweed gives it an advantage when it comes to dispersing itself and establishing in new locations, but this is not the only trait that makes baby’s breath a successful weed. Its substantial taproot, tolerance to drought and a variety of soil conditions, and proclivity to grow along roadsides, in ditches, and abandoned fields also make it a formidable opponent. Mowing the plant down does little to stop it, as it grows right back from the crown. Best bets for control are repeated chemical treatments or digging out the top portion of the taproots. Luckily its seeds are fairly short-lived in the soil, so vigilant removal of seedlings and not allowing the plant to reproduce can help keep it in check. Baby’s breath doesn’t persist in regularly disturbed soil, so it’s generally not a problem in locations that are often cultivated like agricultural fields and gardens.

The first introductions of baby’s breath to North America occurred in the 1800’s. It was planted as an ornamental, but it wasn’t long before reports of its weedy nature were being made. One source lists Manitoba in 1887 as the location and year of the first report. It is now found growing wild across North America and is featured in the noxious weed lists in a few states, including Washington and California. It has been a particular problem on sand dunes in northwest Michigan, where it has been so successful in establishing itself that surveys have reported that 80% of all vegetation in certain areas is composed of baby’s breath.

baby’s breath in the wild (via wikimedia commons)

Invading sand dune habitats is particularly problematic because extensive stands of such a deep-rooted plant can over-stabilize the soil in an ecosystem adapted to regular wind disturbance. Plants native to the sand dunes can be negatively affected by the lack of soil movement. One species of particular concern is Pitcher’s thistle (Cirsium pitcheri), a federally threatened plant native to sand dunes along the upper Great Lakes. Much of the research on the invasive nature of baby’s breath and its removal comes from research being done in this region.

Among numerous concerns that invasive plants raise are the affects they can have on pollinator activity. Will introduced plants draw pollinators away from native plants or in some other way limit their reproductive success? Or might they help increase the number of pollinators in the area, which in turn could benefit native plants (something known as the magnet species effect)? The flowers of baby’s breath rarely self-pollinate; they require insect visitors to help move their pollen and are highly attractive to pollinating insects. A study published in the International Journal of Plant Sciences found that sand dune sites invaded by baby’s breath attracted significantly more pollinators compared to uninvaded sites, yet this did not result in more pollinator visits to Pitcher’s thistle. According to the researchers, “a reduction in pollinator visitation does not directly translate to a reduction in reproductive success,” but the findings are still a concern when it comes to the future of this threatened thistle.

Perhaps it’s no surprise that a plant commonly found in flower arrangements is also an invasive species, as so many of the plants we’ve grown for our own pleasure or use have gone on to cause problems in areas where they’ve been introduced. However, could the demand for this flower actually be a new business opportunity? Noxious weed flower bouquets anyone?

Related Posts:

Drought Tolerant Plants: Yellowhorn

A drought tolerant garden doesn’t have to be treeless. While the pickings are slim, there is a selection of trees that, once established, are well adapted to deal with extended bouts of little to no water. One such tree is yellowhorn, a species that demands to be considered for any waterwise landscape. Yellowhorn is rare in cultivation – and also restricted in its natural distribution – but perhaps that will change as word gets around about this beautiful and resilient tree.

Xanthoceras sorbifolium is native to several provinces in northern China and has been cultivated in a number of places outside of China since at least the 1800’s. Its ethnobotanical value is well understood in China. Its leaves, flowers, and seeds are edible and medicinal, and the high oil content of its seeds make them useful for the production of biofuels. Researchers are also investigating the use of yellowhorn for ecological restoration in arid habitats where desertification is a concern.

yellowhorn in bloom

Yellowhorn is the only species in the genus Xanthoceras, but is one in a long list of trees and shrubs in the Sapindaceae family – a family that now includes maples and horse chestnuts. It is considered both a large shrub and a small, multi-stemmed tree. It reaches a maximum height of about 25 feet, but arrives there at a relatively slow pace. It tolerates a variety of soil types, but like most other drought tolerant plants, it prefers soils that don’t become waterlogged easily. Its leaves are long, glossy green, and compound, consisting of 9 – 17 leaflets. The leaves persist late into the year and turn yellow in the fall. However, late spring, when the tree is covered in flowers, is when this tree puts on its real show.

Large white flowers with yellow-green centers that turn maroon or red-orange as they age are produced on racemes at the ends of branches. Small, yellow, hornlike appendages between each of the five petals of the flowers are what gives the tree its common name. Flowering lasts for a couple weeks, after which fruits form, which are about 2.5 inches wide, tough, leathery, and somewhat pear shaped. In my experience, most of the fruits are eaten by squirrels long before they get a chance to reach maturity. The ones the squirrels don’t get will persist on the tree, harden, and eventually split open to reveal several large, dark, round seeds nestled in chambers within the fruit.

To truly appreciate this tree, it must be seen in person, especially in bloom. At that point you will demand to have one (or more) in your garden. The seeds are said to be delicious, so you should give them a try if you can beat the squirrels to them. For a more thorough overview of yellowhorn, check out this article from Temperate Climate Permaculture, and for more photos of yellowhorn in bloom, check out this post from Rotary Botanical Gardens.

Squirrel nesting in yellowhorn, getting ready to go after more fruits.

All photos in this post were taken at Idaho Botanical Garden in Boise, Idaho.

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More Drought Tolerant Plants Posts:

Idaho’s Native Milkweeds (Updated)

As David Epstein said in an interview on Longform Podcast, “Any time you write about science, somethings is going to be wrong; the problem is you don’t know what it is yet, so you better be ready to update your beliefs as you learn more.” Thanks to the newly published Guide to the Native Milkweeds of Idaho by Cecilia Lynn Kinter, lead botanist for Idaho Department of Fish and Game, I’ve been made aware of some things I got wrong in the first version of this post. I appreciate being corrected though, because I want to get things right. What follows is an updated version of the original post. The most substantial change is that there are actually five milkweed species native to Idaho rather than six. Be sure to check out Kinter’s free guide to learn more about this remarkable group of plants.

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Concern for monarch butterflies has resulted in increased interest in milkweeds. Understandably so, as they are the host plants and food source for the larval stage of these migrating butterflies. But milkweeds are an impressive group of plants in their own right, and their ecological role extends far beyond a single charismatic insect. Work to save the monarch butterfly, which requires halting milkweed losses and restoring milkweed populations, will in turn provide habitat for countless other organisms. A patch of milkweed teems with life, and our pursuits to protect a single caterpillar invite us to explore that.

Asclepias – also known as the milkweeds – is a genus consisting of around 140 species, 72 of which are native to the United States and Canada. Alaska and Hawaii are the only states in the U.S. that don’t have a native species of milkweed. The ranges of some species native to the United States extend down into Mexico where there are numerous other milkweed species. Central America and South America are also home to many distinct milkweed species. Asclepias species found in southern Africa are considered by many to actually belong in the genus Gomphocarpus.

The habitats milkweeds occupy are about as diverse as the genus itself – from wetlands to prairies, from deserts to forests, and practically anywhere in between. Some species occupy disturbed and/or neglected sites like roadsides, agricultural fields, and vacant lots. For this reason they are frequently viewed as a weed; however, such populations are easily managed, and with such an important ecological role to play, they don’t deserve to be vilified in this way.

Milkweed species are not distributed across the United States evenly. Texas and Arizona are home to the highest diversity with 37 and 29 species respectively. Idaho, my home state, is on the low end with five native species. The most abundant species found in Idaho is Asclepias speciosa, commonly known as showy milkweed.

showy milkweed (Asclepias speciosa)

Showy milkweed is distributed from central U.S. westward and can be found in all western states. It occurs throughout Idaho and is easily the best place to look for monarch caterpillars. In fact, the monarch butterfly is Idaho’s state insect, thanks in part to the abundance of showy milkweed, which is frequently found growing in large colonies due to its ability to reproduce vegetatively via adventitious shoots produced on lateral roots or underground stems. Only a handful of milkweed species reproduce this way. Showy milkweed reaches up to five feet tall and has large ovate, gray-green leaves. Like all milkweed species except one (Asclepias tuberosa), its stems and leaves contain milky, latex sap. In early summer, the stems are topped with large umbrella-shaped inflorescences composed of pale pink to pink-purple flowers.

The flowers of milkweed deserve a close examination. Right away you will notice unique features not seen on most other flowers. The petals of milkweed flowers bend backwards, which would otherwise allow easy access to the flower’s sex parts if it wasn’t for a series of hoods and horns protecting them. Collectively, these hoods and horns are called the corona, which houses glands that produce abundant nectar and has a series of slits where the anthers are exposed. The pollen grains of milkweed are contained in waxy sacs called pollinia. Two pollinia are connected together by a corpusculum giving this structure a wishbone appearance. An insect visiting the flower for nectar slips its leg into the slit, and the pollen sacs become attached with the help of the corpusculum. When the insect leaves, the pollen sacs follow. Pollination is successful when the pollen sacs are inadvertently deposited on the stigmas of another flower.

Milkweed flowers are not self-fertile, so they require assistance by insects to sexually reproduce. They are not picky about who does it either, and their profuse nectar draws in all kinds of insects including bees, butterflies, moths, beetles, wasps, and ants. Certain insects – like bumble bees and other large bees – are more efficient pollinators than others. Once pollinated, seeds are formed inside a pod-like fruit called a follicle. The follicles of showy milkweed can be around 5 inches long and house dozens to hundreds of seeds. When the follicle matures, it splits open to release the seeds, which are small, brown, papery disks with a tuft of soft, white, silky hair attached. The seeds of showy milkweed go airborne in late summer.

follicles forming on showy milkweed (Asclepias speciosa)

Whorled or narrowleaf milkweed (Asclepias fascicularis) occurs across western and southern Idaho. Its distribution continues into neighboring states. It is adapted to dry locations, but can be found in a variety of habitats. Like showy milkweed, it spreads rhizomatously as well as by seed. It’s a whispy plant that reaches one to three feet tall and occasionally taller. It has long, narrow leaves and produces tight clusters of greenish-white to pink-purple flowers. Its seed pods are long and slender and its seeds are about 1/4 inch long.

flowers of narrowleaf milkweed (Asclepias fascicularis)

seeds escaping from the follicle of narrowleaf milkweed (Asclepias fascicularis)

Swamp or rose milkweed (Asclepias incarnata) is more common east of Idaho, but occurs occasionally in southwestern Idaho. As its common name suggests, it prefers moist soils and is found in wetlands, wet meadows, and along streambanks. It can spread rhizomatously, but generally doesn’t spread very far. It reaches up to four feet tall, has deep green, lance-shaped leaves, and produces attractive, fragrant, pink to mauve, dome-shaped flower heads at the tops of its stems. Its seed pods are narrow and around 3 inches long.

swamp milkweed (Asclepias incarnata)

Asclepias cryptoceras ssp. davisii, or Davis’s milkweed, is a low-growing, drought-adapted, diminutive species that occurs in southwestern Idaho. It has round or oval-shaped leaves and produces flowers on a short stalk. The flowers have white or cream-colored petals and pink-purple hoods. The range of Asclepias cryptoceras – commonly known as pallid milkweed or jewel milkweed – extends beyond Idaho’s borders into Oregon and Nevada, creeping north into Washington and south into California. Another subspecies – cryptoceras – can be found in Nevada, Utah, and their bordering states.

Davis’s milkweed (Asclepias cryptoceras ssp. davisii)

The final species is rare in Idaho, as Idaho sits at the top of its native range. Asclepias asperula ssp. asperula, or spider milkweed, has a single documented location in Franklin County (southeastern Idaho). Keep your eyes peeled though, because this plant may occur elsewhere, either in Franklin County or neighboring counties. It grows up to two feet tall with an upright or sprawling habit and produces clusters of white to green-yellow flowers with maroon highlights. Its common name comes from the crab spiders frequently found hunting in its flower heads.

A sixth species, horsetail milkweed (Asclepias subverticillata), has been falsely reported in Idaho. Collections previously labeled as A. subverticillata have been determined to actually be the similar looking A. fascicularis.

Using Weeds: Soapwort

Over the past year or so I have written about several edible weeds in an effort to highlight useful weeds. However, weeds don’t have to be edible to be useful. In fact, many weeds are most certainly not edible, but that doesn’t mean they are of no use to humans. Soapwort, for example, is poisonous, and while it does have a history of being used internally as medicine, ingesting it is not advised and should only be done under the direction of a doctor. A much less risky activity would be to make soap out of it.

soapwort (Saponaria officinalis)

Saponaria officinalis, commonly known as bouncing bet, hedge pink, fuller’s herb, scourwort, and soapweed or soapwort, is an herbaceous perennial native to Europe. It has been planted widely in flower beds and herb gardens outside of its native range, desired both for its beauty and utility. Capitalizing on our appreciation for it, soapwort has expanded beyond our garden borders and into natural areas, as well as vacant lots, roadsides, and other neglected spaces. Even in a garden setting it can be a bit of a bully, especially if ignored for a season or two.

The stems of soapwort grow to about two feet tall, are unbranched, and sometimes tinged with pink, purple, or red. The leaves are oblong and oppositely-arranged, and their bases form prominent collars around the stems. Showy clusters of flowers are found atop the stems throughout the summer. Like other flowers in the pink family (Caryophyllaceae), they are cigar-shaped at the base and opened wide at the end, showing off 5 distinct petals with notches at their tips. The petals of soapwort flowers bend backwards, with their sex parts protruding outwards. In his description of the flowers, John Eastman remarks in The Book of Field and Roadside that “the reflexed petals surrounding the sexual organs give the impression of flagrant thrust; this is a gaudy, unshy flower.”

collared stem of soapwort (Saponaria officinalis)

The fragrant flowers are pink to white in color. They open in the evening and remain open for a few short days. In an individual flower, pollen matures and is mostly shed before the stigma is ready to accept it. This helps reduce the chance of self-pollination. Cross pollination occurs with the assistance of moths who visit the flowers at night, as well as bees and other flower-visiting insects that come along during the daytime. Soapwort fruits are oval capsules containing as many as 500 kidney-shaped seeds. Seeds aren’t essential to the plants spread though, as much of its colonization occurs via vigorous rhizomes.

In fact, vegetative reproduction is the means by which soapwort forms such expansive, thick patches. It also helps that it’s poisonous. The saponins – its soap making compounds – that it produces in its roots, shoots, and leaves deter most insects and other animals from eating it. It has a reputation for poisoning horses, cows, and other livestock, and so is unwelcome in pastures and rangelands. Saponins are also poisonous to fish, so growing soapwort near fish ponds is not advised.

soapwort (Saponaria officinalis)

Soapwort occurs in a variety of soils including sandy, dry, and rocky sites and is surprisingly drough-tolerant, fine qualities to have when colonizing neglected sites. While most other organisms ignore soapwort, it has a friend in humans. Eastman sums this up well: “Soapwort’s most important associate – as is true of most plants we label weeds – is undoubtedly humankind, without whose helpful interventions the plant would surely be much rarer than it is.”

I made a soapy liquid out of soapwort by following a recipe that can be found on various blogs and websites by searching “saponaria soap recipe.” Basically it’s a cup of fresh leaves and stems along with a cup of dried leaves and stems added to a quart of distilled water brought to a boil. After simmering for 15 minutes and then allowing it to cool, strain the mixture through cheese cloth, and it’s ready to go.

This gentle but effective soap can be used for cleaning countertops and other surfaces, as well as dishes, fabrics, and skin. Several sources say it is particularly useful for cleaning delicate fabrics. Sierra and I both found it to have a cooked cabbage or spinach scent to it. This can be masked by adding a few drops of essential oil. Despite its odd aroma, both Sierra and I were impressed by its cleansing power and plan to use it more often.

dried leaves of soapwort

soapwort soap