When Acorn Masts, Rodents, and Lyme Disease Collide

“‘Mast years’ is an old term used to describe years when beeches and oaks set seed. In these years of plenty, wild boar can triple their birth rate because they find enough to eat in the forestes over the winter… The year following a mast year, wild boar numbers usually crash because the beeches and oaks are taking a time-out and the forest floor is bare once again.” — The Hidden Life of Trees by Peter Wohlleben

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When a plant population’s annual production of seeds is highly variable and synchronous, it is considered a masting or mast seeding species. Why and how masting happens is a bit of a mystery, and efforts are underway to better understand this phenomenon. One thing is clear, boom and bust cycles can have dramatic effects on animals that use the fruits and seeds of these plants for food. Acorn production in oaks provides a stark example. As Koenig, et al. describe in Ecology (2015), a “variable acorn crop initiates a ‘chain reaction’ of responses that cascades through the ecosystem, affecting densities of deer, mice, ground-nesting birds, gypsy moths, and the tick vectors of Lyme disease.” The connection between mast seeding oaks and the prevalence of tick-borne pathogens is of particular interest considering the risks posed to humans.

Lyme disease is an infectious diesease caused by a bacterium vectored by ticks in the genus Ixodes. The life-cycle of a tick is generally 2 to 3 years, beginning after a larva hatches from an egg. From there the larva develops into a nymph and later an egg-laying adult, taking a blood meal each step of the way. Tick larvae feed on the blood of small rodents and birds, which is where they can pick up the bacterium that causes Lyme disease. After feeding, they develop into nymphs and go in search of another blood meal, perhaps another rodent or maybe something larger like a deer or a human. It is in their nymphal and adult stages that ticks transmit Lyme disease to humans. Nymphs tend to transmit the disease more frequently, partly because they go undetected more easily.

The risk to humans of being infected with Lyme disease varies year to year and is dependent largely on how many infected ticks are present. For this reason, it is important to understand the factors affecting the density of infected nymphs. In a study published in PLoS Biology (2006), Ostfeld, et al. collected data over a 13 year period in plots located in deciduous forests in the state of New York, a hotspot for Lyme disease. The predictors they considered included temperature, precipitation, acorn crop, and deer, white-footed mouse, and chipmunk abundances. Deer abundance and weather conditions had long been considered important in predicting the prevalence of ticks, but little attention had been paid to small mammals – the larval hosts for ticks – and the variability of acorn crops – an important food source for rodents.

deer tick (Ixodes scapularis) — via PhyloPic; user Mathilde Cordellier

The results of their study revealed a clear pathway – more acorns leads to more rodents which leads to more Lyme disease carrying nymphs. The process takes a couple of years. First, oak trees experience a mast year, flooding rodent populations with food. In the following year, the numbers of mice and chipmunks is unusually high. The year after that, there are lots and lots of nymphal ticks infected with Lyme-disease. The relationship is so direct that Richard Ostfeld claims, based on his research, that he can predict the incidence of Lyme disease among residents of New York and Connecticut based on when a mast year occurs. In a summer when there is an abundance of 2 -year-old oak seedlings in the surrounding forests, expect the infection rate of Lyme disease to be high.

Lyme disease also occurs in regions where oak trees are not present or are uncommon, so variability in acorn crops isn’t always the best predictor. The researchers acknowledge that acorn abundance is not going to be “a universal predictor of risk;” instead, anything that leads to an increase in rodent populations, whether it is some other food source or a lack of predators, may be a key indicator since rodents are reservoir hosts of Lyme disease.

A study published in Parasites and Vectors (2020) looked at the effects of rodent density on a number of tick-borne pathogens. They confirmed that an “increase in rodent density positively affects populations of nymphal ticks in the following year;” yet, they could not confirm that rodent density is the sole predictor of disease risk. Other factors come into play depending on the disease in question, and further research is needed to improve models that predict tick-borne diseases. They did, however, confirm that, by flooding the food supply with acorns, mast years can boost populations of a variety of rodents.

white-footed mouse (Peromyscus leucopus) — via wikimedia commons; USGS

A fear of ticks is justified. They suck your blood after all, and besides that, they can transmit some pretty serious diseases. Arm yourself by educating yourself. One place to do that is with The Field Guides podcast. Their tick two-parter is well worth the listen (part one and part two). Not only will it give you valuable information in protecting yourself against ticks, it may also give you an appreciation for their prowess. Just maybe. See also their You Tube video demonstrating how to sample for ticks.

The Weeds in Your Bird Seed

With February comes the return of the Great Backyard Bird Count, a weekend-long, worldwide, bird counting event that Sierra and I have enjoyed participating in for the past few years. While you can choose to count birds anywhere birds are found, part of the appeal of the event is that it can be done from the comfort of one’s own home simply by watching for birds to appear right outside the window. If there are bird feeders in your yard, your chances of seeing birds are obviously improved. Watch for at least fifteen minutes, record the number and species of birds you see, then report your sightings online. It’s for science!

Feeding and watching birds are popular activities. In the United States alone, as many as 57 million households put out food for birds, spending more than $4 billion annually to do so. While there are a variety of things one can purchase to feed birds – suet, berries, mealworms, etc. – the bulk of that money is likely spent on bags of bird seed (also referred to as bird feed). Bird seed is a relatively cheap and easy way to feed a wide variety of birds. Unfortunately, it’s also a great way to introduce new weeds to your yard.

Bird seed contaminated with noxious weed seeds is not a new problem. It has been a concern for decades, and some countries have taken regulatory steps to address the issue. In the United States, however, there are no governmental regulations that address weed seed contamination in bird seed.  With this thought in mind, researchers at the University of Missouri screened a large sampling of bird seed mixes to determine the number and species of weed seeds they harbored, as well as their viability and herbicide resistance. Their results were published last year in Invasive Plant Science and Management.

The researchers examined 98 different bird seed mixes purchased from retail locations in states across the eastern half of the U.S. The seeds of 29 weed species were recovered from the bags, including at least eight species of grasses and several annual and perennial broadleaf weeds. 96% of the mixes contained one or more species of Amaranthus, including Palmer’s amaranth (Amaranthus palmeri), which was found in 27 mixes and which the researchers refer to as “the most troublesome weed species in agroecosystems today.” About 19% of amaranth seeds recovered germinated readily, and five of the seed mixes contained A. tuberculatus and A. palmeri seeds that, once grown out, were found to be resistant to glyphosate, the active ingredient in a commonly used herbicide.

Redroot pigweed (Amaranthus retroflexus) is one of several weedy amaranth species commonly found in bird seed mixes (illustration credit: wikimedia commons)

The seeds of grass weeds were found in 76% of the bird seed mixes and included three species of foxtail (Setaria spp.), as well as other common grasses like large crabgrass (Digitaria sanguinalis) and barnyardgrass (Echinochloa crus-galli). Bird seed ingredients that seemed to favor grass seed contamination included wheat, grain sorghum, and proso millet, three crops that are also in the grass family. No surprise, as grass weeds are difficult to control in crop fields when the crop being grown is also a grass.

After amaranths and grasses, ragweed (Ambrosia artemisiifolia) was the third most common weed found in the mixes. This was a troubling discovery since populations of this species have shown resistance to a number of different herbicides. Moving ragweed to new locations via bird seed could mean that the genes that give ragweed its herbicide resistance can also be moved to new locations. Kochia (Bassia scoparia), another weed on the Weed Science Society of America’s list of top ten most troublesome weeds, was also found in certain bird seed mixes, particularly when safflower was an ingredient in the feed.

A similar study carried out several years earlier at Oregon State University found the seeds of more than fifty different weed species in ten brands of bird feed commonly sold at retail stores. Ten of the weeds recovered from the mixes are on Oregon’s noxious weed list. Both studies demonstrate how bird seed can be a vector for spreading weed seeds – and even new weed species and herbicide-resistant genes – to new locations. Weeds found sprouting below bird feeders can then potentially be moved beyond the feeders by wind and other dispersal agents. Weed seeds might also be moved to new locations inside the stomachs of birds.

Addressing this issue can be tackled from several different angles. Growers and processors can improve their management of weed species in the fields where bird seed is grown and do a better job at removing weed seeds from the mixes after they are harvested. Government regulations can be put in place that restrict the type and quantity of weed seeds allowed in bird feed. Further processing of ingredients such as chopping or shelling seeds or baking seed mixes can help reduce the presence and viability of weed seeds.

Processed bird feed like suet is less likely to harbor viable weed seeds (photo credit: wikimedia commons)

Consumers can help by choosing bird feed that is processed or seedless like sunflower hearts, dried fruit, peanuts, suet cakes, and mealworms, and can avoid seed mixes with a large percentage of filler ingredients like milo, red millet, and flax. Attaching trays below feeders can help collect fallen seeds before they reach the ground. Bird seed can also be avoided all together, and feeding birds can instead be done by intentionally growing plants in your yard that produce food for birds. By including bird-friendly plants in your yard, you will also have a better chance of seeing a wide variety of birds during the Great Backyard Bird Count.

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Winter Trees and Shrubs: Northern Catalpa

The names of plants often contain clues that can either help with identification or that tell something about the plant’s history or use. The name, catalpa, is said to be derived from the Muscogee word, katałpa, meaning “winged head,” presumably referring to the tree’s winged seeds. Or maybe, as one writer speculates, it refers to the large, heart-shaped, floppy leaves that can make it look like the tree is “ready to take flight.” Or perhaps it’s a reference to the fluted, fused petals of the tree’s large, tubular flowers. I suppose it could mean any number of things, but I’m sticking with its seeds, which are packed by the dozens in the tree’s long, slender, bean-like fruits. The seeds are flat, pale brown, and equipped with paper thin, fringed appendages on either side that assist in wind dispersal – wings, in other words.

winged seeds of northern catalpa (Catalpa speciosa)

Catalpa speciosa, or northern catalpa, is a relatively fast growing, short-lived tree native to the Midwest and one of only two species in the genus Catalpa found in the United States. Its distribution prior to the arrival of Europeans appears to have been restricted to a portion of the central Mississippi River valley, extending west into Arkansas, east into Tennessee, and north into Illinois and Indiana. It has since been widely planted outside of its native range, naturalizing in areas across the Midwest and eastern US. Early colonizers planted northern catalpa for use as fence posts, railroad ties, and firewood. Its popularity as an ornamental tree is not what it once was a century ago, but it is still occasionally planted in urban areas as a shade tree. Its messiness – littering the ground below with large leaves, flowers, and seed capsules – and its tendency to spread outside of cultivation into natural areas are reasons why it has fallen out of favor with some people.

The oval to heart-shaped, 8 to 12 inch long leaves with long petioles rotting on the ground below the tree are one sure sign that you’ve encountered a catalpa in the winter time. The leaves are some of the first to fall at the end of the growing season, briefly turning an unmemorable yellow before dropping.

leaf of northern catalpa (Catalpa speciosa) in the winter with soft hairs on the underside still visible

The leaf arrangement on northern catalpa is whorled and sometimes opposite. The twigs are easy to identify due to several unique features. They are stout, round, and grayish brown with prominent lenticels. The leaf scars are large, rounded, and raised up on the twig, looking a bit like little suction cups. They are arranged in whorls of three, with one scar considerably smaller than the other two. A series of bundle traces inside the scar form an ellipse. The leaf buds are tiny compared to the scar and are protected by loose, pointed, brown bud scales. Northern catalpa twigs lack a terminal bud. In the winter, seed capsules or the stalk of an old inflorescence often remain attached to the terminal end of the twig. The pith inside of the twig is thick, white, and solid.

twig of northern catalpa (Catalpa speciosa)

pith inside twig of northern catalpa (Catalpa speciosa)

Another common name for Catalpa speciosa is cigar tree, a name that comes from its up to 18 inch long, cigar-like seed capsules that hang from the otherwise naked tree throughout the winter. The sturdy, cylindrical pod starts out green in the summer and turns dark brown by late fall. Seed pods that haven’t fallen or already split open will dehisce in the spring time, releasing their papery seeds to the wind.

fruits of northern catalpa (Catalpa speciosa) hanging from the tree in the winter

The young bark of northern catalpa is thin and easily damaged. As it matures, it becomes furrowed with either scaly ridges or blocky plates. Mature trees are generally twisted at the base but otherwise grow straight, reaching 30 to 60 feet tall (sometimes taller) with an open-rounded to narrow-oval crown.

maturing bark of northern catalpa (Catalpa speciosa)

Northern catalpa is one of the last trees to leaf out in the spring. In late spring or early summer, 10 inch long clusters of white, tubular flowers are produced at the tips of stems. Before the flowers open, they look a bit like popped popcorn, reminding me of a song from my childhood (which I will reluctantly leave right here). The margins of its trumpet-shaped petals are ruffled and there is yellow, orange, and/or purple spotting or streaking on the inside of the tubes.

flower of northern catalpa (Catalpa speciosa) just before it opens

More Winter Trees and Shrubs on Awkward Botany:

Seed Shattering Lost – The Story of Foxtail Millet

For a plant to disperse its seeds, it must first let go of them. Sounds obvious, but it is a key step in the dispersal process and an act that is actually coded in a plant’s DNA. As fruits ripen and seeds mature, an abscission layer is formed that separates the seed-bearing fruits from the plant. No longer attached to their parents, seeds are left to their own devices. If all goes well, they will find themselves in a suitable location where they can germinate and grow into a whole new plant, fully equipped to make seed babies of their own.

The releasing of mature seeds is known as shattering, a term most commonly used in reference to grasses and plants with dehiscent seed pods (i.e. fruits that split open when ripe, such as those in the bean and mustard families). In grasses, seeds form along a central stem called a rachis. As the seeds ripen, they separate from the rachis and drop from the plant. In some cases, the rachis is brittle and a section of it breaks off with each seed that falls.

Seed shattering is not a desirable trait when it comes to food crops. It’s easy to see how yields can be poor if seeds disperse before they are harvested. Thus, an essential step in domesticating certain agricultural crops was selecting plants that lacked this particular trait. Instead of dropping mature seeds, such plants hold on to them, making them easy to collect. A simple and naturally occurring mutation in the genes of these plants allowed early farmers to select varieties that were more ideal for agriculture than their wild progenitors.

Genetic studies of agricultural crops have located genes in a number of species that code for seed shattering, confirming that domestication in many cases involved selecting plants with this gene turned off. A recent study, published in Nature Biotechnology (October 2020), took a different route in locating this gene, looking instead at a weedy, wild relative of a crop that was domesticated at least 8000 years ago. Green foxtail (Setaria viridis) is the wild antecedent of foxtail millet (Setaria italica), a crop that, while still commonly grown for food in parts of Asia, is mostly grown for hay, silage, and bird seed in North America. Recently, interest in foxtail millet and other millets (a term used to refer to the grains of several different species of grasses) is on the rise due to the ability of these crops to tolerate drought and heat.

Illustration of three Setaria species from Selected Weeds of the United States (Agriculture Handbook No. 366) published in 1970

Setaria viridis is an abundant, widespread weed adapted to human disturbance. It’s of Eurasian origin but has been present in North America since the early 1800’s and was likely introduced both intentionally and accidentally. It’s an annual grass with prominent, bristly flowerheads that are easily recognizable and the reason for its common name, green foxtail. A handful of other closely related, similar-looking species are also common weeds in North America. Due to useful traits including its short life cycle, small genome, and self-fertility, S. viridis has been used frequently as a model species to carry out a variety of scientific studies. The study referred to above aimed to further enhance the use of green foxtail, particularly when it comes to crop science.

Researchers traveled across the United States collecting nearly 600 samples of green foxtail in order to better understand its genome. They found that the North American population of green foxtail is composed of multiple introductions and that, as the species has followed humans around, it has quickly adapted to diverse climates found across the continent. In examining the genome, they were able to identify the genetic underpinnings for three traits that have importance to agriculture: response to climate, leaf angle (which is used as a predictor of yield in grain crops), and seed shattering.

foxtail millet (Setaria italica) via wikimedia commons

The seed shattering gene – which the researchers named Less Shattering 1 (SvLes1) – was an especially interesting discovery. When compared to the orthologous gene found in foxtail millet, they found that a frameshift mutation had caused a disruption in the gene, turning it off. Using CRISPR (a gene editing tool) they were able to recreate a similar interruption in green foxtail, which resulted in a loss of seed shattering similar to that of foxtail millet. It became clear that selecting plants with this mutation was an essential step in the domestication of this ancient grain.

An excerpt about seed shattering from Fruit from the Sands by Robert N. Spengler III: 

In many of the world’s domesticated grains, especially those from the founder crops of southwest Asia (i.e. wheat and barley), the earliest phenotypical trait of domestication that archaeobotanists look for is a tough rachis, the small stem by which an individual grain or small cluster of grains is attached to the ear. In their wild form, most grains are programmed to detach easily after the grain ripens; however, in domesticated cereals, the grains remain attached to the ear throughout the harvesting process. This change is an inadvertent result of human harvesting with sickles: as people reap their harvest, the grains with a brittle rachis are dropped and those with a tough rachis are collected, stored, and replanted for successive harvests.

Further Reading:

The Hidden Flowers of Viola

Violas keep a secret hidden below their foliage. Sometimes they even bury it shallowly in the soil near their roots. I suppose it’s not a secret really, just something out of sight. There isn’t a reason to show it off, after all. Showy flowers are showy for the sole purpose of attracting pollinators. If pollinators are unnecessary, there is no reason for showy flowers, or to even show your flowers at all. That’s the story behind the cleistogamous flowers of violas. They are a secret only because unless you know to look for them, you would have no idea they were there at all.

Cleistogamy means closed marriage, and it describes a self-pollinating flower whose petals remain sealed shut. The opposite of cleistogamy is chasmogamy (open marriage). Most of the flowers we are familiar with are chasmogamous. They open and expose their sex parts in order to allow for cross-pollination (self-pollination can also occur in such flowers). Violas have chasmogamous flowers too. They are the familiar five-petaled flowers raised up on slender stalks above the green foliage. Cross-pollination occurs in these flowers, and seed-bearing fruits are the result. Perhaps as a way to ensure reproduction, violas also produce cleistogamous flowers, buried below their leaves.

an illustration of the cleistogamous flower of Viola sylvatica opened to reveal its sex parts — via wikimedia commons

Flowers are expensive things to make, especially when the goal is to attract pollinators. Colorful petals, nectar, nutritious pollen, and other features that help advertise to potential pollinators all require significant resources. All this effort is worth it when it results in the ample production of viable seeds, but what if it doesn’t? Having a method for self-pollination ensures that reproduction will proceed in the absence of pollinators or in the event that floral visitors don’t get the job done. A downside, of course, is that a seed produced via self-pollination is essentially a clone of the parent plant. There will be no mixing of genes with other individuals. This isn’t necessarily bad, at least in the short term, but it has its downsides. A good strategy is a mixture of both cross- and self-pollination – a strategy that violas employ.

The cleistogamous flowers of violas generally appear in the summer or fall, after the chasmogamous flowers have done their thing. The fruits they form split open when mature and deposit their seeds directly below the parent plant. Some are also carried away by ants and dispersed to new locations. Seeds produced in these hidden flowers are generally superior and more abundant compared to those produced by their showy counterparts. People who find violas to be a troublesome lawn weed – expanding far and wide to the exclusion of turfgrass – have these hidden flowers to blame.

That being said, there is a defense for violas. In the book The Living Landscape by Rick Darke and Doug Tallamy, Tallamy writes: “Plants such as the common blue violet (Viola sororia), long dismissed by gardeners as a weed, can be reconstituted as desirable components of the herbaceous layer when their ecosystem functionality is re-evaluated. Violets are the sole larval food source for fritillary butterflies. Eliminating violets eliminates fritillaries, but finding ways to incorporate violets in garden design supports fritillaries.”

sweet violet (Viola odorata)

In my search for the cleistogamous flowers of viola, I dug up a sweet violet (Viola odorata). I was too late to catch it in bloom, but the product of its flowers – round, purple, fuzzy fruits – were revealed as I uprooted the plant. Some of the fruits were already opening, exposing shiny, light brown seeds with prominent, white elaiosomes, there to tempt ants into aiding in their dispersal. I may have missed getting to see what John Eastman calls “violet’s most important flowers,” but the product of these flowers was certainly worth the effort.

Fruits formed from the cleistogamous flowers of sweet violet (Viola odorata)

Up close and personal with the fruit of a cleistogamous flower

The seeds (elaiosomes included) produced by the cleistogamous flower of sweet violet (Viola odorata)

See Also:

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.

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:

Camel Crickets and the Dust Seeds of Parasitic Plants

A common way for plants to disperse their seeds is to entice animals to eat their seed-bearing fruits – a strategy known as endozoochory. Undigested seeds have the potential to travel long distances in the belly of an animal, and when they are finally deposited, a bit of fertilizer joins them. Discussions surrounding this method of seed dispersal usually have birds and mammals playing the starring roles – vertebrates, in other words. But what about invertebrates like insects? Do they have a role to play in transporting seeds within themselves?

Certain insects are absolutely important in the dispersal of seeds, particularly ants. But ants aren’t known to eat fruits and then poop out seeds. Instead they carry seeds to new locations, and some of these seeds go on to grow into new plants. In certain cases there is an elaisome attached to the seed, which is a nutritious treat that ants are particularly interested in eating. Elaisomes or arils have also been known to attract other insects like wasps and crickets, which may then become agents of seed dispersal. But endozoochory in insects, at first, seems unlikely. How would seeds survive not being crushed by an insect’s mandibles or otherwise destroyed in the digestion process?

camel crickets eating fruits of parasitic plants (via New Phytologist)

While observing parasitic plants in Japan, Kenji Suetsugu wanted to know how their seeds were dispersed. Many parasitic plants rely on wind dispersal, thus their seeds are minuscule, dust-like, and often winged. However, the seeds of the plants Suetsugu was observing, while tiny, were housed in fleshy fruits that don’t split open when ripe (i.e. indehiscent). This isn’t particularly unusual as other species of parasitic plants are known to have similar fruits, and Suetsugu was aware of studies that found rodents to be potential seed disperers for one species, birds to be dispersers of another, and even one instance of beetle endozoochory in a parasitic plant with fleshy, indehiscent fruit. With this in mind, he set out to identify the seed dispersers in his study.

Suetsugu observed three achlorophyllous, holoparisitic plants – Yoania amagiensis, Monotropastrum humile, and Phacellanthus tubiflorus. While their lifestyles are similar, they are not at all closely related and represent three different families (Orchidaceae,  Ericaceae, and Orobanchaceae respectively). All of these plants grow very low to the ground in deep shade below the canopy of trees. Air movement is at a minimum at their level, so seed dispersal by wind is not likely to be very effective. Using remote cameras, Suetsugu captured dozens of hours of footage and found camel crickets and ground beetles to be the main consumers of the fruits, with camel crickets being “the most voracious of the invertebrates.” This lead to the next question – did the feces of the fruit-eating camel crickets and ground beetles contain viable seeds?

Monotropastrum humile via wikimedia commons

After collecting a number of fecal pellets from the insects, Suetsugu determined that the seeds of all three species were “not robust enough to withstand mastication by the mandibles of the ground beetles.” On the other hand, the seeds passed through the camel crickets unscathed. A seed viability test confirmed that they were viable. Camel crickets were dispersing intact seeds of all three parasitic plants via their poop. The minuscule size of the seeds as well as their tough seed coat (compared to wind dispersed seeds of similar species) allowed for safe passage through the digestive system of this common ground insect.

In a later study, Suetsugu observed another mycoheterotrophic orchid, Yoania japonica, and also found camel crickets to be a common consumer of its fleshy, indehiscent fruits. Viable seeds were again found in the insect’s frass and were observed germinating in their natural habitat. Seutsugu noted that all of the fruits in his studies consumed by camel crickets are white or translucent, easily accessible to ground dwelling insects, and give off a fermented scent to which insects like camel crickets are known to be attracted. Camel crickets also spend their time foraging in areas suitable for the growth of these plants. All of this suggests co-evolutionary adaptations that have led to camel cricket-mediated seed dispersal.

Yoania japonica via wikimedia commons

Insect endozoochory may be an uncommon phenomenon, but perhaps it’s not as rare as we once presumed. As mentioned above, an instance of endozoochory by a beetle has been reported, as has one by a species of cockroach. Certainly the most well known example involves the wetas of New Zealand, which are large, flightless insects in the same order as grasshoppers and crickets and sometimes referred to as “invertebrate mice.” New Zealand lacks native ground-dwelling mammals, and wetas appear to have taken on the seed dispersal role that such mammals often play.

Where seeds are small enough and seed coats tough enough, insects have the potential to be agents of seed dispersal via ingestion. Further investigation will reveal additional instances where this is the case. Of course, effective seed dispersal means seeds must ultimately find themselves in locations suitable for germination in numbers that maintain healthy populations, which for the dust seeds of parasitic plants is quite specific since they require a host organism to root into. Thus, effective seed dispersal in these scenarios is also worth a more detailed look.

Further Reading:

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For more stories of seed dispersal check out the first issue of my new zine, Dispersal Stories.

2019: Year in Review

It’s the start of a new decade and the beginning of another year of Awkward Botany. As we’ve done in years prior, it’s time to look back at what we’ve been up to this past year and look forward to what’s coming in the year ahead. Thank you for sticking with us as we head into our eighth year exploring and celebrating the world of plants.

The most exciting news of 2019 (as far as Awkward Botany is concerned) is the release of the first issue of our new zine, Dispersal Stories. It’s a compilation of (updated) writing that originally appeared on Awkward Botany about seeds and seed dispersal and is the start of what I hope will be a larger project exploring the ways in which plants get around. Look forward to the second issue coming to a mailbox near you sometime in 2020.

Also new to our Etsy Shop is a sticker reminding us to always be botanizing, including while riding a bike. Stay safe out there, but also take a look at all the plants while you’re cruising around on your bike or some other human-powered, wheeled vehicle. Whether you’re in a natural area or out on the streets in an urban or rural setting, there are nearly always plants around worth getting to know.

This year we also started a Ko-fi page, which gives readers another avenue to follow us and support what we do. Check us out there if Ko-fi is your thing.

Buy Me a Coffee at ko-fi.com

We also still have our donorbox page for those who would like to support us monetarily. As always you can stay in touch with us by liking and following our various social media accounts (Facebook, Twitter, Tumblr, and our currently inactive, but that could change at any moment Instagram). Sharing is caring, so please be sure to tell your friends about Awkward Botany in whatever way you choose. We are always thrilled when you do.

Below are 2019 posts that are part of new and ongoing series. You can access all other posts via the Archives widget. 2019 saw a significant drop in guest posts, so if you’d like to submit a post for consideration, please visit our Contact page and let me know what you’d like to write about. Guest writers don’t receive much in return but my praise and adulation, but if that sounds like reward enough to you, then writing something for Awkward Botany might just be your thing. And while we’re on the topic of guest posts, check out this post I wrote recently for Wisconsin Fast Plants.

Happy Reading and Plant Hunting in 2020!

Inside of a Seed & Seed Oddities:

Podcast Review:

Poisonous Plants:

Tiny Plants:

Eating Weeds:

Using Weeds:

Drought Tolerant Plants:

Tea Time:

Field Trip:

Awkward Botanical Sketches:

Guest Posts:

Out Now! Dispersal Stories #1

Before I started this blog, I had spent 16 years publishing zines at a steady clip and sending them to all corners of the world through the mail. I had never really meant to abandon zines altogether, and in some ways, putting all my writing efforts into a blog felt a little like a betrayal. My intention had always been to one day put together another zine. Now, six and a half years later, I’m happy to report that day has come.

Rather than bring an old zine back from the grave, I decided to make a new zine. Thus, Dispersal Stories #1. It’s quite a bit different from zines I’ve made in the past, which were generally more personal and, I guess, ranty. In fact, Dispersal Stories is very much like this blog, largely because it is mostly made up of writing that originally appeared here, but also because its main focus (for now) is plants. What sets it apart is that, unlike this blog, it zeroes in on a specific aspect of plants. As the title suggests, it’s all about dispersal. For much of their life, plants are essentially sessile. Once they are rooted in place, they rarely go anywhere else. But as seeds, spores, or some other sort of propagule they are actually able to move around quite a bit. The world is their oyster. What’s happening during this period of their lives is the focus of Dispersal Stories.

But why do a zine about this? Apart from just wanting to do another zine after all these years, my hope is that Dispersal Stories will be the start of a much more ambitious project. A book perhaps. My interest in dispersal was born out of my interest in weeds, and there is so much that I would like to learn and share about both of these subjects – so much so that the blog just doesn’t really cut it. So, I’m expanding the Awkward Botany empire. First a zine, then a book, then … who knows? I’m an oyster! (Or something like that.)

Dispersal Stories #1 is available in our etsy shop, or you can contact me here and we can work something out. While you’re at it, check out our new sticker.

If you love looking at plants and learning their names, then you probably enjoy doing it any chance you get. Usually it’s an activity you do while walking, but who says you can’t botanize while riding a bike? This sticker is inspired by a friend who once said that while mountain biking you get to “see three times as many flowers in half the time!” Stick it on your bike or in some other prominent location to remind yourself and others that we can botanize anytime anywhere.

Your purchase of one or both of these items helps support what we do. You can also support us by buying us a ko-fi or putting money in our donorbox. Sharing these posts also helps us out. If you get a copy of the zine, let us know what you think by sending us an email, a message on twitter or facebook, or by leaving a comment below. As always, thanks for reading.

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