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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.

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Poisonous Plants: Red Squill

Humans have been at war with rats since time immemorial. Ridding ourselves of their nuisance behavior is increasingly unlikely, and in fact, some scientists believe that, following human extinction, rats will be poised to take our place as the most dominant species on earth. Despite being thwarted repeatedly, we make tireless attempts to control rat populations. One major weapon in our arsenal is poison, and one of the most popular rat poisons was derived from a plant with a formidable bulb.

Urginea maritima (known synonymously as Drimia maritima, among other Latin names) is a geophyte native to the Mediterranean Basin, where it survives the hot, dry summer months by going dormant, waiting things out underground. Growth occurs in the cooler months, its bulb expanding annually before it finally flowers late one year after reaching at least 6 years old. Its flower stalk rises to as tall as 2 meters, extending heavenward from a bulb that can weigh as much as a kilogram. Its inflorescence is long, narrow, and loaded with small flowers that are generally white, but sometimes pink or red.

The oversized bulb of Urginea maritima — via wikimedia commons

Urginea maritima is commonly known as red squill or white squill (and sometimes simply, squill). Other common names include sea onion, sea squill, and giant squill. It is related the squill referred to in the Harry Potter universe, which is known botanically as Scilla. However, plants in the genus Scilla are much more dimunutive and generally flower in the spring rather than the fall. Like red squill, Scilla species are known to be poisonous; however, they don’t have the reputation for producing deadly rat poison that red squill does.

Like so many poisonous plants, red squill has a long history of being used medicinally to treat all sorts of ailments. As with any folk remedy or natural medicine, a doctor should be consulted before attempting to treat oneself or others. A 1995 report tells of a woman who ate red squill bulbs to treat her arthritic pain. She exhibited symptoms characteristic of ingesting cardiac glycosides – the toxic compound found in red squill – including nausea, vomiting, and seizures. She died 30 hours after eating the bulbs.

red squill (Urginea maritima) — via wikimedia commons

Toxic compounds are found throughout the plant, but are particularly concentrated in the bulb (especially its core) and the roots. Toxicity is at its highest during summer dormancy and when the plant is flowering and fruiting. The compound used to poison rats is called scilliroside. Bulbs are harvested in the summer, chopped up, and dried. The chips are then ground down to a powder and added to rat bait. Results are highly variable, so to increase its effectiveness, a concentrate can be made by isolating the toxic compound using solvents.

Red squill was introduced to southern California in the 1940’s as a potential agricultural crop. The region’s Mediterranean climate and the plant’s drought tolerance made it ideal for the area. The bulbs can be grown for manufacturing rat poison, and the flowers harvested for the cut flower industry. Breeding efforts have been made to produce highly toxic varieties of red squill for rat poison production.

the flowers of red squill (Urginea maritima) — via wikimedia commons

Around the time red squill was being evaluated as an agricultural crop, studies were done not only on its toxicity to rats, but to other animals as well. A 1949 article details trials of a red squill derived poison called Silmurine. It was fed to rats as well as a selection of farm animals.  Results of the study where “not wholly satisfactory” when it came to poisoning rats. Silmurine was less effective on Rattus rattus than it was on Rattus norvegicus. Thankfully, however, it was found to be relatively safe for the domestic animals it was administered to. Most puked it up or avoided it. Two humans accidentally became part of the study when they inadvertently inhaled the poison powder. Ten hours later they experienced headaches, vomiting, and diarrhea, “followed by lethargy and loss of appetite,” but “no prolonged effects.”

Vomiting is key. Ingesting scilliroside induces vomiting, which helps expel the poison. However, rodents can’t vomit (surprisingly), which is why the poison is generally effective on them.

Today, squill is available as an ornamental plant for the adventurous gardener. For more about that, check out this video featuring a squill farmer:

More Poisonous Plants posts on Awkward Botany:

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Seed Dispersal via Caching – The Story of Antelope Bitterbrush

Generally speaking, individual plants produce an enormous amount of seeds. This may seem like a huge waste of resources, but the reality is that while each seed has the potential to grow into an adult plant that will one day produce seeds of its own, relatively few may achieve this. Some seeds will be eaten before they get a chance to germinate. Others germinate and soon die from lack of water, disease, or herbivory. Those that make it past the seedling stage continue to face similar pressures. Reaching adulthood, then, is a remarkable achievement.

Antelope bitterbrush is a shrub that produces hundreds of seeds per individual. Each seed is about the size of an apple seed. Some seeds may be eaten right away. Others fall to the ground and are ignored. But a large number are collected by rodents and either stored in burrows (larder hoarding) or in shallow depressions in the soil (scatter hoarding). It is through caching that antelope bitterbrush seeds are best dispersed. When rodents fail to return to caches during the winter, the seeds are free to sprout in the spring. Some of the seedlings will dry out and others will be eaten, but a few will survive, making the effort to produce all those seeds worth it in the end.

Fruits forming on antelope bitterbrush (Purshia tridentata)

Antelope bitterbrush (Purshia tridentata) is in the rose family and is often simply referred to as bitterbrush. It occurs in grasslands, shrub steppes, and dry woodlands throughout large sections of western North America. It is a deciduous shrub that generally reaches between three and nine feet tall but can grow up to twelve feet. It has wedge-shaped leaves that are green on top, grayish on bottom, and three-lobed. Flowers are yellow, strongly fragrant, and similar in appearance to others in the rose family. Flowering occurs mid-spring to early summer. Fruits are achenes – single seeds surrounded by papery or leathery coverings. The covering must rot away or be removed by animals before the seed can germinate.

Bitterbrush is an important species for wildlife. It is browsed by mule deer, pronghorn antelope, bighorn sheep, and other ungulates, including livestock. It provides cover for birds, rodents, reptiles, and ungulates. Its seeds are collected by harvester ants and rodents, its foliage is consumed by tent caterpillars and other insects, and its flowers are visited by a suite of pollinators. For all that it offers to the animal kingdom, it also relies on it for pollination and seed dispersal. The flowers of bitterbrush are self-incompatible, and if it wasn’t for ants and rodents, the heavy seeds – left to rely on wind and gravity – would have trouble getting any further than just a few feet from the parent plant.

Antelope bitterbrush (Purshia tridentata) in full bloom – photo credit: wikimedia commons

In a study published in The American Naturalist (February 1993), Stephen Vander Wall reported that yellow pine chipmunks were the primary dispersal agents of bitterbrush seeds in his Sierra Nevada study area. The optimal depth for seedling establishment was between 10-30 millimeters. Seeds that are cached too near the surface risk being pushed out of the ground during freeze and thaw cycles where they can desiccate upon germination. Cached bitterbrush seeds benefit when there are several seeds per cache because, as Vander Wall notes, “clumps of seedlings are better able to push through the soil and can establish from greater depths than single seedlings.”

Another study by Vander Wall, published in Ecology (October 1994), reiterated the importance of seed caching by yellow pine chipmunks in the establishment of bitterbrush seedlings. Seed caches, which consisted of anywhere from two to over a hundred seeds, were located as far as 25 meters from the parent plant. Cached seeds are occasionally moved to another location, but Vander Wall found that even these secondary caches produce seedlings. Of course, not all of the seedlings that sprout grow to maturity. Vander Wall states, “attrition over the years gradually reduces the number of seedlings within clumps.” Yet, more than half of the mature shrubs he observed in his study consisted of two or more individuals, leading him to conclude that “they arose from rodent caches.”

A study published in the Journal of Range Management (January 1996) looked at the herbivory of bitterbrush seedlings by rodents. In the introduction the authors discuss how “rodents [may] not only benefit from antelope bitterbrush seed caches as a future seed source, but also benefit from the sprouting of their caches as they return to graze the cotyledons of germinating seeds.”  In this study, Ord’s kangaroo rats, deer mice, and Great Basin pocket mice were all observed consuming bitterbrush seedlings, preferring them even when millet was offered as an alternative. The two species of mice also dug up seedlings, possibly searching for ungerminated seeds. Despite seed dispersal via caching, an overabundance of rodents can result in few bitterbrush seedlings reaching maturity.

A cluster of antelope bitterbrush seedlings that has been browsed. “Succulent, young seedlings are thought to be important in the diets of rodents during early spring because of the nutrients and water they contain.” — Vander Wall (1994)

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Photos of antelope bitterbrush seedling clusters were taken at Idaho Botanical Garden, where numerous clusters are presently on display along the pathways of the native plant gardens and the adjoining natural areas.