Barbara Ertter

While most sensible plants hunker down during the cold winter season, awaiting the return of warmth and sunshine, there are some that flourish at this time of year.  This is particularly likely in parts of the world where most of the yearly precipitation falls during the period when temperatures vacillate between freezing and thawing, with intermittent blanketing by several inches of snow; in other words, very much like lower elevations in the Boise Front.  Plants that grow here have to choose:  grow when there is plenty of sun and warmth, but very limited and uncertain water, or grow when there is more reliable water, even if it means spending a lot of time being frozen and snowed under?  Very few animals can survive being frozen, but for many cold-adapted plants it’s no big deal, even to the extent of daily cycles of freezing and thawing.  Surviving the hot, dry summer can be the greater challenge, with a variety of dormancy options to choose among.

At the head of the pack of winter-loving plants are true mosses, members of an ancient group of plants called bryophytes that were among the first lineages to colonize the earth’s surface.  Lacking true roots and vascular tissue, mosses have no recourse except dormancy (or death) in the absence of abundant water.  Mosses that thrive locally are those that have perfected not only the capacity to survive both freezing during the depths of winter and desiccation during the searing summer, but also the amazing ability to rapidly pick up where they left off during the intermittent periods of favorable growing conditions.

As a result, winter is actually the primary growing period for mosses in the Boise Front, including during the cool moist conditions when temperatures hover above freezing during the daytime.  Not only does most vegetative growth occur at these times, when plants are bright green, but this is also when sexual reproduction takes places (by spores, not flowers and seeds).  In summer, the exact same plants are dried and brown, appearing dead but actually in a state of dormancy that can be quickly broken when wet conditions return.  Counterintuitively, mosses are extremely vulnerable at this stage; if artificially rehydrated in the heat of the full summer sun, plants that easily withstood both freezing and desiccation can bake to death if too hot when wet.

Some flowering plants also rely heavily on the intermittent favorable growing conditions during the winter months, primarily in preparation for flowering as early as possible once spring conditions arrive.  Sagebrush buttercup (Ranunculus glaberrimus) is one of the champions in this regard.  Plants evade the summer’s heat and drought by retreating underground as drought-tolerant swollen roots.  As soon as the late fall/early winter rains saturate the soil, however, new leaves emerge and continue to grow during intermittent periods of favorable weather throughout the winter.  Flower buds also develop, often opening as early as mid-February locally, coinciding with the emergence of the earliest pollinators (mostly flies).

This strategy comes with risks, however; if the rains do not begin before the ground freezes solid, and if the ground then stays frozen solid, the buttercups lose out on their prime vegetative growing season.  Even worse is if prolonged freezing is coupled with an absence of protective snow cover; in such conditions, tender new leaves can easily dry out, causing the plant to skip a growing season, or even die.  This happened at lower elevations in the Boise Front in the winter of 2022/23, when an unusually mild fall shifted abruptly to a hard freeze that lasted much of the winter; as a result, sagebrush buttercup flowers were less abundant than they were in more favorable years.

Another common strategy for local flowering plants is that of the fall-germinating annual.  After a brief few months of luxuriant growth during the brief spring period when water and warmth coincide, mature plants simply throw in the towel and die rather than trying to survive the searing summer heat.  Instead, they rely on resistant seeds to make it through to the next favorable growing season, which might be several years down the road.  Numerous seeds develop during favorable years, fewer when conditions are less favorable, all potentially accumulating and hanging out in the subsurface seedbank until the right combination of temperature, moisture, and day length trigger germination.

If all works as planned, the reward for those enterprising annuals that germinate in fall and then tough it out over the winter months is getting a good jump on slug-a-bed species that wait until the warm days of spring before germinating.  Unfortunately, some of our most invasive non-natives like cheatgrass (Bromus tectorum) and bachelor buttons (Centaurea cyanus) use this is the same strategy.  Worse, they often do this even more successfully than our natives do, which get crowded out when invasive non-natives beat them out for favorable growing sites and resources.  On the other hand, fall-germinating annuals, both native and non-native, are gambling that fall rains that trigger germination are not followed by a dry spell prolonged enough to cause vulnerable seedlings to shrivel and dry.  While this can significantly reduce the invasive non-natives that survive until spring, it can be equally devastating to our diminishing, diminutive natives.

Take time on your winter walks to look for these plants that also love winter, while also taking care not to trample vulnerable rosettes and tender mosses.  Take delight in experiencing what each season has to offer, with these early signs reminding us that the wildflowers are already getting ready for a new year!

For more information on some of our local mosses, see “Common Mosses and Ground Lichens of the Boise Foothills“.  To learn more about the wonders of mosses in general, I highly recommend Gathering Moss: A Natural and Cultural History of Mosses (2003) by Robin Wall Kimmerer.  This is the first book written by RWK, introducing many of the concepts and themes further developed in her inspirational book Braiding Sweetgrass (which I also highly recommend!)

Even for those of us inured to the inevitability of name changes, and fully versed on the perfectly good reasons behind them, the nearly complete turnover in the nomenclature of our local biscuitroots (genus Lomatium) can be mind-boggling.  For those of you who haven’t been keeping track, here’s a current synopsis, with the what’s and why’s, including a behind-the-scenes glimpse at what all those pesky plant taxonomists are up to.

First, it is important to note that this group of plants is extremely important, both for food and medicine, to most of the Native American tribes within whose traditional territories the many different species of biscuitroot occur (note: although the English common name of “biscuitroot” is sometimes restricted to only those species with enlarged edible roots, which were, and still are, a significant food source for many tribes, I am using this as the common name for the genus Lomatium in general; desert parsley is an alternate common name).  Each tribal language group has its own name, or names, for subunits of Lomatium, with the subunits often based more on specific use than necessarily coinciding with current scientific nomenclature. One of these names, “cous” (rhymes with “house”), is one of the few examples where a Native American name was incorporated into the scientific name:  Lomatium cous, which does not occur in the Boise Front.  The names given by the Shoshone, Bannock, Paiute, and other tribes to those species that do occur in the Boise Front, and the uses of these plants, should be respected as their traditional intellectual property.

As currently defined by science, Lomatium is a large genus that has undergone rapid evolutionary radiation in western and central North America.  Over the last century, taxonomic treatments of the genus have gone through several cycles of expansion and contraction, with major chunks sometimes treated as separate genera (e.g., Cogswellia, Leptotaenia, Peucedanum).  The number of species within the currently inclusive Lomatium is a moving target, with new species being described (or “resurrected” from synonymy) on a regular basis.  In 1997, Intermountain Flora provided a conservative estimate of “about 70”, but over 100 are currently in the draft treatment for Flora of North America (B. Wilson, pers. comm. 2023).

A major reason for all this nomenclatural activity is because several independent teams have recently homed in on Lomatium as a worthy research topic, rich in previously under-appreciated diversity that is amenable to analysis with the burgeoning new tools in the taxonomic toolkit.  One of these teams is centered here in the Treasure Valley, spearheaded by Dr. James F. Smith at Boise State University and Dr. Don Mansfield at The College of Idaho.  Their students and other collaborators have integral roles in the research, currently aided by a major NSF grant.

The genus itself is replete with narrow endemics (i.e., species that occur in a very restricted geographic area, often confined to a specific geologic substrate or soil type), many of conservation concern.  However, many of these species look very similar superficially, with clusters of small yellow flowers and leaves divided into narrow lobes; their morphological distinctiveness only becomes apparent when other features are carefully examined.  Molecular phylogenetic analysis has accordingly been invaluable, especially when combined with morphometric evaluation, extensive field studies, soil analyses, and other techniques.  None of these are “magic bullets” by themselves, but when combined with one another, and with traditional taxonomic approaches, they can bring biodiversity patterns to light that had not previously been evident, including new species.

With this background in hand, what exactly has been going on with biscuitroots in the Boise Front?  Here’s the run-down:

What’s the story with Nineleaf Biscuitroot?

Most wildflower guides and apps will lead you to believe that the most common biscuitroot in the Boise Front is the nine-leaf biscuitroot, Lomatium triternatum.  Until recently, the primary technical floras (e.g., Vascular Plants of the Pacific Northwest, Intermountain Flora) also placed most of these plants in Lomatium triternatum, but specifically in the subspecies platycarpum.  This is the most commonly encountered biscuitroot in relatively sandy and loamy soils, easily recognized by its  blue-green leaves that are commonly divided into nine narrow segments.

As happens, initial molecular phylogenetic analysis showed that ssp. platycarpum was not all that closely related to ssp. triternatum, and the two should therefore be treated as separate species — which is what some even earlier floras (e.g., Illustrated Flora of the Pacific States) had actually done.  Because of the rules of botanical nomenclature, the correct name for these plants at the species rank is Lomatium simplex, not platycarpum, and so this is now the correct name for most of what we had been accustomed to calling L. triternatum, at least in the Boise Front.  For the record, yes, we really do use the term “resurrecting” when an existing name that had been relegated to synonymy under an older inclusive name is deemed to be distinct after all.

Ok, that takes care of ssp. platycarpum, but what about the rest of Lomatium triternatum?  Well, now the story really gets tricky, in that there turns out to be a lot of subtle variation within what had been ssp. triternatum, which had actually been divided into two varieties:  triternatum and anomalum.  Oh, and let’s not forget the closely related Lomatium ambiguum from central Idaho and the recently described L. packardiae from nearby Oregon. As a further complication, it took a significant amount of sleuthing and fieldwork to nail down exactly which of the multiple possibilities was the REAL L. triternatum, all based on a scrappy specimen collected by the Lewis & Clark Expedition somewhere on the “Kooskooske” (Clearwater) River. (See Smith et al. 2018 and “Lewis’s Lost Lomatium Found” by Michael Ottenlips).

On top of which, it’s been a toss-up as to whether one of our other common biscuitroots in the lower Foothills, which often forms dense stands on heavy clay soils (e.g., Chief Eagle Eye Reserve, Foothills East Reserve), is best treated as L. triternatum var. anomalum or L. ambiguum (with the names themselves giving a clue to the challenges of biscuitroot identification!)  Well, guess what?  It turns out to be neither one, but instead a previously overlooked species!  Given the prominence of the new species in the hills overlooking the state’s largest city, and the bottleneck of available funding to continue research on the genus, the team decided to try an innovative fund-raising option that had been used successfully elsewhere: they auctioned off the naming rights for the new species. The winning bid chose to use the opportunity to honor Cecil D. Andrus, so this element of the baffling Lomatium triternatum complex is now officially known as Lomatium andrusianum, or Andrus’s biscuitroot (Stevens et al. 2018).

Next Up:  Butterflies as Botanists

Another conspicuous biscuitroot in the Boise Front, especially along Currant Creek north of Hidden Springs, is easily recognized by its finely divided bluish green leaves with a strong cilantro-scent.  Wildflower guides and apps will probably tell you this is Gray’s biscuitroot (Lomatium grayi), and this was in fact correct . . . until 2018.  That’s how long it took for a different team working on Lomatium to realize that at least three distinct species were shoehorned together under the same name (Alexander et al. 2018).

Well, at least that’s how long it took humans to figure it out; swallowtail butterflies had already known there were several species involved, and they used some but not others as host plants for their caterpillars.  It was actually an entomologist who first homed in on this lepidopteran behavior and cued the botanists in.  The name chosen for the most widespread component of chopped-up Lomatium grayi, which is the one occurring in the Boise Front, is Lomatium papilioniferum, with the epithet translating as “butterfly-bearing” (Papilio being the genus name for swallowtail butterflies).  Although iNaturalist uses “pungent desert parsley” as a common name, “butterfly biscuitroot” is both more euphonious and a tie to the delightful backstory.

As for Lomatium grayi itself, it’s still a perfectly good species, just not the one occurring in the Boise Front.  In fact, it only barely enters southeastern Idaho, with most populations occurring further south and/or east.  As you can see from this photos, leaves are greener, shorter, and less “fluffy” than in L. papilioniferum, at least in this population north of Preston.  How fun that it took butterflies to make us take a more careful look!

Mostly the Same . . . For Now

The other biscuitroots occurring in the Boise Front still retain their previous names, with some reshuffling between varietal and species names.  This, however, could easily change, depending on all the new results flooding in from the various Lomatium teams; all I can say at this time is, hold on to your hats!  For now, here’s the most up-to-date synopsis I’m aware of:

1. Barestem biscuitroot (Lomatium nudicaule):  This is a another relatively common species throughout the Foothills, and the easiest to identify with its distinctive round blue-green leaflets.  A delightful alternate common name is pestle-parsnip, probably due to the swollen apex of the stem.  If any nomenclatural surprises are in the works, I’m not aware of them.
2. Fernleaf biscuitroot:  Previously known as Lomatium dissectum var. multifidum, now just plain L. multifidum.  This is the largest of our locally occurring biscuitroots, with reddish brown flowers that fade with age.  It is mostly missing from the Foothills proper but still fairly common at higher elevations, and particularly abundant in the basalt talus along Highway 21/Mores Creek.  Although fernleaf biscuit is currently in vogue for both culinary and medicinal purposes, please keep in mind that any thoughts of harvesting should be balanced against the continued existence of our local populations already in decline from other threats, any legal restrictions on collecting, and the priority of indigenous rights to a culturally significant plant.
3.   Gumbo or Wasatch biscuitroot:  The jury is still out as to whether this is best treated as Lomatium bicolor var. leptocarpum (which is what is what I’m using for now) and plain Lomatium leptocarpum (as used in Intermountain Flora).  Locally it is only known from south of the Boise River, especially along the Oregon Trail below Bonneville Point.  It differs from the more widespread Lomatium simplex in having more irregularly divided leaves with shorter, greener segments, and also in its more spreading habit.
4. Bigseed biscuitroot (Lomatium macrocarpum):  This is another species known locally only from south of the Boise River, most readily found in the Oregon Trail Recreation Area.  It is easily recognized, at least when in bloom, by the large white inflorescences and lacy blue-green leaves.  Nomenclaturally stable, at least for now.

And One to Add!

The diminutive turkey-pea is one of the contenders for the earliest wildflowers to bloom in later winter, with sagebrush buttercup (Ranunculus glaberrimus) as the primary competition.  The inconspicuous low-lying clusters of white flowers and leaves divided into a few narrow leaflets are within the range of morphological variation of Lomatium, but the globular fruit of turkey-pea differ from the diagnostic flattened fruit of typical Lomatium, with the result that turkey-pea had traditionally been placed in a separate genus as Orogenia linearifolia.  However, one of the side results of the molecular phylogenetic revolution has been an essentially philosophical shift as well, emphasizing cladistic relationships as paramount in classification independent of unique evolutionary adaptions in an embedded lineage, or how disruptive its inclusion might be to an otherwise morphologically coherent genus (sorry, no way to explain this in simpler terms without an entire separate essay!)  Unsurprisingly, the evidence now unequivocally shows that Orogenia linearifolia is nested within (or originated from) the Lomatium clade, and by the currently ascendant philosophy is therefore a Lomatium.  As a result, we can add Lomatium linearifolium to the list of biscuitroots present in the Boise Front, though I will continue using “orogenia” as a common names, along with turkey-pea.

Anyway, that’s what I can tell you about biscuitroots in the Boise Front, at least for now, which I hope has also served as a window into why names keep changing.  In essence, as much as we crave to have nature presented to us as tidy, definitely named chunks, the reality is that a) nature doesn’t really care what we want, and b) our scientific knowledge of exactly what is “out there”, and how best to package it for general consumption, is constantly being refined.  Although not generally appreciated as such, species are fundamentally hypotheses, which are often initially proposed on scanty evidence (e.g., “Lewis’s Lost Lomatium”) and then subsequently supported, undermined, and/or modified as new information comes to light, ranging from basic observational data in the field to sophisticated technological analysis in the laboratory.  The result being that you, the user community, has the joy, and frustration, of a seemingly never-ending bounty of new species and names to try to keep on top of; stay tuned, and go with the flow!

Alexander, J. A., W. Whaley, & N. Blain. 2018. The Lomatium grayi complex (Apiaceae) of the western United States: a taxonomic revision based on morphometric oil composition and larva-host coevolution studies. Journal of the Botanical Research Institute of Texas 12(2): 387-444.

Ottenlips, M.  2019.  Lewis’s Lost Lomatium Found.  Sage Notes 41(1): 1, 7.  https://idahonativeplants.org/wp-content/uploads/2019/03/SageNotesMarch2019.pdf

Stevens, M., D.H. Mansfield, J.F. Smith, and M.E. Feist.  2018.  Resolving the anomaly of Lomatium anomalum: Discovery of a new species in southwestern Idaho (U.S.A.), Lomatium andrusianum (Apiaceae).  J. Bot. Res. Inst. Texas 12(1): 1 – 15.

Mulford’s Milkvetch (Astragalus mulfordiae), one of Idaho’s rarest plants that is currently in peak bloom in the Boise Front, comes with a wonderful back-story involving a woman who was both one of the earliest plant collectors in the young state of Idaho and also the first student to obtain a Ph.D. from the joint doctoral program between Washington University and Missouri Botanical Garden in St. Louis.  This was Dr. Anna Isabel Mulford, with the following biography based on a combination of decades-long sleuthing by my colleague Carol Prentice and a recent on-line biography by Cassidy Moody.

Carol’s interest was piqued in 1979, when Dr. Patricia Packard mentioned that no one in Idaho knew who Astragalus mulfordiae was named for.  The “-iae” ending was a clue that the species was named for a woman (the ending would have been “-ii” if named for a man), but Carol didn’t know this at the time.  It therefore came as a surprise to learn that Marcus E. Jones, who formally described the species in 1898, named it honor of a woman who had first collected it in Idaho.  This led Carol to track down a separate publication by Mulford herself, describing her travels to the wild West in 1892, and several of the plants she found during her visit.  This was all before the internet, adding to Carol’s challenge of tracking down the relevant publications and other sources.

Mulford’s article, published in 1894 in the Botanical Gazette, begins with “In the summer of 1892 I made a very interesting trip in the northwest.  The months of June, July and August were spent in Idaho.  My work in the state was done in the sage  brush of the southern part, in the Owyhee mountains to the southwest, and in the highlands, foothills and mountains of the eastern portions.  I traveled by railroad to the principal points, and by stage, carriage, or horseback to more distant ones.  Of course I walked a good deal.  I was well provided with letters of introduction, and there was seldom any difficulty in procuring company whenever desirable, which was necessarily a great part of the time.”

Among the letters of introduction was evidently one that led to Mary Hallock Foote, well-known locally for her writings and illustrations that capture an early era of Boise’s history.  Foote’s memoirs, posthumously published as A Victorian Gentlewoman in the Far West, describe the travails experienced by the family as her husband, Arthur Foote, pursued various engineering schemes  (including what would eventually become the New York Canal) in the western United States and Mexico.  The memoirs also provided inspiration, and fodder, for Wallace Stegner’s controversial Angle of Repose.

Mulford’s interaction with Foote is documented by one of the several species that Mulford described from her time in Idaho, Scutellaria footeana.  “This was found at about 3,500 ft near Black Cañon, Boise River, June 18th. I have named the plant in honor of Mrs. Mary Hallock Foote, who planned, and accompanied me upon the pleasant expedition which led to its discovery.”  The specific reference to the Boise River makes the basalt-rimmed stretch of the river east of Boise the probable site of Mulford’s “Black Cañon,” and it would have made perfect sense for Foote to take her visitor botanizing in the area where the Foote family had lived for several years. 

Alas, Foote’s botanical claim to fame was short-lived, with Scutellaria footeana soon disappearing into the synonymy of the relatively widespread Scutellaria nana.  Three other plants described by Mulford in the same publication have likewise been absorbed by previously named species: Oenothera idahoensis into O. caespitosa ssp. marginata, Frasera caerulea into F. albicaulis var. cusickii, and Gilia grandiflora var. diffusa into an undifferentiated Collomia grandiflora.  The only species Mulford described from Idaho that has stood the test of time is Frasera montana, whose lovely white blooms grace the mountains of central Idaho.

Somewhat surprisingly, Mulford was already 44 years old at the time of her Idaho excursion, six years after taking a post-graduate course at Vassar College that qualified her to be a professor of botany.  She eventually went on to obtain a Ph.D. at Washington University in 1895, age 47.  Her doctoral dissertation was on the genus Agave, working with Garden director William Trelease.  Although this choice of topic leaves unanswered the impetus for her collecting trip to Idaho, it did create an opportunity for her to interact with another early Idaho collector, Timothy E. Wilcox, who had been Assistant Surgeon at Fort Boise from 1879 to 1882.  At the time of Mulford’s doctoral research, Wilcox was posted at Fort Huachuca in southern Arizona, where Agave was abundant.  Mulford credits Wilcox with assistance and for photos, specifically noting the value of his “habit photos,” and also quotes Wilcox describing which species of agave that cows will eat in Arizona.

Little is known of Mulford’s early years, other than growing up in East Orange, New Jersey, and attending Trenton Normal School (now The College of New Jersey).  Obituaries indicate that her father, Timothy Mulford, was a wheelwright who “manufactured wheels used by Union forces in the Civil War.”  After obtaining her Ph.D., Mulford had a long teaching career, including high schools in St. Louis (and possibly Wake Forest University in North Carolina). Census records show that she had moved back home to East Orange by 1930.  She evidently never married, retaining her maiden name at the time of her death in 1943, age 95.

Astragalus mulfordiae is one of three species named after Isabel Mulford, and the only one still in active use. The other two are Agave mulfordiana Trelease (now a synonym of A. schottii) and Viola mulfordiae Pollard (now used for the hybrid of V. brittoniana × V. sagittata).

Mulford’s milkvetch is endemic to the western Snake River Plain in southwestern Idaho and adjacent Oregon, restricted to loose, sandy substrates of former Lake Idaho sediments.  It has an S2/Imperiled conservation status in Idaho, due to the ongoing loss and habitat degradation of its scattered populations from development, invasion by cheatgrass and other non-natives, and careless recreation.  Federal protection would easily be warranted if this species were a mammal or bird, but the abundance of plants with equivalent status would overwhelm an already overloaded system out of favor with competing agendas.  If the decline of this delicate species is not reversed, we will lose not only one of the special plants that still grace the Boise Front, but also a tangible tie to an intrepid woman who is also nearly forgotten.

Mulford, A. Isabel. 1894. Notes upon the northwestern and Rocky Mountain flora. Botanical Gazette 19: 117-120.

Mulford, A. I. 1895. A Study of the Agaves of the United States. Missouri Botanical Garden Annual Report 7: 47-100.

With “zombie fungus” currently in the public eye, thanks to the popularity of “The Last of Us”, let me introduce you to a botanical counterpart that’s easily observed in the Boise Front in spring.  As is so often the case, what nature comes up with can be more bizarre than what human minds could conceive of on their own, which is why we often turn to the natural world for inspiration in our fiction, and even our technology.

The two plants in this photo are in all likelihood different shoots of the same plant, connected by underground branches.  The one on the right is a normal early-season shoot of phlox, either prickly-leaf phlox (Phlox aculeata) or long-leaf phlox (Phlox longifolia).  The two closely related species are relatively distinct and easy to tell apart in the portions of their respective ranges that don’t overlap on one another, but a full gamut of morphological intermediates occurs in the Boise Front, and young shoots with leaves just developing are particularly problematic.

In contrast, the shoot on the left is so different that it appears to be an unrelated species, with much wider, yellowish leaves.  These differences are caused by a rust fungus that has infected this shoot, hijacked its metabolic controls, and forced the plant to grow into a form that benefits this fungus, not the plant.  I’m guessing the rust fungus in question is a species of Puccinia, though if there are any existing studies on this particular rust/flower combination, I’m not aware of them.

What intrigues me most is the similarity of the resultant morphology to a well-studied counterpart in some other genera, notably rockcresses (Boechera and Arabis).  Rust–infected shoots in these genera terminate in conspicuous yellow clusters of modified leaves that not only look enough like flowers to attract flies and other pollinators, but can even produce fragrances and sugary nectars (see “Fungus Is a Flowerlike Con Artist“).  It is probably no coincidence that these “pseudoflowers” can most often be found in early spring, when actual flowers are still in short supply, and when buttercups (Ranunculus) and other bright yellow flowers are particularly noticeable.

The set-up works great for the fungus, which takes advantage of the existing flower-pollinator collaborative system for dispersing its own spores.  These spores, and usually(?) a sweet nectar, are produced in abundance in tiny cups on the undersides of the modified leaves.  Flies, which are often the most abundant early spring pollinators, seem perfectly satisfied with the pseudoflower nectar, and may even prefer it over whatever the real flowers that are in bloom have to offer.

The loser, alas, is the poor infected plant itself.  Not only is the modified shoot prevented from blooming, but even normal-looking adjacent shoots appear to be less likely to develop flowers, at least in the phlox I’ve been observing in the Boise Front.  My guess is that energy and nutrients are diverted to the greedy zombie shoots via underground stems, depleting what would otherwise be available to the rest of the plant.  It’s even possible that other spring-blooming species lose out, if pollinators are in short supply and preferentially visit the scamming pseudoflowers.

Of course, a major difference between zombies in pop culture, and zombie-esque fungi in real life, is that the latter don’t end up threatening the continued existence of the host species, either by directly killing them off or by simply disrupting their reproductive capacity.  Any fungus or other parasite that wiped out its host species would end up wiping out itself, and while we can’t be certain this never happens, it does mean that any fungus so poorly adapted would quickly disappear after a very brief existence.  In the case of phlox, although a few shoots in a population might be infected, and some individual plants weakened beyond recovery, the population itself generally continues just fine, blooming merrily away as the season progresses.  And by peak bloom, the zombie shoots have often finished their fungal reproduction and disappeared, waiting to reappear in the sequel year.

Given that this website is based on blogging software, I thought I’d try an actual blog post, inspired by this poor struggling sagebrush buttercup (Ranunculus glaberrimus) that I photographed on this morning’s walk.  It was one of the few lonely survivors in a small isolated population along the Corrals Trail, not far north of the east end of Bob’s Trail.  This is a surprisingly popular trail for hikers as well as cyclists, given that it is nearly 10 miles to do as a loop (rather than the wimpy out-and-back from the 8th Street Road that I opted for), or so I was told by one set of hikers.

It was a fine day to be out in the central Foothills, especially after a agonizingly protracted winter.  However, while I was certainly sharing the general enjoyment of the expansive views and just being in nature that my fellow trail users were presumably relishing, I was also suffering the curse of being a botanist, incapable of closing my eyes to the biological poverty and degradation of too much of the area I was walking through.  I refer to this portion of the Foothills as “Low-Diversity Mid-Elevation Pastureland”, characterized by a dominance of non-native perennial grasses and only a smattering of the most common native wildflowers.

The reasons for this biological impoverishment began with the intense unregulated grazing by cattle and sheep in the early days of Euroamerican settlement, which both reduced the more vulnerable wildflowers and created prime conditions for the spread of non-native annual grasses such as cheatgrass (Bromus tectorum). Although grazing became moderated as the Foothills became carved up among privately owned ranches and leased areas of public lands, management of these lands was specifically as pastureland for livestock and big game animals (aka “range improvement”). Erosion control has also been a major concern, especially in the aftermath of devastating fires that repeatedly swept across the Foothills in late summer, driven by hot dry winds (and increasingly sparked by human carelessness). The default management tool for both grazing purposes and erosion control/post-fire rehabilitation has been seeding with a selection of perennial grasses that thrive in the Boise Front.  Selection of species and cultivars has been based on performance and availability; only recently have native species been promoted as the desirable option, and even then local gene pool is rarely an option (WAY too costly!)

In the Boise Foothills, one of the most commonly and widely planted grasses for range improvement and post-fire rehabilitation is intermediate wheatgrass (Thinopyrum intermedium, alternatively treated as Agropyron or Elytrigia). Although sometimes mistakenly assumed to be native, intermediate wheatgrass was introduced from Eurasia in the early 1900s and is now one of the most esteemed grasses in land management arsenals.

Unfortunately, the features that make intermediate wheatgrass superb for the intended uses can also come at the expense of native diversity, not only wildflowers but also moss, lichens, many insects, and even some smaller mammals and birds.  This is because this Eurasian species does so well that it displaces, replaces, the shrub-steppe habitat that our unique native plant, animal, fungal, and microbial diversity evolved in and is adapted to. The characteristic feature of the shrub-steppe habitat is an open scattering of shrubs, tufted bunch-grasses, and tufted forbs (i.e., wildflowers) in a matrix that we tend to dismiss as “bare ground”, but which is actually a complex micro-ecosystem of mosses, lichens, and microbes called biotic crust.  This crust provides an essential germination site and nursery for both annual and perennial wildflowers, which in turn provide critical food and other requirements for many animal species.  The biotic crust most likely plays an integral role in nutrient cycling and water retention as well, though this is still under investigation.

Enter intermediate wheatgrass, which differs from our native bunch-grasses in being an aggressively rhizomatous perennial, capable of forming a solid turf that fills in the “bare ground” and eliminates the previous matrix habitat.  Some of the more hardy native perennials, such as the buttercup in the above photo, may persist for a number of years, but the population will eventually wink out if suitable conditions for seedling germination and development no longer exist.  And the previously established mature plants might also face a shortened life span, in the face of competition for light and nutrients from such a successful invader. The story is different for species that co-evolved with turf grasses, especially in areas with summer rainfall such as the Great Plains and African savannas, but this is not that habitat, or our story.

To add insult to injury, the dried thatch of non-native grasses evidently accumulates more than that produced by native grasses, creating the conditions in the above photo.  If there is any research on why this happens, it’s not something I’m not familiar with; I can’t help but suspect, however, that our native decomposers are not well-equipped to deal with Eurasian grasses, for whatever reason. This could possibly change over time, given how evolution operates, but what we have is the current situation.

Whether there is any hope for restoring native diversity in areas already dominated by intermediate wheatgrass and other non-native grasses is a problematic question.  Unsurprisingly, the standard range-management solution to thatch build-up is . . . more grazing!  While intensive grazing will indeed reduce thatch, it comes with costs that are seldom mentioned.  For a start, while there are indeed some less palatable native species that thrive under moderate grazing pressure, others (like our only native peony, Paeonia brownii) are preferentially eaten as “cattle candy” and will quickly disappear from even moderately grazed sites. Browsers like goats can be even worse, tending to eat dried grass only after more desirable plants, including bitterbrush (Purshia tridentata) have been decimated.  Furthermore, the trampling can be devastating to the critical biotic crust, especially on steep hillsides.  The benefits ascribed to herds of large herbivores in summer-rain grassland habitats do not automatically carry over to semi-arid shrub-steppe habitats, which evolved with smaller and sparser herbivores (and without having to compete with aggressive non-natives).

So, what, if anything, can be done?  Paradoxically, my primary response is to say:  do nothing.  This comment is primarily intended to counter our deep-rooted tendency to do SOMETHING:  to continue modifying declining habitat to suit our insatiable needs, to implement drastic remediation measures that often cause more harm than simply allowing burned or otherwise impacted habitat to recover on its own, even when there is no significant threat to human lives or property.  Stop planting more intermediate wheatgrass and other non-native species, at least as the one-size-fits-all solution to just about everything. Beyond that, follow a Hippocratic oath for ecological practitioners, beginning with “Do no harm”.  Proceed cautiously and tailor treatment to individual situations, taking note of what works and what doesn’t.  Above all, pay attention to the entire “patient” (i.e., the full diversity of a unique biological community), not just the “disease”; outcomes where “the operation was a success, but the patient died” are not success stories.  Cherish the buttercups that still remain in the Foothills, and give them and the rest of our bountiful native diversity the best chance possible to continue sharing this wonderful area we all call home.

(in collaboration with Carol Prentice)

Our sole native true primrose is officially known as Cusick’s primrose (Primula cusickiana), but locally it is sometimes referred to as Wilcox’s primrose, or even wilcoxiana.  The reason why is intertwined with the early history of Boise.

The story begins with Captain Timothy Erastus Wilcox (1840-1932), who was posted as Assistant Surgeon at Fort Boise from June 1879 to August 1882.  With the Fort conveniently situated at the base of the Boise Foothills, near the mouth of Cottonwood Creek, Wilcox was able to indulge his naturalistic bent, and may in fact be the first Boise resident to have made plant collections that ended up in permanent herbaria.  Among his claims to fame is a paper that he published in Nature calling attention to the absence of earthworms in Boise; evidently they did not occur here until introduced by settlers.  He is also credited with introducing pheasant and bobwhite in Oregon and Idaho (Ewan & Ewan 1981).  As part of his official duties, Wilcox was even delegated with providing daily weather observations, a task required by U.S. Army Post Surgeons during Wilcox’s tenure at Fort Boise (Smith 2017).*

The plants Wilcox collected in Boise were sent to botanists in New York, where they triggered this write-up in the July 1881 issue of Bulletin of the Torrey Botanical Club:

“Idaho Plants.—The acknowledgements of the Torry Club are due to Dr. T. E. Wilcox, U.S.A., of Boisee[sic] City, Idaho, for several packages of plants of that vicinity collected in April and May of this year.  Many interesting species are represented; among them, Delphinium decorum var. Nevadense, Watson; Brodiaea laxa, Watson; Fritillaria pudica, Spreng.; Allium nevadense, Watson; Balsamorrhiza[sic] Hookeri, Nutt.; Antennaria dimorpha, T. & G.; Crepis occidentalis, Nutt.; Lupinus Chamissonis, Eschs.; Mertensia oblongifolia, DC.; Plectritis congesta, DC.; also a dwarf primrose of the same general size and habit as Primula angustifolia, but having a from 1–5-flowered scape and showing also marked differences in the inflorescence from that described by Dr. Gray.  Prof. Wood regarded this as a variety of P. Parryi, Gray, and named it var. Wilcoxiana.  It appears to be the same as var. Cusickiana, Gray, of P. angustifolia (N. A. Flora, p. 393) except that the last-named variety is described as only 2-flowered.  P. angustifolia flourishes in its typical, 1-flowered form, on high mountain elevations, such as Gray’s and Pike’s Peak, at 11,000 ft. to 13,000 ft., where we have found it abundant.  The many-flowered forms grow at much lower elevations (where P. parryi, a much larger species, is found) but the foliage shows no marks of any greater vigor in the plants.  It is possibly a distinct species.”

The “Prof. Wood” who proposed naming the Primula in honor of Wilcox is Alphonso Wood (1810-1881), the principal of Brooklyn Female Academy and the first to use dichotomous keys in his  “Class-Book of Botany”; most of Wood’s own herbarium is now at New York Botanical Garden.  However, neither Wood nor any other botanist ever actually followed through on formally describing Wilcox’s collection of Primula as either a distinct variety or a species, though the exact relationship between plants in the Boise Front (“wilcoxiana”) and plants in the Wallowa Mountains of northeastern Oregon has been an ongoing question.  The latter location is where William C. Cusick made the collection that Asa Gray described first as Primula angustifolia var. cusickiana, and then elevated to a full species.

The use of “Wilcoxiana” as a local common is confirmed in Bernice Bjornson’s 1946 Key to the Spring Flora of Southwestern Idaho, a mimeographed collector’s item that I myself used in a biology class in Boise High School.  In it, Bjornson notes that:

“A few places in the West are fortunate in having the attractive little primrose known in southern Idaho as Wilcoxiana, Primula cusickiana. Its leaves are simple, entire, and basal. Its flowers borne on a leafless stalk vary in color from pale blue to dark purple with a yellow eye. They are fragrant. People who lived in southwestern Idaho in the early 1900’s tell us that in spring the foothills used to be a mass of color with these flowers. Surely this is a flower which needs protection; one has to hunt for it nowadays.”

Although the primrose was never formally published, at least one plant was officially named in Wilcox’s honor:  Quercus wilcoxii Rydb., from Arizona (now included within Quercus chrysolepis s.l., canyon live oak).  The cactus genus Wilcoxia Britton & Rose (now a section of Echinocereus) is also named after him, from the period he was posted in Arizona.  However, the majority of plants with the epithet wilcoxii or wilcoxiana are named after other plant-collecting Wilcoxes.  As a prime example, Wilcox’s penstemon (Penstemon wilcoxii), whose lovely blue flowers grace higher elevations in the Boise Front, is named in honor of Earley Vernon Wilcox, a biology professor at University of Montana 1896-1899 (Ewan & Ewan 1981).

More information on Wilcox’s life can be found in his obituary (Eggleston, 1933).  He was born in North Litchfield, New York, on 26 April 1840, and received his M.D. from Albany Medical College in 1864.  Noteworthy events in his career included attending Jefferson Davis during his detention at Fortress Monroe after the Civil War.  He went to Cuba in 1898 (i.e., during the Spanish-American War) as lieutenant-colonel chief surgeon and was honorably discharged the following year.  He joined the Torrey Botanical Club in 1880, after being proposed for membership by Alphonso Wood, and was made a life member in 1930.  From 1917 onward he was nearly blind, but his mind was clear until his death in 1932.  Wilcox is buried in Arlington Cemetery, along with his wife Clara Brainard Brown.

Extended version published in Sage Notes:

Ertter, B. & C. Prentice.  Dr. Wilcox—Army Doctor, Botanist—and his primrose: Part 1. Sage Notes 44(2): 1, 4–5, 9.

Prentice, C. & B. Ertter. 2023. Dr. Wilcox—Army Doctor, Botanist—and his primrose: Part II. Sage Notes 44(3): 1, 4–6, 13.  (C. Prentice & B. Ertter).

Prentice, C. & B. Ertter. 2022.  Dr. Wilcox—Army Doctor, Botanist—and his primrose: Part III.  Sage Notes 44(4): 1, 4–7.

Recording of talk to Pahove Chapter, Idaho Native Plant Society, on 7 Oct 2023: “Timothy E. Wilcox: Army Surgeon at Fort Boise, Boise’s first resident Botanist/Naturalist” https://youtu.be/ZyEnwPc5rtQ

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*Reports of local weather at stations throughout the country were telegraphed to a central location in Washington, D.C., where they were compiled and analyzed as the foundation to understanding broad-scale climatic patterns.  As of 1877, the inclusion of weather reporting in the duties of medical personnel at Fort Boise overlapped those of U.S. Signal Service Observers.  The first of these, Sergeant Barnet Edward Light, was my great-grandfather, and the beginning of my own roots in the Boise Front.

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References:

Brown, A.  1881.  Idaho Plants. Bulletin of the Torrey Botanical Club 8(7): 81-83.

Bjornson, Bernice.  1946.  A Key to the Spring Flora of Southwestern Idaho.  mimeographed, 121 pp.  Copyright by the author.

Eggleston, W. W.  1933.  Obituary: Timothy E. Wilcox.  Science 77 (1995): 300.

Ewan, J., and N. D. Ewan.  1981.  Biographical Dictionary of Rocky Mountain Naturalists.  Regnum Vegetabile vol. 107.  citing Auk 2 (1885): 315.

Smith, J. D.  2017.  The History of Weather Observation — Boise Idaho.  NOAA Technical Memorandum NWS WR-291. https://www.weather.gov/media/wrh/online_publications/TMs/TM-291.pdf

The recording of Barbara Ertter’s talk on “New Discoveries and What’s Cool in the Boise Front”, given to the Pahove Chapter of the Idaho Native Plant Society on 14 December 2021, can now be enjoyed on YouTube (click here).