6. BUTTERCUP FAMILY (Ranunculaceae)

LARKSPUR (Delphinium barbeyi) AND DWARF LARKSPUR (D. Nuttalianum)

GETTING A LIVING 

Tall larkspur (D. barbeyi) is a highly resilient species, living up to 75 years, thriving in moist rich soils in subalpine meadows and nearby forest edges, and attracting an abundance of pollinators, especially bumblebees and hummingbirds. Dwarf larkspur (D. nuttallianum) pops up quite early after snowmelt in dry, rocky upland meadows and mixes in with sagebrush. When hiking high in the central Colorado Rockies around Crested Butte in early July, Tall larkspur, D. barbeyi (sometimes called subalpine larkspur), can't be missed nor should it be. It can rise to as much as two meters in height and bears a multitude of beautiful blue flowers with their distinct spurs extending out the back end. The flower is made up of five petal-like sepals and four true petals. The sepals come together at the back of the flower to form the pointed spur that encloses the plants nectar storage pocket (or nectary), and at the front end each sepals flare out to form the face of the flower. The sepals enclose four inconspicuous true petals, the top two of which form two separate nectar-contains spurs that are in turn wrapped together by a folding over of the top of the five sepals.  This flower complexity serves the purpose of getting pollinators going after nectar to dump their pollen from other flowers on the pistil to bring about fertilization. The result is a flower of great beauty and interest to the human eye. There is nothing like flower sexuality to bring about patterns of great beauty that inadvertently bring human joy. The leaves of a larkspur are deeply lobed and have a palmate shape. The botanical term "palmate" refers to a leaf with lobes or leaflets that emanate from its base, much like an open hand.  A larkspur leaf can have from three to five pointed, toothed lobes. Each plant supports anywhere from 20 to 150 stalks and each stalk boasts up to 50 flowers.

Dwarf Larkspur looks like a miniature version of its tall cousin, although with a lesser flower density per plant.

DEFENSE

Tall larkspur's (D. barbeyi) big challenge today, along with numerous of it's fellow high-mountain species, is to survive global warming. Because climate change moves up the timing of Spring  snowmelt in the high mountain Rockies, Larkspur, like many of its co-blooming species, now emerges much earlier than its historic norm. One might think a longer growing season would be a benefit for larkspur, giving it more time to grow and reproduce, but moving snowmelt up actually increases the probability of damage to plant parts from nighttime freezes trumping the benefit of a longer season. A later snowmelt retards plant survivals until further in the season when night freezes become less likely. To reiterate a truly important point, climatic warming advances daytime high temperatures more quickly than nighttime lows. Tall larkspur suffers a special vulnerability to night temperature plunges because its flower buds possess an extraordinary sensitivity to frosts. Paradoxically,  lower June nighttime temperatures continue to occur at high elevations and latitudes even though annual snowmelt dates are receding along with warming daytime temperatures.

REPRODUCTION 

Historical records show a positive relationship between flower abundance in dwarf larkspur and snowpack in the Colorado Rockies near Crested Butte. A warming experiment cause both plant and flower abundance per plant to diminish in this species due to increase water stress. This plant is an important early nectar source for queen bumblebees and hummingbirds who both depend on the sequential blooming of dwarf and and tall larkspur. Unlike the later, the former possess buds that are immune to freezes, but can suffer from damage to early emerging flowers inhibiting reproduction, and drought from early soil drying. Later flowering Tall larkspur (D. barbeyi) and earlier flowering dwarf larkspur (D. Nuttalianum) are advancing at about the same pace in flowering time relative to snow-melt, and both species experience less floral abundance in early snow-melt years and, as a result, diminished prospects for reproduction. 

5. LILY FAMILY (Liliaceae)

GLACIER LILY (Erythronium grandiflorum) 

GETTING A LIVING

The life of a Glacier Lily (Erythronium) begins below ground with its long (33-55mm), slender, deeply planted, corm, or in ordinary gardening parlance, its bulb. Out of this springs  stems and two lance-shaped (lanceolate) green to greenish-gray long leaves (up to 25 cm) connected to the stem at ground level (basal leaves botanically speaking). From each corm will grow one to five separate stems 15-40 cm long, and from each of these as many as six nodding yellow flowers will emerge, although just one is common.

Where Erythronium likes to reside offers clues about how it lives and prospers. In the Colorado Rockies, Erythronium  can be found between 6,500 and 11,000 feet elevation and prefers cool, wet locations with fertile soils. It especially favors wet meadows and slopes that abut streams, lakes, or seeps, and is most common in snow shoots and late lying snow banks. This impatient flower doesn't wait until snowmelt is done but will pop up through the snow itself. Among mountain wildflowers it is one of the first to appear (late May or early June), and in doing so it gains the special advantages of a monopoly on pollinators and access to sunshine, moisture, and nutrients. By showing up early, this species has few worries about competition for resources--light, soil, nutrients, water, and pollinators.

DEFENSE 

The early arrival strategy brings along with it a serious risk from early snow melt. Over the 33-year period ending in 2008, because of climate change the snowmelt date advanced by an average of 4.14 days per decade. At the same time the mean summer temperature increased by 0.38°C per decade. Despite warmer temperatures, average summer precipitation hasn't changed much, but the first, peak, and last flowering dates for Erythronium advanced an average of 3.2 days per decade. Even before the snow is entirely gone, Erythronium flowers pop up to get an early start, but arriving early carries with it the danger of exposure to night frosts that commonly occur early in the spring, even when daytime temperatures are well above freezing. Flowers arriving at the party early may be stuck out in the freezing cold at night, against which plant parts may lack defenses. Fruit production for Erythronium has been found to be lower in years when late frosts occur. In many wildflower species, more frost means fewer healthy flowers for flower-lusting bees and hummingbirds to do the work of moving pollen from one flower to another completing the reproductive cycle. For Erythronium, however, earlier snow melts doesn't seem to have hurt overall flower abundance even though fruits are sometimes damaged by frost.  It's early blooming propensity could well have caused it  to genetically adapt to cold night temperatures. Because of its early spring appearance,  Erythronium  is important as food when little else is available for mule deers arriving from their wintering grounds and for pollinators such as early arriving queen bumble bees and migratory humming birds. Conceivably, if Erythronium moves its blooming peak forward enough, animals dependent on the species for food could arrive too late for a full meal. Changes in the presence of  one species can spill over to others in an ecologically interconnected world. 

Early snowmelt in mountain settings is so important for wildflowers in general that we should be fully aware of its causes. The steady buildup of human-produced greenhouse gases such as CO2 in the earth's atmosphere has caused global temperature averages to advance in recent years according to the predominance of of scientific opinion. Scientists also acknowledge that climate warming will be felt more extensively than average at higher elevations and latitudes around the world. This means earlier snowmelt and higher summer temperatures in the subalpine Rocky Mountains to the detriment of climate sensitive wildflowers. The irony is that a cold-adapted, early blooming species such as Erythronium may actually be able to handle climate change because of its resilience to night frost exposure that increases as snow melt, daytime temperatures, and and peak flower blooms advance more quickly than incidence of night frosts recedes.  Those species that bloom a bit later, such as the Aspen sunflower (Hellianthella quinquenervis), are less likely to be pre-adapted to night frosts and more likely to suffer significant damage to their plant parts from freezing temperatures.

REPRODUCTION

On each Erythronium stem, all the flowers together are referred to as the inflorescence, and each individual is a perfect flower possessing both male and female parts and is capable of self-pollination. In short, a perfect flower is a sexual cross dresser that can make itself pregnant, a hermaphrodite, a beneficial strategy if pollinators are scarce, which could well be the case for an early bloomer whose fruits periodically suffer frost damage.  Erythronium has  perfection but not completeness with tepals for flower parts but not not both petals and sepals. Tepals mean a lack of distinction between petals and sepals.  

4. PEA FAMILY (Fabaceae)

BIG-LEAVED LUPINE Lupinus polyphyllus

GETTING A LIVING 

Many of us know about lupine, a common wildflower found in a variety of habitats, from reading "The Legend of Bluebonnet" to our children, a story about a young Comanche girl who sacrificed her favorite toy to save her tribe from starvation. Lupines are easily recognized as mostly large and shrubby plants with beautiful blue or purple spikes of pea-like flowers and distinct palmate shape leaves that look a bit like fingers emerging from the palm of a hand.  These fold up at night to minimize water and heat loss. Big-leaved lupine (Lupinus polyphyllus) favors montane and subalpine woodland edges as its prime habitat.

 DEFENSE 

Big-leaf lupine (L. polyphyllus) experiences reduced flower populations in early snowmelt year caused in part by frost damage to its flower buds. Early snowmelt is also correlated to a lengthening of a mid-season drought in June and early July that could reduce flower populations as well. In this instance, both frost and drought may paradoxically pose a threat as a consequence of climate warnings. Lupine alkaloids are poisonous for sheep, but not for birds, bear, elk, or rodents that feed on the plant's seed pods and roots. Alkaloids often defend against herbivores, but not so much for L. polyphyllus.

REPRODUCTION  

Pinch the keel of the lower flower part of most lupines, and out pops the stamens with  pollen squirting outward. Bees accomplish this same task with their buzzing and are on the receiving end of the  pollen shot that sticks to their belly and gets transported to the next flower. Once a flower is pollinated, its upper scoop-like banner turns from blue to white, telling the bee not to bother landing for nectar that's no longer available. An adaptation that eases the pollinator's task will  increase visits to other flowers on the same plant improving reproductive prospects.  Nature's bag of tricks focuses heavily on  competition and survival of the fittest, but also cooperative synergies .  

While different species compete for pollinators, multiple blooming species can support and attract more pollinators than otherwise. Pollinators experience reproductive benefits from overlapping blooms of different species that extend the total time in which flowers are blooming and supplying pollen. The reproductive lifespan of a pollinator can extend beyond the blooming period of any single plant species, and benefits from multiple sequential bloomers. This is the deal for L. polyphyllus (big-leaved lupine) and Lathyrus lanszwertii (peavine), given a normal linked blooming season. For these two species, the snowmelt date affects flowering overlap. In early snowmelt years, Lathyrus lanszwertii (peavine) moves its peak flowering date forward much more than L. polyphyllus (big-leaved lupine). The worry is that diminished co-flowering of such plants that share pollinators will have a negative impact on pollinator populations that could feed back on plant populations. If species flower separately in time, the gap between could lead to pollinator starvation and ultimately a reduction of  pollinator populations. This reduction in turn will mean less flower pollination and diminished plant reproduction.

PEAVINE (Lathyrus lanszwertii)

GETTING A LIVING  

Showing up normally in the same habitat with L. Pollyphylus in mid-June, Peavine (Lathyrus lanszwertii) possesses attractive classic pea-shaped creamy white flowers with bright pink nectar guides leading pollinators to their sweet reward and completion of the floral mating game. Lathyrus  occupies a variety of habitat types including sage-dominated dry meadows,  high-elevation subalpine meadows, and aspen woodlands.

DEFENSE. With snowmelt advancing steadily in subalpine mountain meadows,  Lathyrus is moving its dates of first and peak blooming ahead in time without any apparent loss in in its flower populations. It's ability to do so suggests that it is well defended against increase exposure to night frosts. For its co-flowering partner, Lupinus polyphyllus, as already noted, this is apparently not the case. As snowmelt advances, this species reduces the volume of its flower production and keeps its flowering peak out of the night frost time zone. A byproduct of this change is the decline of co-flowering by these two species as already explained.

REPRODUCTION 

Lathyrus shifted its blooming time forward with early snowmelt without apparently suffer substantially from night freezes, but it does face the danger of blooming prior to pollinators being very active or even emerging. Night frosts can slow bumblebee queen as well as solitary bee activity and reproduction and cause a delay in daily pollinating until the air has warmed sufficiently. Fewer pollinator visits can in turn result in reduced seed set and reproduction for  Lathyrus.  

Our two habitat-sharing pea species, big-leaved lupine and peavine, plants that play a key part in the beauty and diversity  of the high-mountain, subalpine summer wildflower bloom, give us hints about the complexity of climate change's likely effects on that plantlife that makes a special contribution to the wonders of our earthly existence. Climatic warming does more than simply cause droughts. It throws out of sync the timing (the phenology) of interacting species that depend on one another. Timing matters. Paradoxically, climatic warming also exposes plant life to increased threat from freezing temperatures.

3. THE BORAGE FAMILY (BORAGINACEAE)

DWARF BLUEBELLS (Mertensia fusiform) 

TALL BLUEBELLS (Mertensia ciliata)

GETTING A LIVING  

Dwarf bluebells (M. fusiformis) is among the earliest blooming wildflowers in the Rocky Mountain subalpine meadows near Crested Butte, Colorado, emerging and flowering just one or two weeks after snowmelt.  A plant of this species possesses thick leaves covered with stiff hairs that give it a measure of protection against frosty nights. The plant typically reaches peak blooming in early June, meaning that it emerges in May, a sometimes chilly month when nighttime freezes are common. The special advantage of early-blooming is an absence of competition from very many other species for resources and pollinators, more moisture, and less shading by other vegetation.  The danger of arriving early is damage to plant parts from night frosts and a scarcity of pollinating bumble bees whose own emergence and activity can be retarded by frosty nights. As emphasized in earlier posts, a warming climate induces early snowmelt and warmer early spring days, but leaves the timing of night frosts largely unchanged.

By the time M. fusiformis has gone to seed, it's late arriving tall bluebell (M. ciliata) cousin is in bloom, with its peak flowering occurring sometime in the first or second week of July. This species achieves a height of a meter or more with a single plant often covered with a lineup of multiple hanging flowers against a lush background of soft, hairless green leaves. It expands its coverage locally through clonal colonies created by networks of rhizomes, and favors wet meadows and stream-sides at subalpine and lower alpine elevations. 

DEFENSE

A changing climate affects both dwarf M. Fusiformis and tall M. Ciliate, moving peak blooming forward in time for both species as the date of spring snowmelt advances. The early blooming M. Fusiformis defends itself against night frosts with preformed underground ice-resistant flower buds and thick, ice resistant, hairy leaves, and avoids substantial losses of flower populations in years of early snowmelt and increased exposure to freezing night temperatures. Simply put, this species is pre-adapted to freezing from early blooming's normal increased exposure to frost. Its later blooming relation, M. Ciliata, does, however, suffer lower flower populations in early snowmelt years, but frost is unlikely to be the cause because it blooms so late in the summer. The problem for this species is likely to be a shortage of moisture caused by a lengthening and intensifying of the usual summer drought between snowmelt in May and the arrival of the summer monsoon rains in July and August. The earlier the snowmelt, the longer the drought.

REPRODUCTION 

Plants adapt to early season environmental variability in the form of frost damage or shortage of pollinators by producing flowers that develop later or last longer if un-pollinated giving them a shot at delayed pollination. Some also develop a capacity for early self-pollination, but this is not the case for outcrossing M. Fusiformis. For early blooming M. Fusiformis, a late snow melt in 2008 led to a short but intense and productive flowering period. An early snow melt in 2007 caused a longer flowering season and more late opening flowers. The total number of flowers didn't differ between years but the flowering peak shifted forward in 2007 and early arriving flowers did get exposed to some frost damage. Extended longevity of unpollinated flowers and the production of numerous late season flowers may be giving this species some resilience against climatic warming and the damage of early snowmelt. Possibly because of plant damage from hard frosts, seed set was nonetheless lower in 2007 than 2008.  

As already noted, climate change and early snow melt causes peak blooming to jump farther forward for M. Fusiform than it does for its later blooming taller relation, M. Ciliata. This growing gap could leave bumble bees in a lurch without a source of nectar and pollen in some localities, potentially diminishing their populations. This could in turn create a pollinator shortage for the late blooming M. Ciliata and eventually cause its population to decline. Fortunately, so far researchers don't find any evidence of an actual pollinator shortage. Whatever the cause, in early snowmelt years, the total number of flowers for late blooming M. Ciliata  drops substantially, and some locations plant populations disappear completely where they have been observed in the past. The most likely culprit is a lengthened and intensified midsummer drought caused by earlier snow melts.

To sum up, so far the early-blooming M. Fusiformis has stood up to the challenge of climatic warming, probably because of its pre-adaptation to the night frost problem, but late-blooming  M. Ciliata populations and reproduction appear to be suffering from midsummer moisture shortages occurring because of a warming climate.

2. THE SUNFLOWER FAMILY (ASTERACEAE)

YARROW (Achillea Millefolium) 

GETTING A LIVING

Wherever one goes in the montane and subalpine Rockies, Yarrow (Achillea) seems to be there. It likes slightly disturbed soils in both meadows and forests and doesn't mind a fairly dry landscape. This is a widely distributed species that occurs from sea level up to 12,000 feet elevation.

DEFENSE  

Given its status as a “weedy” species, a warming climate will likely do little harm to this plant given its capacity to flourish in dry meadows right along side such prairie species as big leaf sage. Its been introduced in Australia and New Zealand where it has become an aggressive weedy invasive. Achillea contains inflammation fighting flavonoids and has been used historically as a folk remedy around the world. The adaptive value of flavonoids could be as a defense against fungal pathogens. 

REPRODUCTION

Achillea is a member of the sunflower family with from four to eight ray flowers. It’s flower heads are small, perhaps a centimeter across and the flowers themselves are miniscule. Yarrow is self-compatible, meaning that a flower can become fertile using its own pollen, maximizing its reproductive ability where pollinators are scarce and there is a potential for quick, opportunistic spreading into disturbed soils. A wide range of insect species can pollinate this plant and are attracted by a nectar reward, increasing further its likelihood of reproductive success. Its ubiquity is further assisted by an ability to expand and propagate through spreading rhizomes.

BIG SAGEBRUSH (Artemisia tridentata)

GETTING A LIVING

Big Sagebrush (Artemisia) flourishes in the semi-arid habitats of the Great Basin and Rocky Mountains receiving from 20 to 40 cm (6-16 inches) of rain annually. Artemisia is especially abundant in the central Rockies around Gunnison, Colorado where it provides winter habitat to the threatened Gunnison Sage Grouse. In this area, the plant reaches its highest elevations in the drier subalpine meadows above Crested Butte.The plant's big advantage in dry habitats is its 1-4 meter taproot reaching down into the damper soils unavailable to many other high-mountain meadow plants. Artemisia reproduces by way of both seedlings and shoots charging up from rhizomes that give it the ability to easily spread from existing plants out over the landscape. Artemisia is threatened throughout much its range by the invasive, fire-prone cheatgrass that displaces it through periodic burns, and from being uprooted for grazing and other kinds of land development. Nonetheless, the plant is poised to expand its range at the expense of showy mountain wildflowers in the Crested Butte area if recent research findings stand the test of time. Pollen deposits over the past 10,000 years reveal a hint of what can happen to montane meadows in the mountain west under a warmer climate. Pollen records show that as the tree-line moved upward historically during periods  of climatic warming, Artemisia pollen becomes more abundant. In short, as climate warmed in the distant past, Artemisia flourished.

Does this mean that a warmer climate today will cause Artemisia  to outcompete and displace showy forbs (i.e. wildflowers) common currently in high-mountain meadows?  We will only know this for sure in a future warmer world, but biologists can tell us what that future might be like by simulating climatic warming in a montane meadow by using overhead infrared heating lamps on test plots and comparing plant growth for different species with untouched adjacent control plots. This is exactly what scientists at the Rocky Mountain Biological Lab in Gothic, Colorado having been doing now for two decades on a meadow containing more than 100 plant species, the vast majority of which are those wildflowers that absorb the attention of so many of us who visit the area for the enjoyment of high mountain beauty. On this meadow, researchers have carefully kept track of soil temperatures, moisture, and other measures over the full period of time for both test and control plots, and they have measured above ground biomass for shrubs and forbs to measure responses to artificial heating. The meadow includes a dry upland portion as well as a slope between and a wet lowland segment. In the relatively dry areas of the site, Artemisia  has expanded its biomass at the expense of forms in the warmed plots in comparison to controls, while in the wet areas shrubby cinquefoil (Pentaphylloides floribunda) did the same thing. In the warmed plots, sagebrush also expanded its seedling establishment. These results together infer that showy wildflowers will likely be displaced by better-adapted shrubs in a warmer world. Instead of diverse meadows in the Crested Butte wildflower wonderland, we will see instead an expanded sage scrubland.

DEFENSE

Artemisia is a visually satisfying silver-gray evergreen shrub with small three-lobed leaves and a pungent, wonderful, addictive odor emanating from its inner complements of terpenoid compounds. Pick a few leaves and squish them in your hand and breath in; you will love it! Artemisia is pictured here with Wyoming Paintbrush, an attractive but threatening root parasite that sucks up nutrients. Artemisia nonetheless overcomes this threat and is a highly successful prairie plant that does well in an arid environment. One reason for its success is its capacity to engage in chemical warfare. That wonderful “sage” smell that smashed Artemisia gives off comes from those very terpenoids that give the plant a special advantage in competing with seedlings and young plants from other species. The terpenoids inhibit the growth of seedlings and lowers the respiration rate of juvenile plants giving Artemisia a huge advantage in its competitive struggle for spatial dominance. These same chemicals also ward off some of the plant's herbivorous predators.

REPRODUCTION

In the world of wildflowers, pollinators typically do the work of getting pollen from one plant to another. For Artemisia, this work is done by the wind. Wind pollination is common among trees but less so for shrubs. The advantage is that the plant has no worries about a shortage of pollinators; the wind will invariably blow. A wind pollination strategy could be an advantage in droughty landscapes where a long dry spells periodically depress pollinator populations. Drought could thus be to the benefit of Artemisia relative to wildflowers. In a world of climate warming, Artemisia looks to have a bright future in high elevation Rocky Mountain habitats. This is a faint repayment for the stunning losses of sagebrush habitat to chaining, burning, and herbicide spraying to make room for cattle forage in the western great plains. Millions of acres of sage brush habitat have been taken over by agriculture, roads, urbanization, energy development, exotic plants, and woodlands to the detriment of threatened species such as the sage grouse. Millions more acres are threatened by cheat grass invasion. The upward extension of Artemisia into meadow habitat will be a minor compensation for past harms, and is probably not worth the habitat loss that will result for mountain wildflowers and their dependent herbivores and pollinators.

SHOWY FLEABANE (Erigeron  specious)

GETTING A LIVING

 Fleabane (Erigeron) spreads itself widely through the high mountain landscape all the way from lower elevation sagebrush prairies up to the highest subalpine meadows splashing shades of purple and lavender all over the place. In spreading it is helped along by rhizomes in its root structure. The contrast of its orange disk flowers with its lavender rays can't help but get the attention of any wildflower lover, especially a photographer. Erigeron flourishes alongside sagebrush in dry, rocky prairies the lower elevations, but it also does well in high-elevation wet or dry meadows alongside a variety other showy wildflowers. 

DEFENSE

Erigeron survives the challenges of summer drought pretty effectively once it comes into bloom and for this reason could be up to the challenge of climate warming. The problem is, its one vulnerability against which it lacks a defense, and that is climate change-induced increased exposure to night freezes that kill its flower buds. Early snowmelt results in Erigeron emerging ahead of schedule and facing increased exposure to freezing temperatures at night. Even though climate warming raises daytime temperatures and moves average Rocky Mountain snowmelt dates forward in time, the timing of below freezing night temperatures remains essentially unchanged. Since the 1970s, the average snowmelt date per decade has advanced by about 4 days. With snow melt dates advancing rapidly in the Rocky Mountains and the initial emergence of Erigeron following suit, more and more plants get exposed to night frost that damages the plant’s flower parts and leaves and frequently killing flower buds. In some years with early snowmelt in the Rocky Mountain Biological Lab survey plots, most or all flower buds are killed by frost. The abundance of Erigeron flowers on these plots between 1975 and 2006 has diminished  as the snowmelt date moved forward in time. 

REPRODUCTION

A key pollinator for Erigeron is a butterfly, Mormon fritillary (Speyeria mormonia). This same pollinator in turn depends heavily on Erigeron for nectar and successful reproduction. In years when Erigeron populations are low because of exposure to night freezing, Speyeria populations suffer and can’t lay as many healthy eggs as otherwise and populations of the next generation in the following year are reduced. Not only do night freezes harm Erigeron and Speyeria reproduction, such freezes also damage Speyeria larvae and pupae reduces their chances for survival to adulthood. A reduction in Speyeria populations may have its own cascading negative effects on other species it normally visits such as orange sneezeweed (Dugaldia hoopsii), curly golden weed (Pyrrocoma crocus), and showy goldeneye (Heliomeris multiflora). These three wildflower species will see their populations suffer reductions in pollination by Speyeria. 

ASPEN SUNFLOWER (Hellianthella quinquenervis)

GETTING A LIVING 

Aspen sunflower (Hellianthella) resides in high elevation montane and subalpine meadows and open woodlands. It is one of those big, showy, yellow spring and early summer wildflower that stands out and advertises to the world, and especially to pollinators, "here I am." Hellianthella ironically means "little sunflower," but these guys are by no means little, reaching as much as 1-2 meters in height and sporting always east-facing, single flower heads that reach 10cm in diameter. Their mostly basil leaves have five veins running through them if you need some security about having the plant correctly identified. 

DEFENSE

Some countries have historically hired mercenary armies to defend themselves against their enemies. In a certain sense, some plants do the same thing. Hellianthella does something very much like this to defend itself against egg-laying flies whose larvae prey on its seeds. Hellianthella produces nectar in its tiny flower bracts on its flower head that is continuously harvested by ants throughout the summer. In the process of collecting nectar, ants chase off flies that would otherwise lay their eggs. The ants get a living and Hellianthella gets its seeds protected and increases its potential for reproduction. Unfortunately, Hellianthella is not so well defended against night freezes.

The single most important force driving biological timing at high elevations is the date of snowmelt. If snows melt earlier, plants will emerge and bloom earlier, gaining a longer season in which to grow and reproduce. The plus of a longer growing season can be more than trumped by a dangerous stress: increased exposure to night frost from early emergence due to a shift backward in the snowmelt date and stability in the nighttime freezing temperature profile. Hellianthella lacks defenses against frost. Because it emerges and develops typically after snowmelt, and snowmelt historically occurred mostly after the danger of nighttime freezes, such defenses were unnecessary. A warming climate can cause daytime temperatures to rise above freezing earlier in the spring but works more slowly on pushing back nighttime freezing temperature. Early snowmelt moves up the timing of first emergence for Hellianthella, exposing flowers to a greater danger of frost damage. Seventy-four  percent of  flower buds have been damaged by frost annually on survey plots near the Rocky Mountain Biological Laboratory in recent years. The resulting loss of flowers and the seeds they produce dampens recruitment to the  plant's populations and diminishes food supplies for pollinators, herbivores, and seed predators. 

REPRODUCTION

A "killing frost" reeks havoc on plant parts essential to survival and and reproduction. Older plant parts such as stems and mature leaves are better able to resist frost then newer leaves and buds. A plant's reproductive organs are usually the most frost sensitive of all. Flowers and ovaries in Colorado Rocky's wildflowers, such as the Hellianthella, frequently suffer damage from early season frosts even though leaves and stems survive. Below freezing, ice crystals can form within and between cells destroying cell walls and causing plant solutes to leak away. Frost damage can also become a pathway for infection by pathogens, and freezing in soils can cause soil heaving that can damage newly emergent plants. 

Over the long haul, the picture on frost and plant population  health is murkier. An early spring killing frost in one year will reduce seedlings in the next but also put a dent in future seed predator populations who will lack a fruitful site to lay their eggs. With less predation in the next year, seed survival will increase and a boom in seedlings will occur in year two. Predator populations will take more time to recover than seedlings giving wildflower populations such as Hellianthella a window for population recovery. Here is a plus offered by climate change--starved pathogens. Species prone to frost exposure because of early blooming may well genetically adapt with an in-species variation in timing for emergence and blooming, and this could be the case for Hellianthella. If early spring killing frosts become the norm, those plants within the species genetically prone to a later emergence and blooming time will selectively become more predominant. This won't be a free lunch because increasing populations of late bloomers will be constrained by competition for space, resources, and  pollinators with suites of other species already adapted for late blooming. Evolutionary adaptation may offer some wiggle room for dealing with climate change but how much is unclear.

1. INTRODUCTION: THE LIVES OF HIGH MOUNTAIN WILDFLOWERS IN A WARMER WORLD

THE BEING OF WILDFLOWERS

To exist for an organic being is to live. And to live is to gain sustenance from the earth's material resources, to defend oneself against the perils of being, and to reproduce one's kind. This is what life is all about at its simplest. Existence as a living thing just "is." And the continuation of that "is", to repeat, requires  the getting of a living, self-defense, and reproduction. These imperatives form the basis of life for anything organic, wildflowers included.  To meet these imperatives is no piece of cake for the vast majority of organic life. Earth's resources are limited and everything ultimately must compete against everything else for access to the necessities of life. Nature is a tough task master. Success in the competitive race is a question of evolutionary adaptation to a resource niche. Occupy a niche to flourish. Only the well adapted survive. 

The exception to the resource scarcity rule is modern humanity. By virtue of our capacity to produce beyond biological necessity, we have time left over in the day to do something other than obeying the biological imperatives. This is called ecological release. We are released from the scarcity imperative. In short, we human beings have time to engage in experiences in the world for their own sake. We can tell stories, dance, play softball, or write philosophical tracts. We can contemplate the wonders of being and the beauties of nature. We can do work simply for the purpose of satisfying our creative impulses. And we can sit around trying to figure out the meaning of life on earth. These things wildflowers and  other biotic organisms can't do. 

The odd thing is that despite the blessing of ecological release humanity in the modern world has gotten stuck in the trio of life's imperatives and has become as a result a bull in nature's global china shop. We have become paranoid about getting a living and defending ourselves and spend way to much time at producing material possessions we do not need, absorbing way too much of nature's resources in the process. We collectively possess a will to material power that defeats our underlying desire to experience the wonders of life and to fulfill our creative impulses. The fact that we have ecological release leads us to the "bull in a china shop" manipulation of the natural world requiring huge amounts of energy from fossil fuels that shoot excessive volumes of greenhouse gases into the earths atmosphere. These gases (mostly carbon dioxide) are transparent to the incoming light rays from the sun, but absorb a proportion of the reflected infrared energy that bounces back into the outer space. Their accumulation results in a warming of of the earth's atmosphere and a change in climatic conditions faced by its plant and animal species. This result is a diminished capacity for some wildflower species to defend themselves from perils they face, to get an adequate living in the form of solar energy, nutrients, gases such as CO2, and water, and to produce more of their own kind. Nature would be better off if we took ecological release seriously as something to be treasured and used for qualitative experiences rather than an expansion of our power for material production. We have ecological release but we produce as if we are perpetually threatened with material poverty, as if there is no tomorrow without more material stuff.

Why should we as human beings care at all about anything so frivolous as the lives of high mountain wildflowers? Doing so is unlikely to expand our productive powers, make us wealthier, or solve the problem of unemployment. The best I can do in justifying such care is to claim that doing so may help us wean ourselves off of our current imperative to perpetual economic expansion and its attendant environmental threats. Exploring the lives of wildflowers gives us an endlessly fascinating and aesthetically pleasing  project that will keep us off the treadmill of endlessly expanding material production and give us something to do in the world that is environmentally benign. In short, get out there and enjoy the drama and adventure of the lives of wildflowers.

GETTING A LIVING

Water, mineral nutrients, space, sunlight and CO2 for photosynthesis, and an amenable temperature; these are the mother's milk of getting a livings for a wildflower. Plants, like any living thing, must devote time and energy to the essentials of building and sustaining organic life. The essentials of getting a living is conceptually easy to state, but the devilish reality is in the complexity of the details. Photosynthesis for example, the process by which sunshine, CO2, minerals, and water are converted to energy plants can use to live, is  a complicated business not easily understood.

DEFENSE

For a wildflower, nature is filled with evils that have to be avoided or subdued for survival--herbivores, parasites, damaging weather events, competitors appropriating necessary resources. Organic life is no piece of cake. Wildflowers, in addition to being objects of beauty, have to be tough. Some engage in chemical warfare by producing compounds that repulse or debilitated their enemies. Some develop toughened skins on their plant parts to defend against freezes or droughts. Some engage in symbiotic relationships through quid pro quote deals with other organisms that take up the work of defense on behalf of the flower in exchange for nutrients. Some wildflowers avoid damage from summer downpours by simply closing up their flower parts for business when it gets cloudy. 

REPRODUCTION

Sensuality, connection, and nurture; these are the essentials of wildflower reproduction. On top of being good at self-defense and an aggressive seekers of life's resources,  wildflowers must be objects of attractive beauty in the eyes of their pollinators. It's the pollinator that transfers pollen from male to female flower and thus male sperm to female egg. Only those wildflowers with the ability to self pollinate from their male stamen to their own female style can be ugly ducklings. They don't necessarily need pollinators to reproduce, although they do at some point to avoid the problem of insufficient genetic diversity and inbreeding depression. Flowers nurture their young, but they do so primarily through the legacy of energy they leave behind within the seed's embryo for the seedlings initial emergence. Parental care is limited to perennials who sometimes shade their offspring seedlings in dangerously hot climates.  The same task is often better performed by unrelated shrubs with a comparatively expansive and deep shadow.

UNDERSTANDING BEING 

Understanding is a mental and material process about which we humans lack much real understanding. This lack points up humanity's strictly limited capacity for perception, especially of its own mental functioning. All we have is our perceptual senses as our window to the world. We may well have more luck in understanding the Being of a wildflower than our own. Its Being is driven by real necessity, but ours is driven by a destructive mock necessity. We think that we have to devote the bulk of our day to getting a living, but we don't. We think that continuous economic expansion is essential to employ everyone, but it is not. We think that life's meaning comes from material possessions, but it doesn't. It just might come from getting to know the Being of Wildflowers and the dependency of that Being on humanity's behavior.