Claytonia rubra, what is that? pt. II

Hi, all — I’ve decided to switch things up and simply chat with you a bit about a topic that has apparently been the most popular on my blog so far: Claytonia rubra (Howell) Tidestrom. You may find yourself now asking the question, “Claytonia rubra, what is that?” Well, there’s a blog post for that.

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Claytonia rubra photographed while flowering, plants growing abundantly in the Pinyon-Juniper belt on the Modoc Plateau, northern California.

But Claytonia rubra may be more than meets the eye, especially if you’ve been paying close attention to it where it grows in different areas, or if you’ve been so lucky as to see it from across its entire geospatial distribution. Let’s think about this for a minute, considering first the global range of Claytonia rubra.

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Range map from Flora of North America Treatment by Miller and Chambers (2003).

It is across this wide and ecologically diverse distribution, north to south from British Columbia to near (probably in) Mexico, and west to east from the San Francisco Bay Area to the Black Hills of South Dakota, that the annual Claytonia rubra displays a seriously diverse range of morphologies. Is it phenotypic plasticity? Is it local adaptation? Is it speciation? Is it hybridization? Is it some mix of all of these? Claytonia rubra is a tough cookie to crack!

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Claytonia rubra photographed while flowering on the Modoc Plateau, northern California. Notice the largest leaf blades (outermost in the rosette) are among the first to have emerged in early development — These lay flat on the ground surface (or nearly so)

Starting with identification, something I’ve found to be fairly characteristic of Claytonia rubra is that its older leaves (outermost in the rosette of basal leaves), which often have the largest blades, tend to lay flat on the ground — some close relatives (like certain subspecies of C. perfoliata) do this too, or something like it, so don’t take this for a smoking gun. Furthermore, some races of C. rubra do not have basal rosettes that lay flat, rather they are elevated ever so slightly above the forest floor. Don’t you just love those kinds of dichotomous key breaks? — Is is erect to ascending (sometimes spreading), or ascending to spreading (sometimes erect)?! — Still, if you see something in the mountains of western North America like in the picture below, where the outermost (older) leaves in the basal rosette lay flat, appressed to the ground, you can be fairly confident that you are looking at something that is quite appropriately called Claytonia rubra.

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Vegetative Claytonia rubra photographed at the Tejon Ranch in the Tehachapi Mountains, southern California.

For effect, compare the leaves of Claytonia rubra plants in the above picture with the leaves of the plant in the picture immediately below, C. perfoliata, both of which were photographed at the Tejon Ranch in somewhat close proximity (either side of the same mountain ridge) — see how Claytonia perfoliata (below) tends to elevate its leaves off the ground?

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Vegetative Claytonia perfoliata photographed at the Tejon Ranch in the Tehachapi Mountains, southern California.

The characteristic of Claytonia rubra have low-growing leaves seems pretty useful, until you come across a putative lineage (sometimes in the same mountain range!) that looks like the plant below — these plants have distinctly erect / ascending leaves, although the proximal portion of their petioles are still appressed to the ground. This gives the leaves a sort of ‘S-curved’ look. Believe it or not, these are another subspecies of Claytonia rubra!

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Claytonia rubra photographed while flowering at the Tejon Ranch in the Tehachapi Mountains, southern California.

This is a time where it comes in handy to know a few other characters that, when used in combination, can help to distinguish Claytonia rubra from similar-looking miner’s lettuce.

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Classic, beet-colored (reddish to purplish) undersides of basal leaves of Claytonia rubra. Whenever possible, try to observe this character in multiple plants — Generally, the basal leaves should be deep green adaxially and reddish abaxially in Claytonia rubra.

For example, the plants photographed above demonstrate another conspicuous characteristic of most lineages of Claytonia rubra — betalain pigmentation is generally evident on the abaxial (lower) surfaces of the basal (and sometimes cauline) leaves, giving them a purple/reddish appearance. This is a sort of ‘no-brainer’ character I tend to point people towards, but just as with the characteristic basal leaf orientation I mentioned above, this morphological character doesn’t always hold true. Plants love exceptions WAY MORE than rules: in this particular case there appears to be an environmental element to variation in Claytonia betalain pigmentation, as evidenced by some unpublished observations I have made in common garden experiments involving C. perfoliata, C. rubra, and some perennials in the C. “peirsonii” complex.

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Variegation on foliage characteristic of some lineages of Claytonia rubra.

OK, so the leaves are sort of laying flat, and kinda red undeneath, but you’re still not totally convinced you are looking at Claytonia rubra and not some other similar-looking miner’s lettuce. Well, you’re in luck — I’ve noticed that many races of Claytonia rubra tend to have the look of variegated leaves, possibly related to a breaking up of the cuticle during development. If you see something like the plant in the picture(s) above, where there are variegated streaks on the leaves of a Claytonia, it is VERY likely that you are indeed looking at Claytonia rubra.

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[From left to right, Claytonia parviflora, C. perfoliata, and C. rubra!] Multiple miner’s lettuce species growing in sympatry in Mill Creek Canyon, San Bernardino Mountains, California.

IF you still can’t be sure about which Claytonia you are looking at — just ask me! That’s what I am here for…

Beginning of the end of Silene verecunda sensu lato

Hey, all — I thought I would share with you that my recent manuscript with colleagues (Diana D. Jolles and Heath A. Bartosh) describing a new species of Silene (Caryophyllaceae) has finally been published! You can access our article here, or the whole issue is here. It is a very special issue for the California Fish and Game Journal, #100, and the first ever all botany issue for the journal — some really GREAT articles in there that are worth reading, including an introduction by Governor Jerry Brown. Neat Stuff!

combined_finalAs always, I am interested to know what you think about this topic — the Silene verecunda problem is VERY complex, but fortunately the taxonomy isn’t.

Who is pollinating Claytonia in California?! pt. I

I have been thinking a lot lately about pollination biology in the tuberous perennial species of Claytonia, which often have flowers the size of a penny or even larger. What are they pollinated by? It is said that these flowers are often not open for very long, maybe two or three days sometimes, but that they can be visited by a variety of pollinators while open…

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Thanks to Scott Eliason (credit above picture), Botanist on the San Bernardino National Forest, we are beginning to gather some clues about who is pollinating Claytonia in the San Bernardino Mountains. The insect visiting the Claytonia flower above is a Bristle Fly (Tachinidae). Note also that there is a brown Leafhopper (Cicadellidae) perched nearby.

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Pictured above is my first pollinator observation for the tuberous perennial Claytonia in the southern Sierra Nevada, a couple of soft-winged flower beetles (Melyridae). One of my collaborators, Dr. Emile Fiesler, President of Bioveyda-Innoveyda and member of the North American Dipterists Society, suggests these beetles are among the most productive of pollinators! I hope Emile and I can make more pollinator observations next year for some of the other new species of Claytonia I will be describing soon!!!

Claytonia Rocks!

So these plants are considered to be ‘geophytes’ because they have underground storage organs, but I find this so-called classification to be much more meaningful than that for the tuberous perennial Claytonia. In order to better understand this, let me break it down a little bit.

Phyte – Greek, generally referring to a plant.

Geo – Greek, generally referring to anything of or relating to the Earth.

2013-05-30 11.44.33Pictured here is the habitat of the typical variety for the Claytonia lanceolata complex, C. lanceolata sensu stricto. At least in northern California, this taxon is known only from soils derived from granitic rocks. The suggestion has been that as a generalist, C. lanceolata (as currently circumscribed) occurs in a variety of habitat types across its range and thus is highly polymorphic morphologically. Well, there are some interesting substrates in southern California within very close proximity where one can find members of C. “peirsonii” complex occurring in nearly equivalent niche space across different substrates… Wait a second, am I studying geology or botany?!

2013-04-20 09.57.32Because bragging about how awesome the San Bernardino Mountains are is my favorite thing in the world to do, I’d like to take this opportunity to point to some interesting Claytonia there that occur on sedimentary carbonate rocks on the north slope of Bertha Ridge. This is the ridge that separates Big Bear Valley to the south from Holcomb Valley to the north. This is where I first encountered plants that match the description of Claytonia lanceolata var. peirsonii Munz&Johnston, or any plant treated as Claytonia lanceolata in the Jepson Manual or Flora of North America, while I was working as a Rare Plant Technician for the USDA Forest Service on the San Bernardino National Forest. More pictures of these plants can be found on CalPhotos.

IMG_9508These plants, as mentioned previously, are ridiculously cute (see related articles below and cast your vote!). More importantly for this story, these plants occur on dolomite, which is a particular type of carbonate rock containing the mineral dolomite.

2013-04-20 12.17.49And boy-howdy are these rocky hills STEEP! Unfortunately, this is an aspect of the habitat that many of the tuberous perennial species of Claytonia appear to have conserved across lineages, making field work incredibly physically demanding when sites are remote. But what is my point, exactly?

2013-04-20 14.20.56Pictured here, approximately 7 air-km to the north, there exists another population of tuberous perennial Claytonia that is primarily associated with the same species (Pinus flexilis, P. monophylla and Juniperus) but otherwise occurs on a different kind of carbonate rock than the population at Bertha Ridge. This becomes significant when you look at the morphology of the plants here, which is inconsistent with the plants at the Bertha Ridge population just 7 air-km away.

2013-04-20 14.21.55Not only do the plants at this more northern locality on gray dolomite have pink nectary guides, in contrast to the yellow spots at the base of the petals on plants from Bertha Ridge, but cauline leaf shape appears also to be strikingly different between these two populations. This interesting case of inconsistency in overall morphology, coupled with signs of genetic divergence among these populations (see my most recent phylogeny here), has gotten me very excited about patterns of evolution in this group that is already well known for complex histories of hybridization and polyploidy

ClaytoniaRocksAlthough it is all C. lanceolata for now, soon enough we’ll make some sense out of it!

Claytonia lanceolata species complex

figure1The Claytonia lanceolata complex in California. In northern California, (A) Claytonia lanceolata Pursh on granite, (B) alternate-leaved and undescribed C. “serpenticola” on ultramafic rocks, and (C) an unrecognized taxon on greywacke that fits the description of C. obovata Rydb., unsampled in previous phylogenetic studies. In southern California, (D-H) The Claytonia “peirsonii” complex, with (D) var. “peirsonii” on granite in the San Gabriel Mountains near the type locality of C. lanceolata var. peirsonii Munz & Jtn. New discoveries in this complex within a species complex include (E) the long-petioled C. “peirsonii” var. “panamintense” in the Panamint Mountains on dolomite, (F) light pink-flowered var. “yorkii” on volcanic tuff in the southern Sierra Nevada, and two variants in the San Bernardino Mountains on (G) grey and (H) white dolomite at Furnace Canyon and Bertha Ridge, respectively.