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…

And now something different…

It has been a while since I blabbered about the cotyledons of Claytonia, but I want to let you all in on something I find VERY interesting…IMG_1521… so maybe we should start with a survey, to see what you think. How many cotyledons do you think Claytonia have? Keep in mind, they are dicots.

Claytonia saxosa seedlings2013-12-01 12.10.10If you guessed that Claytonia have 2 cotyledons, you’re right… but you’re not the only one that is right. Technically, those of you that guessed Claytonia have only 1 cotyledon are also correct — that’s right, there is in fact a group of dicots with only 1 cotyledon (probably several, but that question exceeds the scope of this blog post). Claytonia Section Claytonia, otherwise known as the tuberous perennials, lack a second cotyledon present in other species of Claytonia (and all of their closest relatives). To me, this is just another reason why you should believe that Claytonia is the whackiest group of plants this side of the Mississippi River. 😉

IMGP9244So who cares, there has been a loss of one of the cotyledons in this group of plants. One time only evolution, and now these plants simply can’t recover that lost cotyledon — I’m over it… right? WRONG! There is something fishy going on here, and it has to do with a certain caudicose perennial I have mentioned before: Claytonia megarhiza (pictured below).

IMG_4012IMG_4025You can see from the second image (the photo immediately above) that Claytonia megarhiza clearly has two cotyledons, not one like the tuberous, perennial Claytonia species I mentioned before. Thus, you’d expect that this species is more closely related to those other Claytonia that have two cotyledons, right? Well…

Print Trees PreviewAbove is a preliminary phylogenetic tree that I presented at the Botanical Society of America Meeting this year in Edmonton, Alberta, Canada. This tree has been developed from ~500 single nucleotide polymorphisms isolated from the nuclear genome of all of the samples included (using ddRADseq). You can see from the tree that the caudicose perennial C. megarhiza (indicated by [morphologically similar, but anatomically quite different] cartoon carrots ) clearly has a close association with the tuberous, perennial Claytonia, albeit the exact area in the tree where they will stabilize is still yet to be determined. Without question, Claytonia megarhiza is nested somewhere in this clade of otherwise tuberous, perennial Claytonia.

So how did C. megarhiza find itself with 2 cotyledons while all of its closest relatives (including those diverging away from the lot much earlier in evolutionary history) have only 1? If you have the answer… I would love to hear it from ya! From my perspective, it is a question that is ‘to be determined’ but I am hopeful that my dissertation will change things. 😉

Oops, that took embarassingly long: UPDATE on what’s new with western Claytonia

IMG_0850First off (technically second, after the eye candy above), I need to announce that my research on Claytonia (Montiaceae) has been recently funded [yay!] by the National Science Foundation. You can see the abstract here for #DEB1502085:

http://www.nsf.gov/awardsearch/showAward?AWD_ID=1502085

Second (yes, I know, technically third), we should play my favorite game: compare the above eye candy with the below ridiculously beautiful [spring] beauty… SAME? or different. 😉

IMG_2978Now before you answer to yourself, consider that the Claytonia in the above two pictures are considered in the current taxonomy to be the same as the plants in the below picture, all being C. lanceolata [for now]. IMG_2834Check out the variation in those leaves!!! Who cares about the flowers, right? No way…IMG_0865IMG_7563IMG_2825The three photos immediately above correspond respectively with the same three putative taxa [let’s call them operational taxonomic units] in the first three pictures of this blog post. So? SAME or different? Feel free to vote at the bottom, and I welcome any feedback or discussion — I can’t say it all in 140 characters or less.

Third [ok, let’s stop with the sequential nonsense], I took an epic trip across California, Nevada and Oregon during the spring months of 2015 as part of #DEB1502085. While on this trip, I photographed new, mind-boggling [taxonomically speaking], tuberous Claytonia. Almost immediately after that, I took another most epic trip across California, Nevada, Utah, and Colorado (with guest appearances in Arizona and Wyoming) with ‘Los Caryophylleros’. Let’s just say, I can’t wait to get my new morphological and molecular data (RADSeq X 2) up to speed…

Lastly [HA! I win!], I’ll finish this [short] commentary with what has been the most difficult collection of photos I’ve ever had to select from for the purpose of sharing, then I need to finish preparing for presentations at BOTANY and the Caryophyllales meetings…

IMG_1648IMG_1815IMG_2120IMG_2266IMG_2280IMG_2297IMG_2585IMG_2598IMG_2696IMG_2821IMG_2944IMG_2985IMG_3223IMG_3256IMG_3520IMG_3578IMG_3582IMG_3665IMG_3728IMG_3756What a wild ride!!!

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.

Will the REAL Claytonia lanceolata please stand up…

Hey all!

I wanted to take this opportunity to give you a brief overview of my dissertation research, hopefully explaining a bit better how my own research relates to the AWESOME project I’m flying for a limited time only on experiment.com in collaboration with Dr. Ingrid Jordon-Thaden — I know you’re interested in supporting a good cause, so check it out! See the project here.

Many of you may think that you have seen Claytonia lanceolata some time during one of your forays out into the field, but have you REALLY seen it, or just something that looks like it?! Where were you when you saw it? If your answer isn’t Sweetwater, Idaho, than I’ve got some news that could make you want to reconsider what you think you saw — it is possible that you’ve been duped by a lance-leaved imposter!!! Don’t worry, you’re not the only one…

pursh1814_sketch.tiffAbove is an illustration of Claytonia lanceolata from the original description published in the Flora Americae Septentrionalis (Pursh 1814). The illustration is putatively drawn from the type material collected by M. Lewis, although one cannot be entirely sure given how poorly Claytonia seem to make the transition from live plants in the ground to becoming flattened museum specimens — their succulence does not do them any favors in terms of preserving diagnostic morphological characteristics.

Did your plants look like the one in the above illustration from the original description? What about the plants below photographed in the Klamath Region of northern California, last year, growing very near to North Trinity Mountain — did your plants look like these? As far as I can tell, this is the REAL Claytonia lanceolata,or at least something VERY closely related, the best proxy I’ve got (for now) for C. lanceolata sensu stricto.

2013-05-30 11.37.042013-05-30 13.44.13I mention these might be only very near C. lanceolata sensu stricto because these plants have entire petals — C. lanceolata is described as having bifid petals, probably more similar to something like what you see on the flowers of C. sibirica. The basal leaves are also lacking in the plants pictured above from northern California (only observed to be associated with non-flowering individuals in the population), but the sessile, triple-ribbed, ovate cauline leaf pair and the lax, elongate, solitary inflorescence is a dead ringer for C. lanceolata Pursh. The plot thickens when you step off of the granite and onto ultramafic substrates…

2013-05-28 11.53.36 2013-05-28 11.12.53Wait a second — is that a second raceme I see? Lance-linear cauline leaves with a single vein? And are those leaves ALTERNATE?! Surely this is not the same as C. lanceolata sensu stricto — this is one of many lance-leaved imposters masquerading as C. lanceolata Pursh. For better or for worse, the Operational Taxonomic Unit (OTU) above is treated as C. lanceolata in the recent monograph for Claytonia (Miller and Chambers 2006) as well as in current treatments in the Jepson Manual and the Flora of North America. The following plants photographed in populations from throughout California also are ALL treated as C. lanceolata under the current circumscription by Miller and Chambers (2006):

IMG_2684IMG_9512IMG_5528IMG_5450IMG_49592014-02-15 13.41.46Whoa — I feel like this kind of variation might not ALL be related to ‘plastic’ responses to a heterogeneous environment… and preliminary molecular data seems to agree thus far.

This is where the Experiment Project is a TREMENDOUS help — It’d be great to get more samples from outside of California… expanding onto more substrates from across the distribution of C. lanceolata sensu lato, which ranges from Canada to Mexico. Idaho, Montana, and Yukon Territory are on this bill…

The REAL kicker? Check out the last line in Pursh’s 1814 description of C. lanceolata below: pursh1814.tiff“In the collection of A. B. Lambert, Esq. I found a specimen collected by Pallas in the eastern parts of Siberia, perfectly agreeing with the present species.”

Did you see my post about going to Siberia? I’m working on a grant for that right now… I’m really eager to see more plants in the field — as I mentioned previously, these plants don’t preserve all that well! 😦

Here today, gone tomorrow — Doyle and Doyle get a taste of the Desert Southwest Claytonia

IMG_5134IMG_5178Remember the above, beautiful plants photographed flowering a little over two weeks ago in Nevada? If not, see my previous blog post, ¡Holy frijoles!

Well, almost two weeks later Claytonia umbellata is apparently no longer flowering atop Peavine Peak outside of Reno — more snow has come (see below). These poor little guys are now packing on the pounds (and buds) getting very little exercise while they sit and wait for the warmth of a spring (beauty) day. It will come soon. In fact, I’d wager those little monsters are blooming now, as I’m writing from the comfort of my home in southern California. After all, the photographs below are from April 2nd, 2014, and these plants (desert Claytonia) are über-ephemeral. IMG_5739I’m so cold, I couldn’t possibly think of flowering right now…

Dr. Jeff Doyle, Cornell University, searching for the elusive flowering Claytonia.

Dr. Jeff Doyle (Cornell University) trying to stay warm while searching for the elusive flowering Claytonia.

Well, this disappointment came AFTER a morning of heartbreak — searching seemingly suitable habitat without finding plants. Do I really have a good idea of what suitable habitat is? Are they even capable of dispersing here? Let’s back up for a second…

IMG_5221Above is a picture of a location where I collected another population of Claytonia umbellata in the Pine Nut Mountains at Minnehaha Canyon on my spring break trip nearly two weeks ago — the talus I know and love, seemingly characteristic of the Desert Southwest Claytonia. Bearing that in mind, when I see a habitat like the one above, or the one pictured below, and I am very near the type locality of Claytonia umbellata in the area of Mt. Davidson Pass (Nevada), I have to stop and take a look, right? IMG_5724WRONG! There aren’t any Claytonia there (above), not even a C. rubra or C. parviflora ssp. viridis to cheer me up for making a bad call — Nothing! But it looks so good, and the rocks are the same as at Minnehaha Canyon… sample size of one strikes again with these guys. Ever heard the phrase, to each his own? Well, I’m beginning to think these Claytonia take that to the extreme. They are not very good at getting around (dispersal), or they have very picky palettes… or both.

We have just GOT to find some flowering Claytonia, I promised Jeff and Jane! The Doyles joined me for this trip (and to give a seminar at RSABG later), flying all the way out from New York. The Doyles have A LOT of experience with eastern Claytonia, but this western stuff was pretty new for them. We sat in the car on top of Peavine Peak thinking (and not eating) — what were we going to do? That’s it! I know another place we can try, but its further north…

IMG_5772Pictured above, a talus slope of volcanic rocks in the Pah Rah Range near Pyramid Lake, Nevada. Oh no! The sun will be setting in a few hours! We can try to climb that slide and run the risk of not finding any plants, or we can try to race up the canyon to get to a known locality before dark sets in… Jeff and Jane agree, it’ll be worth giving this hillside a good look over before we continue up the canyon.

IMG_5763And thank goodness we did — this area of talus was rich with numerous plants of Claytonia umbellata! We found a new locality for Claytonia umbellata, approximately one mile away from another known locality in the Pah Rah Range AND it was the large-leaved plants, like those seen in the Pine Nut Mountains at Minnehaha Canyon. Compare those to the plants from Peavine Peak (top of page), which happen to grow on different volcanic rocks!

Déjà vu — a feeling of having already experienced the present situation, independent of whether you have or have not experienced the situation or something like it… No idea what I’m talking about? See my “Claytonia rocks!”post — click here for more background information.

I still was unable, though, to fulfill my promise to Jeff and Jane Doyle — we had not yet found flowering Claytonia. There was one more chance to find flowering plants while in Nevada looking for Claytonia umbellata.This was a revisit for me, to a site in the Pine Nut Mountains where I was unable to locate a population on my previous visit. This population is near Bismark Peak, and as far as I can tell, it has not been documented by any herbarium specimen. I found out about the population by scouting CalPhotos for pictures of Claytonia — this population had been photo documented but never vouchered. I want to give a shout out to Larry Crawford, for giving me enough information to find some plants! I was not able to relocate Larry’s ‘secret patch’ of Claytonia umbellata atop Bismark Peak, but the Doyles and I were able to find a new population along the same ridgeline!

IMG_5782Believe it or not, and I’m sure to Larry’s delight, the plants here occur in a really unique habitat — I’m still working on the rock identification (volcanic, or sedimentary?). Crazy thing is, this Claytonia umbellata population consists of only a few plants scattered about in the cracks of the vertically oriented rocks (presumably oriented as such due to frost heaving) — they don’t seem to occur with the other rocks oriented horizontally.

IMG_5786Isn’t that just bizarre! but there were STILL no flowering Claytonia to be found… I was running out of options — a revisit to the population near the Devil’s Punchbowl County Park on Saturday before Jeff and Jane’s departure seemed the most obvious choice. Rather than visiting the Devil’s Backbone (where recent snows may have set plants back from flowering), I decided we should visit a lower elevation site. Would there be flowering Claytonia hanging on at the Devil’s Punchbowl, where I observed plants flowering on February 15th, 2014? (click here to see post, “Yep, it’s Claytonia time…”)

IMG_5816NOPE, it’s not Claytonia time here, at least not really anymore. There were some plants around with buds, but most of the C. lanceolata var. peirsonii here has dispersed its seeds and are beginning to wither back to their underground tubers. We did not find any flowering — I guess the Doyles will have to come back again next year, or meet me somewhere northward in a few more weeks!

We did at least find some flowering Claytonia rubra there, just in the nick of time. It was time to send Jeff and Jane back to New York. Although they may not have gotten to see (much) flowering Claytonia, at least they got to see some flowering California!

Spectacular desert wildflower bloom in the Mojave desert near Piñon Hills, highway 138.

Spectacular wildflower bloom in the Mojave Desert near Piñon Hills, highway 138.

 

 

Claytonia rubra, what is that? pt. I

Possibly my favorite outgroup material, pictured here is Claytonia rubra, a widespread annual species that is a close relative of the group of tuberous perennials that I study. 2013-04-28 11.17.10

This species has been involved in a number of ’bouts’ of hybridization primarily with two other annual Claytonia (C. perfoliata and C. parviflora), making the identification of annual species in this group incredibly challenging. Polyploid lineages have been recognized as varieties for this species and others formerly treated as C. perfoliata sensu lato, creating what appears like a continuum of morphological variation in one ‘species’ resulting from recombination among three unique lineages (pictured below).

2013-04-28 11.34.01Why do they call it Claytonia rubra? Well, because it is REALLY RED on the abaxial side (bottom) of the leaves, a condition resulting from increased betalain production, aside from its darker green adaxial surfaces as seen in the first picture. Claytonia parviflora (left) with linear leaves and C. perfoliata (middle) with spatulate leaves are readily distinguished from C. rubra by this morphological character alone, but subtle floral features exist as well that can be used for this group, including petal shape.

2013-04-28 11.41.08Next spring, when you’re out hunting for melting patches of snow to see the pretty Claytonia that are in bloom next to them, be on the look out for ‘miner’s lettuce’ at slightly lower elevations than the tuberous perennials but still in areas of high moisture. Claytonia rubra is typically found in Mixed Conifer habitats, often in deep shade and at higher elevations than other members of the C. perfoliata sensu lato complex.

The C. perfoliata sensu lato complex (including C. rubra) is incredibly ruderal and can be found nearly anywhere in the montane habitats of California and the Southwest, making my choice to study the tuberous perennials seem a bit silly at times. Polyploidy seems to be intimately linked with the successful dispersal into new and disturbed habitats in this group, and it will be interesting to compare with the evolutionary patterns and relationships among the tuberous perennial Claytonia.

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.