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…

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!!!

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! 😦

Is it Spring (Beauty) time already?! pt. III

Oh yes, I think it is finally here… Spring Beauty time!!!

2014-02-22 10.47.31Pictured above is an emerging Claytonia “yorkii” plant, found growing in the southern Sierra Nevada at Cross Mountain (Jawbone Canyon) in Kern County almost two weeks ago.

2014-02-22 12.27.47Unfortunately, I didn’t find any plants flowering on this windswept ridge (all were emerging or in bud), so I tried my best not to disturb the ones I did find as I proceeded to dig up a few tubers. I managed to safely transport a few plants to their new homes at Rancho Santa Ana Botanic Garden (where I am conducting my research). To my surprise (I don’t have a green thumb by any means), the buds continued to develop and the first flowers are just beginning to open!

2014-03-05 09.46.18Now if I can just manage to get a few more species into the greenhouse in the near future, I might be able to try a crossing study or two. What I’m really interested in is whether or not these ‘strongly out-crossing’ tuberous perennials are capable of self-pollinating like many of their close relatives (such as in the Claytonia perfoliata complex).

2014-03-03 15.59.52So why should you care, exactly, whether or not it is Claytonia time?!

2014-03-04 12.56.14 Well… you can help me study something really fascinating about Claytonia, that’s why! Send me your pictures so we can get to the bottom of this — How much variation is typical in terms of petal shape, size, and color for a given population of Claytonia?! My collaborators and I have noticed quite a bit of variation among populations of different species, but what may be even more interesting is the amount of variation in petal morphology within populations of a species. It might bend your brain the next time you kneel down and take a look around at the Claytonia blooming in your area. Better yet, take a picture of 50-100 flowers from directly facing the flowers (trying to center the gynoecium in the flower as best as possible), and then compare them (or send them my way). Crazy, right?

2014-03-05 09.43.25This could turn out to be a very interesting experiment indeed 😉

…but only if you can get out into the field, and remember to send me your pictures of Claytonia!!! Eastern or Western species of North America, Northeastern Asian species, it doesn’t matter — show off your Spring Beauties!

Who is pollinating Claytonia in California?! pt. II

I am still thinking about pollination biology of the tuberous perennial species of Claytonia in southern California, which often have large flowers compared to the annual species in the area. I am making an effort to observe more pollinators this year after having so few observations in the last two years (see pt. I). I have yet to observe the same pollinator at any given locality in southern California for the five putative taxa in the C. “peirsonii” complex. What are they pollinated by? To me, it is all very interesting stuff and I’m happy to take what I can get here and there… but my collaborator Diana Jolles (credit all photos) set out on our previous hike to shoot as many pollinator photos as possible so we might get to the bottom of this. Thanks to Diana’s efforts, we are gathering more clues about who may be pollinating the tuberous perennial Claytonia of southern California, this time from the San Gabriel Mountains populations.

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The insect visiting the Claytonia flower above is a Greater Bee Fly (Bombylius major), a member of the Bombyliidae. It was spending a lot of time visiting each flower: what a fuzzy fly!

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Pictured above are some of Diana’s photos from our first observation of a solitary bee visiting any tuberous perennial Claytonia in southern California. This gorgeous, green, metallic organism is a member of the genus Osmia (orchard bees) in the Megachilidae. These are in the running for the coolest looking native bees in California in my book. Such amazing coloration!

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Lastly, above are Diana’s photos of a third insect visitor seen on February 15th, 2014 near the Devil’s Punchbowl County Park. This dark butterfly is a Duskywing (Erynnis), a member of the Hesperiidae (skippers). It was hard to get a shot as the butterfly hardly seemed to want to sit still, but Diana managed to sneak in a few good ones — thanks so much for all the pictures! I’d like to thank also one of my collaborators, Dr. Emile Fiesler, for identifications of these beautiful pollinators. I hope we can make more pollinator observations this year for all of the members of the C. “peirsonii” complex. Still no overlap in pollinators observed among the members of this southern California species complex, a group currently included in the broad circumscription of Claytonia lanceolata!

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!

My first targeted Claytonia hike: Eastern San Gabriel Mountains

I was dropped off at the base of the ski lifts for Mt. Baldy Ski resort as the sun was rising over a gorgeous day in May of 2012. It happened to be a shuttle hike approximately 17.5 miles end to end in the Cucamonga Wilderness of the Angeles National Forest; another solo hike… I hope I’ll be able to find this little thing!

I climbed up as quickly as I could towards the ski runs under Thunder Mountain, where what was once treated as Claytonia lanceolata var. peirsonii is reported from the steep, forested ridges between the runs. I hadn’t seen these plants in the San Gabriel Mountains just yet, but by this time I had stumbled across a Claytonia in the San Bernardino Mountains two years prior that answered to the rather vague description of this taxon (var. peirsonii) while working for the USDA Forest Service.

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That’s it! North-facing slopes! It can be found blooming next to melting patches of snow in areas where there is also heavy accumulation of conifer litter, often with associated species such as Pinus lambertiana, Abies concolor, Pyrola dentata and Fritillaria pinetorum. I don’t see any snow over there, but it fits the bill otherwise!

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OK, not quite flowering, but it’ll do. I’ve got two more peaks to ‘bag’ before this day is done if I am to see more Claytonia on this hike. I set off toward Telegraph Peak, where I had to shimmy a bit off trail right over the edge of the north slope down to another historically collected locality for this rare local endemic.

photo copy 10A bit of snow around here, this is more like it! Now I’ve just got to find the little buggers flowering…

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What a beauty! Spring Beauty that is… now I just had to get back up to the top of Telegraph to grab the trail over toward Timber Mountain to complete the ‘Three T’s Hike’ I’ve heard so much about.

photo copy 23The north slope of Timber Mountain seemed to be the most ‘typical’ habitat based upon reports for Claytonia lanceolata var. peirsonii, with persistent snow patches as I had just seen over at Telegraph Peak. Sure enough, I was not disappointed by the beauties there!

photo copy 31I made it down to the trailhead around 6pm at Ice House Canyon, where my car had been hanging out all day waiting to hear the news about my big adventure.

“You’ll never believe me, WaSaabi, but I accidentally stepped right onto a rattlesnake’s head this morning. He didn’t seem too thrilled about it.”

If you’re going to hike 25 miles in one day, it better be worth it…

ImageThe utterly gorgeous Spring Beauty pictured here was a sight for sore eyes after I had a less than pleasant hike this day in March at Death Valley National Park. I drove past Badwater, the low point that most people are familiar with, and grabbed a dirt road that headed toward the Panamint Mountains on the west side of the valley in the direction of Telescope Peak. Little did I know that what I was expecting to be a well-graded dirt road turned out to be a 4-wheel drive only road, and my proposed 9 mile, 3500 ft gain and loss hike had just turned into a 25 mile, 7000 ft gain and loss hike if I wanted to see the Claytonia. Good morning from just above sea level!

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The road was long and grueling, with the trail being only slightly more forgiving. I made it up into the Pinyon/Juniper belt by the afternoon and I really started to feel good about the habitat I was seeing, but I was more than exhausted from the hike, and I had a rapidly approaching turn around time of 5pm. It’s got to be around here…

ImageMy 5pm alarm went off as I dug up the first couple of plants for voucher specimens. I had to work quickly and run down the mountain before dark set in too deeply, with my head on a swivel to watch for mountain lions. Did I forget to mention I was alone? I’ll definitely be back to this and other desert mountain ranges next spring in the very early spring months (Claytonia is often found blooming next to melting patches of snow). But next time, I’m bringing a truck! Long hike…

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no luck with Peirson’s rock cress

I went up to San Gorgonio Mountain last weekend with Diana Jolles (all pictures credit D.D. Jolles) to look for Boechera peirsonii at a locality it has not been found. We looked for it a bit south of Charlton Peak near (just below) Dry Lake View, accessed from Dry Lake. This sits on the west end of the bowl that was once a large glacier on the north slope of San Gorgonio Mountain, only a stone’s throw from the type locality for Boechera peirsonii.

DSC_6887We checked out this super sweet habitat, but didn’t find any Boechera peirsonii just below Dry Lake View, at the toe slope of the north face of San Gorgonio Mountain.

DSC_6890Not to say that we didn’t find any Boechera, we just didn’t find the one we were looking for! Looks like what we call Boechera platysperma in southern California needs some work!

Next week, cirque east of San Gorgonio Mountain above Mineshaft Saddle. Fish Creek Trail? Any takers?