Doesn’t the above (or below) picture look like a fun place to do a plant survey? I thought so… until I did it, that is.
Imagine yourself romping around at 11,000 feet in elevation on a slope as steep as the one in the photo(s) above, searching the nooks and crannies of north-facing cliff faces for a specific plant that grows only in that area, all the while trying not to fall down a snow chute as tumbling rocks move under your every footstep. This is what it takes to call oneself an extreme, alpine botanist, as Marta LeFevre-Levy (USFS San Bernardino National Forest, Big Bear Ranger District) learned last month. “Daredevil Botany,” she called it — I couldn’t have said it better myself.
After all was said and done, we found what we were looking for, and plenty of it! Unfortunately, we were late in the flowering phenology season of Boechera in a drought year, so late that these apomictic plants weren’t flowering; but Boechera peirsonii is distinctive in the San Bernardino Mountains even without flowers. By no means did we search every snow chute across the hillside pictured above, but we surveyed enough to know that if you can get into the right habitat (which, unfortunately, is extremely steep, north-facing cliff and snow chute habitats comprised of primarily metamorphic rock), you’ll find Boechera peirsonii hiding out, almost as common as dirt. Interestingly enough, there isn’t a lot of dirt in this habitat, so you may still want to consider it rare! More extensive surveys should be conducted in the cliffs east of where we found plants by someone with more energy than I, probably younger and more daring, too.
Boechera peirsonii, growing on a cliff face at the head of Whitewater Canyon, just east of ‘The Tarn’ below San Gorgonio Mountain.
Short post, but I wanted to share with everyone that we are making progress on the Conservation Plan for Boechera peirsonii, thanks to monetary support from the California Native Plant Society! Expect a Rancho Santa Ana Botanic Garden Special Publication in 2017.
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.
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!
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.
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?
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!
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.
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.
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.
[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…
Hey, all y’all! I am VERY excited to announce that I am ready and willing to share ddRADSeq data (double digest, Restriction site Associated DNA Sequences) with anyone and everyone that can benefit from using them. I am currently testing outgroup rooting using ddRADSeq. Thus, I have data for some distant samples (phylogenetically speaking, relative to my interest group Claytonia) from Order Caryophyllales that someone out there on the world wide web might be interested in using. Maybe someone just wants to try their hand at playing with some Next Generation Sequencing data, just to see what it is like, or in order to gain some experience for the evermore competitive post-doc opportunities out there…
More specifically, I have these free data to offer up:
In the (likely) case that I made an error during the copy-and-paste-athon above, feel free to follow this link to a folder containing all of the data. Also, let me know where the copy/paste errors are and I’ll work quickly to fix them!
Data details: Genomic DNA was digested using EcoRI and MspI enzymes. Window for size selection was +/- 150 base pairs. Libraries were sequenced on an Illumina HiSeq. Entire dataset includes reads from 2 lanes of single-end (lanes one and two) and 1 lane (lane 3) of paired-end sequencing. These data were generated by Global Biologics.
Anything else you want/need to know in order to use these data in a publication, such as voucher information or having the data served in an alternative format, don’t hesitate to ask!
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…… 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.
If 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. 😉
So 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).
You 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…
Above 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. 😉
First 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:
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. 😉
Now 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]. Check out the variation in those leaves!!! Who cares about the flowers, right? No way…The 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], tuberousClaytonia. 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…
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!
As 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.
Hey, all — I’m following up on a popular blog post from last year, ‘which is the cutest?‘, where I pit some of the cutest Claytonia I know against each other in a winner-takes-all Battle Royale… C. saxosa came out on top with about twice as many votes as the other contestants.
I feel the cards may have been stacked during the last contest, the only logical explanation for how C. saxosa could claim victory in a battle o’ cute by such a large margin of victory, right? I have decided to re-run the analysis, this time with denser sampling and a better explanation of how the voting works: There is a poll at the bottom of this post. You can vote for more than one, but you can only vote once!
(Above) OK, starting it off with the ‘reigning’ champion of cute, Claytonia saxosa standing proud on Anthony Peak in northern California — it is a REAL winner. Definitely a reasonable choice but so many options…
(Above) This is a tough one for me — Claytonia lanceolata var. peirsonii from thenorth slope of Bertha Ridgein the San Bernardino Mountains still holds a place near and dear to my heart. It was my first introduction to the Claytonia lanceolata species complex, and the yellow blotches at the base of its otherwise mostly white flowers are just enough to give some glimmer — pink venation of petals is weak in this population. Southern California might be a little over-represented in this new poll, but please bear with me — they’re cute!
Claytonia lanceolata var. peirsonii at the head of Furnace Canyonin the San Bernardino Mountains (above), on the other hand, has quite a bit of pink in the veins of the flower petals. In addition, there is a proliferation of the pink in the veins in place of the yellow blotches typical of many other Claytonia — this is another serious CUTIE Claytonia!!! I’m not giving my vote(s) away this time, but I’d say that this little bugger has a good shot. Still others, still others…
One such other Claytonia worth mentioning is C. lanceolata var. peirsonii from the Panamint Mountains (above), Death Valley National Park — these plants are truly SPECTACULAR! They have similar flowers to those of the plants on Bertha Ridge in the San Bernardino Mountains, but these plants have much more pink in their petal veins, and they seem to have slightly wider petals. Still other members of the C. “peirsonii” complex (= C. lanceolata var. peirsonii) from southern California deserve mention…
(Above) The REAL owner of the epithet ‘peirsonii’, this is Claytonia lanceolata var. peirsonii from the San Gabriel Mountains (where the variety was first described) — this taxonomic group will see some changes soon (if you didn’t notice already), witha systematic study of the C. “peirsonii” complex representing a serious thrust of my dissertation. I’m working, I promise, but there is a significant problem in that these plants make REALLY awful herbarium specimens… the beauty is almost entirely lost, let alone the diagnostic characters, so remember to soak it in while out in the field!
Rounding out members of the Claytonia “peirsonii” complex represented in this contest, C. lanceolata var. peirsonii from the southern Sierra Nevada is pictured above. I thought this cutie would have done better in the last contest — I’ve thrown in some props this time to help boost its score. These Claytonia have continued to stun me year after year with their beautiful floral displays, but there are Claytonia outside of the C. “peirsonii” complex that are certainly contenders in the cutest contest we’ve got going here…
The Claytonia umbellata species complex is yet another difficult taxonomic group that ranks high on the cuteness scale — it has not yet been a contestant in this little experiment of mine! Pictured above is C. umbellata from the Pine Nut Mountains at Minnehaha Canyon, Nevada. It has pretty awesome flowers and grows in a series of very fascinating habitats from talus to fully stabilized rock cracks and ledges — a HIGHLY variable group indeed!
The population of Claytonia umbellata at Peavine Peak (above), Nevada, is another fabulous population worth mentioning — oh, so CUTE! I tried to show these plants to Jeff and Jane Doyle when they came out west, but it didn’t work out when a late storm rolled through the weekend of their trip… they saw mostly cold, non-flowering Claytonia, but suggested that they had a good time anyhow.
(Above) More Claytonia umbellata species complex from the Pine Nut Mountains at Bismark Peak, Nevada — thanks to Larry Crawford for the two photos above from this location. These plants knocked my socks off when I saw them for the first time — they grow in a very bizarre habitat of frost heaved ‘shingles’ of a sedimentary rock unlike anything I’ve seen for members of the complex. I’m still trying to work out the relationships (part of my dissertation research), but I suspect that C. umbellata is rather closely related to the C. “peirsonii” group.
Speaking of close relatives, I’m hoping to figure out this year just where Claytonia sessilifolia in the central Sierra Nevada (above) fits into the picture — it has remained unsampled in previous molecular studies but I’ll be visiting this population later this week and incorporating it into my dataset. Currently, this taxon is not recognized as anything different from C. lanceolata, but I suspect this may be a stretch. Time (and molecular tests!) will tell, but morphology suggests something unique here, and it already has a name in the literature… Thanks to Larry Crawford again for contributing more photos to the most recent edition of the annual Spring Beauty Pageant.Continuing with more northerly taxa in the state of California, Claytonia obovata from the northern Californiaand Oregon Mountains (above) is another heavy hitter in cutest Claytonia contests past — this is blooming in northern California RIGHT NOW! Although this taxon is currently subsumed under a very broad circumscription of C. lanceolata, it is highly divergent both genetically and morphologically. I’m writing up a manuscript right now about this taxon, arguing why it should be recognized and included in the keys in the Jepson Manual. It will DEFINITELY be in my treatment of Claytonia for the Flora of Oregon Project, which I am collaborating on with Dr. Robin O’Quinn at Eastern Washington University.
Of course, in northern California there is also Claytonia “serpenticola” (above), an endemic to ultramafic substrates in the Klamath Region (including Oregon). It has alternate leaves, a different petal shape, and different leaf venation compared with closely sympatric C. lanceolata and C. obovata, so it shouldn’t be confused with either! Yet another taxon that will creep into my Flora of Oregon Treatment… just have to get it into the literature first! Boy, that one is yet another CUTIE — helps that it grows in particularly beautiful areas, too!
And who could forget the eastern United States and Canadian Claytonia virginica complex (above), photographed here in North Carolina — this is a CHALLENGING group that has perplexed many a botanist before me… I’m treading lightly — but its pretty stunning, and amongst the first to flower in this region, so it is worth a shout out! As I can recall (its been a while since I dug into this) Claytonia virginica exhibits the longest aneuploid series on record for ANY living organism, 2n = 12 to ca. 190 — wow, that is A LOT of chromosomes!
Another representative of the eastern United States and Canada, Claytonia caroliniana complex (above) is yet another contestant on this torturous ‘cutest’ contest — these plants were photographed by Mary Jolles in New Hampshire. Thanks, Mary! This group is a bit less troublesome than C. virginica, but it is still a HIGHLY variable group of species with chromosomal ‘problems’ reported similar to those of C. virginica — the two species even hybridize in close contact… Showy, nonetheless.
Rounding it out, Claytonia lanceolata from northern California (above) — enough said!
Well, I think I’m running out of pictures, and memory on my website here, so I should probably post already… Don’t forget to vote! There is a poll at the bottom of this post. You can vote for more than one (up to 5), but you can only vote once!
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…
Above 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.
I 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…
Wait 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):
Whoa — 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: “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! 😦
(1) I passed my qualifying exams… wow! Next step, actually writing my proposed dissertation. I am thinking something along the lines of this FUN title:
Circumnavigating the Claytonia lanceolata Species Complex: Systematic Studies of Claytonia Section Claytonia (Montiaceae).
(2) If you haven’t heard already — I am involved with another crowd-sourced fundraising effort in collaboration with a colleague at Bucknell University, Dr. Ingrid Jordon-Thaden. This is the type of fundraiser where EVERY contribution counts! Even if you were to pledge $1, you would help to demonstrate just how much interest there is in research projects like this one.
Hopefully, we’ll be successful in raising funds to conduct field research this summer in Idaho, Montana, and the Yukon Territory (Canada)! Show your support today — you’re NOT CHARGED ANYTHING unless we meet our full fundraising goal!