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.

IMG_2004

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.

Untitled

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!

IMG_2270

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.

IMG_5856

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?

IMG_5868

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!

IMG_7912

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.

IMG_7434

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.

2013-04-28 11.41.08

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.

2013-04-28 11.34.01

[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!!!

Hybridization — are generalist pollinators the key to success?

I’ve been on a bit of a rant lately concerning hybridization in plant species complexes, which recently resulted in my last post suggesting hybridization could be a potential ‘solution’ to the problem of an ever-changing climate. I want to dig a little deeper now and ask the questions — can generalist pollinators have anything to do with the maintenance of a generalized floral form in Claytonia? If so, can this indirectly act as a selective force for increased hybridization among distantly (and closely) related species? If you have seen the ‘face shots’ from a recent ‘Spring Beauty Pageant‘ than you know what I’m talking about. Maybe you’ve even suggested to a friend, because Claytonia grow like weeds where you are from, that you believe there to be little variation in floral form among species in the genus. I’d agree with you, to a certain extent… but then again, maybe there is so much intra-population variation in floral morphology (coloration, petal shape) that we are blind to see the true pattern of differentiation in floral form among closely related taxa. I’ve touched upon the subject of within-population variation of the flowers of one member of the C. “peirsonii” complex from the southern Sierra Nevada, but now I want to expand on this subject with photographs taken within a single population of C. lanceolata sensu lato on the Great Continental Divide in Montana. As you can easily tell from the first picture below, in which all of the white spots are Claytonia flowers, it is not difficult to conduct this sort of ‘experiment’ yourself. In other words, it is easy to get a lot of face-on photographs of Claytonia flowers because they occur in extremely dense populations. This spring, try to see how much your favorite Claytonia varies in its floral form — you may be surprised at what you find!IMG_8497IMG_8570IMG_8449IMG_8582 IMG_8453IMG_8458IMG_8477IMG_8478IMG_8531IMG_8547IMG_8516     Keep in mind that all of the above pictures are of plants that occur in a single population of C. lanceolata sensu lato on the Great Continental Divide in Montana. As evidenced by the last photograph, even merosity can go haywire from time to time. If I showed you just that last picture, you might even confuse the plants for Lewisia. That is a lot of variation! Are these hybrid plants of C. lanceolata (which has retuse petal apices) crossed with another species in the area with entire petals? Good question! Let me get back to you on that one…

All I know is that multiple Claytonia species can occur in sympatry, and intermediate forms can be found in those areas. This is especially true of the annual species of Claytonia. I haven’t observed putative hybrids among the annuals (i.e., the miner’s lettuces) and tuberous perennials (i.e., C. lanceolata species complex), although I have observed them sharing pollinators in southern California. Pictured below are soft-wing flower beetles (Melyridae) visiting both C. rubra (section Limnia, first picture below) and C. “peirsonii (section Claytonia, second picture below) at the same location on the same day in the southern Sierra Nevada, California. You might remember from one of my very first posts: these pollen eaters are quite effective pollinators.

IMG_5457 IMG_5402Too many questions for a single dissertation to address, but I’ll see what I can do! 😉

THANKS (Experimental Success!)… and a report on my Northern California and Southern Oregon trip to sympatric Claytonia sites

IMG_7744Above, two members of the genus Claytonia from different Sections — C. “serpenticola” (an undescribed species in Section Claytonia) on the left, and C. rubra (Section Limnia) on the right. The two occur in close sympatry in the Lassic Mountain Wilderness, practically high-fiving their cauline leaves. This was a highly productive trip to the north, but before I get to that…

 

I want to say THANKS to all who supported my recent fundraising endeavor with Dr. Ingrid Jordon-Thaden on Experiment.com — we are going to be able to do some much needed fieldwork and conduct fundamental biodiversity research with the money you have all helped us to raise! Thanks so very much, and stay tuned for updates (Lab Notes and Blog Posts) on this ‘open access’ alpine research.

 

OK, back to talking about my recent trip to Oregon and California — I was on the hunt to find locations where multiple Claytonia occur in close sympatry, particularly areas where species may be hybridizing. I am interested in knowing just how well species can maintain their identities in these situations…

IMG_6819I started the trip by meeting up with Larry Crawford for some botanizing in the Sierra Nevada — we found Claytonia sessilifolia (Torrey) Henshaw(pictured above) flowering profusely near Carson Pass in an area that Larry had previously scouted. This plant was featured in the recent Spring Beauty Pageant hosted on my website — it isn’t too late for you to cast a vote for your favorite(s)! In this population it seems like there is only one species present (C. sessilifolia), but this taxon occurs in an interesting habitat (pictured below, with Diana Jolles and Larry Crawford). It also occurs next to some other fantastic spring ephemerals, such as Dicentra uniflora (pictured below) — the hike was well worth it!

Image

Larry Crawford and Diana Jolles enjoying the habitat on Meiss Ridge

IMG_6704

Following some rest and recovery at Larry’s house (thanks, Larry and Suzanne!), my colleagues and I continued north to Lassen Volcanic National Park, catching a wonderful view of Mt. Lassen (pictured below) — this was my first time taking in such a beautiful sight. I also have never (knowingly) been this close to a stratovolcano. I can’t say that I was nervous that the mountain might erupt, but it actually hasn’t been long since something like that happened (click here to read more about the geologically recent eruption of Lassen Peak). Unfortunately we didn’t find any Claytonia in this area, but we might have also been a tad on the early side for the area we looked — several feet of snow in some drifts. It has been a bit of a whacky weather year…

IMG_6885Having been thwarted by a lack of Claytonia at Lassen Volcanic National Park and lower elevation sites in Lassen National Forest, I decided to book it northward to Crater Lake National Park in an attempt to catch up with a rapidly fading spring. Many of the areas in California where Claytonia have previously been reported as abundant are looking quite parched — the Spring Beauties are apparently less than pleased about it. At Crater Lake, though, we found C. sessilifolia plants blooming where else but in the parking lot (pictured below)!Snagging a few quick pictures of the cinder cone “Wizard Island” (pictured below) in the middle of Crater Lake was just a bonus of doing fieldwork in the area. 😉IMG_6959

IMG_7012

We found some more plants a few miles north of Crater Lake NP on the Rogue River-Siskiyou National Forest, and I also happened to snap a quick picture of the BEAUTIFUL Mt. Thielsen (pictured below) on my way out of Crater Lake National Park — this is another place I’ve got my eye on visiting. I’ve heard the Claytonia on Mt. Thielsen are quite interesting… IMG_7019

Next was Abbott Butte (west of Crater Lake National Park) — I ABSOLUTELY had to visit that area this year, as it is the type locality for Claytonia obovata Rydberg. Ray Davis chose to sink this taxon into synonymy in the 1960’s — it has been treated as C. lanceolata since that time. Ray mentioned in his 1966 North American Perennial Species of Claytonia that he collected plants answering the original descriptions of C. obovata, C. multicaulis, and C. chrysantha, as well as C. lanceolata at Abbott Butte. Although he didn’t recognize C. obovata, tubers he collected of that taxon from around the type locality that he grew in a ‘common garden’ retained their rounded leaf shape. What did I find at Abbott Butte? Claytonia lanceolata (pictured below, lacking bifid petals), but not any C. obovata — did it not come up this year? I’ve certainly observed this happening before with C. “peirsonii” in southern California… at least Julian (pictured below) found some delicous black morels (Morchella elata-M. angusticeps-M. conica complex). = YUMMY.IMG_7177

Julian enjoys yummy morels...

Julian enjoys yummy morels…

What do you do when you can’t find a species at its type locality? You write a paper about it  — look for Stoughton et al. (in prep.) coming to a Madroño near you. Fortunately, I have seen C. obovata previously with Heath Bartosh at Hull Mountain — this area is also mentioned in the protologue for C. obovata Rydberg. I did find C. obovata with Diana Jolles and Julian Roberts later during our trip near Cory Peak and Mount Eddy on the Shasta-Trinity National Forest in the Klamath Region of California, but I want to tell you first about the next stop on our road trip at the Siskiyou Mountains of southern Oregon — Observation Peak (pictured below) is an area where C. “serpenticola” (pictured below, and at the beginning of this post) is known to occur.IMG_7263

IMG_7259

After seeing C. “serpenticola” (pictured above) more or less ‘alone’ in the Siskiyou Mountains of southern Oregon on serpentinite and peridotite, we next saw C. obovata (pictured below) similarly secluded in the Scott Mountains of California near Cory Peak — I now had my search images for a hike that was a major part of the trip. The two species occur in sympatry at Deadfall Lakes just below Mount Eddy, which is where we were headed next. I REALLY want to measure levels of hybridization in this population on Mount Eddy — obvious intermediates exist in the area in seemingly marginal habitats.

IMG_7414

What do hybrids between C. obovata and C. “serpenticola” look like? They probably look something like the pictures below — the two species occur in very close proximity in this area…IMG_7574

IMG_7685

SUCH A COOL SPOT — I will revisit this area next year when there is hopefully a bit better snow pack (which generally = many happy Claytonia). Who is coming with me?! I’m climbing up to the top again for sure. Claytonia “serpenticola” (pictured below) grows very near to the summit, and the view of Mount Shasta (pictured below) from atop Mount Eddy is SPECTACULAR.IMG_7698

IMG_7721

After poking around Mount Eddy with Dana York and Julie Nelson, and a much needed night of rest (Thanks, Dana and Eva!), Diana and Julian headed with me south to the Lassic Mountain Wilderness (Six Rivers National Forest) and Anthony Peak (Mendocino National Forest) to see C. “serpenticola” (pictured below) and C. obovata (pictured below) at additional areas where they occur in allopatry — plants were pretty crispy in northern California, but they were still identifiable!

IMG_7743IMG_7795

I am VERY excited to see how things shake out in the molecular phylogeny I am developing for this group… Stay Tuned!!!

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.

 

 

What are those stamens doing?

1

(Above) How weird is that?! The filaments curl back onto the petals and the anthers shed pollen at the same time that the stigma is receptive in some species of tuberous perennial Claytonia!

4

(Above) In others, the filaments begin to curl back when the anthers start shedding pollen, with the stigma become receptive shortly thereafter. Excellent!

(Below) In this species, the filaments curl back and the stigma becomes receptive after most of the pollen has seemingly been released. Gnarlatron!

2

(Below) And still in others it seems like the filaments don’t reflex much toward the petals at all. Whoa!

5But what does it all mean?! That is a good question… I’ll have to get back to you on that one.

😉

Which is the cutest?

You’re really going to ask me which Claytonia I think is the cutest?! I find the entire genus to be cute… It’s a really tough choice!

Image

This one from the Southern Sierra is a real crowd pleaser, being found growing in talus of volcanic rocks (Tuff). It appears to be in the C. parviflora complex (C. perfoliata sensu lato), putatively a local endemic based upon evidence from morphology and patterns in tuberous perennial Claytonia in the area. Although not my all-time favorite Claytonia, this cute little annual always makes for an entertaining photo shoot.

Image

This member of the C. virginica complex is also quite stunning, being found on nearly every flood plain of the major rivers draining toward the U.S. Eastern seaboard (this one photographed in North Carolina). The reason it isn’t my favorite? Well, poison ivy (Toxicodendron radicans) also grows on nearly every flood plain draining toward the Eastern seaboard. What is the significance of this distribution you ask? Well, when I go collecting Claytonia and nearly everything else is un-identifiable due to the extremely early flowering season of Claytonia, I end up getting exposed to Toxicodendron a LOT! I now have a systemic reaction whenever I get exposed to Toxicodendron or any other member of the Anacardiaceae… so also when I eat mangos or cashews. I (used to) love mangos!!! Dang…

IMG_2674OK, now we are talking cute here… This thing took my breath away the first time I saw it in the North Coast Ranges of northern California, where it can be found on Franciscan Graywacke (a type of sedimentary rock). Formerly treated as C. obovata Rydb., this taxon is currently treated as C. lanceolata despite its morphological dissimilarity with the latter taxon. And You’re saying this isn’t the cutest?! Why not? Well, I think this little guy was ruined for me when some hooligans decided to start shooting various guns in the air very close to where I was. It was a bit scary, and I think it spoiled the good time I was having at the time ripping these beauties out of the ground (for the sake of science, of course).

Image

Now this one absolutely has to be the cutest, right? Wrong! It is cute, oh yes, but not the cutest. This beauty is just another narrow endemic that is not currently recognized (but hopefully will be soon!)… Levels of cuteness approach the maximum score, but this little beauty (only a few centimeters tall) just isn’t the cream of the crop. It grows in a crummy habitat (steep, loose talus of volcanic rocks), for one, and it also lives behind a locked gate in an area that is proposed for wind energy development. Until I find populations of this unique taxon outside of the Jawbone Canyon in the southern Sierra Nevada, it sits in the same category as the C. virginica complex because it comes with too much baggage… NEXT!

Image

Oh yeah! If you’re cute and you know it, be super small… This is the first member of the C. lanceolata complex I ever found (in the San Bernardino Mountains of southern California on white dolomite). You might say it is the reason I got into this mess in the first place, and thus it holds a special place in my heart. It is amongst the smallest members of what I am calling the C. “peirsonii” complex, a southern California species complex nested within the C. lanceolata complex (which truly is summarized as the ENTIRE clade of tuberous perennial Claytonia, e.g. Claytonia sect. Claytonia). It is very VERY cute, but still not the absolute cutest. Tough competition, right?! This is very close to my all-time favorite Claytonia, but there is one species that takes the cake…

Image

Wow! What a show-stopper! Claytonia saxosa is indeed the cutest Claytonia I have seen to date, but there may just be some species out there that are even cuter than this one, including yellow and pink-flowered tuberous perennial Claytonia that I hope to see in 2014. Until then, this is the cream of the crop, king of the hill, or whatever you want to call it. It is just dang CUTE! Also growing on Franciscan Graywacke in northern California (in close sympatry with C. obovata), this annual species is said to occur on serpentine rocks as well. I’m interested to see if the serpentine plants are really the same taxon, given what patterns have been observed in the tuberous perennials, but until I can get the material I must consider it the same species across its range. I guess I’ll just have to get out there and see it elsewhere… I recommend you get out there and see it too! Anthony Peak (Mendocino National Forest) in the North Coast Ranges of California has a healthy population that is ripe and ready for photographs in the month of May.

In order to get to the bottom of this, I encourage you to cast your vote! Cast your vote for more than one, whatever, but you can only vote once! I want to know if you agree, that C. saxosa is indeed the cutest of all Claytonia-land…

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.