Claytonia rubra, what is that? pt. II

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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Variegation on foliage characteristic of some lineages of Claytonia rubra.

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

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

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

And now something different…

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

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

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

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

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

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

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

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

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

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

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

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

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

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Beginning of the end of Silene verecunda sensu lato

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

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

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!

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Larry Crawford and Diana Jolles enjoying the habitat on Meiss Ridge

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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

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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

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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.

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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

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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

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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!

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

Can I borrow $20? I’ll pay you back with publications…

Hey all — two quick ‘news’ items:

(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.

https://experiment.com/projects/alpine-plant-biodiversity-why-is-it-so-high

 

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!

Floral morphology in Claytonia: How much variation is ‘normal’ for a species?

Ok, so you may remember this recent post, where I asked for interested people to send me pictures of as many flowers as they could photograph…

The premise is simple: How much variation is considered the ‘normal’ or ‘typical’ amount of variation for a species? How much is typical in terms of petal shape, size, and color for individuals in a given population of tuberous perennial 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 within populations. It might bend your brain the next time you kneel down and take a look around at the tuberous perennial Claytonia blooming in your area (right now!!!)… Better yet, take a picture of 50-100 different flowers from directly facing the flowers (trying to center the gynoecium in the flower as best as possible). Compare all your photos, or send them my way. We’ll get to the bottom of this!

Enter stage left as ‘proof of concept’, a member of the C. “peirsonii” complex from the southern Sierra Nevada — all of the below pictures are from different flowering individuals within a single population, taken on the same day within about 45 minutesIMG_5421 IMG_5418  IMG_5404Note the visitors above: an ichneumonoid wasp (likely a parasitoid braconid) on the left, and a soft-winged flower beetle (Melyridae) on the right. We saw these same beetles last year, and I’ve also mentioned them in the ‘who’s pollinating Claytonia?‘ series — special thanks to Dr. Emile Fiesler for help with the identifications!IMG_5432IMG_5398IMG_5382IMG_5387IMG_5324IMG_5331IMG_5396The variation is crazy, right?!

IMG_5340I really need to get a new ‘cutest Claytonia‘ contest going soon…

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!

Name a new species by making a $3000 tax-deductible donation!

Hey, everyone!

First, I want to thank several of you out there for the noble contributions you’ve already made. I encourage you to keep the momentum up and share with a friend so we can Fund the Sciences! It is all C. lanceolata at this point, but we can make some changes…

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I am so passionate about conserving rare plants that I wanted to announce to you that I have made the offer for donors to name a new species by making a $3000 tax-deductible donation to this project! I will be describing as new to the scientific community a few rare species of Claytonia (and new species in other genera in North America) as part of my dissertation research. Not only will you receive the super sweet reward of a color reprint for tax-deductible donations, but donate $3000 or more to my Research Fund at RSABG and I will name a new plant species in your honor (or take suggestions)l. With all of your help we can get this research project off the ground!

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CLIMATE CHANGE: Local student seeks funds for plant research

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Hey Everyone! My project on Microryza.com is being featured by the Press Enterprise. Check out this short blog from Janet Zimmerman:

http://blog.pe.com/environment/2013/09/16/climate-change-local-student-seeks-funds-for-plant-research/