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. 😉

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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 — problems, or solutions?

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Claytonia perfoliata species complex on the Tejon Ranch — this is only the beginning of your Montiaceae-related nightmares…

The taxonomy for the annual species of Claytonia (section Limnia) is VERY difficult to work with, mostly because diagnostic characters are not well preserved on herbarium specimens. Unfortunately, a molecular phylogeny that spells out the relationships among taxa in this group is still lacking, but I have added many samples as out-group taxa for my own dissertation studies on section Claytonia (i.e., the tuberous perennial Claytonia)… Is the cause for confusion in field identification due to phenotypic plasticity, hybridization, or both? Is it something else altogether, such as the rapid formation of localized ‘races’ due to poor dispersal capabilities? If the difficulty of accurate field identification is due to hybridization, is this a problem? For taxonomy, the morphological consequences of hybridization are an issue that must be addressed, but for the longevity of lineages, hybridization is probably not a ‘problem’ but rather a solution — it might just be the key to sustaining genetic diversity and promoting adaptation in a forever changing climate — evolutionary success! Before I get too deep into the hole I am digging, we’d better take a look at some nice plant images to really smell what I am stepping in 😉

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Claytonia parviflora subsp. viridis — Tejon Ranch. Note the linear basal and cauline leaves of this mildly succulent annual. Were you to collect, you could rest easy with ID of this plant.

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Claytonia parviflora subsp. viridis X C. rubra ?!?!– Tejon Ranch. Note the non-linear basal and cauline leaves of this moderately succulent annual, and the floral morphology of C. rubra. Yet, this plant retains the fully separate leaf pair of C. parviflora subsp. viridis.

Claytonia parviflora subsp. viridis X C. perfoliata ?!?! -- Tejon Ranch. Note the non-linear basal and cauline leaves of this mildly succulent annual.

Claytonia parviflora subsp. viridis X C. perfoliata ?!?! — Tejon Ranch. Note the spatulate basal leaves. I certainly wouldn’t want to be doing a flora of this area, as I’m sure this isn’t the only group with these kinds of ‘problems’ with hybridization 😉

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Claytonia exigua (left) and C. exigua X C. perfoliata (right) ?!?! — Tejon Ranch. Note the glaucus basal leaves of the plant on the right, which should otherwise look like the plant in the first picture of this post. Yikes!

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More fun with sympatry on Tejon Ranch!!! Left to right, Claytonia exigua, a diminutive C. perfoliata, and something that approaches C. rubra in morphology — section Limnia is tough!

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Could this possibly even be the same as any taxon we’ve seen elsewhere in this blog post? I’m really not sure, but I would lean toward calling it C. perfoliata-like. The C. perfoliata complex, including also C. rubra and C. parviflora, is probably the most difficult group in section Limnia.

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This is also Claytonia perfoliata! Its first leaves are narrow like these. Under drought conditions it can ‘bypass’ switching to the more mature spatulate leaf condition by going directly into flowering and therefore only producing the perfoliate cauline leaves.

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Contrast (or compare, in case of my suggested hybrids) the above pictures with this one of Claytonia rubra — this appears to be a good example of this species which at one point was treated as C. perfoliata but has been found to be a unique lineage and is treated as such.

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Now, compare the picture immediately above (of Claytonia rubra subsp. rubra) with this one — same or different?! Same! Well, sorta… This is ALSO C. rubra, presumably subsp. depressa, which apparently also occurs on the Tejon Ranch — how exciting!

Okay, so interestingly enough, I took all of these photos (and many, many, many more) in only one day on Tejon Ranch with Nick Jensen, a colleague at Rancho Santa Ana Botanic Garden doing a floristic study of the area. His website can be found here. This is a fascinating area because it serves (or, served) as a biogeographic traffic signal for the Sierra Nevada, the Transverse Ranges AND the San Joaquin Valley and the Mojave Desert. I suspect the evidence for hybridization exhibited by the numerous sympatric Claytonia here is mirrored in other plant groups on the Tejon Ranch where multiple taxa co-occur.

But that isn’t all, of course — the evidence of hybridization reappears in every mountain range I have explored in southern California! It seems to be a battle among divergent lineages to survive and compete for overlapping and limited niche space near the southern part of the geographic distribution of these plants, but maybe a little cooperation wouldn’t hurt for the ‘greater good’ of Claytonia if it increases genetic diversity… In the San Bernardino Mountains, you can find members of the C. perfoliata species complex in sympatry. The evidence for hybridization in this area mirrors what I observed at Tejon Ranch.

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From left to right, Claytonia parviflora, C. perfoliata and C. rubra, all members of the C. perfoliata species complex occurring together in Mill Creek Canyon, San Bernardino Mountains.

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A good example of Claytonia parviflora; note the erect, linear basal leaves that are almost terete in shape, and the cleft in the disc-like cauline leaf-pair resulting from incomplete fusion of two leaves.

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Compared with the picture immediately above, this one looks like a fusion of C. parviflora and presumably one (or both?) of the other two annual species in the area. I tend to think C. perfoliata may have played a role in the formation of this hybrid, but we can only be sure by DNA sequencing or by extensive cytological screening. Note the erect basal leaves that are no longer linear in shape but instead have rather distinct blades — this is NOT C. parviflora in the strictest sense of the name.

In the San Jacinto Mountains, you can find more of the same — members of the C. perfoliata species complex in sympatry and evidence for hybridization that may make you want to tear your hair out if you are working to identify plant specimens collected here. For instance, the pictures below from the San Jacinto Mountains are more or less taken from within ~1 contiguous square mile in Garner Valley.

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An excellent example of C. parviflora subsp. viridis is strutting its stuff roadside in the San Jacinto Mountains. Here, you have more action in the ring coming from the C. parviflora corner than from elsewhere in the C. perfoliata complex — I’m not sure the reason but think it is related to the locality I visited, a recently burned area in Garner Valley with little overstory cover.

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Compare the picture immediately above (of C. parviflora subsp. viridis) with this one of C. parviflora subsp. parviflora — similarly erect basal leaves that approach a terete-linear shape, but a contrastingly perfoliate cauline leaf-pair that is almost completely fused (as opposed to the free cauline leaves of C. parviflora subsp. viridis photographed above). The two appear to have different coloration under the same conditions here as well in both the flowers and foliage. — Garner Valley, San Jacinto Mountains.

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Now, compare the picture immediately above (of C. parviflora subsp. parviflora) with this one of some sort of C. parviflora hybrid — similarly erect basal leaves but these appear to be forming distinct blade and petiolar regions unlike the other two photos above from the same general location… tempting to call them the same, but I imagine this fishiness has both rhyme and reason to it. — Garner Valley, San Jacinto Mountains.

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This plant is OBVIOUSLY different from the rest, but what do you call it?! Under some Jeffrey pines that survived the fire, these plants approach the morphology of C. perfoliata subsp. mexicana but the smaller leaves in the center of the rosette are suggestive of certain races of C. rubra. — Garner Valley, San Jacinto Mountains.

What a wild and wacky world we live in! Claytonia will keep me busy (and happy) for many years to come…

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…

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.