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. ūüėČ

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!!!

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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! ūüėČ

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! ūüė¶

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 √ľber-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!

D√©j√† 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‚Äôs Claytonia¬†time‚Ķ”)

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 Pi√Īon Hills, highway 138.

Spectacular wildflower bloom in the Mojave Desert near Pi√Īon Hills, highway 138.

 

 

¬°Holy frijoles! You are just too cute, and too close to home…

OK, so I might just have a new favorite Claytonia after my most recent trip into the desert in northwestern Nevada. Let me just tell you something — in the Desert Southwest, spring has sprung! Just like my collaborator at Eastern Washington University (Dr. Robin O’Quinn), I think I am falling in love with the tuberous perennial ‘Great Basin Spring Beauty’ (C. umbellata) —¬†photographed here on March 21st, 2014.

IMG_5178IMG_5195IMG_5134As Stevie Wonder might say, isn’t she lovely?!

It gets better… Also photographed on March 21, 2014, below are some pictures of plants from another population of C. umbellata that occurs on a different substrate in another mountain range nearby in Nevada.

IMG_5221IMG_5227IMG_5230There’s something funny going on here, and I’m going to get to the bottom of it…

Question for readers out there: Who thinks the above plants look similar to the below plants from southern California? Just wonderin’ about evolutionary relationships ūüėČ

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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 ūüėČ

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

Who is pollinating Claytonia in California?! pt. II

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

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

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

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

Yep, it’s Claytonia time…

That’s right! Below are my first pictures of the 2014 Field Season of¬†Claytonia “peirsonii”,¬†collected just¬†west of Devil’s Punchbowl County Park (Feb 15th, 2014). This population was documented for the first time today, now the westernmost occurrence of C. “peirsonii”¬†that I know.

2014-02-15 14.12.46How many Claytonia do you see in the image below? I’ve counted about 100 individuals… wowza!

2014-02-15 14.12.36

Off to see the southern Sierra Nevada¬†C. “yorkii”¬†next weekend! Jawbone, here we come ūüėČ

Claytonia Rocks!

So these plants are considered to be ‘geophytes’ because they have underground storage organs, but I find this so-called classification to be much more meaningful than that for the tuberous perennial Claytonia. In order to better understand this, let me break it down a little bit.

Phyte – Greek, generally referring to a plant.

Geo – Greek, generally referring to anything of or relating to the Earth.

2013-05-30 11.44.33Pictured here is the habitat of the typical variety for the Claytonia lanceolata complex, C. lanceolata¬†sensu stricto. At least in northern California, this taxon is known only from soils derived from granitic rocks. The suggestion has been that as a generalist, C. lanceolata (as currently circumscribed) occurs in a variety of habitat types across its range and thus is highly polymorphic morphologically. Well, there are some interesting substrates in southern California within very close proximity where one can find members of C. “peirsonii” complex occurring in nearly equivalent niche space across different substrates… Wait a second, am I studying geology or botany?!

2013-04-20 09.57.32Because bragging about how awesome the San Bernardino Mountains are is my favorite thing in the world to do, I’d like to take this opportunity to point to some interesting Claytonia there that occur on sedimentary carbonate rocks on the north slope of Bertha Ridge. This is the ridge that separates Big Bear Valley to the south from Holcomb Valley to the north. This is where I first encountered plants that match the description of Claytonia lanceolata var. peirsonii¬†Munz&Johnston, or any plant treated as Claytonia lanceolata in the Jepson Manual or Flora of North America, while I was working as a Rare Plant Technician for the USDA Forest Service on the San Bernardino National Forest. More pictures of these plants can be found on CalPhotos.

IMG_9508These plants, as mentioned previously, are ridiculously cute (see related articles below and cast your vote!). More importantly for this story, these plants occur on dolomite, which is a particular type of carbonate rock containing the mineral dolomite.

2013-04-20 12.17.49And boy-howdy are these rocky hills STEEP! Unfortunately, this is an aspect of the habitat that many of the tuberous perennial species of Claytonia appear to have conserved across lineages, making field work incredibly physically demanding when sites are remote. But what is my point, exactly?

2013-04-20 14.20.56Pictured here, approximately 7 air-km to the north, there exists another population of tuberous perennial Claytonia that is primarily associated with the same species (Pinus flexilis, P. monophylla and Juniperus) but otherwise occurs on a different kind of carbonate rock than the population at Bertha Ridge. This becomes significant when you look at the morphology of the plants here, which is inconsistent with the plants at the Bertha Ridge population just 7 air-km away.

2013-04-20 14.21.55Not only do the plants at this more northern locality on gray dolomite have pink nectary guides, in contrast to the yellow spots at the base of the petals on plants from Bertha Ridge, but cauline leaf shape appears also to be strikingly different between these two populations. This interesting case of inconsistency in overall morphology, coupled with signs of genetic divergence among these populations (see my most recent phylogeny here), has gotten me very excited about patterns of evolution in this group that is already well known for complex histories of hybridization and polyploidy

ClaytoniaRocksAlthough it is all C. lanceolata for now, soon enough we’ll make some sense out of it!