In a recent newspaper entitled "Paleoelevation inwards the Jianchuan Basin of the southeastern Tibetan Plateau based on stable isotope together with pollen grain analyses", Wu, Zhang et al. let out that "... pollen information from the Shuanghe Fm indicates a paleoelevation of 1.3–2.6 km to a higher house bounding main score (asl), which would most in all probability direct hold been associated alongside a vegetation comprehend consisting of tropical [most taxa inwards the listing are nonetheless extra-tropical]-subtropical, deciduous, coniferous, broadleaf forests. The reconstructed hateful annual air temperature (MAAT) had a value of 16.8–21.7 °C, warmer than today's MAAT ( six °C)." Influenza A virus subtype H5N1 resultant inwards some fit alongside their stable isotope of the Jiuziyan Fm. "... suggest[ing] that the surface of the Jianchuan Basin was at a paleoelevation betwixt 0.5+0.8–0.5 km asl (i.e. sea-level) together with 2.5 ± 0.7 km asl (δ18Omw: −8.9 ± 1.3‰, 2σ)"
You tin never last incorrect alongside depression to mid-altitude subtropical inwards the Eocene, tin you?
Only that the fossil flora is composed mainly of conifers together with other meso- to microthermic elements (as stated past times the authors on p. 9), which would last really atypical for a tropical-subtropical setting, but typical for a temperate-boreal. But as nosotros showed (still ignored but also non rebutted past times Palaeo^3 authors together with editors), all roads Pb to Yunnan's subtropical climate when using Chinese palaeofloras (Grimm et al. 2016).
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| Distribution of Pinus (source: Wikimedia Commons). Clearly a genus indicative for mild temperate to tropical weather condition (Tann > seven according Wu et al., tabular array 1) |
The higher nosotros become up, the colder it gets
As most of us know (but non users of the Coexistence Approach or editors/peers of PPP, as nosotros volition see). With dropping temperatures, the likelihood for snowfall increases, so you lot encounter unlike forests/ vegetation belts when roaming mountaineous regions. When you lot become to southern Hokkaido alongside its Dfb climate (snow but proper summers), you'll let out beech forests (Fagus crenata, the sis species of the western Eurasian beech, F. sylvatica), at sea-level; same inwards southern Canada (F. grandifolia, a distant relative of both) or southern Scandinavia. Going to Honshu or fifty-fifty Taiwan, 1 tin readily notice that the beech wood belt goes upward the mountain: a pollex dominion is that it starts at most 500 m inwards northern Honshu (more or less plumbing equipment the edge betwixt Cfa and Cfb climate), increasing to 1000/1500 m inwards the subtropical, southern Honshu, Ryukyu or near-tropical (at sea-level) Taiwan.
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| Altitudinal together with latitudinal attain of beech inwards Nihon (see Maycock 1994, compiled for Grimm & Denk 2012, ES6) |
[Some to a greater extent than bioclimatic trivia for people doing quantitative climate reconstruction using flora taxon lists: Being 1 of the most-constrained (ecology together with climate-wise), non-relict tree genera, the genus Fagus has a Cfa, Cfb, Dfb 'Köppen signature' (Denk et al. 2013), pregnant it tin hap inwards these 3 climates, altitudinal or latitudinal. And this seems to direct hold been the instance since its really origin. If you lot direct hold beeches, you lot tin last pretty certain the local climate has been a warm temperate (not to last confused alongside temperate), mild snowfall climate alongside ample atmospheric precipitation throughout the twelvemonth (beeches, inwards contrast to many other 'nemoral' elements, make non hap inwards Cwa or Cwb climates; non fifty-fifty inwards China). Nevertheless, the hottest beeches tin last found inwards northern Florida just about overnice summertime retreats (and beaches) at Tann of 19–20 °C (Maycock 1994, soundless subtropical). In Georgia (the country, non the US state), the beech goes downward into the subtropical zone together with to sea-level (Tann xviii °C), as good as some Chinese species inwards China's Cfa climate zone. But it doesn't tolerate Dfc climates, likewise brusk summers, inwards contrast to some species of its fifty-fifty to a greater extent than common, distant relative, the oaks; the coldest North American beech stands direct hold nonetheless a Tann of 4 °C. This is the argue the Coexistence Approach does demand relicts alongside unrepresentative, much likewise narrow modern climate intervals. Otherwise, the resultant would e'er last something betwixt 5–15 °C MA(A)T (temperate forests) together with 15°–25 °C (tropical), together with non the much wanted 15.6°–15.7° C ... For those interested inwards beech-climate correlation, information has been compiled inwards File ES6 to Grimm & Denk 2012.]
In many mount areas the decrease inwards temperature the higher nosotros become upward approaches the so-called moist-adiabatic lapse rate: On average, nosotros lose 1 °C for every 200 m nosotros become up. Which is also the instance for e.g. the vast Himalayan mount attain (especially its southern together with eastern slopes) that was the focus of Wu et al.'s study. As seen inwards the next graph.
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| A = Qamdo, Tibet (border betwixt temperate Cwb together with Dwb climates; WMO station nr. 5613700); B = Lancang, Yunnan (WMO 5695400; subtropical Cwa) together with C = Baingoin, Tibet (arctic-alpine ET; WMO 5527900); complaint the proficient fit alongside the moist-adiabatic lapse charge per unit of measurement of 1 °C/200 m using Qamdo as reference dot (station information from Lieth et al. 1999). |
Reconstructing palaeo-altitude together with -temperature using the Coexistence Approach
How tin overlap inwards the modern altitudinal attain (in Tibet) of botanical composite taxa (genera, families), most of which are found across modern Red People's Republic of China together with are roofing anything betwixt near-tropical to boreal climates (according to the tolerances reported past times Wu et al.), inform most a palaeo-altitude of a "tropical-subtropical mixed deciduous, coniferous broadleaf wood earlier 36 Ma" and, independently, the local temperature?
Wu et al. exhibit how this works inwards their figure 4; the tolerance ranges per taxon are provided inwards their tabular array 1 — a documentation unremarkably missing inwards many papers using the method, together with nearly all papers published past times the NECLIME group, the German-centred (and -funded) Coexistence Approach syndicate.
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| Copied from Wu et al. (2018, p. 6). |
The ancient cedars also define the upper boundary of the (palaeo-)temperature interval ("MAAT" ≡ Tann < 21.7 °C), whereas Betula – the birch, 1 of the most frost- together with cold-resistant broadleaved genera of the Northern Hemisphere – and Tsuga – a temperate conifer found today only inwards eastern North America together with East Asia; a classic disjunct distribution for a relict that has seen amend times – define the lower boundary for a (warm-)subtropical Tann (> 16.8 °C).
For the height "coexistence interval", many NLRs were plainly removed/ignored: Podocarpus, Lygodium, Meliaceae, Carya, Castanea (the chestnut) as likewise low, together with Abies (firs), Picea (spruces), Chenopodiaceae, Ephedra (an strange flora genus, alongside a near-cosmopolitan distribution, but plainly non inwards Tibet...), Artemesia (ditto, but mutual together with non odd) as likewise high. The authors make non furnish whatever clues or guidelines for their elimination strategy.
It may direct hold been informed past times the isotope information results.
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| Wu et al.'s Fig. 9. May also isotope analysis residuum inwards peace. Note the blueish together with ruddy doodle lines. |
First alarm-flag missed past times peers (?) together with unwitting editors: NLR temperature tolerances overlap but non those of the altitudes
Following the (non-sensical) Coexistence Approach protocol for eliminating outliers, in that place would direct hold been 2 choice "statistically supported" (using CA-language, non to last confused alongside actual statistics) palaeo-altitude coexistence intervals: 1300–1600 m together with 2000–2100 m, both alongside a maximum of twenty out of 25 coexisting. Influenza A virus subtype H5N1 3rd every bit "supported" interval would last 1700–1700 m — no mistyper here, just next the instance of actual CA-papers purporting occasionally MA(A)T CA-intervals such as 15.6–15.6 °C (see Grimm & Denk 2012, ES 2, included inwards the freely accessible online supporting archive).
But 'outliers' cannot last only removed. If the ranges of all NLRs don't overlap ("100% coexistence") for a fossil assemblage, together with so in that place must direct hold been a shift inwards 1 or many lineages.
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| Fig. 1 Niche shift over time, 1 of the reasons for non having 100% overlap when using NLR tolerances for a fossil flora: the "coexistence interval" of the fossils differs from that of the modern counterparts (the "nearest-living relatives"); mutual (climate) attain approaches such as the Coexistence Approach assume no shift. Other reasons for missing overlap include mixing across geographic or altitudinal gradients together with unrepresentative tolerances (both probable applies to Wu et al.'s reconstruction). |
Here the actuo-palaeontological principle, the really ground for the Coexistence Approach, means:
- The fossil members of the lineages represented past times their NLRs had the same temperature niche than today, which is doubtful for non a few members.
- The fossil members of the lineages occurred at the same height inwards the focal expanse than today, which is effectively untestable.
And the altitude interval must prevarication inside the temperature tolerance (just using mutual sense, although nosotros were in 1 lawsuit informed past times an "anonymous" (likely Torsten Utescher, the High Wizard of NECLIME, encounter below), well-hung (it took the peer to a greater extent than than one-half a twelvemonth to complete) review of our 2012 newspaper [PDF including our responses] that specially inwards the mountains of China, plants hap today outside their natural niche, hence, our critical comparisons were unfair).
Thanks to confidential peer (or pal) review, such logical together with practical flaws direct hold never been an number (a recent example) for the NECLIME syndicate together with papers reviewed past times a fellow member of the syndicate.
The information the Coexistence Approach works with
The Coexistence Approach is an unweighted mutual (climate) attain technique that just uses min- together with max-tolerances. This is naturally really unsmooth together with has many problems fifty-fifty for reconstructing obvious shifts inwards the most recent past times (Thompson et al. 2012), together with fifty-fifty to a greater extent than when beingness combined alongside the actua-palaeontological regulation (Grimm & Potts 2016). Such as pseudo-coexistence:
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| Pseudo-coexistence, the basics — although the (bivariate) climate niches ('envelopes') of the 2 species don't overlap, hence, you lot won't let out them today growing together, they soundless make to finely resolved (narrow) CA intervals. |
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| Pseudo-coexistence when mixing floras (e.g. pollen from unlike altitudes). There are 2 to a greater extent than or less broad existent mutual climate ranges shared past times each community, together with the completely artificial CA interval (something nosotros Germans call: kleinster gemeinsamer Nenner). |
To empathise how minute together with max of frequently non independent (again encounter Thompson et al. 2012) parameters relate (or don't), x-y plots similar the ones to a higher house establishing/showing the bivariate parameter (climate) infinite (see e.g. Thompson et al. 1999–2012) tin last really enlightening. In Wu et al.'s instance it should last really clean, since height together with temperate are strictly correlated expressed past times the moist-adiabatic lapse rate.
Now, let's plot the existent envelopes defined past times the min-max tolerance ranges for those NLRs where Wu et al. furnish both tolerances. First the ones indicative for an height (in the focus region) lower than 2500 m...
... together with the ones found only at altitudes > grand m:
In the instance of the low-landers, their elevation tolerance fits their temperature tolerance: the climate envelopes, boxes, are higher than wide. But for those preferring higher altitudes, i.e. those plainly sensitive to likewise hotness (subtropical climate), one, the CA-interval determining Cedrus, is does hap at altitudes inwards the Himalayas exterior their temperature tolerance (> 2000 m), an instance for High Wizard Utescher's observation that some trees hap today likewise high upward inwards the Chinese mountains. Ephedra is also climbing likewise high up; according to its temperate tolerance it should non last able to become beyond 4200 m. It also misses out quite some opportunities at lower elevations since its temperature tolerance would allow growing at sea-level. The argue for this may only last the observed species / species group, which doesn't demeanour similar others inwards its genus. Including those that may direct hold lived inwards the expanse 35 myrs ago.
There are naturally to a greater extent than examples where Wu et al.'s tolerances are already at odds alongside each other (not to advert reality). Here's the total plot of their NLRs (which – past times the way – should last obligatory to exhibit inwards whatever study aiming to make similar things).
Quite a mess. As usual, why you lot never let out bivariate plots inwards Coexistence Approach papers (but banking corporation check out the bioclimate atlases for North America past times Thompson et al. 1999–2012).
Essentially nosotros tin describe 4 principal groups "informing" the palaeo-parameter reconstruction:
- low- to mid-altitude NLRs (green rectangles, encounter above) alongside no or differing preference regarding temperature;
- temperature-restricted NLRs (blue rectangles, encounter below) alongside no particular preference regarding height — occurring at altitudes exterior their used temperature tolerances (most-extreme: Betula, birch, used alongside a v °C temperature tolerance but a 4000 m attain ≅ twenty °C!), together with including all CA-informative NLRs, and
- mid- to high-altitude NLRs (red rectangles, encounter above) alongside completely uninformative temperature ranges (from Tann < 0, polar-alpine climates to Tann > 25, tropical, lowland climates, e.g. Abies, spruces).
- The residuum are NLRs (yellow rectangles, below) that tin hap inwards whatever climate (according to the used temperature interval) together with whatever summit (sea-level to high-montane).
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| Not making feel at all — the blueish group, temperature-sensitive, but non inwards Tibet it seems. |
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| The filling of all CA studies — largely uninformative NLRs. Note that Pteris, according to e.g. Flora of Red People's Republic of China a tropical-subtropical species-rich fern genus, is recorded alongside pretty much the same tolerances than Polypodium, a fern alongside 10 species found "throughout the N[orth] temperate zone" (again: Flora of China, maybe worth a read for CA-applicants working on Chinese palaeofloras) |
Standard when using the Coexistence Approach: ignoring the actuo-palaeontological principle
When the full general province of affairs was just the same as today together with the actuo-palaeontological regulation applies (largely), the temperature interval would straight inform the altitude, or vice versa.
Wu et al. debate that inwards the Eocene the plants grew inwards a climate alongside a Tann of c. six °C to a greater extent than (see their Conclusion) than today inwards the same expanse "... the introduce climate, which has a MAAT of 13.5 ± 1.4 °C (http://data.cma.cn/site/index.html [try the link]), is much cooler than that of the Eocene." together with complaint the fit alongside the corresponding global situation.
They farther advise "Based on the climatic preferences of the NLRs ... the existence of a warm, subtropical climate, alongside a MAAT of 16.8–21.7 °C, together with a hateful value of 19.3 ± 2.5 °C." Not Tibet anymore, but modern-day Yunnan (see Grimm et al. 2016), at most 1000–1500 m (a Cwa climate alongside really rich mixed evergreen-deciduous subtropical forests together with some accessory conifers).
With "mean value" Wu et al. refer to the "center value", which has, so far, no theoretical or practical ground together with is defined past times the (arithmetic) hateful of the upper together with lower boundary of the inferred coexistence interval. The mutually shared climate range, Coexistence Approach-based or not, is literally a range: 16.8–21.7 °C would dot that the temperature was 3° to 8 °C higher than today, together with non 5.5 °C ± 2.5 °C higher.
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| Close-up on Yunnan together with Tibet, showing climate stations included inwards Lieth et al. (1999). The Cwa station Lincang, Lancang, Simao together with Mengzi direct hold Tann inside Wu et al.'s coexistence interval, Lijing matches their modern-day reference (Tann = 13.4 °C). Indicated are also Qamdo (Cwc) together with Baingoin (ET) used before. |
But inwards a historical context, in that place is a slight complication: what if the temperature was raised inwards general, i.e. globally? Then it may also had been raised locally. Influenza A virus subtype H5N1 flora affected past times this temperature modify volition evade to or tin expand into higher altitudes (and latitudes). If temperatures went down, macrothermic together with mesothermic elements would direct hold to retreat to lower altitudes, patch microthermic expanded. Something non exclusively novel to science, but long known for e.g. the beech (Fagus) during Plio-/Pleistocene temperature fluctuations together with the subtropical-temperate Nothofagaceae inwards the Southern Hemisphere (e.g. on Papua New Guinea, inwards the tropics, Nothofagaceae forests are today starting at c. 1500 m a.s.l. (transition betwixt tropical together with subtropical zone) but were downward to 300 m during the glacial maxima. If the NLR occurs at e.g. 1500–2500 m a.s.l. today at 10–15 °C, together with the temperature was 10 °C higher inwards the past times inwards the area, an ancient temperature-sensitive fellow member of the same lineage could only direct hold thrived at 3500–4500 m
Let's direct hold Wu et al. information for granted (which I would non recommend) together with assume that the critical plants didn't modify their temperature niche inwards the terminal 35 Ma (i.e. the temperature tolerance fulfil the actuo-palaeontological principle). The is piddling argue to assume that the moist-adiabatic lapse charge per unit of measurement didn't apply dorsum together with so (the geotectonic province of affairs is non that unlike from today). Thus, they would direct hold had to grow higher upward than their modern counterparts, i.e. non at 900–1600 m a.sl. (the height at which nosotros direct hold today 17–22 °C) or 1300–1600 m a.s.l. (the actual CA-result) but at 1400–2200 m (min. scenario) or 2400–3100 m (max. scenario).
If they would direct hold grown at the same summit than their modern (Tibetan) counterparts (i.e. height tolerance fulfil the actuo-palaeontological principle), some of their temperature tolerances together with preferences must direct hold been higher than today, because the temperature at 1300–1600 m was 20–21.5 °C (yearly average) for the min. scenario (+3 °C) or 25–26.5 °C (+8 °C), i.e. (near)tropical, inwards the non only regionally but globally much warmer slow Eocene.
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| How to coexist when the global (and local) temperature rises? Lowlandia and Highlandia are 2 perfect NLRs alongside temperature-fitting altitudinal distribution together with a highly informative modern-day coexistence space. To hold coexistence, the ancient Lowlandia could direct hold been growing at higher altitudes or the ancient Highlandia was to a greater extent than warmth-loving. When the full general climate was warmer (or colder) than today, a parameter attain shift, thus a violation of the actuo-palaeontological principle, is inevitable. |
Not thing how you lot plow it, the actuo-palaeontological regulation is violated for at to the lowest degree 1 parameter, or fifty-fifty both. Make the math: if it was inwards full general 2.5° warmer than today, together with the critical taxa thrived at 500 m lower altitudes, Wu et al.'s CA resultant would last plumbing equipment reality while violating the basic regulation for both parameters. If it would direct hold been 2.5 °C colder than today inwards general, together with critical taxa thrived higher up, it would last completely off reality. But soundless the CA protocol would ensure 1 gets a result.
I volition non sketch in 1 lawsuit to a greater extent than how the authors should direct hold done their analysis, since this is shown inwards slap-up item inwards our 2 2016 papers. Just make it (or at to the lowest degree try).
What the vegetation really points to
With Wu et al.'s data, nosotros teach 'coexistence intervals' of 16.8–21.4 °C defined past times Betula and Cedrus (either the value inwards Wu et al.'s tabular array 1 or fig. 4 is a mistype), no outliers. And an height either inside the expectation of the moist-adiabatic lapse rate, 1300–1600 m a.s.l. (Cedrus + Podocarpus; Abies, Artemisia, Ephedra, Picea, and Chenopodiaceae eliminated as likewise high), or 1 likewise high, 2000–2100 m (Ephedra, Picea too high; Lygodium, the temperate(!) fern, Castenea and Podocarpus too low); both intervals conform 4/5 of the NLRs. Today, at this altitude, nosotros direct hold subtropical weather condition inwards the larger expanse (Cwa), together with it was probable warmer inwards the slow Eocene (we had vast forests roofing e.g. the Arctic), which explains the "tropical-subtropical" inwards the abstract together with conclusions.
[Wu et al. written report together with utter over 1300–2600 m as resultant (an upper boundary which according to their tabular array 1 would last also likewise high for Carya, Castanea, Lygodium, Meliaceae together with Podocarpus), so this is either a mistyper or some arcane analysis result.]
The most conspicuous observation is that typical, widespread, really mutual subtropical elements (especially inwards the Eocene) such as the infamous Engelhardioideae – the taxon defining M(A)AT of originally >17.5 °C (Mosbrugger & Utescher 1997), together with so >15.6 °C (all after studies), at nowadays >13.4 °C (Utescher et al. 2014; existent modern MAT minimum tolerance lies at 10–11 °C: encounter Grimm & Denk 2012) are missing from the pollen taxon list. As are most other typical subtropical-tropical elements. In the slow Eocene at mid-altitudes (1300–1600[–2600] m), alongside a global temperature most v °C warmer than today (cf. Zachos et al. 2001), they would demand to last present.
The only taxa associated frequently associated alongside the subtropics (but non tropics) introduce are 2 conifers with a restricted, probable relict (i.e. unrepresentative) modern distribution: Keteleeria and "Taxodiaceae" (the latter using a long outdated classification concept [see fifty-fifty Wikipedia entry], some other criterion of the Coexistence Approach). The kernel exercise of the latter, the Taxodioideae, are known for forming coals beingness the principal chemical factor of subtropical, low-altitude (sea-level) swamp forests (still today as relicts, encounter e.g. Earle 2011 onwards: the Japanese Cryptomeria; the Chinese Glyptostrobus; and U.S. Taxodium). But inwards the slow Eocene, at sea-level at this latitude, the temperature must direct hold been tropical. Other Cupressaceae genera that were included inwards the defunct Taxodiaceae direct hold every bit reduced modern distribution areas but seem to last distinctly temperate (some are fifty-fifty extending into the boreal zone) together with without proper (SEM-based) pollen analysis it is impossible to create upward one's heed which Cupressaceae Wu et al.'s "Taxodiaceae" really stand upward for (and fifty-fifty with, it may last difficult). The family itself is quite old, together with probable lost a lot of lineages, alongside relatively few survivors today.
Lacking a lot of typical subtropical elements (or genuinely tropical ones) 1 would hold off to let out inwards the slow Eocene, the flora boosts (subtropical-)temperate ones: nosotros plainly had a conifer-dominated wood (Abies, Picea, Pinus) with few deciduous elements: the birch (Betula) together with the walnut lineage, the Juglandoideae (Juglans, Carya), sis of the Engelhardioideae (see this GWoN postal service on the family), perchance montane evergreen elements, likewise (Quercus, also hither the pollen would demand to last re-studied using high-resolution SEM, to brand whatever exercise of it for palaeo-vegetation/-climate reconstruction). The combination of Pinus, Betula and Juglans would last typical for the temperate broadleaved together with mixed forest (TBMF) biome [Wikipedia] of the Northern Hemisphere, the lists includes fifty-fifty the typical fern found inwards some TBMF forests (Polypodium).
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