the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Treeline species Betula ermanii are more adaptable to alpine environments than non-treeline species Picea jezoensis: evidence from leaf functional traits
Abstract. Understanding functional trait differences between treeline and non-treeline species is key to exploring their adaptive strategies under environmental stress and predicting subalpine forest dynamics. On Changbai Mountain, Betula ermanii dominates over 90 % of the treeline zone, while Picea jezoensis accounts for over 70 % of the lower elevation zone. It remains unclear whether P. jezoensis, a treeline genus elsewhere, would eventually shift upward and replace B. ermanii. We thus investigated leaf functional traits, their intraspecific variation, and inter-trait relationships for both species along the elevational gradient. B. ermanii exhibited higher LDMC, N, P, and gs, but lower WUE and δ18O at higher elevations, with the greatest intraspecific variability in photosynthetic and hydraulic traits, and tighter linkages among traits. In contrast, P. jezoensis exhibited an increase in δ13C and a decrease in SLA with elevation, accompanied by the greatest intraspecific variability in photosynthetic traits and weaker correlations among traits. Overall, B. ermanii employs a resource-acquisition strategy enabling it to occupy resources and space, while P. jezoensis adopts a resource-conserving strategy by emphasizing shade and drought-tolerance, resource conservation, and long-term adaptation at lower elevation, limiting its ability of upward range expansion. These findings enhance our understanding of their adaptive strategies and responses to elevational change, informing predictions of subalpine forest dynamics.
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Status: open (until 02 May 2025)
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RC1: 'Comment on egusphere-2025-369', Anonymous Referee #1, 31 Mar 2025
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The manuscript has an important scientific significance by giving new information about functional traits and their inter- and intraspecific variation of two tree species in an elevation gradient in Northeast China: a deciduous treeline species Betula ermanii and a conifer tree species Picea jezoensis ( non-treeline species), and a discussion on the species adaptive strategies and ability of upward range expansion. However, there are some important issues to be fixed, regarding the analyses, explanations of the obtained results and result interpretation. After these have been fixed, the manuscript is of good quality to be published in EGUsphere, according to my opinion. Below are my detailed comments, which are also marked as comments to the pdf file attached:
L23-24: What is expected from these in relation to treeline environment?
L33-34: This was against expectations, right?
L36-37: Why these traits are expected to relate adaptation to lower elevation? I would expect an opposite effect, as high elevation environments are sometimes also dry, and resource conservation strategy may be beneficial there due to harsh conditions.
L61: Do you refer here inter- or intraspecific variation in traits? Or both?
L71-73: Isn't resource conservation strategy something that would be beneficial at the treeline due to harsh environmental, not the resource acquisition strategy?
L83-85: How about the evolutionary aspect here? Physiological traits, such as photosynthesis, is an old trait in evolutionary perspective, and is therefore expected to be rather fixed, i.e. not very plastic, due to the conservative inheritance of the trait caused by its complex genetic basis.
L121-123: Why we are expecting reduced leaf trait connectivity when the construction costs need to be minimized? Could you give a short clarification for this.
L147-148: How much of leaf trait differences are explained by the fact that the other sp is conifer and other is deciduous? A short discussion on this would be good to have.
L216-218_ What were used here as fixed and random factors?
L221: Can you explain here what Q3 and Q1 are?
L251: This is an unclear statement.
L263-264: Adaptation to higher elevation (colder or drier conditions)?
L266-267: The datapoints in each elevation are not independent, as they are from the same tree. Therefore, a simple linear regression is not appropriate here. Mean values per tree should be used, but this is also a problem as there is only 6 datapoints then. Thus, some other method than simple lm should be used. Maybe using a tree as a random factor (?)
L269-270: Can you give a little bit more explanation here, e.g. was the other analysis done combining all the data together (both species)?
L280-283: I am wondering how much of this variation is due to measurement challenges, as these traits, especially photosynthetic traits, are very sensible, and therefore their measurement usually need several repetitions? A discussion about this would be good to add.
L312: Surprising that there is no relationship between A and SLA, and only a weak relationship between A and gs in P. jezwoensis and no relationship at all in B. ermanii ? There is a discussion about this in the discussion section., but it should be extended and made more clear.
L313: This is confusing as the same color combination is used to separate the species.
L317: Does this figure give significantly more information than fig 3 and 4? If so, can you explain a bit more what does each point (a, b and c) mean?
L324-325: What does the radical water use strategy mean and how was that observed? This is discussed later on (in 4.2) a bit more, but if it is mentioned here it needs some explanation.
L328-329: But there was no connection between these traits, which is surprising (fig 4). This is discussed a bit more later on in 4.2, but same as above: if it is mentioned here, it needs some explanation.
L331-335: I think this is a bit controversial, as harsh treeline environments are not resource-rich. Can you explain better this idea?
L337-339: What kind of mechanisms is this based on? Can you explain better, so that the next sentence would be more justified.
L351-352: But on the other hand, the traits showed more intraspecific variation, indicating that there is potential.
L355-356: It may limit a rapid expansion, but on the other hand, it can better ensure survival in harsh conditions. This should be also discussed.
L362-363: Can you explain what is the reasoning behind this statement?
L364-365: Is this due to a stronger adaptation to a certain type of environment (cold /dry)?
L365: Is there something missing from this sentence?
L367-371: I think that this part is somehow aiming to give an explanation for my questions /comments (regarding e.g. the lines 324-325 and 328-329), but it could be a bit more clear, expanded and combined with the statements in the corresponding lines.
L374-376: I think that this statement is not so clear, especially what comes to photosynthesis, which is an old plant trait and thus expended to be quite fixed.
What do you mean by anisotropic traits?
L377-378: Maybe some of them, but photosynthesis is an old trait and therefore expected to be more fixed.
L382-384: This makes sense, indicating that this is thus more as a species-specific difference than environmental adaptation.
L397-399: This sentence seems incomplete, or something is missing.
L417-418: What do you mean by "the central traits"?
L421-427: Could you explain these interesting mechanisms a bit more. For example, why an increase in photosynthetic rate was not observed in B. ermanii when stomatal opening increased? How the independent regulation of water and carbon is expected to work?
L433-435: This was not tested with an experimental setting. So, it is difficult to claim that the differences were driven only by the environment and not by species-specific differences. And the difference in WUE seemed to be species-specific difference anyway. This aspect should not be forgotten from the discussion.
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