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Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change

Published online by Cambridge University Press:  05 November 2026

Munib Khanyari*
Affiliation:
Nature Conservation Foundation, India Institut de Ciència i Tecnologia Ambientals , Spain Snow Leopard Trust, USA
Deepti Bajaj
Affiliation:
Nature Conservation Foundation, India Ashoka University , India
Manjot Kaur Jamwal
Affiliation:
Nature Conservation Foundation, India
Shahid Hameed
Affiliation:
Nature Conservation Foundation, India
Herman Ramesh
Affiliation:
Nature Conservation Foundation, India
Kulbhushansingh Ramesh Suryawanshi
Affiliation:
Nature Conservation Foundation, India Snow Leopard Trust, USA CIFAR, Future Flourishing Program, Canada
*
Corresponding author: Munib Khanyari; Email: munib@ncf-india.org
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Abstract

Content of image described in text.

The Western Himalaya harbours distinctive and rich mountain mammal assemblages, yet the persistence of species may conceal underlying ecological decline. We propose the concept of deceptive diversity, describing the period during which extinction debt accumulates but remains largely undetected as species continue to persist. Synthesising evidence from long-term population monitoring, distributional assessments, genetic studies, and recent observations of climate- and development-driven ecological change, we show that many mammals in the Western Himalaya persist as small, fragmented and peripheral populations exhibiting restricted distributions and isolation. Emerging consequences include novel species interactions, declining connectivity, reduced dispersal opportunities, and increased extinction debt, despite the continued presence of many species. We argue that biodiversity assessments based solely on occupancy or species richness risk overlooking these hidden processes of ecological erosion. Recognising deceptive diversity shifts conservation attention from documenting extinctions to identifying the early demographic and ecological warning signals that precede them. This provides a framework for proactive conservation of populations before declines become irreversible; a proposition which is relevant to other important mountain biodiversity hotspots globally.

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Type
Perspective
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
Copyright
© The Author(s), 2026. Published by Cambridge University Press
Figure 0

Figure 1. Global change as a driver of deceptive diversity, eventually accumulating extinction debt, in the Western Himalaya. Climate change and accelerating infrastructure development interact to compress habitats, increase fragmentation, facilitate novel species interactions, and reduce connectivity among naturally small and isolated range-edge populations. These processes diminish demographic resilience and adaptive capacity while species remain present, generating an extinction debt that may remain undetected until local extirpations occur. [Digitally illustrated by Manini Bansal originally for this manuscript].Figure 1. Long description.

Figure 1

Figure 2. Map showing select species exhibiting fragmented populations in the Western Indian Himalaya.Figure 2. Long description.

Author comment: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R0/PR1

Comments

Dear Editors,

Please find attached our manuscript entitled “Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change ” for consideration as a Perspective Article in Cambridge Prisms: Extinction.

The Western Himalaya supports one of the world’s most distinctive mountain mammal assemblages, yet many species persist as small, fragmented, and isolated populations that appear stable despite increasing ecological pressures. In this manuscript, we synthesise evidence from long-term population monitoring, distributional studies, genetics, and recent observations of climate and development driven ecological change to argue that biodiversity assessments based solely on species presence or richness may overlook early warning signs of decline. We introduce the concept of deceptive diversity as a framework for describing the period during which populations remain present while the demographic, ecological, and genetic processes necessary for long-term persistence progressively erode, potentially leading to the accumulation of extinction debt.

Rather than documenting extinctions after they occur, our review advocates for detecting the hidden processes that precede biodiversity loss. We discuss how interacting climate and development pressures, i.e. global change, limits the ability of mountain species to respond to environmental change and illustrate these ideas using examples from the Western Himalaya. Although centered on this region, we argue that the framework has broader relevance for mountain ecosystems worldwide where species persistence may mask declining population resilience.

We believe this manuscript is well suited to Prisms: Extinction because it combines empirical evidence with conceptual synthesis to address a central challenge in extinction science, recognizing biodiversity loss before local extirpations occur. We hope the paper will stimulate discussion on how extinction risk is assessed and encourage greater emphasis on long-term demographic, genetic, and ecological monitoring in conservation.

This manuscript is original, has not been published previously, and is not under consideration elsewhere. All authors have approved the manuscript and have no conflicts of interest to declare.

Thank you for your consideration. We look forward to hearing from you.

Sincerely,

Munib Khanyari

On behalf of all co-authors

Review: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R0/PR2

Conflict of interest statement

Reviewer declares none.

Comments

I read with great interest the manuscript entitled “Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change”. The manuscript highlights the ways in which looking at diversity alone can mask ecological decline in the face of global change through mechanisms such as accumulating extinction debt, range-edge vulnerability, and biotic homogenization, specifically focusing on medium-large mammals in the Western Himalayas. I found the manuscript to be very relevant and timely, emphasizing the urgent need for more intensive and long-term monitoring of mammalian species in this region to detect the early-warning signs of population decline in order to prevent local extirpation. However, I believe the authors could improve the quality of the manuscript and have some recommendations to that effect:

Major comments:

1) The manuscript will greatly benefit from a round of proofreading and editing. Some instances of awkward phrasing of sentences, grammatical errors, and inconsistent usage of terminology are listed in the Minor Comments section. Addressing these will improve the clarity and readability of the manuscript.

2) In Figure 1, many of the points being made are redundant. For example, the concept of population decline is emphasized under ecological mechanisms (point #5), inside the blue box under “Community appears intact…”, as well as under the Early Warning Signs subheading. The same can be said for the points being made about altered species interactions, reduced connectivity, etc. I think this figure can be made much more concise and these redundancies removed.

3) In p6-L3-45 (page 6, lines 3-45), the manuscript makes the point that a population being peripheral doesn’t necessarily indicate greater extinction risk, and that peripheral populations become vulnerable when they are also small, fragmented and demographically isolated. Then some species are listed as falling into these criteria and being “geographically restricted peripheral populations that are isolated from the core of their global distributions”, with a reference to Figure 2. On looking at Fig. 2, the ranges presented in 2A are very clearly fragmented, and it is easy to see why these populations might be vulnerable. However, when looking at Fig.2B, which plots ranges for peripheral (but seemingly not geographically restricted) populations, I am unsure as to what the point being made is. Are these populations less vulnerable compared to the species in 2A on account of the ranges being more contiguous and much larger? Are they equally vulnerable because the Indian populations are peripheral and disconnected from the core of their global distribution? If it is the latter, I don’t think 2B adequately conveys that message.

4) In Figure 2 and the parts of the manuscript referencing Fig. 2 (p6-L30-40), some species like wild yak are mentioned in the text but do not feature in Fig. 2. Conversely, the figure includes ranges for species like blue sheep and Tibetan Argali that are not mentioned in the manuscript.

Minor Comments:

-Line numbers start afresh on every page, which makes it confusing to reference particular lines. Continuous line numbering is preferable.

-p3-L17: Sharma et al. 2025 cited here is not included in the reference list.

-p7-L19: Replace “unable to increase” with “fails to increase in size”

-p7-L38-39: “...32% (16-53%)...” I presume the brackets contain 95% confidence intervals, but it would be good to state that explicitly.

-Inconsistent capitalization of “urial” or “Urial” throughout the manuscript

-Inconsistent use of either Tibetan antelope or chiru to refer to the same species.

p8-L35-40: This sentence is phrased in a confusing manner and could use some reworking/editing.

-p10-L3-8: “demonstrate”, not “demonstrating”. It would also be good to give a couple of examples of climate change hotspots that have glacier retreat rates comparable to the trans-Himalaya to emphasize this point.

-p10-L16: Military infrastructure is framed as affecting mountains worldwide; however, it seems to me to be a more local issue mostly restricted to the study area.

-p10-L20-21: I presume this is referring to the government of India, but it would be good to state that explicitly.

-p10-L23: The conversion from km2 to hectares is only offered here and nowhere else in the manuscript. I would recommend not including it here for consistency.

-p12-L23-24: The sentence “Roads alongside…” is phrased awkwardly and can be reworked for clarity.

-In the subheading for Figure1, the meaning of the phrase “...but ecological processes are shifting beneath the surface” is not very direct and clear. I think this can be reworded for clarity.

-Figure 1 also mentions concepts like ecosystem functions that are not mentioned/addressed at all in the manuscript.

-The colors used for Ladakh urial and Markhor in Figure 2A are very similar; a different color choice would be preferable.

-The manuscript refers to the same species as Hangul in the text and as Kashmir stag in Fig. 2, which is confusing.

Recommendation: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R0/PR3

Comments

I was only able to secure one review of this manuscript, but that review plus my own reading of the manuscript were collectively sufficient to make an assessment of “Major Revision.”

Reviewer 1 sees value and novelty in the manuscript, but notes several areas for improvement, which I endorse. In particular, the manuscript suffers from a disconcertingly high level of internal inconsistencies that frequently crop up with regard to terminology, context, and species names. The authors must attend to all of these clean-up issues, which will ultimately pay off by easing readers’ understanding and adoption of the ideas proposed.

Beyond the points made by Reviewer 1, I would also add that the manuscript, though well intentioned, is critically lacking in context, both with regard to important foundational publications and recent papers on basically the same problem (just in different parts of the world)

For example, the idea that a variety of forces interact synergistically to drive a species toward extinction (or a population toward extirpation) are well embodied in the concept of the extinction vortex (Gilpin and Soule 1986). That classic reference makes the key point, echoed in this manuscript, that ‘what you see is not what there really is.’ Their primary context was in population abundance, but the idea that a suite of synergistic interactions carries over to drive extinction dynamics exists with regard to spatial occupancy data as noted in this manuscript.

More recently, Falaschi et al (2025) investigate directly how different predictions of extinction risk are when based on occupancy data versus abundance data. This paper is highly relevant to the authors’ point of view and should be cited.

Related points are also made by Ruiz-Gutiérrez and Zipkin (2011) and, in a geographically similar region, by Collen et al. (2006).

Other comments:

Latin binomials need to be italicized on all usages, but they are not, especially in the Literature Cited. Please correct this.

References mentioned:

Falaschi, M., Lo Parrino, E., Manenti, R. and Ficetola, G.F., 2025. Differences in estimates of extinction risk between occupancy and abundance data. Conservation Biology, 39(4), p.e70020.

Collen, B., Bykova, E., Ling, S., Milner-Gulland, E.J. and Purvis, A., 2006. Extinction risk: a comparative analysis of Central Asian vertebrates. Biodiversity & Conservation, 15(6), pp.1859-1871.

Gilpin, M.E. & Soule´, M.E. (1986). Minimum viable populations: processes of extinction. In: Conservation Biology: The Science of Scarcity and Diversity (ed. Soule´, M.E.). Sinauer Associates, Sunderland, MA, pp. 19–34.

Harnik, P.G., Simpson, C. and Payne, J.L., 2012. Long-term differences in extinction risk among the seven forms of rarity. Proceedings of the Royal Society B: Biological Sciences, 279(1749), p.4969.

Ruiz-Gutiérrez, V. and Zipkin, E.F., 2011. Detection biases yield misleading patterns of species persistence and colonization in fragmented landscapes. Ecosphere, 2(5), pp.1-14.

Decision: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R0/PR4

Comments

No accompanying comment.

Author comment: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R1/PR5

Comments

No accompanying comment.

Recommendation: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R1/PR6

Comments

I have read the manuscript and the authors' responses to the reviews. Overall, I found an earnest effort to address the long list of critiques that the reviewer and I raised with the first submission. The authors have improved the contextualization of their manuscript via citations to classical literature that was pertinent to their main points and by correcting / standardizing terminology that non-specialists would have likely found confusing with regard to species naming conventions and regional references.

With these modifications, I am recommending acceptance.

Bill Fagan

Decision: Deceptive diversity: Mammals in the Indian Western Himalaya face extinction debt due to global change — R1/PR7

Comments

No accompanying comment.