SPECIES AND FUNCTIONAL DIVERSITY OF THE VALLEY FORESTS OF THE KARKARALY MOUNTAINS (CENTRAL KAZAKHSTAN)

       N. V. Ivanova^1*, M. Yu. Ishmuratova², M. P. Shashkov²

 

¹Institute of Mathematical Problems of Biology, Keldysh Institute of Applied Mathematics, Russian Academy of Sciences

Professor Vitkevich st. 1, Pushchino, 142290 Moscow Region, Russian Federation

²Karaganda National Research University named after academician Ye. A. Buketov

Universitetskaya st. 28/3, 10002 Karaganda, Kazakhstan

*E-mail: natalya.dryomys@gmail.com

Received: 10.06.2026

Revised: 26.06.2026

Accepted: 17.08.2026

The forests of the Kazakh Uplands (Saryarka) are island pines (and secondary small-leaved stands) with boreal and other forest herb species considerably separated from their range geographically. The aim of this study is to quantitatively assess the species and functional plant diversity of the valley forests of the Karkaraly Mountain Range. Two groups of communities were studied: (1) forests in the Kendara intermountain valley and (2) forests growing on the outer edge of the Karkaraly mountain range. A total of 33 vegetation relevés were conducted in the forests of Group 1 and 14 in the forests of Group 2. Alpha diversity was assessed using the species richness index and randomisation tests. To analyse gamma diversity, we calculated Hill indices and plotted cumulative diversity curves. Beta diversity was assessed using the Jaccard distance. Functional diversity was assessed based on the diversity of ecological-coenotic groups (ECG). Species indicator values were calculated for each group. The results of the study showed that the groups of communities are divided into two groups in the NMDS diagram. The species richness of the herbaceous layer in the forests of the Kendara Valley is significantly higher than in the forests growing outside the mountain range (24±5 and 21±5 species, respectively). The forests of the Kendara Valley were somewhat poorer in terms of species diversity (113 and 118 species, respectively); significant differences were observed in the effective species numbers for the Shannon and Simpson indices. The Jaccard distance between the analysed groups of relevés was 0.63. An assessment of functional diversity revealed that meadow-steppe, boreal, and nemoral species prevailed in the communities. In the Kendara Valley, boreal and nemoral species together represented 52% of the total; in the forests outside the mountain range, their proportion decreased as the proportion of nitrophilous, meadow-steppe, pine forest, and wetland ecological communities increased. For forests in the second group, a greater diversity of ecological-coenotic groups among significant indicator species has been observed. The results confirm that the forests in the Kendara Valley are in a better state of preservation. The potential impact of recent invasions of non-native earthworm species (Lumbricidae) and the box elder (Acer negundo) on the ecosystems of the Karkaraly forests is discussed.

Keywords: Hill numbers, ecological-coenotic groups, Kazakh Uplands, Karkaraly State National Natural Park, Acer negundo

 REFERENCES

Auelbekova A. K., Shorin S. S., Kyzdarova D. K., Musina R. T., Esengalieva A. D., Formirovanie podrosta i drevostoya na mestakh gorel’nikov v Karkaralinskom gornolesnom massive (Regeneration of undergrowth and tree stands in burnt areas in the Karkaraly Mountain Forest Massif), Ekologo-geograficheskie problemy regionov Rossii. VIII vserossiiskaya nauchno-prakticheskaya konferentsia s mezhdunarodnym uchastiem (Environmental and Geographical Issues in Russian Regions. The 8th All-Russian Scientific and Practical Conference with International Participation), Samara, January 15, 2017, Samara: Samarskii gosudarstvennyi sotsial’no-pedagogicheskii universitet, 2017, pp. 90–93.

Vinogradova Yu. K., Maiorov S. R., Kostina M. V., Klen yasenelistnyi (Acer negundo L.): morfologiya, biologiya i otsenka invazivnosti (Box elder (Acer negundo L.): morphology, biology, and assessment of invasiveness), Moscow: Tovarishchestvo nauchnykh izdanii KMK, 218 p.

Vinogradova Yu. K., Formirovanie vtorichnogo areala i izmenchivost’ invazionnykh populyatsii klena yasenelistnogo (Acer negundo L.) (The formation of a secondary range and the variability of invasive populations of Box Elder (Acer negundo L.)), Byulleten’ Glavnogo Botanicheskogo sada, 2006, Vol. 190, pp. 25–48.

Gvozdetskii N. A., Mikhailov N. I., Fizicheskaya geografiya SSSR. Aziatskaya chast’ (Physical Geography of the USSR: The Asian Part), Moscow: Vysshaya Shkola, 1987, 448 р.

Gorchakovskii P. L., Lesnye oazisy Kazakhskogo melkosopochnika (Forest oases of the Kazakh Lowland), Moskva: Nauka, 1987, 157 p.

Gribanov N. G., Stepnye bory Altaiskogo kraya Kazakhstana (Steppe pine forests of the Altai Region of Kazakhstan), Moscow-Leningrad: Goslesbumizdat, 1960, 155 p.

Dancheva A. V., Zalesov S. V., Raionirovanie sosnovykh lesov Respubliki Kazakhstan (Zoning of pine forests in the Republic of Kazakhstan), Lesa Rossii i khozyaistvo v nikh,  2024,  No 1 (88), pp.  4–28, DOI: 10.51318/FRET.2023.88.1.001

Dikareva T. V., Leonova N. B., Fitoraznoobrazie sopok Karkaralinskogo nauional’nogo parka (Respublika Kazakhstan) (Plant diversity of the hills in the Karkaraly National Park (Republic of Kazakhstan)), Aridnye ekosistemy, 2014, Vol. 20, No 4 (61), pp. 105–114.

Kozina V. V., K istorii demograficheskogo razvitija Karagandinskoj oblasti (On the history of demographic development in the Karaganda Region), Vestnik Karagandinskogo universiteta. Serija: istorija, filosofija, pravo, 2006, No 3 (43), pp. 81–92.

Ishmuratova M. Yu., Myrzaly G. Zh., Ivlev V. I., Matveev A. N., Flora Ulytau (Tsentral’nyi Kazakhstan) (Flora Ulytau (Central Kazakhstan)), Karaganda: Bolashak-Baspa, 2016, 127 p.

Ishmuratova M. Yu., Tleukenova S. A., Gavril’kova E. A., Tyrzhanova S. S., Shashkov M. P., … & Hapilina O. N., Sosudistye rasteniya Karagandinskoi oblasti (konspekt, analiz i DNK-barkodirovanie flory) (Vascular Plants of the Karaganda Region (synopsis, analysis, and DNA barcoding of the flora)), Astana: “Tsentr Elit”, 2024, 592 p.

Ivanova N., Shashkov M., Species and functional diversity of forest communities with wild fruits plants in Central Kazakhstan, Fundamental and Experimental Biology, 2025, Vol. 11930, No 3, pp. 159–167.

Kupriyanov A. N., Konspekt flory Kazakhskogo melkosopochnika (Synopsis of the flora of the Kazakh Lowland), Novosibirsk: Akademicheskoe izd-vo “Geo”, 2020, 423 p., DOI: 10.21782/B978-5-6043021-8-7

Maksutova P. A., Dyusekeeva Sh. E., Kulmaganbetova A. O., Fizicheskaya geografiya Karagandinskoi oblasti (Physical geography of the Karaganda Region), Karaganda: KarGU, 2005, 59 p.

Malinovskikh A. A., Vliyanie klena yasenelistnogo (Acer negundo L.) na estestvennoe vozobnovlenie sosny obyknovennoi v Barnaul’skom lentochnom boru (The effect of Box Elder (Acer negundo L.) on the natural regeneration of Scots Pine in the Barnaul strip pine forest), Lesnoi vestnik. Forestry Bulletin, 2023, Vol. 27, No 3, pp. 48–56, DOI: 10.18698/2542-1468-2023-3-48-56

https://www.pogodaiklimat.ru/weather.php?id=36335, (2026, 1 April).

Popov Yu. G., Ryazantsev V. P., Kornilovy, Karbyshevy, Ryazantsevy i drugie kazaki stanitsa Karkaralinskoi. Shtrikhi kraevedcheskoi letopisi (The Kornilovs, Karbyshevs, Ryazantsevs, and other cossacks of the Karkaralinskaya stanitsa. Notes from a local history chronicle), Novosibirsk: Arta, 2012, 142 p.

Popov Yu. G., Khraniteli Karkaralinskogo bora (sud’ba Yakova Bernatskogo) (The guardians of the Karkaralinsky Forest (the fate of Yakov Bernatsky)), Botanicheskie issledovaniya Sibiri i Kazakhstana, 2008, Vol. 14, pp. 100–111.

Rachkovskaya E. I., Opustynennye polynno-dernovinnozlakovye stepi Tsentral’nogo Kazakhstana (Desert sagebrush-bunchgrass steppes of Central Kazakhstan), Rastitel’nost’ Rossii, 2016, Vol. 28, pp. 108–124, DOI: 10.31111/vegrus/2016.28.108

Slavchenko V. S., Slavchenko N. P., Analiz fauny dozhdevykh chervei (Lumbricidae, Oligochaeta) Karkaralinskogo prirodnogo parka (Kazakhskoe nagor’e) (Analysis of the earthworm fauna (Lumbricidae, Oligochaeta) in the Karkaraly Natural Park (Kazakh Highlands)), Vliyanie genotipa i kompleksa ekologicheskikh faktorov na zhiznedeyatel’nost’ organizmov (The influence of genotype and a complex of environmental factors on the vital functions of organisms), Karaganda: Karagandinskii gosudarstvennyi universitet, 1989, pp. 107–112.

Smirnov V. E., Khanina L. G., Boborovskii M. V., Otsenka vidovogo raznoobraziya rastitel’nosti na osnove integral’nogo statisticheskogo podkhoda v usloviyakh neodnorodnykh dannykh (na primere zapovednika “Kaluzhskie zaseki”) (Assessment of vegetation species diversity based on the integrated statistical approach for heterogeneous data (a case study for the “Kaluzhskie zaseki” State Nature Reserve)), Tomsk State University Journal of Biology, 2014, No 4 (28), pp. 70–87, DOI: 10.17223/19988591/28/5

Smirnov V. E., Khanina L. G., Bobrovskii M. V., Obosnovanie sistemy ekologo-tsenoticheskikh grupp vidov rastenii lesnoi zony Evropeiskoi Rossii na osnove ekologicheskikh shkal, geobotanicheskikh opisanii i statisticheskogo analiza (A Framework for ecological-coenotic groups of plant species in the forest zone of European Russia based on ecological scales, vegetation relevés, and statistical analysis), Bulletin of Moscow Society of Naturalists. Biological Series, 2006, Vol. 111, No 2, pp. 36–47.

Smirnov V. E., Khanina L. G., Bobrovskii M. V., Rasshirennaya sistema ekologo-tsenoticheskikh grupp vidov sosudistykh rastenii dlya boreal’noi, gemiboreal’noi i umerennoi lesnykh zon Evropeiskoi Rossii (Validation of the ecological-coenotic groups of vascular plants for European Russian forests on the basis of ecological indicator values, vegetation releves and statistical analysis), 2008, available at: https://golnk.ru/5KDQK (2026, 1 April).

Terekhina T. A., Ovcharova N. V., Elesova N. V., Biologicheskaya i strukturnaya otsenka populyatsii klena yasenelistnogo (Acer negundo L.) v Barnaul’skom lentochnom boru (Biological and structural assessment of Box Elder (Acer negundo L.) populations in the Barnaul strip pine forest), Problemy botaniki Yuzhnoi Sibiri i Mongolii, 2020, Vol. 19, No 2, pp. 374–379.

Khanina L. G., Smirnov V. E., Bobrovskii M. V., Otsenka vidovogo raznoobraziya rastitel’nykh soobshchestv zapovednika “Kaluzhskie zaseki” na osnove indeksov Shennona, Simpsona i chisel Khilla (Species diversity of plant communities of Kaluga Zaseki Reserve: estimation based on Shannon and Simpson indices and Hill’s numbers), Lesovedenie, 2016, No 1, pp. 65–73.

Shashkov M. P., Ivanova N. V., Abukenova V. S., Ishmuratova M. Yu., Otsenka faktorov, limitiruyushchikh rasprostranenie dozhdevykh chervei v lesnykh soobshchestvakh Tsentral’nogo Kazakhstana, po geobotanicheskim dannym (An assessment of factors limiting the distribution of earthworms in forest communities of Central Kazakhstan based on vegetation data), Vserossiiskaya nauchnaya konferentsiya s mezhdunarodnym uchastiem “80 let ekologicheskoi nauke na Urale” (All-Russian Scientific Conference with International Participation: “80 Years of Environmental Science in the Urals”), Ekaterinburg, 11–15 November 2024, Ekaterinburg: Reekshen, 2024, pp. 255–257.

Acer negundo occurrences from Karkaraly mountains, GBIF Occurrence Download, 2026, available at: https://doi.org/10.15468/dl.6qfgag (2026, 1 April).

Alexander G., Almendinger J., White P., The long-term effects of invasive earthworms on plant community composition and diversity in a hardwood forest in northern Minnesota, Plant-Environment Interactions, 2022, Vol. 3, pp. 89–102, DOI: 10.1002/pei3.10075

Aporrectodea caliginosa occurrences from Karkaraly mountains, GBIF Occurrence Download, 2026, available at: https://doi.org/10.15468/dl.9tqfs4 (2026, 1 April).

Chao A., Gotelli N. J., Hsieh T. C., Sander E. L., et al., Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies, Ecological Monographs, 2014, Vol. 84, pp. 45–67. DOI: 10.1890/13-0133.1

Core Team. R: A Language and Environment for Statistical Computing, R   Foundation for Statistical Computing, Vienna, Austria, 2025, available at:  https://www.R-project.org/ (2026, 1 April).

De Cáceres M., Legendre P., Associations between species and groups of sites: indices and statistical inference, Ecology, 2009, Vol. 90, I. 12, pp. 3566–3574, DOI: 10.1890/08-1823.1

Hsieh T. C., Ma K. H., Chao A., iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill numbers), Methods in Ecology and Evolution, 2016, Vol. 7, pp. 1451–1456. DOI: 10.1111/2041-210X.12613

Ishmuratova M. Yu., Ivanova N. V., First Vegetation Plot Data from Forest Islands of Central Kazakhstan, Sampling event dataset, Karaganda Buketov University, 2024, available at: https://doi.org/10.15468/wwtfde (2026, 1 April).

            Ivanova N. V., Ishmuratova M. Yu., Shashkov M. P., Fedorova A. A., Vegetation data from Kendara Valley (Karkaraly National Park, Kazakhstan), Sampling event dataset, Karaganda Buketov University, 2026, available at: https://doi.org/10.15468/ju7pnw (2026, 1 April).

Thouvenot L., Ferlian O., Craven D., Johnson E. A., … & Eisenhauer N., Invasive earthworms can change understory plant community traits and reduce plant functional diversity, iScience, 2024, Vol. 27, No 3, Article ID: 109036, DOI: 10.1016/j.isci.2024.109036

Manly B. F. J., Randomization, bootstrap and Monte Carlo methods in biology, New York: Chapman & Hall CRC, 2007, 480 p., DOI: 10.1201/9781315273075

Mathieu J., Reynolds J. W., Fragoso C., Hadly E., Multiple invasion routes have led to the pervasive introduction of earthworms in North America, Nature Ecology & Evolution, 2024, Vol. 8, pp. 489–499, DOI: 10.1038/s41559-023-02310-7

Oksanen J., Simpson G., Blanchet F., Vegan: Community Ecology Package, R package version 2.6-8, 2024, available at: https://CRAN.R-project.org/package=vegan (2026, 1 April).

Rossi J.-P., rich: An R Package to Analyse Species Richness, Diversity, 2011, Vol. 3(1), pp. 112–120, DOI: 10.3390/d3010112

Shashkov M. P., Ivanova N. V., Abukenova V., Earthworm (Lumbricidae) field occurrences from Central Kazakhstan, Karaganda Buketov University, Occurrence dataset, 2025, available at: https://doi.org/10.15468/d4k8cg (2026, 1 April).