Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758)
Ecological niche modeling of the wood mouse Sylvaemus sylvaticus, using machine learning method and a space of 35 bioclimatic factors, revealed that the most favorable conditions for this species are found in Western Europe. Th e primary core distribution areas are concentrated there, with the proba...
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Mezhzherin, S.V. Tytar, V.M. Tereshchenko, V.O. 2025-03-07T11:16:38Z 2023 Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) / S.V. Mezhzherin, V.M. Tytar, V.O. Tereshchenko // Доповіді Національної академії наук України. — 2023. — № 5. — С. 66-72. — Бібліогр.: 13 назв. — англ. 1025-6415 https://nasplib.isofts.kiev.ua/handle/123456789/202234 599.323.45:591.9 DOI: doi.org/10.15407/dopovidi2023.05.066 Ecological niche modeling of the wood mouse Sylvaemus sylvaticus, using machine learning method and a space of 35 bioclimatic factors, revealed that the most favorable conditions for this species are found in Western Europe. Th e primary core distribution areas are concentrated there, with the probability of encountering this species in habitable landscapes reaching 60 %. In contrast, the bioclimatic conditions of Eastern Europe are pessimistic: the habitat of the species is significantly fragmented, the cores are absent, the probability of encounters does not exceed 40 %, most often being less than 10 percentiles. It is evident that among the bioclimatic factors, climate continentality plays a central role, indirectly influencing the eastern border of the S. s. sylvaticus range and shaping specific vegetation types. Direct effects of individual factors and interspecies relations are possible and do occur, limiting the species’ eastern expansion, primarily in the southern part of its eastern range borde. Моделювання екологічної ніші лісової миші Sylvaemus sylvaticus методом машинного навчання в просторі 35 біокліматичних факторів показало, що найсприятливішими для виду є абіотичні умови Західної Європи, де зосереджені головні ядра ареалу і ймовірність знахідок у придатних ландшафтах сягає 60 %. Біокліматичні обставини Східної Європи є песимальними: зона перебування виду значно фрагментована, ядра відсутні, вірогідність стрівальності не перевищує 40 %, найчастіше менш ніж 10 процентилів. Очевидно, що серед біокліматичних чинників головне значення має континентальність клімату, що опосередковано впливає на розташування східної межі ареалу S. s. sylvaticus і формування певних типів рослинності. Безпосередня дія окремих факторів і міжвидові відносини можливі і мають місце, певним чином обмежуючи просування виду на схід, однак лише на південному фасі східної межі ареалу. en Видавничий дім "Академперіодика" НАН України Доповіді НАН України Біологія Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) Моделювання екологічної ніші і фактори, що визначають ареал лісової миші Sylvaemus sylvaticus (Linnaeus, 1758) Article published earlier |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine |
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| title |
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) |
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Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) Mezhzherin, S.V. Tytar, V.M. Tereshchenko, V.O. Біологія |
| title_short |
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) |
| title_full |
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) |
| title_fullStr |
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) |
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Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) |
| title_sort |
ecological niche modeling and factors determining the range of the wood mouse sylvaemus sylvaticus (linnaeus, 1758) |
| author |
Mezhzherin, S.V. Tytar, V.M. Tereshchenko, V.O. |
| author_facet |
Mezhzherin, S.V. Tytar, V.M. Tereshchenko, V.O. |
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Біологія |
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Біологія |
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2023 |
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Доповіді НАН України |
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Видавничий дім "Академперіодика" НАН України |
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Article |
| title_alt |
Моделювання екологічної ніші і фактори, що визначають ареал лісової миші Sylvaemus sylvaticus (Linnaeus, 1758) |
| description |
Ecological niche modeling of the wood mouse Sylvaemus sylvaticus, using machine learning method and a space of 35 bioclimatic factors, revealed that the most favorable conditions for this species are found in Western Europe. Th e primary core distribution areas are concentrated there, with the probability of encountering this species in habitable landscapes reaching 60 %. In contrast, the bioclimatic conditions of Eastern Europe are pessimistic: the habitat of the species is significantly fragmented, the cores are absent, the probability of encounters does not exceed 40 %, most often being less than 10 percentiles. It is evident that among the bioclimatic factors, climate continentality plays a central role, indirectly influencing the eastern border of the S. s. sylvaticus range and shaping specific vegetation types. Direct effects of individual factors and interspecies relations are possible and do occur, limiting the species’ eastern expansion, primarily in the southern part of its eastern range borde.
Моделювання екологічної ніші лісової миші Sylvaemus sylvaticus методом машинного навчання в просторі 35 біокліматичних факторів показало, що найсприятливішими для виду є абіотичні умови Західної Європи, де зосереджені головні ядра ареалу і ймовірність знахідок у придатних ландшафтах сягає 60 %. Біокліматичні обставини Східної Європи є песимальними: зона перебування виду значно фрагментована, ядра відсутні, вірогідність стрівальності не перевищує 40 %, найчастіше менш ніж 10 процентилів. Очевидно, що серед біокліматичних чинників головне значення має континентальність клімату, що опосередковано впливає на розташування східної межі ареалу S. s. sylvaticus і формування певних типів рослинності. Безпосередня дія окремих факторів і міжвидові відносини можливі і мають місце, певним чином обмежуючи просування виду на схід, однак лише на південному фасі східної межі ареалу.
|
| issn |
1025-6415 |
| url |
https://nasplib.isofts.kiev.ua/handle/123456789/202234 |
| citation_txt |
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) / S.V. Mezhzherin, V.M. Tytar, V.O. Tereshchenko // Доповіді Національної академії наук України. — 2023. — № 5. — С. 66-72. — Бібліогр.: 13 назв. — англ. |
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66
ОПОВІДІ
НАЦІОНАЛЬНОЇ
АКАДЕМІЇ НАУК
УКРАЇНИ
ISSN 1025-6415. Dopov. Nac. akad. nauk Ukr. 2023. No 5: 66—72
C i t a t i o n: Mezhzherin S.V., Tytar V.M., Tereshchenko V.O. Ecological niche modeling and factors determining the
range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758). Допов. Нац. акад. наук Укр. 2023. № 5. С. 66—72.
https://doi.org/10.15407/dopovidi2023.05.066
© Видавець ВД «Академперіодика» НАН України, 2023. Стаття опублікована за умовами відкритого доступу за
ліцензією CC BY-NC-ND (https://creativecommons.org/licenses/by-nc-nd/4.0/)
БІОЛОГІЯ
BIOLOGY
https://doi.org/10.15407/dopovidi2023.05.066
UDC 599.323.45:591.9
S.V. Mezhzherin, https://orcid.org/0000-0003-2905-5235
V.M. Tytar, https://orcid.org/0000-0002-0864-2548
V.O. Tereshchenko, https://orcid.org/0000-0001-6671-0265
I.I. Schmalhausen Institute of Zoology of the NAS of Ukraine, Kyiv
E-mail: smezhzherin@gmail.com
Ecological niche modeling and factors
determining the range of the wood mouse
Sylvaemus sylvaticus (Linnaeus, 1758)
Ecological niche modeling of the wood mouse Sylvaemus sylvaticus, using machine learning method and a space of
35 bioclimatic factors, revealed that the most favorable conditions for this species are found in Western Europe. Th e
primary core distribution areas are concentrated there, with the probability of encountering this species in habitable
landscapes reaching 60 %. In contrast, the bioclimatic conditions of Eastern Europe are pessimistic: the habitat of the
species is signifi cantly fragmented, the cores are absent, the probability of encounters does not exceed 40 %, most oft en
being less than 10 percentiles. It is evident that among the bioclimatic factors, climate continentality plays a central
role, indirectly infl uencing the eastern border of the S. s. sylvaticus range and shaping specifi c vegetation types. Direct
eff ects of individual factors and interspecies relations are possible and do occur, limiting the species’ eastern expansion,
primarily in the southern part of its eastern range borde.
Key words: modeling, Sylvaemus sylvaticus, ecological niche, habitat fragmentation.
Th e wood mouse Sylvaemus sylvaticus (Linnaeus, 1758) is a widespread species with a large Eu-
ropean range. Comprehensive studies have designated it as a model organism for evolutionary-
geographic analyses. Karyotyping [1], allozyme analysis [2], and phylogeographic investigations
[3-5] have provided a basis for classifying this species as polytypic, represented by two subspecies:
S. sylvaticus sylvaticus and S. s. dichrus. Th is classifi cation is attributed to the existence of two refu-
gia during the last glacial maximum [4]: a western refugium covering the Pyrenees and Southern
France, and an eastern refugium encompassing the Apennine-Balkan region. From these refugia,
two cases of prochoresis were developed, forming the modern range of the species. Moreover, the
67ISSN 1025-6415. Допов. Нац. акад. наук Укр. 2023. № 5
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758)
scale of the western prochoresis signifi cantly exceeds the eastern. It extends along the Atlantic
coast from the zone of subtropical dry forests of North Africa to the boreal forest zone of Norway
and Iceland inclusively, and outreaches to the east up to the North of Ukraine. Th e territories of
Austria, Romania, Southern and Central Ukraine lie beyond the Apennine-Balkan prochoresis.
Consequently, the eastern border of the S. sylvaticus habitat spans from the Baltic Sea to the Black
Sea, through the south of Lithuania, the center of Belarus, and adjacent regions of northeastern
Ukraine, including the Russian Federation and eastern Ukraine [6, 7].
What restricts the species’ eastward expansion? Landscape and biotope factors, which typi-
cally determine the stability of the range, are not apparent in this case. Th e eastern border crosses
through relatively homogeneous habitats, which contrasts with the clear climatic diff erences be-
tween the northern and southern limits of the species’ distribution.
In addition to bioclimatic factors, the expansion of the range to the east can be limited by
competitive interactions with related species, such as voles of the Microtus aravalis — M. levis
group [8].
Th e purpose of this study was to address the following questions: 1) whether bioclimatic
factors limit the eastward distribution of S. sylvaticus; 2) whether their eff ect is direct or indi-
rect; 3) can the interaction of S. sylvaticus with related species be considered a limiting factor
for its distribution.
Th ese questions can be answered by modeling the ecological niche of S. sylvaticus using bio-
climatic variables. As a result, not only the most favorable zones for the existence of this species,
but also the bioclimatic factors that limit its geographical distribution will be established.
Th e material for modeling were a number of individuals of S. sylvaticus, the species affi liation
was established by the sequences of the cytb gene taken from GenBank, as well as a number of
samples labeled at the allozyme level [2]. A total of 115 points were used, which evenly covered
the species’ range (Fig. 1).
Th e work uses the method of machine learning based on Bayesian additive regression trees
(BART), and is implemented in the R soft ware environment (“embarcadero” package [9]). A
distribution map of predicted environmental suitability values for a species tied to geographic
coordinates is simultaneously produced, varying from 0 to 100 %. Th e quality of the model
was evaluated using ROC analysis. Th e quantitative interpretation of ROC is given by the AUC
indicator — the area bounded by the ROC curve and the axis representing the share of false-
positive classifi cations. Th e higher the AUC, the better the classifi er. Models with an AUC value
above 0.7 are considered acceptable. Th e program also
automatically determines the contribution of each pre-
dictor to the construction of the fi nal model and draws
dependence curves describing the changes in the predic-
tive values of environmental suitability as a function of
individual predictors.
Th e predictors are retreived from the CliMond data-
base [10], which contains bioclimatic indices taking into
Fig. 1. Localization of samples involved in modeling: white circles
are S. s. sylvaticus, black circles are S. s. dichrus. Habitat scheme ac-
cording to [7]
68 ISSN 1025-6415. Dopov. Nac. akad. nauk Ukr. 2023. No 5
S.V. Mezhzherin, V.M. Tytar, V.O. Tereshchenko
account temperature, humidity, and solar radiation indicators. A total of 35 parameters character-
izing the modern climate are used.
Ecological niche modeling allowed to determine the optimal areas for the habitat of S. syl-
vaticus (Fig. 2). In this case, the AUC is 0.86. Th e most favorable area for the wood mouse is the
territory of Western Europe, including France, Belgium, the Netherlands, and the western and
southern parts of Germany and Switzerland. In these well-fi tting landscapes, the probability of
56°
48°
40°
48°
40°
56°
8° 0° 8° 32°16° 24°
8° 0° 8° 32°16° 24°
56°
48°
40°
48°
40°
56°
8° 0° 8° 32°16° 24°
8° 0° 8° 32°16° 24°
Fig. 2. Territories with a set of bioclimatic factors that are suitable to varying
degrees for the habitat of S. sylvaticus. Detection probability: gradient from
dark red to orange — 50-65 % and from yellow to light green — 35-50 %. The
red contour line marks the 10th percentile threshold. Places of material collec-
tion are indicated by circles
Fig. 3. Fragmentation degree of S. sylvaticus range. Cores (non-fragmented or
slightly fragmented areas of the range) are shown in dark blue; the average lev-
el of fragmentation is depicted in blue; highly fragmented areas are is red
69ISSN 1025-6415. Допов. Нац. акад. наук Укр. 2023. № 5
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758)
detecting this species reaches 64 %. Th e mountain systems of the Alps and the Apennines belong
to zones with a detection probability of less than 10 percentiles. Southern and Northern Europe
are less favorable, with a probability of detection of 50 % occurring only in coastal areas. Eastern
Europe is the least suitable for the wood mouse, with the probability of encounters, even in the
most favorable places, being less than 50 %, and in most areas, the detection frequency of S. syl-
vaticus is up to 10 %.
Th e evaluation of the degree of fragmentation presented on the map yields similar results.
Cores are non-fragmented or slightly fragmented areas that are not scattered by zones unsuitable
for S. sylvaticus habitat. Th ey make up at least 10 % of the area range (Fig. 3). Areas with an aver-
age level of fragmentation are usually adjacent to the cores, but they also occur in isolation. Th e
rest of the distribution range is covered by fragmented territories, where areas suitable for the
species give way to unsuitable areas. Th e main cores are within the nominative range of S. s. syl-
vaticus, explaining its extensive distribution area. Th e largest cores are concentrated in Western
Europe, with one in the southwest of France, including the Pyrenees, and the second covering the
Northeast of France, the western part of Germany, Belgium, Switzerland, and the western part of
Austria. Th ere are cores in the South (the western part of the Iberian Peninsula), the North (the
southwestern part of the Scandinavian Peninsula, the northern part of Britain), and in the Center
of Europe, where the core is formed in the south of Poland and the east of the Czech Republic.
Smaller cores are concentrated within the distribution zone of S. s. dichrurus, located in the
Mediterranean, including the north of the Apennine Peninsula (with adjacent islands), to a lesser
extent the Western and Southern Balkans, and part of Asia Minor around the Sea of Marmara.
Th ere are small cores in Romania (Southern Carpathians and Transylvania). Th ere are no cores
in the territory of Eastern Europe.
Th e BART algorithm determined the likely impact of fi ve climate predictors. Th ese are tem-
perature factors (Fig. 4): the diff erence between the highest and lowest temperatures per year
(conditional climate continentality) (X5m_07); the maximum temperature of the warmest month
(X5m_05), as well as humidity indicators (precipitation seasonality (X5m_15), precipitation levels
during the driest (X5m_17) and most humid (X5m_16) months). It should be noted that the con-
nection between the factors and the response to their actions is non-linear, and therefore, there
are certain zones of optimal factor eff ects. Th us, regions with a fairly humid temperate climate
Re
sp
on
se
0.6
0.5
0.4
0.3
0.2
0.1
12 16 20 24 28
2
1
32 36 40
Degrees
Fig. 4. Response curves of pre-
dictors on the gradation of bio-
climatic factors: 1 — maximum
temperature of the warmest
month (X5m_05); 2 — the dif-
ference between the highest and
lowest temperature of the year
(X5m_07)
70 ISSN 1025-6415. Dopov. Nac. akad. nauk Ukr. 2023. No 5
S.V. Mezhzherin, V.M. Tytar, V.O. Tereshchenko
and high summer temperatures are the most favorable for the wood mouse. Th ese regions include
Western and Southern Europe and the southern part of Northern Europe. Th e continental and
fairly dry climate of Eastern Europe does not favor this species.
Th e established bioclimatic factors do not directly aff ect the wood mouse population, limit-
ing the species’ distribution to the East. Most likely, these are mainly abiotic circumstances that
form the type of vegetation. Th e wood mouse, whose prohoresis evolved from the Mediterranean,
avoids both forests with dense canopies and bushy grassy vegetation. Th erefore, in the territory
of Eastern Europe, where the primary types of natural vegetation are coniferous forests, meadows
and steppes, S. sylvaticus has a limited number of optimal stations. Th ese are mainly areas with
a dense soil surface, where it is easy to collect small seeds: low-growing shrubs and thickets of
ruderal vegetation. Such places are concentrated in anthropogenically altered landscapes, which
serve as corridors for the species’ expansion to the East. In the north of the eastern border of the
range, the infl uence of the vegetation factor is very evident since S. sylvaticus is present exclusively
in agricultural lands. In the south, where open landscapes predominate, there are more plant com-
munities suitable for the wood mouse. It is no coincidence that in this zone, the range extends
much further to the east than in the north. An additional limiting factor here may be low winter
temperatures and the presence of common species in eastern Ukraine, such as S. uralensis, which
have a symbiotic interaction with S. sylvaticus.
A case of an ambiguous infl uence of climatic factors on the area is represented in subspecies
of house mice Mus musculus musculus / M. m. domesticus, with their hybrid zone passing through
the south of Denmark and further across Southern Germany, Western Czech Republic, Western
Austria and Northern Balkans [11-13]. Th is division of Europe into two climatic zones is provid-
ed: a southwestern zone with a temperate climate and Mediterranean as M. m. domesticus habitat,
and the northeastern part with a continental climate, belonging to Mus musculus musculus range.
Intriguingly, these are synanthropic mice, which deal with adverse climatic factors by hibernating
in buildings. Th is suggests that, as with the wood mouse, indirect circumstances play a decisive
role. Apparently, the type of vegetation relevant in the summer, when house mice leave their hu-
man settlements, is signifi cant.
Conclusion. As a result of the research, the questions posed in the work can be answered
as follows. 1. Th e eastern limit of the range of S. sylvaticus is determined by bioclimatic factors.
2. Th eir eff ect is indirect and is linked to the formation of a certain type of vegetation. Th e factor
that directly limits the expansion of the range to the east may be only the degree of continental-
ity of the climate, which is infl uenced by low winter temperatures. 3. Competitive interactions
with other species of mice may be present, but only at the southern border of the eastern limit of
the range and are of secondary importance as a factor in restraining the distribution of the wood
mouse to the East.
71ISSN 1025-6415. Допов. Нац. акад. наук Укр. 2023. № 5
Ecological niche modeling and factors determining the range of the wood mouse Sylvaemus sylvaticus (Linnaeus, 1758)
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Received 15.02.2023
72 ISSN 1025-6415. Dopov. Nac. akad. nauk Ukr. 2023. No 5
S.V. Mezhzherin, V.M. Tytar, V.O. Tereshchenko
С.В. Межжерін, https://orcid.org/0000-0003-2905-5235
В.М. Титар, https://orcid.org/0000-0002-0864-2548
В.О. Терещенко, https://orcid.org/0000-0001-6671-0265
Інститут зоології ім. І.І. Шмальгаузена НАН України, Київ
E-mail: smezhzherin@gmail.com
МОДЕЛЮВАННЯ ЕКОЛОГІЧНОЇ НІШІ І ФАКТОРИ, ЩО ВИЗНАЧАЮТЬ АРЕАЛ
ЛІСОВОЇ МИШІ SYLVAEMUS SYLVATICUS (LINNAEUS, 1758)
Моделювання екологічної ніші лісової миші Sylvaemus sylvaticus методом машинного навчання в просторі
35 біокліматичних факторів показало, що найсприятливішими для виду є абіотичні умови Західної Євро-
пи, де зосереджені головні ядра ареалу і ймовірність знахідок у придатних ландшафтах сягає 60 %. Біоклі-
матичні обставини Східної Європи є песимальними: зона перебування виду значно фрагментована, ядра
відсутні, вірогідність стрівальності не перевищує 40 %, найчастіше менш ніж 10 процентилів. Очевидно,
що серед біокліматичних чинників головне значення має континентальність клімату, що опосередковано
впливає на розташування східної межі ареалу S. s. sylvaticus і формування певних типів рослинності. Без-
посередня дія окремих факторів і міжвидові відносини можливі і мають місце, певним чином обмежуючи
просування виду на схід, однак лише на південному фасі східної межі ареалу.
Ключові слова: моделювання, Sylvaemus sylvaticus, екологічна ніша, фрагментація ареалу.
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