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DISTRIBUTION PATTERNS OF PTILIUM CRISTA-CASTRENSIS (BRYOPHYTA, HYPNACEAE) IN THE EAST EUROPEAN PLAIN AND EASTERN FENNOSCANDIA
Sergey Yu. Popov*, Yulia A. Makukha
Lomonosov Moscow State University, Russia *e-mail: s_yu_popov@rambler.ru
Received: 04.06.2018. Revised: 12.01.2019. Accepted: 15.01.2019.
Ptilium crista-castrensis is one of the most common moss species in the forest zone. It is dominant in the moss layer of the blueberry and cowberry forests. At least with low abundance, it occurs almost in every forest type and mires. In addition, it is a component of the moss layer in tundra. We compiled the published annotated lists of regional bryophyte floras into a generalised dataset. We then created a map of the P. crista-castrensis model range using the kriging method. We included data from 179 locations, with data points in 39 Protected Areas. We determined climatic preferences of the species by comparing the species occurrence and climatic factors on locations. The comparison of the created model map with the map of vegetation zones allowed us to analyse the spatial distribution of P. crista-castrensis. We could demonstrate a sharp decrease of P. crista-castrensis abundance at the borders of the forest and steppe zones. By the range boundaries, this moss is a rarely occurring species, where it grows exclusively in limited pine forest patches. This species is completely lacking in the steppe zone. Its abundance has maximal values in the northern taiga subzone considered as the climatic optimum of P. crista-castrensis. In the forest zone, the occurrence of P. crista-castrensis varies from sporadically to commonly. At the northern and southern borders of the forest zone, the species occurrence is characterised as rare. A too low average temperature and precipitation are unfavourable for P. crista-castrensis in the north of the study area. At the same time, a too high temperature and too low precipitation and humidity have the same effect in the south of the study area.
Key words: biogeography, distribution range, geostatistical techniques, species' climatic optimum, species' occurrence
Ptilium crista-castrensis (Hedw.) De Not. is a feather moss. It is a widely distributed moss in boreal ecosystems. This species grows together with other bryophyte species of boreal spruce forests, like Pleurozium schreberi (Brid.) Mitt. and Hy-locomium splendens (Hedw.) Schimp. (Bonan & Shugart, 1989; Esseen et al., 1997). Ptilium crista-castrensis occurs predominantly in the forest zone, rarer in tundra. Spruce forests of the Myrtillus-type and forested mires of the taiga zone are the most typical plant communities with P. crista-castrensis participation. In Europe, it occurs predominantly in spruce forests and rarely in heathlands (association Vaccinio-Callunetum vulgaris) (Dierssen, 2001). This moss inhabits frequently the forest soil, along the tree trunk bases, on rock-covered cliffs and boulders, in meadows. To the south of the taiga zone this moss is considered as a rare species (Kurnaev, 1980; Ignatov & Ignatova, 2004; Belyaeva & Neshataeva, 2017). Ptilium crista-castrensis can easily be identified in the field. Its ecology is very well known.
The geographic distribution of Ptilium crista-castrensis is also well known. This species is dis-
tributed in Western and Central Europe, European Russia, Caucasus, Urals, and in the Far East (Ig-natov & Ignatova, 2004). There are many recent publications on regional bryophyte floras. It allows us to generalise these data and identify the range zone of P crista-castrensis. The development of GIS-based spatial analysis methods has enabled attaining this goal. Moreover, modern GIS approaches provide the tools for studying climatic and other biogeographic patterns of species distribution (Mateo et al., 2013).
The approach to the current study of the P crista-castrensis distribution is based on Geostatis-tical techniques. In this paper, we aimed 1) to close the gap in the P. crista-castrensis biogeography in the East European Plain and Eastern Fennoscandia (EEPEF); 2) to visualise its distribution range; 3) to elucidate the influence of climatic factors on the species spatial distribution.
Material and Methods
We have analysed the published annotated lists of bryophyte floras of different regions (European Russia, Baltic countries, Ukraine,
Belarus, Moldova) to study the distribution of Ptilium crista-castrensis (Fig. 1). Some dots have been chosen outside the area of interest (e.g. Romania, Poland, Kazakhstan, Caucasus, the eastern mountainside of the Urals) to correct possible errors of extrapolation at the area boundaries (Demyanov & Savelyeva, 2010; Popov, 2016). We applied the basic principles of area modelling, using geostatistics methods and analysing the distribution range of the moss species by using the kriging method published previously (Popov, 2017).
After generalisation of all available data, we evaluated the Ptilium crista-castrensis occurrence using the following six-point scale of species occurrence: 0 - absent (0 records), 1 - very rare (1-2 records), 2 - rare (3-7 records), 3 - sporadic (more than seven records, but not everywhere), 4 - frequent (common species, but sometimes not found in suitable phytocoenoses), 5 - common (common and coenotically active species within the study area). According to this scale, we created a continuous coverage map with a resolution of 10 km per 1 pixel using the kriging method according to Demyanov & Savelyeva (2010).
• study sites - boundary of the studied area boundaries of vegetation zones
Fig. 1. Vegetation zones with indicated locations used in this study: I - tundra; II -forest-tundra; III - northern taiga; IV - middle taiga; V - southern taiga; VI - mixed forests; VII - broad-leaved forests; VIII - forest-steppe; IX - steppe; X - semi-desert; XI - desert. Boundaries of vegetation zones are given according to Ahti et al. (1968) and Kurnaev (1973).
In total, we used data from 179 locations to create a continuous coverage map (Fig. 1), Of which 39 are in the following Protected Areas: Shichengsky Landscape Sanctuary, Basegi State Nature Reserve, Bolshaya Kokshaga State Nature Reserve, Bryansky Les State Nature Reserve, Caucasian State Nature Biosphere Reserve, Central Forest State Nature Biosphere Reserve, Darwin State Nature Reserve, Denezhkin Kamen State Nature Reserve, Kandalakshsa State Nature Reserve, Kerzhensky State Nature Reserve, Kivach State Nature Reserve, Kologrivsky Les State Nature Reserve, Kostomuksha State Nature Reserve, Lapland State Nature Reserve, Mordovia State Nature Reserve, Niznesvirsky State Nature Reserve, Oksky State Nature Reserve, Orenburg State Nature Reserve, Pasvik State Reserve, Pechora-Ilych State Nature Reserve, Pinega State Nature Reserve, Prisursky State Nature Reserve, Privolzhskaya Lesostep' State Nature Reserve, Shulgan-Tash State Nature Reserve, Teberda State Nature Reserve, Vishersky State Nature Reserve, Visimsky State Nature Biosphere Reserve, Vitim State Nature Reserve, Volzhsko-Kamsky State Nature Biosphere Reserve, Voronezh State Nature Reserve, Zhiguli State Nature Biosphere Reserve, Iremel Nature Park, Kulikovo Pole Nature-Historical Reserve, Losiny Ostrov National Park, Narew National Park, Russky Sever National Park, National Park «Smolny», Vodlozersky National Park, Yugyd Va National Park. References to published sources with annotated species lists for 179 locations are listed in Popov (2018). In this paper, we could add two more locations (Silae-va et al., 2011; Chernyadyeva et al., 2017).
The verification of the continuous coverage was performed using a cross-validation method with SAGA GIS software. In geostatistics, the index of cross-validation quality is the coefficient of determination (R2) (Demyanov & Savelyeva, 2010). In this study, this indicator value was 0.912.
In our study, we also used continuous coverage of climatic factors. We used the dataset proposed by the authors of the WORLDCLIM program (BIOCLIM, 2009). In total, we used 23 climatic variables, including monthly temperature (seven variables), annual temperature (one variable), annual precipitation (one variable), monthly precipitation (seven variables), and relative humidity (seven variables), extracted for April - October. For our study, we selected only months of the vegetation season. The coverage of climatic factors and species occurrence were combined into a single spatial database, which has been transformed into the table consisting of 24 variables (23 climatic factors and one species' occurrence) and 49 557 cases (number of pixels). A correlation analysis was per-
formed using the Statistica 6.0 software. The coverages were created and verified using SAGA software. The intersection of vector layers and areas' calculation were performed using the ArcGIS software. All these techniques were explained in Popov (2017).
Results and Discussion
Fig. 2 shows the model map of the P. crista-castrensis' distribution range. Evidently, the area of the common species occurrence («com») coincides approximately with the northern taiga subzone. The occurrence level of P. crista-castrensis decreased to the south and to the north of this zone. In general, the frequent occurrence area coincides with the taiga zone. This is not surprising, because P. crista-castrensis is one of the dominant species in coniferous forests (Esseen et al., 1997; Ignatov & Ignatova, 2004). This species is rare in the tundra zone (e.g. in the Kola Peninsula). Ptilium crista-castrensis is also present in the mainland tundra, having, however, a low abundance. The species completely disappears by the northern limit of the study area. For example, there are only a few locations with P. crista-castren-sis in the Yamal Peninsula, and it is completely absent on Novaya Zemlya (Afonina & Chernyadyeva, 1995; Chernyadyeva, 2001). Outside the study area, the distribution of spruce forests coincides with the P. crista-castrensis distribution in Scandinavia and in the mountains of continental Europe (Soder-strom, 1998; Dierssen, 2001; GBIF, 2018). This fact is in good agreement with its higher occurrence in the northern taiga of Russia (Fig. 2).
In the southern part of the study area, P. crista-castrensis has a sporadic occurrence in the vegetation zones of mixed forests and broad-leaved forests. The species is rare in the forest-steppe zone. Further, P. crista-castrensis disappears in the vegetation cover in the northern steppe subzone (Fig. 2). The species inhabits sporadically distributed and limited pine plantations in the steppe and in forest-steppe vegetation zones (Gapon, 1997; Popova, 2002).
The occurrence of P. crista-castrensis depends upon the autumn - summer precipitation and the relative air humidity (Table 1). Annual precipitation produces a moderate effect. The temperature factor looks to be the most important for P. crista-castren-sis. It includes both the annual temperature and the monthly temperature during the vegetation season (Table 1). We could state that P. crista-castrensis's optimum is in the northern taiga (Fig. 2), taking into account that according to biogeographic principles (Grinnell, 1917), the climatic optimum of a species is in the area where it has the maximal occurrence.
Fig. 2. Model range of Ptilium crista-castrensis. Black lines indicate the boundaries of vegetation zones. Occurrence zones: abs - species is absent; vr - species is very rare; r - species is rare; sp - species is sporadic; fr - species is frequent; com - species is common. The letters (n) and (s) placed after the abbreviation of the occurrence zones indicate respectively the relative north and south of the study area.
Table 1. The Spearman correlation coefficient between values of climatic factors and species occurrence. Values of r > 0.5 in absolute value are highlighted in bold. All values are statistically significant (p < 0.05)
№
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Climatic factor
amt
pr04
pr05
pr06
pr07
pr08
pr09
pr10
pr_a
reh04
reh05
reh06
reh07
reh08
reh09
reh10
tm04
tm05
tm06
tm07
tm08
tm09
tm10
Correlation coefficient
-0.82
-0.02
0.06
0.36
0.13
0.72
0.79
0.75
0.40
0.38
0.45
0.36
0.53
0.75
0.80
0.84
-0.86
-0.87
-0.86
-0.82
-0.87
-0.87
-0.79
Note: Climatic factors: amt - annual amount of precipitation; pr01 - pr12 - monthly amount of precipitation in January - December; pr_a - average of annual precipitation; reh4 - reh10 - relative humidity in April - October; tm04 - tm10 - monthly average temperature in April - October.
1
2
3
4
5
6
7
8
9
Table 2 shows the climatic optimum and pes-simum values for P. crista-castrensis in the study area. According to Table 2 (column «com»), the species requires an average annual temperature of about -1°C and a monthly temperature of+11°C to +15°C during the summer period. In other words, apparently P. crista-castrensis is better adapted to areas with cool summers. However, this species requires a relatively high humidity (ca. 80%) in the summer period (Table 2). The climatic conditions of the northern taiga meet these requirements to the greatest extent (Kurnaev, 1973).
A low temperature and precipitation are the main unfavourable conditions for Ptilium crista-castrensis in the north. On the contrary, a high temperature, low precipitation and low humidity restrict its spread to the south (Table 2). If the average annual temperature exceeds +7°C and the annual rainfall is less than 500 mm, P. cris-ta-castrensis is disappearing. Thus, the distribution pattern of P. crista-castrensis corresponds
to Shelford's rule (Tolmachev, 1974; Vtorov & Drozdov, 2001). The conducted analysis showed the maximal and minimal values of the limiting factors (Table 2). These are considered to be of the greatest impact on the distribution of P. cris-ta-castrensis. In Table 1, these factors have correlation coefficients over 0.5 (absolute values).
The model distribution map clearly showed that the major part of the EEPEF area is located in the southern parts of the frequent and sporadic occurrence zones (Table 3). The zone of common occurrence occupies only 15.3% of the total area (Table 3). The zones of rare and very rare occurrence (both in the north and in south parts of the study area) have very small percentage values of the study area (Table 3). Accordingly, the species is quite widespread in the central part of the study area. Then, it gradually disappears at the borders both in the north and in the south. The absence zone covers 17.3% of the EEPEF area (Table 3).
Table 2. Average values of climatic factors in the occurrence zones of Ptilium crista-castrensis
Occurrence zones Average
Climatic factor abs vr(s) r(s) sp(s) fr(s) com fr(n) sp(n) r(n) vr(n)
amt 8.7 6.3 5.6 4.9 2.0 -0.9 -2.3 -3.7 -4.2 -6.2 3.7
pr04 31.7 35.7 38.5 38.6 37.3 31.9 26.4 24.9 22.8 19.9 35.1
pr05 40.7 44.0 48.3 50.1 48.4 41.8 35.0 33.8 30.7 28.4 45.3
pr06 50.5 59.6 65.3 69.0 64.8 57.4 48.6 46.2 41.6 36.3 60.9
pr07 46.2 62.8 75.0 79.9 78.5 68.0 59.3 55.6 50.6 46.4 69.1
pr08 37.4 48.9 59.0 68.6 74.2 71.7 63.8 61.6 58.2 54.0 62.6
pr09 33.9 42.0 48.8 58.5 64.5 62.2 57.6 56.3 54.7 54.2 54.3
pr10 28.7 38.2 43.9 51.9 60.7 54.9 50.4 47.6 46.1 41.7 48.7
pr_a 436.6 529.9 577.6 612.3 626.4 561.7 485.1 453.7 421.1 357.6 560.5
reh04 64.8 65.8 67.9 70.7 71.8 76.2 83.2 87.2 89.9 94.8 71.2
reh05 57.6 56.6 58.9 64.0 63.7 68.7 75.5 78.2 80.6 83.4 63.5
reh06 58.8 60.8 64.2 68.6 66.9 67.5 71.9 73.2 75.2 76.2 65.8
reh07 57.8 62.3 66.9 72.0 71.8 71.8 74.5 74.8 76.1 75.8 68.7
reh08 57.6 61.6 66.6 73.7 76.9 78.7 80.7 81.3 82.4 82.8 71.6
reh09 62.7 65.7 70.6 77.7 81.8 83.6 84.5 84.9 85.9 86.5 76.1
reh10 73.7 76.0 79.3 83.5 87.2 89.4 90.6 92.5 93.5 96.7 83.3
tm04 9.8 8.0 7.0 5.5 2.2 -2.1 -5.0 -7.2 -8.3 -11.2 3.9
tm05 16.6 15.2 14.4 12.8 9.5 4.8 1.6 -0.2 -1.2 -3.3 11.0
tm06 20.5 18.7 17.8 16.4 14.5 11.4 8.6 7.4 6.5 5.4 15.6
tm07 22.8 20.5 19.3 18.0 17.0 14.8 13.1 12.5 11.8 11.3 18.0
tm08 21.6 19.3 18.1 16.6 14.6 12.1 10.6 9.9 9.4 8.6 16.2
tm09 16.3 14.0 12.8 11.6 9.1 6.8 5.8 5.2 4.9 4.0 11.0
tm10 8.8 6.6 5.9 5.4 2.4 -0.2 -1.4 -2.6 -2.9 -5.0 4.1
Note: see Fig. 2 for abbreviations of the occurrence zones.
Table 3. Occurrence zones of Ptilium crista-castrensis by vegetation zones, km2
Vegetation zone abs vr r sp fr com Total
Tundra - 3418.4 92822.8 65335.1 16161.8 14020.1 191758.2
Forest-tundra - - 126.1 28440.5 59630.6 13894.7 102091.9
Northern taiga - - - - 31146.4 519581.3 550727.7
Middle taiga - - - - 545491.0 202528.6 748019.6
Southern taiga - - - 31060.0 502545.3 6368.5 539973.8
Mixed forests - - - 742694.8 71629.5 - 814324.3
Deciduous forests 8027.5 27955.9 92921.4 382382.4 5863.3 - 517150.5
Forest-steppe 39467.3 103734.0 307901.5 71600.1 - - 522702.9
Steppe 549057.8 145742.1 11809.6 1973.2 - - 708582.7
Semi-desert 204708.3 51.7 - - - - 204760
Desert 54863.8 - - - - - 54863.8
Total, km2 856124.7 280902.2 505581.5 1323486.0 1232468.1 756393.1 4954955.4
Total, % 17.3 5.7 10.2 26.7 24.9 15.3 100
Conclusions
The abundance of Ptilium crista-castrensis is correlated with the occurrence of coniferous forests (taiga zone) in such a large area as EEPEF, where 11 vegetation zones occur (Fig. 1). The species' occurrence is maximal in the northern taiga. This region is characterised by a specific climatic optimum for P crista-castrensis, considered as an average annual temperature of about -1°C and an average monthly humidity of about 80%. In the forest zone, the P. crista-castrensis occurrence varied from sporadically to commonly. This species is rare both to the north and to the south of the forest zone.
References
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РАСПРОСТРАНЕНИЕ PTILIUM CRISTA-CASTRENSIS НА ТЕРРИТОРИИ ВОСТОЧНО-ЕВРОПЕЙСКОЙ РАВНИНЫ И ВОСТОЧНОЙ ФЕННОСКАНДИИ
С. Ю. Попов*, Ю. А. Макуха
Московский государственный университет имени М.В. Ломоносова, Россия *e-mail: s_yu_popov@rambler.ru
Ptilium crista-castrensis является одним из наиболее распространенных видов в лесной зоне. Он является обычным видом мохового яруса в черничных лесах. В небольших количествах он может произрастать почти во всех типах леса, даже на болотах и лугах. Он также является компонентом мохового яруса в сообществах тундры. На основе сведения литературных источников с аннотированными списками локальных бриофлор в единую географическую базу данных, построена карта модельного ареала P. crista-castrensis методом кригинга. Всего было использовано 179 точек, среди которых 39 относится к заповедникам и национальным паркам. На основании наложения непрерывных покрытий встречаемости вида и климатических факторов установлены климатические предпочтения вида. Наложение модельной карты на карту растительных зон позволило провести анализ его пространственного распространения. На границе между лесной и степной зонами встречаемость P. crista-castrensis резко падает. На юге он становится довольно редким видом, растущим преимущественно в небольших массивах сосновых насаждений. В открытой степи этот вид исчезает. Максимум его встречаемости приходится на подзону Северной тайги. Здесь находится климатический оптимум вида. В численных значениях климатический оптимум соответствует годовой температуре около -1°C и влажности воздуха около 80%. В лесной зоне P. crista-castrensis имеет встречаемость от спорадической до широкой. Наихудшими климатическими условиями для P. crista-castrensis на севере являются низкие температуры и низкое количество осадков, несмотря на высокую влажность воздуха. На юге он перестает расти из-за высоких летних температур и низкого количества осадков.
Ключевые слова: ареал, биогеография, встречаемость вида, климатический оптимум вида, методы геостатистики
