The epiphytic bryophyte succession of Buxus sempervirens forests in the Fırtına Valley, Rize (North Türkiye)

Abstract

In this study, the epiphytic bryophyte succession of the Buxus sempervirens L. forests in Fırtına Valley (Çamlıhemşin-Rize, North Türkiye), one of the nine biodiversity hotspots in Türkiye, were investigated. For this purpose, a total of 60 sampling plots were taken from the live trunks of the B. sempervirens trees of different ages. Twenty-nine epiphytic bryophyte species were determined (24 mosses and 5 liverworts) within the sample plots. Also, six different life form types and four different habitat affinity categories were determined. The mat type life form is in the first place with 34.4% whereas the and cortico-saxicolous species are the most common with 51.7%. Two-Way Indicator Species Analysis (TWINSPAN) classified the epiphytic bryophyte communities on the trunks of the B. sempervirens at the second level into two main clusters (A and B) and three sub-clusters (A1, B1 and, B2). Detrended Correspondence Analysis (DCA) axis 1 was interpreted as gradient along the height of the epiphytic habitat (from the lower base to the upper zone) on trunks and the DCA axis 2 was interpreted as gradient of moisture (from mesic to xeric). Exsertotheca crispa (Hedw.) S.Olsson, Enroth & D.Quandt was the species with the highest Index of Ecological Significance (IES) value on the lower bases of the aged trees. Species diversity and epiphytic cover in the upper zones were lower than in the basal and middle zones in the study area. While Metzgeria furcata, (L.) Corda, Oxyrrhynchium hians (Hedw.) Loeske, Plagiothecium nemorale (Mitt.) A.Jaeger, and Radula lindenbergiana Gottsche ex C.Hartm were only found on old trees, Ctenidium molluscum (Hedw.) Mitt. and Pseudoleskeella nervosa (Brid.) Nyholm were only found on middle-aged trees.

Full text

10 ACTA BOT. CROAT. 82 (1), 2023
Acta Bot. Croat. 82 (1), 10–19, 2023 CODEN: ABCRA 25
DOI: 10.37427/botcro-2023-002 ISSN 0365-0588
eISSN 1847-8476
e epiphytic bryophyte succession of Buxus
sempervirens forests in the Fırtına Valley, Rize
(North Türkiye)
Tülay Ezer1, Mevlüt Alataş2, Nevzat Batan3, Hüseyin Erata4*
1 Niğde Ömer Halisdemir University, Faculty of Architecture, Department of Landscape Architecture, 51100, Niğde, Türkiye
2 Munzur University, Tunceli Vocational School, 62000, Tunceli, Türkiye
3 Karadeniz Technical University, Faculty of Science, Department of Molecular Biology and Genetics, 61080, Trabzon, Türkiye
4 Gümüşhane University, Kürtün Vocational School, 29810, Gümüşhane, Türkiye
Abstract – In this study, the epiphytic bryophyte succession of the Buxus sempervirens L. forests in Fırtına Valley
(Çamlıhemşin-Rize, North Türkiye), one of the nine biodiversity hotspots in Türkiye, was investigated. For this pur-
pose, a total of 60 sampling plots were taken from the live trunks of B. sempervirens trees of dierent ages. Twenty-nine
epiphytic bryophyte species were determined (24 mosses and 5 liverworts) within the sample plots. Also, six dierent
life form types and four dierent habitat anity categories were determined. Among them, the mat type life form is in
rst place with 34.4% and the cortico-saxicolous species are the most common with 51.7%. Two-way indicator species
analysis (TWINSPAN) classied the epiphytic bryophyte communities on B. sempervirens trunks at the second level
into two main clusters (A and B) and three sub-clusters (A1, B1 and, B2). Detrended correspondence analysis (DCA)
axis 1 was interpreted as gradient along the height of the epiphytic habitat (from the lower base to the upper zone) on
trunks and the DCA axis 2 was interpreted as gradient of moisture (from mesic to xeric). Exsertotheca crispa (Hedw.) S.
Olsson, Enroth & D. Quandt was the species with the highest index of ecological signicance (IES) value on the lower
bases of the aged trees. Species diversity and epiphytic cover in the upper zones were lower than in the basal and middle
zones in the study area. While Metzgeria furcata, (L.) Corda, Oxyrrhynchium hians (Hedw.) Loeske, Plagiothecium
nemorale (Mitt.) A.Jaeger, and Radula lindenbergiana Gottsche ex C.Hartm were only found on old trees, Ctenidium
molluscum (Hedw.) Mitt. and Pseudoleskeella nervosa (Brid.) Nyholm were only found on middle-aged trees.
Keywords: community, index of ecological signicance, liverworts, mosses, ordination analyses
Introduction
Bryophytes, the pioneer plants of dierent substrate
types, are one of the most important component of forest
ecosystems (Longton 1992, Baldwin and Bradeld 2005,
Ezer 2017, Mellado-Mansilla et al. 2017). Bryophytes are
poikilohydric organisms whose their moisture content rap-
idly equilibrating with environmental conditions (Green
and Lange 1994). Therefore, they are highly sensitive to en-
vironmental factors (Schoeld 2001). In particular, abiotic
ecologica l factors such as humidit y directly or indirectly af-
fect the colonization and the distribution of bryophytes in
the epiphytic habitat (Mazimpaka and Lara 1995, Schoeld
2001, Mishler 2003, Mazimpaka et al. 2009). In addition to
environmental drought, phorophyte-type, physical and
chemical properties of bark characteristics such as rugosity,
water retention capacity, bark pH, and dust deposition are
also important for the spatial distribution of bryophytes on
epiphy tic habitats (Lara and Mazimpaka 1998, Mazimpaka
et al. 2010, Ezer 2017).
Some studies on the succession of epiphytic bryophyte
communities have revealed that the succession gradient of
epiphytes is highly complex due to changes in positive and
negative interactions among species within epiphytic com-
munities as trees age (Mazimpaka et al. 2010, Ódor et al.
2013, Bargali et al. 2014, Ezer 2017, Ezer et al. 2019).
Although phytosociological studies on epiphytic bryo-
phytes in Türkiye have made progress in the last decade
* Corresponding author e-mail: huseyin__erata@hotmail.com

SUCCESSION OF EPIPHYTIC BRYOPHYTES IN TÜRKIYE
ACTA BOT. CROAT. 82 (1), 2023 11
(Alataş et al. 2017, 2021, Alataş 2018, Can Gözcü et al. 2018),
few studies have been done on the spatial distributions and
community composition of epiphytic bryophytes in succes-
sional stages (Ezer and Kara 2013, Ezer 2017, Ezer et al.
2019).
The present study focuses on the vertical distribution
patterns of epiphytic bryophytes and community com-
position in the successional stages on trunks of Buxus
sempervirens trees in Fırtına Valley. This study aimed to
reveal the successional trends of bryophyte communities on
the epiphytic habitats of box wood trees and to contribute to
bryo-ecological studies in Türkiye.
e study area
The Çamlıhemşin (Rize, Türkiye) District, in which the
study area is located, is surrounded by Pazar and Ardeşen
to the north, Çayeli, Hemşin and İkizdere districts to the
west, and Artvin Province Yusufeli district to the east. The
distance of the district to Rize city center is 62 km. The B.
sempervirens forests, located in the Fırt ına Valley and with-
in the boundaries of the Çamlıhemşin District, are located
within the A4 square according to the Henderson (1961)
grid-square system and are in the colchic zone of the Euro-
Siberian phytogeographic region (Anşin 1983, Fig. 1).
The Fırtına Valley, which exhibits a unique phytological
diversity, hosts many rare species due to the presence of all
the main habitats of the region. Buxus sempervirens, which
has a very wide distribution in the Euro-Siberian Phytogeo-
graphic Region, forms remarkable communities in the val-
ley (Kurdoğlu et al. 2004). Moreover, in 1999, the WWW
(World Wildlife Fund) identied Europe’s 100 forest areas
valuable in terms of biodiversity and in urgent need of pro-
tection (Myers et al. 2000). Nine of these areas, which are
called “Hot Spots of European Forests”, are located in Tür-
kiye (Satar and Güneş 2014). One of these nine hot spots is
the Fırtına Valley. Only one study on bryophytes has been
conducted in the valley so far (Abay et al. 2006).
Fırtına Valley, like the whole of the Eastern Black Sea
Region, is an area very open to natural disasters such as
landslides, rockfalls and avalanches due to the very steep
slopes, rainy climatic conditions and, soil cover (Tunçel
1990).
Lithologically, there are units from almost all geological
times in the valley (Bayrakdar 2006). In Türkiye, not rich
in current glaciers, Kaçkar Mountains (3932 m) located to
the south of the Fırtına Valley are one of the regions where
current glaciers are found (Çiner 2003). The Kaçkar Moun-
tains, the Verçenik (Üçdoruk) Mountain, the Bulut Moun-
tains and, the Altınparmak Mountains are mountain rang-
es that limit the study area in the SW-NE direction. In the
valley, which is also very rich in terms of rivers, Fırtına
Stream is divided into branches at various degrees and
forms the smaller Tunca, Hala, Palovit, Haçivanak and
Hemşin creeks. These creeks also forked among themselves
and surround the valley like a net system.
Fig. 1. The Fırtına Valley where epiphytic bryophyte species were investigated (A), its location in Türkiye (B) and in the Henderson’s
grid system (1961) (C).

EZER T., ALATAŞ M., BATAN N., ERATA H.
12 ACTA BOT. CROAT. 82 (1), 2023
The forest formation, which starts at 20 0 m in the nort h
of the study area, is very rich in terms of under-forest ora,
depending on the oceanic and temperate climatic condi-
tions in all seasons. This forest formation, which is domi-
nated by broadleaf trees, loses its colchic feature with the
increase in altitude and gives way to mixed forests and then
to coniferous forests (Özçağlar et al. 2006). The vegetation
in the Fırtına Valley shows a distinct dierence from other
valleys, especially with its particular forest formations and
ora richness. There are basically three types of forest for-
mation such as alluvial forests, hardwood forests found in
river oodplains and regularly ooded for a portion of the
growing season; boxwood forests; and old growth forest for-
mations in the Fırtına Valley (Kurdoğlu et al. 2004).
The B. sempervirens forests spread along the Fırtına
Stream and its tributaries, between 200-1500 meters of the
study area. Although these forests are seen along the
streams, they are also found in large groups on the valley
slopes. These forests are widely found between 900-1300
meters along the Çamlıhemşin-Meydan road, in Şimşirlik
place, within the Gito Forests and Palovit Valley.
The study area generally has a temperate Oceanic cli-
mate. The annual average precipitation is 2192 mm and, the
annual average temperature is 8.3 ºC in Çamlıhemşin. The
hottest month of the year is August and the coldest is Janu-
ary. The absence of a dry season indicates that the Fırtına
Valley is under the inuence of an Oceanic climate (Akman
2011).
Materials and methods
Field sampling procedures
The epiphytic bryophyte specimens were collected from
the living trunks of B. sempervirens in Fırtına Valley duri ng
eld studies in 2020. The locality details are given in Tab. 1.
A total of 60 sampling plots were taken from the trunks of
20 boxwood trees of dierent ages in the valley. Spatial dis-
tributions and community structures of epiphytic bryo-
phytes in successional stages on boxwood trees were inves-
tigated according to tree diameter at breast height (dbh) and
tree age. The trees were divided into three age classes using
an indirect method as young (dbh 20-35 cm, 21 plots), mid-
dle-aged (dbh 40-50 cm, 18 plots), and aged trees (dbh 60-80
cm, 21 plots). Each boxwood tree was divided into the low-
er base zone (0 - 40 cm from the ground), the middle zone
(40 - 120 cm), and the upper zone (120 - 180 cm) as proposed
by Moe and Botnen (2000).
Sampling plots from tree zones were dened by 20 × 20
cm
2
, determined according to species diversity on the living
trunks of B. sempervirens. In the present study the branch-
es of the boxwood trees did not provide sample plots, only
the trunks. The percentage cover of the species within the
each sample plot was visua lly estimated and ecological data
such as humidity, aspect and light of the epiphytic habitat
were recorded.
The nomenclature of the epiphytic bryophyte species de-
termined within the sample plots follows Ros et al. (2013)
and Hodgetts et al. (2020) (Tab. 2). Habitat anity types of
bryophytes were established following Mazimpaka and Lara
(1995) and Draper et al. (2003). Life forms of the taxa were
determined according to Mägdefrau (1982). Voucher speci-
mens were deposited in the herbarium of Niğde Ömer Hal-
isdemir University.
Data analysis
The relative frequency of each taxon in the sample plots
was determined by the index of ecological signicance (IES)
described by Lara and Mazimpaka (1998), Albertos et al.
(2001) and Mazimpaka et al. (2009). The formula used to
calculate the IES values for each taxon is as follows:
IES = F (1 + C)
 C=Σci/x
where F is the relative frequency (100 x / n), and C is the
cover of the taxon (Σ ci / x), while x represents the number
of sample plots containing the taxon, n is total number of
sample plots, and ci is cover class assigned to the taxon in
each sample plot.
Cover classes of taxa were established using the six-point
Lara and Mazimpa ka (1998) scale: [0.5 (< 1%), 1 (1-5%), 2 (6-
25%), 3 (26-50%), 4 (51-75%) and, 5 (76-100%)]. In addition,
the IES values were combined in the following abundance
classes: very scarce (< 25), scarce (26-50), moderately abun-
dant (51-100), abundant (101-200), and dominant (> 200).
Tab. 1. Sampling localities from where epiphytic bryophyte specimens were collected and their characteristics.
Localities Altitude (m) Date GPS coordinates
1Çat Valley, Doğa Village 1275 17.05.2020 N 40º51’48.95” E 40º55’58.08”
2Between Çatköy and Meydanköy 1231 17.05.2020 N 40º51’59.44” E 40º55’46.26”
3Meydanköy Exit 1176 17.05.2020 N 40º52’13.82” E 40º55’37.52”
4Meydanköy 1108 26.08.2020 N 40º52’48.29” E 40º55’43.37”
5Meydanköy Entrance 1061 26.08.2020 N 40º53’20.49” E 40º55’51.11”
6Meydanköy, Pul Place 1026 26.08.2020 N 40º53’40.83” E 40º56’31.71”
7Gito Plateau- Southeast slopes 1018 09.10.2020 N 40º54’18.08” E 40º56’52.30”
8Between Zilkale and Meydanköy 1004 09.10.2020 N 40º54’27.30” E 40º56’53.71”
9Zilkale Place 956 09.10.2020 N 40º54’47.61” E 40º56’52.38”

SUCCESSION OF EPIPHYTIC BRYOPHYTES IN TÜRKIYE
ACTA BOT. CROAT. 82 (1), 2023 13
study area (Tab. 2). Exsertotheca crispa (Hedw.) S.Olsson,
Enroth & D.Quandt is the most common species in the B.
sempervirens forests of Fırtına Valley.
Ecological results
Six dierent life forms were determined. Among them,
the mat li fe form prevailed (34.4%), followed by the weft life
form (24.1%). The dendroid life form was negligible (3.4%)
(Tab. 2). In addition, four dierent habitat anity categories
as cortico-saxicolous, indierent, customary epiphyte and
preferentially not corticolous were determined belonging to
the species. While cortico-saxicolous species were the most
common with 51.7% within the sample plots, the preferen-
tially not corticolous type habitat anity is least common
with one species (Tab. 2).
Lower base zone
The spatial distributions and community structures
analyses showed that a total of 11 species, all of them are
mosses, were found on the base zone of young boxwood trees
(dbh 20-35 cm). Among of the mosses E. crispa was the most
Here, TWINSPAN and DECORANA (Hill 1979, Seaby
and Henderson 2007) were used to explore the community
composition and spatial patterns of epiphytic bryophyte
communities and their relationship with the associated en-
vironmental factors of the epiphytic habitat. In this context,
TWINSPAN and DECORANA were applied to the matrix
of cover in 60 sample plots according to the computer pro-
gram CAP (Community Analysis Package-5) of Seaby and
Henderson (2007).
Results
Floristical results
Twenty-nine species belonging to 17 families and 25
genera were determined as a result of the identication of
362 specimens. Among them 24 are mosses (21 pleurocar-
pous and 3 acrocarpous), and 5 are liverworts. Neckeraceae
(6 species, 20.6%) and Brachytheciaceae (5 species, 17.2%),
both pleurocarpous moss fami lies, are the most species-rich
families found in epiphy tic habitats on boxwood trees in the
Tab. 2. List of epiphytic bryophyte species found on the Buxus sempervirens trees, their families and life form types and anity for
epiphytic habitats (Mägdefrau 1982, Mazimpaka and Lara 1995, Draper et al. 2003).
Species Family Life
form
Anity for
epiphytic habitats
Frequency
(%)
MOSSES
Alleniella besseri (Lobarz.) S.Olsson, Enroth & D.Quandt Neckeraceae fan Cortico-saxicolous 65
Alleniella complanata (Hedw.) S.Olsson, Enroth & D.Quandt Neckeraceae fan Cortico-saxicolous 76.6
Anomodon viticulosus (Hedw.) Hook. & Taylor Anomodontaceae tail Cortico-saxicolous 3.3
Brachythecium rutabulum (Hedw.) Schimp. Brachytheciaceae weft Preferantially not corticolous 5
Ctenidium molluscum (Hedw.) Mitt. Myuriaceae weft Indierent 3.3
Exsertotheca crispa (Hedw.) S.Olsson, Enroth & D.Quandt Neckeraceae fan Cortico-saxicolous 93.3
Fissidens serrulatus Müll.Hal. Fissidentaceae fan Indierent 5
Homalia trichomanoides (Hedw.) Brid. Neckeraceae fan Cortico-saxicolous 16.6
Hypnum cupressiforme Hedw. Hypnaceae weft Indierent 20
Isothecium alopecuroides (Lam. ex Dubois) Isov. Lembophyllaceae mat Cortico-saxicolous 28.3
Leucodon sciuroides (Hedw.) Schwägr. Leucodontaceae tail Cortico-saxicolous 33.3
Orthotrichum pumilum Sw. ex anon. Orthotrichaceae cushion Customary epiphyte 30
Oxyrrhynchium hians (Hedw.) Loeske Brachytheciaceae weft Indierent 1.6
Palamocladium euchloron (Müll.Hal.) Wijk & Margad. Brachytheciaceae tail Cortico-saxicolous 15
Plagiothecium nemorale (Mitt.) A. Jaeger Plagiotheciaceae mat Indierent 1.6
Pseudanomodon attenuatus (Hedw.) Ignatov & Fedosov Neckeraceae mat Cortico-saxicolous 31.6
Pseudoamblystegium subtile (Hedw.) Vanderp. & Hedenäs Amblystegiaceae weft Cortico-saxicolous 6.6
Pseudoleskeella nervosa (Brid.) Nyholm Pseudoleskeellaceae mat Cortico-saxicolous 1.6
Pterigynandrum liforme Hedw. Pterigynandraceae tail Cortico-saxicolous 6.6
Sciuro-hypnum otowianum (Sendtn.) Ignatov & Huttunen Brachytheciaceae mat Cortico-saxicolous 35
Sciuro-hypnum populeum (Hedw.) Ignatov & Huttunen Brachytheciaceae weft Indierent 3.3
amnobryum alopecurum (Hedw.) Gangulee Neckeraceae dendroid Indierent 5
uidium delicatulum (Hedw.) Schimp. Thuidiaceae weft Indierent 6.6
Ulota crispa (Hedw.) Brid. Orthotrichaceae cushion Customary epiphyte 31.6
LIVERWORTS
Frullania dilatata (L.) Dumort Frullaniaceae mat Cortico-saxicolous 10
Frullania tamarisci (L.) Dumort. Frullaniaceae mat Cortico-saxicolous 8.3
Metzgeria furcata (L.) Dumort. Metzgeriaceae mat Indierent 5
Radula complanata (L.) Dumort Radulaceae mat Customary epiphyte 51.6
Radula lindenbergiana Gottsche ex C. Hartm. Radulaceae mat Customary epiphyte 1.6

EZER T., ALATAŞ M., BATAN N., ERATA H.
14 ACTA BOT. CROAT. 82 (1), 2023
frequent and the most dominantm with the highest IES val-
ue (271). Isothecium alopecuroides was co-dominant with 214
IES values. Homalia trichomanoides and Sciuro-hypnum
otowianum were abundant species with 171 and 143 IES
values on the lower base of trunks of B. sempervirens.
Brachythecium rutabulum, S. populeum and amnobryum
alopecurum which have the lowest IES values (43) were
scarce on the lower base (Tab. 3). Whereas the weft life form
was the most dominant (36.3%) in the base zone, the life
forms mat and fan were co-dominant (27.2%). Also, the cor-
tico-saxicolous type habitat anity of the species was the
most common with 54.5% on the base zones of young box-
wood trees.
Fifteen mosses were collected from the lower base of the
middle-aged B. sempervirens (dbh 40-50 cm). Among them,
E. crispa (266 IES value) and H. trichomanoides (216 IES
value) were the two most dominant species. Sciuro-hypnum
otowianum was the most abundant on the base zones of
the middle-aged trees with the 183 IES value. Anomodon
viticulosus, Pseudoleskeella nervosa and S. populeum were
scarce with the lowest IES values (33). Moreover, Ctenidium
molluscum and P. nervosa were only found on the lower
bases of middle-aged trees (Tab. 3). While the fan life form
is the most dominant (33.3%), the life forms mat and weft
were co-dominant (26.6%) on the lower base of middle-aged
boxwood trees. Cortico-saxicolous type anity of epiphyt-
ic habitats (60%) were the most common on the basal zone
of middle-aged trees.
Twelve species (11 mosses, one liverwort) were deter-
mined on the lower bases of aged boxwood trees (dbh 60-80
cm). Exsertotheca crispa was still the most frequent and the
most dominant with the highest IES value (371). This value
is also the highest among all tree-size groups (Tab. 3). While,
Hypnum cupressiforme, I. alopecuroides, Pseudanomodon
attenuates and, S. otowianum were abundant, Fissidens
serrulatus, Oxyrrhynchium hians and, Plagiothecium
Tab. 3. Index of ecological signicance (IES) values in each tree-size groups according to tree diameter at breast height (dbh) and tree
age at lower base zone, middle zone and upper zone.
Species Young trees
(dbh 20–35 cm)
Middle-aged trees
(dbh 40–50 cm)
Aged trees
(dbh 60–80 cm)
Lower
base
Middle
zone
Upper
zone
Lower
base
Middle
zone
Upper
zone
Lower
base
Middle
zone
Upper
zone
MOSSES
Alleniella besseri –328 271 50 316 233 –314 300
Alleniella complanata 100 328 285 115 266 250 71 342 342
Anomodon viticulosus –43 –33 –––––
Brachythecium rutabulum 43 – – 50 – – 57 – –
Ctenidium molluscum –––67 –––––
Exsertotheca crispa 271 314 357 266 216 366 371 314 328
Fissidens serrulatus –––67 – – 28 – –
Homalia trichomanoides 171 – – 216 50 –57 – –
Hypnum cupressiforme 71 71 ––––128 114 28
Isothecium alopecuroides 214 43 –130 33 33 171 85 –
Leucodon sciuroides – – 257 –50 300 – – 300
Orthotrichum pumilum – – 200 – – 100 –28 200
Oxyrrhynchium hians ––––––28 – –
Palamacladium euchloron –128 – – 150 – – 85 28
Plagiothecium nemorale ––––––28 – –
Pseudanomodon attenuatus 85 128 71 100 150 33 114 57 –
Pseudoamblystegium subtile –28 – – 33 – – 43 28
Pseudoleskeella nervosa –––33 –––––
Pterygynandrum liforme –71 –––––57 –
Sciuro-hypnum otowianum 143 86 28 183 116 33 128 28 –
Sciuro-hypnum populeum 43 – – 33 ––––
amnobryum alopecurum 43 – – 83 –––––
uidium delicatulum 85 – – 100 –––––
Ulota crispa – – 200 – – 167 – – 200
LIVERWORTS
Frullania dilatata – – 57 – – 33 –28 57
Frullania tamarisci ––––33 – – 86 43
Metzgeria furcata –––––––57 28
Radula complanata –200 114 –133 100 71 171 143
Radula lindenbergiana –––––––28 –

SUCCESSION OF EPIPHYTIC BRYOPHYTES IN TÜRKIYE
ACTA BOT. CROAT. 82 (1), 2023 15
nemorale usually not epiphy tic, were scarce with the lowest
IES values (28). The mat life form was the most dominant
with 41.6% and cortico-saxicolous species (50%) were still
the most common on base zones of aged B. sempervirens.
Middle zone
Twelve species (11 mosses, one liverwort) were deter-
mined on the middle zones of the young boxwood trees.
Alleniella besseri and A. complanata were the most frequent
and the most dominant with the highest IES values (328).
While E. crispa was co-dominant with 314 IES values, Radula
complanata (20 0), P. attenuatus (128) and, Palamacladium
euchloron (128) were abundant on the middle parts of the
trunks of the young B. sempervirens (Tab. 3). The mat life
form wit h 33.3% and the cor tico-saxicolous type habitat af-
nity with 83.3% were still most dominant on the middle
parts of trunks of young boxwood trees.
Twelve species were collected from the middle zones of
the middle-aged boxwood trees. Alleniella besseri was the
most dominant with the highest IES values (316). Alleniella
complanata and E. crispa were co-dominant, both with
266 IES values. Frullania tamarisci, I. alopecuroides and,
Pseudoamblystegium subtile were scarce with the same IES
values (33) on the middle zones of middle-aged trees (Tab.
3). The life form mat was the most dominant with 41.6% and
cortico-saxicolous species were conspicuously the most
common with the rate of 91.6% on middle parts of the
middle-aged B. sempervirens.
A total of sixteen species, ve of which were liverworts,
were determined in the middle zones of the aged boxwood
trees. All of the liverworts, which were determined in the
epiphytic habitats of the boxwood forests, were found on the
middle parts of the old trees. While A. complanata was the
most frequent and the most dominant, with the highest IES
value (342), E. crispa and A. besseri were co-dominant with
the same IES values (314) (Tab. 3). The mat life form (50%)
and the cortico-saxicolous type anity (68.75%) were the
most dominant on the middle zones of aged trees.
Upper zone
Ten species were found on the upper zones of young
trees. Among them, eight were mosses and, two were
l i v e r w o r t s . Exsertotheca crispa was still the most frequent
and the most dominant with the highest IES value (357).
Alleniella complanata, A. besseri and, Leucodon sciuroides
were co-dominant with the higher IES values (>200) on
the upper zones of the young B. sempervirens. While
Orthotrichum pumilum (IES value 200), Ulota crispa (200)
and, R. complanata (114) were abundant on these zones, S.
otowianum (28) was scarce with the least IES value (Tab.
3). The mat life form (40%) and the habitat anity type
cortico-saxicolous (70%) were the most common on the
upper parts of young trees.
Eleven species (nine mosses, two liverworts) were col-
lected from upper zones of the middle-aged boxwood trees.
While E. crispa was still the most frequent and the most
dominant with the highest IES value (366), A. besseri, A.
complanata and, L. sciuroides were co-dominant (> 200).
And also, U. crispa (167) was abundant on the upper zones
of the middle-aged B. sempervirens (Tab. 3). The mats
(45.45%) and cortico-saxicolous species (72.72%) were the
most dominant.
A total of thirteen species were determined on the upper
zones of the old B. sempervirens. Among them, four were
liverworts and nine were mosses. Alleniella complanata was
the most frequent and the most dominant with the highest
IES value (342). While A. besseri (IES value 300), E. crispa
(328) and, L. sciuroides (300) were co-dominant with the
higher IES values, O. pumilum (200), R. complanata (143)
and, U. crispa (200) were abundant on the upper parts of
Fig. 2. Classication of TWINSPAN based on the matrix of 29 epiphytic bryophyte species over 60 sample plots.

EZER T., ALATAŞ M., BATAN N., ERATA H.
16 ACTA BOT. CROAT. 82 (1), 2023
aged boxwood trees (Tab. 3). The life form mat (30.7%) and
the habitat anity type cortico-saxicolous (53.8%) were the
most common again on the upper parts of old trees.
TWINSPAN classication
TWINSPAN classied the epiphytic bryophyte com-
munities on the trunks of B. sempervirens in the Fırtına
Valley at the second level into two main clusters (A and B)
and three sub-clusters (A1, B1 and, B2) (Fig. 2). These main
and sub-clusters were named according to the dominant,
co-dominant and abundant species which were the distinc-
tive species within the communities. The main cluster, A,
occurred in lower-base communities and it was character-
ized by dominant species E. crispa and H. trichomanoides,
co-dominant I. alopecuroides and abundant P. attenuatus
and S. otowianum. The second main cluster, B, occurred
in middle and upper zone communities. Cluster B was
characterised by dominant species A. besseri and A.
complanata, co-dominant E. crispa and L. sciuroides, abun-
dant species O. pumilum, P. euchloron, R. complanata, and
U. crispa.
Epiphytic bryophyte communities
The A1 community was named Exsertotheca crispa-
Isothecium alopecuroides due to the frequency, constancy
and, dominancy of these species within the lower-base
community. Both E. crispa and I. alopecuroides had the
highest IES value on the lower bases of all tree-size groups
(young, middle-aged and old boxwood trees) (Tab. 3). The
community was represented with 19 moss species in a total of
20 sample plots. Moderately abundant R. complanata was the
only liverwort in the lower-base community. Exsertotheca
crispa-Isothecium alopecuroides community was co-dominat-
ed by H. trichomanoides. In this community, P. attenuatus
and S. otowianum were also abundant. While the domi-
nant life forms within the community were weft and mat
(31.5%), cortico-saxicolous species (47.3%) were dominant
and indierent type anity was also co-dominant (42.1%).
The B1 community was named Alleniella complanata-
Exsertotheca crispa according to its dominant and co-dom-
inant species. It was represented by 19 species (15 mosses, 4
liverworts) in a total of 17 sample plots from the middle parts
of the young, middle-aged and old boxwood trees. The liv-
erwort R. complanata was still present in the middle parts of
the trunks with relatively high IES values. Also, P. euchloron
was abundant in the community B1 on the middle zones,
particularly of young and middle-aged trees particularly
(Tab. 3). While mats were the most dominant with the rate
of 42.1% within the middle parts of community B1, cortico-
saxicolous species were the most common with 63.1%.
The B2 community was named Alleniella besseri-
Leucodon sciuroides according to its co-dominant species.
This community, consisting of 17 species (13 mosses and
four liverworts), was found on the upper zones of boxwood
trees. The community B2 was represented by a total of 23
sample plots. Orthotrichum pumilum, R. complanata, and U.
crispa were abundant species in the upper zones community.
While mats were the most dominant with the rate of 41.1%
within the middle parts of community B2, the cortico-saxic-
olous type habitat anity was the most common with 70.5%.
DECORANA ordination
DECORANA grouped the sample plots on axis 1 and
axis 2 according to the similarity and the environmental
gradients (Fig. 3). While the DCA axis 1 was interpreted as
Fig. 3. The relationship between the three epiphytic bryophyte groups generated after the application of TWINSPAN classication
technique on 60 sample plots and the distribution of the groups along the environmental gradient on the rst and second axes of
DECOR ANA. A–1: Exsertotheca crispa-Isothecium alopecuroides, B–1: Alleniella complanata-Exsertotheca crispa, B–2: Alleniella
besseri-Leucodon sciuroides, axis 1: the gradient of height of the epiphytic habitat, axis 2: the gradient of moisture.

SUCCESSION OF EPIPHYTIC BRYOPHYTES IN TÜRKIYE
ACTA BOT. CROAT. 82 (1), 2023 17
gradient along the height of the epiphytic habitat (from the
lower base to the upper zone) on trunks, the DCA axis 2 was
interpreted as gradient of moisture (from mesic to xeric)
(Fig. 3).
Discussion
When the epiphytic habitats on the boxwood trees in the
Fırtına Valley were examined in terms of species diversity
and species composition, the lower bases and the middle-
parts were the richest zones with equal numbers of species
(19 species), while the upper zones contain 17 epiphytic species.
Large pleurocarp mosses such as E. cri spa, H. trichomanoides,
and I. alopecuroides were common on the basal parts of the
trunks of B. sempervirens. These strong competitor members
of pleurocarpous ty pe mosses are more sensitive to drought,
have a faster growth habit than acrocarpous mosses and
spread horizontally in a carpet-like appearance on the sub-
stratum (Schoeld 2001, Ezer 2017). The basal parts of the
trunks are usually more humid and more nutrient rich than
the other parts (middle and upper) due to soil proximity.
Therefore, basal zones allow early establishment and rapid
colonization of bryophytes due to higher water retention ca-
pacity, higher soi l humid ity, low evapotranspiration rate and
low insolation (Lara and Mazimpaka 1998, Mazimpaka et
al. 2009, Ezer and Kara 2013, Ezer 2017). In this respect, the
lower base parts of the trees can be considered an extension
of the forest oor (groundlayer) environment. For this rea-
son, species such as B. rutabulum, C. molluscum, O. hians,
P. nemorale, P. nervosa , S. populeum, T. alpecurum, uidium
delicatulum, and F. serrulatus, which usually grows on the
soil, were encountered only in this part. This caused the weft
life form and indierent type habitat anity to co-dominate
with the mats and cortico-saxicolous species were the strong
competitor of robust pleurocarpous in basal parts of box-
wood trees.
Alleniella besseri, A. complanata and E. crispa were most
dominant on the middle zones of the boxwood trees. Also,
the mesophytic species P. euchloron, which was not present
in the base zone, was abundant in the middle zone. There-
fore, P. euchloron can be considered a characteristic and dis-
tinct ive species of the communities in the middle zones. The
customary epiphyte xerophytic species O. pumilum, which
in the present study was generally abundant in the upper
zones, in the present study was found for the rst time only
in the middle zones of old trees. As the middle and upper
parts of the trunks move away from the soil eect, they are
periodically exposed to higher insolation and desiccation
that makes colonization of species on epiphytic habitats dif-
cult (Moe and Botnen 2000). Therefore, as in the present
study, small cushion-type mosses which have xerophytic
characters such as Orthotrichaceae members and photoph-
ilous or heliophilous species such as A. besseri, A. complan ata,
and L. sciuroides are most common species within the mid-
dle and upper zone communities. Mat type life forms were
predominant in the middle parts of old trees, due to the pres-
ence of cortico-saxicolous liverworts.
Although mesoscale climatic factors such as seasonal cli-
matic variables are among the most important determinants
of bryophyte species richness and species diversity, forest
structure and habitat characteristics are also important for
community compositions and spatial distributions of epi-
phytic bryophytes on epiphytic habitats (Medina et al. 2014,
Ezer 2017). Species diversity and epiphytic cover on the up-
per zones were found to be lower than in the base and middle
zones. The xerophytic robust pleurocarpic species L. sciuroides
was dominant on the upper parts of all tree-size groups.
Besides A. besseri, A. complanata and E. crispa there were
other dominant pleurocarpic species in the upper zones of
boxwood trees in the Fırtına Valley. Therefore, weak com-
petitor species such as liverworts Frullania dilatata and
Metzgeria furcata and small cushion-type mosses (such as
O. pumilum and U. crispa) remain under these large pleuro-
carpous species and decreased or disappeared from the
epiphytic habitats in the valley.
Some studies on the succession of epiphytic bryophyte
communities have demonstrated that variables of the epi-
phytic bryophyte composition in the successional stages are
closely related to tree age, trunk height (basal, middle and
upper zones) and bark characteristics (Lara and Mazimpaka
1998, Mazimpaka et al. 2010, Ódor et al 2013, Bargali et al.
2014, Ezer 2017). However, in the present study, trunk height
rather than tree age and bark characteristics were eective
in the variability of epiphytic bryophyte composition in the
per successional stage. Hygrophytic pleurocarpous species
that usually grow on soil, such as B. rutabulum, C. molluscum,
F. serrulatus, O. hians, P. nemorale P. nervosa, S. populeum,
T. alpecurum, and T. delicatulum were particularly the pio-
neer colonizers in the early successional stages on the basal
parts particularly of middle aged B. sempervirens. While the
xerophytic small cushions O. pumilum and U. crispa were
the pioneer colonizers in the early successional stages, the
large pleurocarpous L. sciuroides was a secondary colonizer
in the advanced stages on the upper parts of all tree-size
groups. In addition, cortico-saxicolous species P. euchloron,
a characteristic and distinctive species on the middle zones,
was pioneer colonizer in the early successional stages on the
middle parts of all tree-size groups. In this study, A. besseri,
A. complanata, and E. crispa were other colonizers in the
advanced successional stages on the middle zones. Consid-
ering the morphological physiognomy of the bryophyte
communities on trunks of B. sempervirens in the Fır tına Val-
ley; all communities were dominated by large pleurocarpous
mosses. E. crispa was in particula r the most constant and the
most dominant within all communities in the B. sempervirens
forests. In sum, the succession of the epiphytic bryophyte
communities of the boxwood forests in the Fırtına Valley
has reached the climax.
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