159
Key words: Chara, Sphaerochara, 
Tolypella, floristic novelties, 
protection status
Ključne besede: Chara, 
Sphaerochara, Tolypella, floristične 
novosti, varstveni status
Corresponding author:
Roman Romanov
E-mail: streptophytes@gmail.com
Received: 21. 12. 2024
Accepted: 8. 1. 2025
New national species records of 
charophytes (Characeae) for Montenegro, 
Croatia, and Cyprus: update of rare 
species distribution in Southeast Europe
Abstract
Field studies in 2022–2024 and the checking of collection of the Natural 
History Museum of Montenegro revealed seven new national species records for 
Montenegro: Chara corfuensis Groves ex Fil., C. galioides DC., C. squamosa Desf., 
Sphaerochara intricata (Trentep. ex Roth) Feist-Castel & N. Grambast, Tolypella 
hispanica Nordst. ex Allen, T. nidifica (O. F. Müll.) A. Braun and “Mediterranean 
Chara”. The checking of the specimens stored in ZE Botanischer Garten und 
Botanisches Museum, Freie Universität Berlin, Komarov Botanical Institute of 
RAS , and Naturhistorisches Museum Wien revealed old specimens of C. galioides, 
a new species for Croatia, and new sites of T. hispanica for Greece. The field 
research in Cyprus resulted in a new national species record of C. squamosa. New 
data enriched knowledge about the distribution of charophytes in Central and 
Eastern Mediterranean and Southeast Europe and allowed tentative suggestions 
of important charophyte areas in Montenegro. The tentative evaluation of species 
protection statuses was implemented here. The updated species list of Montenegrin 
charophytes counted 37 species making Montenegro one of the charophyte species-
rich countries in Europe.
Izvleček
S terenskimi raziskavami med letoma 2022 in 2024 in pregledom zbirk v 
Prirodoslovnem muzeju Črne gore smo odkrili za Črno goro sedem novih vrst: 
Chara corfuensis Groves ex Fil., C. galioides DC., C. squamosa Desf., Sphaerochara 
intricata (Trentep. ex Roth) Feist-Castel & N. Grambast, Tolypella hispanica 
Nordst. ex Allen, T. nidifica (O. F. Müll.) A. Braun in “Sredozemska Chara”. 
S pregledom primerkov, shranjenih v zbirkah ZE Botanischer Garten und 
Botanisches Museum, Freie Universität Berlin, Komarov Botanical Institute 
of RAS in Naturhistorisches Museum Wien, smo odkrili star primerek vrste 
C. galioides, nove vrste za Hrvaško in nove lokalitetete vrste T. hispanica v Grčiji. 
S terenskimi raziskavami na Cipru pa smo odkrili novo lokaliteto vrste C. 
squamosa. Novi podatki so obogatili poznavanje razširjenosti parožic v srednjem in 
vzhodnem Sredozemlju ter jugovzhodni Evropi, hkrati pa izpostavili pomembna 
območja parožic v Črni gori. V članku smo ovrednotili začasni varstveni status 
vrst. Posodobljen seznam sedaj vsebuje 37 vrst in Črna gora je tako ena od držav v 
Evropi z največ vrstami parožic.
Roman Romanov1, Vera Biberdžić2 , Snežana Dragićević3 &  
František Bednár4 
DOI: 10.3986/hacq-2025-001224/2 • 2025, 159–170
1 Komarov Botanical Institute of the Russian Academy of Sciences, St. Peterburg, Russia; current adress: Nišice, Dobra Voda, Bar, Montenegro.
2 Natural History Museum of Montenegro, Podgorica, Montenegro.
3 Montenegrin Academy of Sciences and Arts, Podgorica, Montenegro.
4 Švošov, Slovakia.
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New charophytes for Montenegro, Croatia, and Cyprus
Introduction
The charophytes (Charophyta, Charophyceae, Characeae) 
are a widely distributed group of macroscopic algae asso-
ciated with continental water bodies. Plant arrangement 
allows easy recognition of this group from first glance. 
However, species identification still can be an inspiring 
challenge because of insufficiently known variability of 
some species and still debated species concepts (Schubert 
et al., 2024). Besides the importance of charophytes as 
keystone species for some types of ecosystems, they are 
important species in threatened and protected habitats, 
e.g. “3140. Hard oligo-mesotrophic waters with benthic 
stoneworts Chara spp.”, “3170. Mediterranean temporary 
ponds”, and “3190. Lakes of gypsum karst” of the Habi-
tat Directive by European Union and “C1.2a. Permanent 
oligotrophic to mesotrophic waterbody with Characeae”, 
a vulnerable habitat type according to the European Red 
List of Habitats (European Commission, 2017). Medi-
terranean temporary ponds appear to be one of the most 
precious habitat types in the south of Europe because they 
harbor rare and peculiar species (Bagella, 2023). Charo-
phytes are one of the most threatened groups of plants in 
Europe (Baastrup-Spohr et al., 2013), which can be easily 
illustrated by the high proportions of endangered species 
in many national and preliminary European Red Lists 
(Becker, 2016; Stewart, 2024). The key areas important 
for protecting of vulnerable and endangered species were 
outlined in a few European countries only (e.g. Blaženčić 
et al., 1998; Stewart, 2004). Therefore, accurate and reli-
able datasets about species distribution in time and space 
are essential for monitoring and designing natural pro-
tected areas avoiding already known irreversible loss of 
important sites (Langangen, 2017; Troia, 2020). Despite 
long history of research on charophyte species distribu-
tion in Central Mediterranean and Southeast Europe 
(see Blaženčić et al., 1997, 1998, 2006a, b; Biberdžić 
et al., 2006; Bazzichelli & Abdelahad, 2009; Biberdžić 
& Blaženčić, 2013; Blaženčić, 2014; Troia et al., 2018; 
Becker, 2019) even national species lists are not com-
plete. In the last decade, new national species records are 
added almost every year (Zeneli & Kashta, 2016; Troia 
et al., 2018; Becker, 2019; Romanov, 2019; Romanov 
et al., 2019; Trbojević et al., 2020, 2024; Becker et al., 
2021; Koletić et al., 2021; Casanova & Becker, 2022; 
Biberdžić et al., 2023; Schubert et al., 2024). Montene-
gro was already recognized as one of the important areas 
of charophyte species diversity in Europe (Blaženčić & 
Stevanović, 2015) partly because of the high species rich-
ness in Lake Skadar (Blaženčić et al., 2018). According to 
the contemporary nomenclature (Schubert et al., 2024), 
30 species are reported for Montenegro.
This research aimed to update the national species lists 
adding seven novel species records for Montenegro, one 
new species for Croatia and Cyprus respectively, enrich-
ing knowledge of charophyte species distribution in Cen-
tral and Eastern Mediterranean and Southeast Europe, 
and assessing their national protection statuses.
Material and Methods
During 2022–2024, a survey of different aquatic habitats 
of Montenegro, mostly on the Adriatic coast from the 
westernmost to easternmost extremities of the country, 
was carried out, as well as an analysis of samples from the 
collection of the Natural History Museum of Montene-
gro (NHMM). Most areas of the Adriatic coast and some 
inland sites looking suitable for charophytes at satellite 
images were checked several times in different seasons. It 
allowed tentative estimation of the duration of the exist-
ence of most temporal water bodies and the stability of 
charophyte populations. The specimens were collected by 
hand and grapnel from water bodies by the authors. The 
rough assessment of the water salinity was based on the 
species and abundance of angiosperms and macroscopic 
algae associated with charophytes in Montenegrin sites. 
Electric conductivity and pH were measured by Thermo 
Scientific™ Elite PCTS (Thermo Fisher Scientific Inc., 
Waltham, MA USA) after 2 days of sampling in Cyprus, 
where some plants were collected by Martin Dobrota. 
They were cultivated at room temperature for 10 weeks 
to trace stem cortex traits. The specimens were pressed 
or stored in 70% ethanol. They were deposited in the 
herbarium of the NHMM. The collections of Naturhis-
torisches Museum Wien (W), ZE Botanischer Garten 
und Botanisches Museum, Freie Universität Berlin (B), 
Naturalis Biodiversity Center (L) and the Komarov Bo-
tanical Institute of the Russian Academy of Sciences (LE) 
were checked by the first author in search of specimens 
from Central and Eastern Mediterranean, and South-
east Europe. The morphological features of the speci-
mens were examined using a stereo microscope Olym-
pus SZX12, and microscopes Olympus BX51 and BH2 
(Olympus Corporation, Shinjuku, Tokyo, Japan). Species 
distribution in Southeast Europe was illustrated with 
maps prepared with SimpleMappr (Shorthouse, 2010). 
Nomenclature follows the monograph “Charophytes of 
Europe” (Schubert et al., 2024). The tentative assess-
ment of protection statuses was implemented according 
to IUCN criteria (IUCN, 2012) and the suggestions fo-
cused at charophytes by Stewart (2024). However, further 
researches in the region are needed for a stable assessment 
of the IUCN categories.
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Results and discussion
During our field survey, more than 220 sites (18 species 
and 5 genera) were found mostly on the Montenegrin 
Adriatic coast. Seven species listed below are new national 
records for Montenegro. One species is a new national 
record for Croatia, and one species is a floristic novelty 
for Cyprus. Some Greek sites for two species missed in 
recent maps of species distribution in Europe (Lambert & 
Schubert, 2024a; Troia et al., 2024) were traced in stud-
ied herbarium collections. Results are here reported by 
taxon, in alphabetical order.
Chara corfuensis Groves ex Fil. (Figure 1a), Montene-
gro, Ulcinj Municipality, northeast vicinities of Velika 
plaža (Long Beach): 1. bare Đurišića, elongated sand pit, 
41.88200 N 19.33840 E, 30.08.2023; 2. deep sand pit, 
41.88600 N 19.33020 E, 30.08.2023; 3–5. small inter-
dunal ponds in a large sand pit with a lot of silty sedi-
ments, 41.88780 N 19.32180 E, 41.88790 N 19.32170 
E, 41.88460 N 19.33140 E, 22.09.2023. 
This species was spotted in stands forming at the deep-
est parts of plain freshwater interdunal ponds and inun-
dated sand pits with prominent seasonal changes in water 
levels increasing during winter and severely dropping dur-
ing summer to ca. 50 cm and less depth. Chara corfuensis 
was found in freshwater water bodies in Montenegro as 
in Croatia, and the North Black Sea region (Blaženčić, 
2006; Romanov et al., unpubl. data; this research) in con-
trast to mostly brackish lagoons and lakes in Albania and 
Greece (Christia et al., 2024). The preference for small 
ponds with highly variable water levels in Montenegro 
can be evidence of the susceptibility of this species to in-
terannual differences in weather and climate change. The 
development of Velika plaža vicinities, including urbani-
zation and spontaneous dumping as well as sand extrac-
tion, resulting in physical disturbance, fragmentation and 
loss of habitats can be recognized as a threatening factor 
suitable for all charophyte species in this area. Chara corf-
uensis can be referred to “Endangered” (IUCN criteria B1, 
2a,b, C2a, D) in Montenegro. It was recognized as “Criti-
cally Endangered” in the Red List of Balkan charophytes 
(Blaženčić et al., 2006b) and “Endangered 
(European and Globally)” in the provisional 
European Red List (Stewart, 2024).
Figure 1: The general habit of species from 
Montenegro: A – Chara corfuensis having a spiny 
outline because of long pointed bract-cells, spine-cells, 
and stipulodes, B – “Mediterranean Chara” having a 
less spiny outline, C – male plants of Chara galioides, 
D – female plants of Chara galioides. Arrowheads 
indicate female plants, subtle and less conspicuous in 
comparison to rough male plants bearing large orange 
antheridia. Photographs by R. E. Romanov.
Slika 1: Splošna podoba vrst v Črni gori: A – Chara 
corfuensis ima trnast obris zaradi dolgih špičastih 
braktejnih celic, trnskih celic in paprilističev,  
B – “Sredozemska Chara” ima manj trnast habitus, 
C – moške rastline vrste Chara galioides, D – ženske 
rastline vrste Chara galioides. Puščice kažejo manj 
opazne ženske rastline, saj imajo moške rastline velike 
oranžne anteridije. Fotografije R. E. Romanov.
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New charophytes for Montenegro, Croatia, and Cyprus
In Southeast Europe, Chara corfuensis is known from 
Albania, Croatia, and Greece from very few sites (Fig-
ure 2a; Blaženčić, 2006; Langangen 2012 and references 
herein; Christia et al., 2024 and references herein). It is 
distributed in Southern Europe, mainly in the Mediter-
ranean, but it is known from the Atlantic coast and North 
Figure 2: Distribution 
of charophyte species in 
Southeast Europe and 
Cyprus based on available 
data according to time 
intervals in the monograph 
“Charophytes of Europe” 
(Schubert et al., 2024; see 
references in the text).  
A – Chara corfuensis,  
B – Chara galioides,  
C – Chara squamosa,  
D – the taxon of uncertain 
affiliation “Mediterranean 
Chara”, E – Tolypella 
hispanica, F – Sphaerochara 
intricata, G – Tolypella nidi-
fica. Black dots are records 
from 1980 on, red dots 
from 1950 to 1979, and 
yellow dots before 1950. 
Slika 2: Razširjenost 
parožic v jugovzhodni 
Evropi in na Cipru, prika-
zana na osnovi podatkov iz 
monografije “Charophytes 
of Europe” (Schubert et al., 
2024; glej reference v tek-
stu) v časovnih intervalih.  
A – Chara corfuensis,  
B – Chara galioides,  
C – Chara squamosa, 
D – takson z nejasno 
pripadnostjo “ Sredozemska 
Chara”, E – Tolypella 
hispanica, F – Sphaerochara 
intricata, G – Tolypella nidi-
fica. Črne točke so podatki 
od leta 1980 naprej, rdeče 
točke med letoma 1950 in 
1979 in rumene točke pred 
letom 1950.
Black Sea region too (Christia et al., 2024 and references 
herein; Romanov et al., unpubl. data). It belongs to the 
subsection Hartmania R.D. Wood, or in other words, to 
the group trickiest for the delineation and identification 
in the Mediterranean (Blindow et al., 2024b). 
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New charophytes for Montenegro, Croatia, and Cyprus
Greece. Attica: in aquis salsuginosis stagnatibus, ad 
Phalerum. Legimus foem. 16 Mai et marem 19 Mai 1886. 
De Heldreich Herbarium Graecum Normale. 995. Det.: 
Chara crinita Wallr., Sydow Charac. p. 52, Nym. Con-
spect. 876, No. 18. forma b. polysperma Al.Br. f. longifolia 
Syd. l. c., laxius aculeata, valde incrustata Sydow! in litt. 
1887. Mas et foemina. Det. U. Raabe: Chara galioides 
12.01.2000. Conf. R. E. Romanov (W 536, W 537).
This is a common species in the south-easternmost 
area of Montenegro, in plain small temporary brackish 
to nearly freshwater water bodies in less disturbed areas 
of Velika plaža and in some inundated crystallization and 
evaporation basins of Ulcinjska Solana. It occurs slightly 
less frequently in water bodies in the northeastern and 
eastern vicinities of Ulcinjska Solana. In contrast, in 
Montenegro, it was not found west of this area. All spot-
ted populations are consisted of annual plants appearing 
during January and February and sterile during this pe-
riod, but richly fertile during spring and summer. The 
gametangia appear not early during plant germination 
and the beginning of growth despite annual populations 
in temporary water bodies mainly disappearing at the end 
of spring and summer. The antheridia appear earlier than 
oogonia. This species can be spotted as mostly dry plants 
at dry bottoms for a short period after water evaporation 
during summer. In Croatia, it is known only from the col-
lection of the 19th century from the small Adriatic island 
of Sveti Petar. Chara fragilis Desv. var. brevibracteata sine 
auctoris was reported from this island. It was spotted on 
the same visit by the same collector (Reichardt, 1863). It 
is unclear if it was based on the same specimens of Chara 
galioides or if it was true Chara globularis Thuill. consider-
ing the small size of this dry Mediterranean island and 
the existence of the specimen of Chara globularis collected 
from the same island on 20 May 1867 (B: 35811). The 
specimens by Reichardt were referred by A. Braun to 
Chara aspera Willd. with some doubts (Leonhardi, 1864). 
The checking of old herbarium specimens stored in W 
allowed adding one site for Greece (Figure 2b) missed on 
the last map of species distribution in Europe (Lambert 
& Schubert, 2024a).
The preference of the species for temporary ponds can 
be evidence of the susceptibility to interannual differences 
in weather and climate change. The urbanization of Ve-
lika plaža vicinities resulting in physical disturbance and 
loss of habitats can be recognized as a threatening fac-
tor. Considering the number of sites in Montenegro in 
comparison to few other sites in Southeast Europe the 
surveyed area has an international importance for the 
protection of this species. Chara galioides can be referred 
to “Near Threatened” (IUCN criteria B2a,b, C2a, D) 
in Montenegro and “Critically Endangered” in Croatia 
Chara galioides DC. (Figure 1c, d), Montenegro,  Ulcinj 
Municipality. Vicinities of Velika plaža, temporary 
ponds: 
1. inundated shallow depression in community of Juncus 
acutus L. near Tamarix sp., ca. 41.87864 N 19.33009 E, 
14.01.2023; 2. inundated shallow depression in the 
community of Juncus acutus, 41.88926 N 19.30924 
E, 11.02.2023; 3. margin of the inundated dirt road, 
41.88821 N 19.31089 E, 11.02.2023; 4. a puddle at 
dirt road, 41.87237 N 19.33935 E, 22.02.2023; 5. in-
undated shallow depression in the community of Juncus 
acutus, 41.87838 N 19.32989 E, 22.02.2023; 6. inun-
dated channel, 41.87873 N 19.33600 E, 22.02.2023; 7. 
inundated shallow depression W of the road, 41.91603 N 
19.33852 E, 23.03.2023; 8. inundated shallow depres-
sion in the community of Juncus acutus, 41.87928 N 
19.32992 E, 17.04.2023; 9, 10. inundated dirt road, 
41.87202 N 19.33910 E and 41.88003 N 19.33110 E, 
25.03.2023; 11–15. vicinity of the beach of Copacaba-
na, 41.88889 N 19.30902 E, 41.88937 N 19.30865 E, 
41.89279 N 19.30923 E, 41.89309 N 19.30940 E, 
and 41.88889 N 19.30902 E, 17.04.2023; 16. inun-
dated sand pit, 41.8820 N 19.3387 E, 30.08.2023. 
Ulcinj municipality, NE of the Nature Park “Ulcinjska 
Solana” (Ulcinj Saline): 17. Kneta ispod Darze, tem-
porary pond, 41.94066 N 19.31667 E, 02.03.2023. 
18, 19. inundated field near the channel, 41.94173 N 
19.32634 E and 41.94165 N 19.32614 E, 01.05.2024. 
Ulcinj municipality, Ulcinjska Solana: 20–35. temporary 
ponds in abandoned crystallization and evaporation ba-
sins: 41.92788 N 19.27491 E, 17.04.2023; 41.91960 N 
19.25330 E, 30.04.2023; 41.91785 N 19.32479 E, 
and 41.92040 N 19.25210 E, 06.05.2023; 41.9114 N 
19.3262 E, 41.9139 N 19.3286 E, 41.9142 N 19.3269 E, 
and 41.9145 N 19.3265 E, 23.05.2023; 41.92155 N 
19.33144 E, 41.92162 N 19.33099 E, 41.92069 N 
19.33163 E, 41.91922 N 19.33114 E, 41.91898 N 
19.33167 E, 41.91895 N 19.33190 E, 41.91864 N 
19.33130 E, and 41.91779 N 19.33040 E, 19.05.2024.
Croatia. 1. In mit süssem Wasser erfüllten *** [illeg-
ible words] Insel San Pietro di Nembi im Quarnero, [24] 
April 1862, H.W. Reichardt. Det. A. Braun: Chara aspera. 
2. Istrien. In einer Süsswasserlake der Insel San Pietro di 
Nembi, 24 April 1862, H.W. Reichardt. Det. [A. Braun]: 
Chara aspera. 3. San Pietro di Nembi im Quarnero, [24 
April 1862], H. W. Reichardt. Det. [A. Braun]: Chara 
aspera. Det. R. E. Romanov: Chara galioides (W, three 
sheets). 4. Quarnero. In einer Süsswasserlake *** [illegible 
words] San Pietro di Nembi, 24 April 1862, H. W. Reich-
ardt. Det. [A. Braun]: Chara aspera. Det. R. E. Romanov: 
Chara galioides (B 35797).
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New charophytes for Montenegro, Croatia, and Cyprus
(IUCN criteria A1, B1, 2a,b, C1). It was recognized as 
“Critically Endangered” in the Red List of Balkan charo-
phytes (Blaženčić et al., 2006b) and “Least Concern” in 
the provisional European Red List (Stewart, 2024).
In Southeast Europe, Chara galioides is known from 
very few sites in Albania (Zeneli & Kashta, 2016; Kashta 
et al., 2023) and Greece (Raabe & Koumpli-Sovantzi, 
2000, 2002) (Figure 2b). It is distributed in coastal re-
gions of Mediterranean mostly, in Europe and North Af-
rica, as well as in Atlantic coast of France, Iberian Pen-
insula, and North Black Sea region, but most records are 
known from Western Mediterranean (Muller et al., 2017; 
Lambert & Schubert, 2024a and references herein).
Chara squamosa Desf. (Figure 3a), Montenegro, Bar 
Municipality, southern slope of Rumija Mountain Ridge: 
1. Vicinity of Tuđemili, well Manduke, at ca. 40 cm 
depth, 42.14605 N 19.13615 E, 10.02.2023. 2. North 
of Stari Bar, small stream – a left tributary of Zeljeznica 
River, 42.14984 N 19.11771 E, 22.05.2023. Bijelo Polje 
Municipality: 3. Šipovica, small stream below the me-
morial fountain in water of Ćamil Sijarić, 43.01773 N 
19.89482 E, 08.07.2024. 4. Vojan grab, small stream, 
43.02098 N 19.85268 E, 08.07.2024. Šavnik Munici-
pality: 5. Bare, small stream near local road, 42.93076 N 
19.25131 E, 15.08.2021.
Cyprus. 1. Agios Minas Chapel, west of Neo Chorio, 
Paphos district: small stream below spring near a road, at 
1–3 cm depth, pH 8.01, electrical conductivity (EC) 880 
μs/cm, total dissolved solids (TDS) 625 ppm, salinity 
0.40 ppt, 35.02400 N, 32.34152 E, 23.10.2024 (https://
www.inaturalist.org/observations/249076970). 2. West 
of Ineia, Paphos district: Paphos, concrete basin, at 50–
100 cm depth, pH 7.35, EC 1233 μs/cm, TDS 868 ppm, 
salinity 0.60 ppt, 34.95555 N, 32.38611 E, 23.10.2024 
(https://www.inaturalist.org/observations/249076974).
Figure 3: Key traits and the general habit of species 
from Cyprus and Montenegro. A – Chara squamosa 
from Cyprus: diplostichous tylacanthous stem cortex 
with short solitary spine cells, ecorticate branchlets 
(arrowhead), B – upper part of Sphaerochara intricata, 
arrowheads indicate typical forked sterile branchlets, 
C – general habit of Tolypella nidifica, D – general 
habit of male plants of Tolypella hispanica with large 
orange antheridia. Photographs: A – F. Bednár, 
B–D – R. E. Romanov.
Slika 3: Ključni znaki in splošen habitus vrst s Cipra 
in Črne gore. A – Chara squamosa s Cipra: dvoslojna 
grbančasta skorja stebla s kratkimi, posamičnimi 
trnskimi celicami, vejice brez skorje (puščica), 
B – zgornji del vrste Sphaerochara intricata, puščice 
kažejo značilne viličaste sterilne vejice, C – splošen 
habitus vrste Tolypella nidifica, D – habitus moške 
rastline vrste Tolypella hispanica z velikimi oranžnimi 
anteridiji. Fotografije: A – F. Bednár,  
B–D – R. E. Romanov.
This species is associated with small fresh-
water mountain streams and artificial basins 
filled with fresh water in regions with lime-
stone outcrops. The records from small wa-
ter bodies can be evidence of the susceptibil-
ity of this species to interannual differences 
in weather and climate change threatening 
groundwater discharge. This species can be 
recognized as “Data Deficient” because its 
distribution in Montenegro and Cyprus 
is poorly known. It was not found among 
specimens from Cyprus collected by other 
botanists despite the targeted search in B, 
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L, LE, and W. It was recognized as “Critically Endan-
gered” in Serbia (Trbojević et al., 2024) and in the Red 
List of Balkan charophytes (as Chara conimbrigensis A. G. 
Cunha; Blaženčić et al., 2006b) and “Data Deficient” in 
the provisional European Red List (Stewart, 2024).
In Southeast Europe, Chara squamosa is known 
from Croatia, Serbia and North Macedonia (Figure 2c; 
Blaženčić & Blaženčić, 1997; Blaženčić et al., 2006b; 
Trbojević et al., 2024). It is distributed mostly in the 
Mediterranean. Chara squamosa is known from Europe, 
North Africa, and West Asia, including Caucasus (Ro-
manov et al., 2023, 2024a and references herein). Consid-
ering its close similarity to Chara gymnophylla A. Braun, 
targeted checking of collections and new field studies 
can change a suggestion about the rare occurrence of this 
species in the south of Europe (Romanov et al., 2024a), 
especially in Southeast Europe, as it was recently shown 
for Serbia (Trbojević et al., 2024). However, this seems 
not to be the case for the Montenegrin Adriatic coast 
where it appears to be a rare species found in mountain 
landscapes only (this research). The same environment 
is notable for new sites from the Dinaric Alps in Mon-
tenegro and two locations from Cyprus (this research). 
The general occurrence of Chara squamosa in mountain-
ous and hilly landscapes was found in Serbia and West 
Asia – Caucasus and Eastern Mediterranean (Trbojević 
et al., 2024; Romanov et al., unpubl. data). Few records 
of Chara squamosa from the Italian peninsula, Sicily, and 
the Balearic Islands fit the same pattern (Romanov et al., 
2019, 2024a; Romanov et al., unpubl. data). Considering 
the re-identification of this species from Sardinia this dis-
tributional pattern seems suitable for this island too (cf. 
Becker, 2019; Romanov et al., 2024a). Therefore, more 
sites of Chara squamosa could be expected in the inland 
part of Montenegro, i.e. in the Dinaric Alps. This sug-
gestion is not supported by the rare occurrence of Chara 
gymnophylla, the most similar to and not easily deline-
able with Chara squamosa (cf. Romanov et al., 2024a), in 
the country (Blaženčić & Stevanović, 2015; unpublished 
data by the authors; NHMM, checked by the authors). 
Genetic studies are implemented for this species group 
clarification (Romanov et al., in progress). 
Taxon of uncertain affiliation “Mediterranean Cha-
ra” (Figure 1b), Montenegro, 1–6. Ulcinj Municipal-
ity, northeast vicinities of Velika plaža, inundated sand 
pits, 41.89706 N 19.29504 E, 41.89740 N 19.29045 E, 
41.90015 N 19.27985 E, 41.89807 N 19.28941 E, 
41.89703 N 19.29504 E, 41.89739 N 19.29044 E, 
27.10.2022. 7–14. Budva Municipality, plain coastal area 
southern of Buljarica, brackish artificial permanent ponds 
and channels of the drainage system, at 20–50 cm depth, 
42.18494 N 18.97176 E, 42.18490 N 18.97163 E, 
42.18455 N 18.97165 E, 42.18668 N 18.97121 E, 
42.18762 N 18.97092 E, 42.18560 N 18.97092 E, 
42.18611 N 18.97170 E,  42.18671 N 18.97132 E, 
24.06.2023. 
This species is annual in Montenegro. The plants start 
germination in November becoming well-developed 
during June to October. The occurrence in only shallow 
water bodies can be evidence of the susceptibility of this 
species to interannual differences in weather and climate 
change. The urbanization of the coastal area in the vi-
cinities of Buljarica and Velika plaža touching the existing 
drainage system and water bodies can be recognized as a 
threatening factor. This species can be referred to “Vul-
nerable” (IUCN criteria B2a,b, C2a, D2) in Montenegro. 
It was recognized as “Critically Endangered” in the Red 
List of Balkan charophytes (as C. baltica (Hartm.) Bruze-
lius; Blaženčić et al., 2006b) and “Data Deficient” in the 
provisional European Red List (Stewart, 2024)
“Mediterranean Chara”, a member of the subsection 
Hartmania R.D. Wood, or, in other words, a part of 
long-lasting “botanicorum crux et scandalum” consider-
ing still debatable species concepts, was tentatively ac-
cepted as a species in the monograph “Charophytes of 
Europe” (Blindow et al., 2024a). It requires further stud-
ies for evaluation. The Montenegrin plants are largely 
useful for this purpose. Some of them have apical parts 
characteristic for Chara corfuensis. However, they were 
growing in stands of plants mostly suitable for “Mediter-
ranean Chara” leaving a space for doubts in their species 
identity. Probably, these taxa are conspecific and Chara 
corfuensis, which has a nomenclatural priority, represents 
a long-spiny, long-bracteate, and long-stipulate morpho-
type with verticillate bract cells. Chara corfuensis grows 
at good insolation without any restraints for the expres-
sion of the great length of bract cells and spine cells. 
Most spiny parts of plants are spotted near the surface 
of the water after the water level drops. It could be hy-
pothesized that both insolation and, less probably, sa-
linity increase can be responsible for this expression of 
morphological traits, essential for species delineation in 
the subsection Hartmania (see Blindow et al., 2024b). In 
Southeast Europe “Mediterranean Chara” was found in 
Slovenia, Albania, and Greece only; it was known there 
as Chara baltica (Hartm.) Bruzelius (Figure 2d; Firbas & 
Al-Sabti, 1995; Zeneli & Kashta, 2016; Blindow et al., 
2024a). This taxon occurs mostly in the Mediterranean, 
but most records are available from the Western Mediter-
ranean. It is known from the Atlantic coast of France too 
(Blindow et al., 2024a).
24/2 • 2025, 159–170
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Romanov et al.
New charophytes for Montenegro, Croatia, and Cyprus
Sphaerochara intricata (Trentep. ex Roth) Feist-Castel 
& N. Grambast (Figure 3b), Montenegro, Ulcinj Mu-
nicipality, vicinity of Fraskanjel, the easternmost part of 
Kneta e Fraskanjel, temporary pond, depth 40–50 cm, 
41.96580 N 19.37476 E, 02.04.2023. 
The annual population appearing during spring only 
was found in the single plain freshwater shallow tempo-
rary water body existing during winter and spring in the 
depression inundated with waters of Lake Šas (Šasko jeze-
ro) and probably Bojana River. It is dry during summer 
and autumn. The finding in the temporary pond can be 
evidence of the susceptibility of this species to interannual 
differences in weather and climate change. This species 
can be referred to “Critically Endangered” (IUCN crite-
ria B2a,b, C2a, D) in Montenegro. It was recognized as 
“Endangered” in the Red List of Balkan charophytes (as 
Tolypella intricata (Trentep. ex Roth) Leonh.; Blaženčić et 
al., 2006b) and “Vulnerable” in the provisional European 
Red List (Stewart, 2024).
This is a sparsely distributed rare species in Southeast 
Europe (Figure 3f ), where it is known from Greece, Bul-
garia, Serbia, and Romania (Blaženčić & Stanković, 2009; 
Vesić et al., 2011; Marković et al., 2023; van de Weyer, 
2024). Sphaerochara intricata is a Holarctic species; its 
distribution area covers Eurasia, North Africa, and North 
America (Pérez et al., 2014; Korsh, 2018).
Tolypella hispanica Nordst. ex Allen (Figure 3d), Mon-
tenegro, Ulcinj Municipality, Nature Park “Ulcinjska 
Solana”, brackish temporary ponds in abandoned crystal-
lization and evaporation basins, and channel at few to 15 
cm depths or spreading at wet clayish soil in the terrestrial 
environment after water evaporation: 1–8. 41.9235 N 
19.2718 E, 17.04.2023; 41.92088 N 19.26240 E, 
28.04.2023; 41.91960 N 19.25330 E, 41.91860 N 
19.25270 E, 41.91860 N 19.25270 E, 30.04.2023; 
41.92040 N 19.25210 E, 41.9109 N 19.3095 E, 
06.05.2023; 41.91079 N 19.31099 E, 23.05.2023.
Greece. 1. Attica: in aquis salsuginosis stagnatibus, ad 
radices m. Kerata, prope Eleusin. 18 April 1886. De Hel-
dreich herbarium graecum normale. Det.: 998. Tolypella 
prolifera (Ziz, Al. Br. Sub Chara) Leonh., Sydow Charae. 
P. 37, Nym. Consp. 879, No. 2, forma typica, fide 
Sydow! In litt. 1887. Det. U. Raabe: Tolypella hispanica 
12.01.2000 (W 544). The same label. Det. R.E. Romanov 
(B 39755). The same label. Det. R.E. Romanov (LE). 2. 
Griechenland: in Graben im Phaleron bei Athen, 19 April 
1911, Schniffer, Algae marinae No. 1319. Det. Schiffner: 
Tolypella nidifica (Müll.) Leonh. Det. U. Raabe: Tolypella 
hispanica 08.01.1999. Conf. R.E. Romanov (B 38624).
Annual populations appearing during only winter and 
spring were spotted in several plain brackish shallow tem-
porary water bodies within Ulcinjska Solana only. The oc-
currence in only temporary ponds can be evidence of its 
high susceptibility to interannual differences in weather 
and climate change. This species can be referred to “Criti-
cally Endangered” (IUCN criteria B2a,b, C2a, D2) in 
Montenegro and Greece. It was recognized as “Criti-
cally Endangered” in the Red List of Balkan charophytes 
(Blaženčić et al., 2006b) and “Least Concern” in the pro-
visional European Red List (Stewart, 2024). 
The checking of old herbarium specimens stored in B, 
LE, and W and recent observations at iNaturalist.org (see 
below) allowed adding a few sites for Greece missed on 
the last map of species distribution in Europe (Troia et 
al., 2024). The part of old collection from Kerata, Greece, 
stored in Stockholm was reported as Tolypella hispanica by 
Horn af Rantzien (1959). In Southeast Europe, Tolypella 
hispanica is known from Croatia, Albania, and Greece 
(Figure 2e; Lovrić et al., 1989; Blaženčić et al., 1998; 
Troia et al., 2024; https://www.inaturalist.org/observa-
tions/111904610; https://www.inaturalist.org/observa-
tions/111515902; https://www.inaturalist.org/observa-
tions/111516902). It is distributed in Mediterranean 
Europe and North Africa (Muller et al., 2017; Troia et al., 
2024 and reference herein), but most records are known 
from the Western Mediterranean. The remote records 
from West Siberia, Iran, and Pakistan (Groves, 1924; 
Romanov, 2019) can belong to the recently described vi-
cariant species, Tolypella mongolica R.E. Romanov, V.S. 
Vishnyakov, V.Yu. Nikulin et A.A. Gontcharov (Ro-
manov et al., 2024b).
Tolypella nidifica (O. F. Müll.) A. Braun (Figure 3c), 
Montenegro, Ulcinj Municipality, Nature Park “Ulcin-
jska Solana”, brackish temporary ponds in abandoned 
crystallization basins, at the depth up to 10 cm and 
spread over wet clayish soil in a terrestrial environment 
after water evaporation: 1–3. 41.92731 N 19.27498 E, 
41.92656 N 19.27403 E, 12.04.2023; 41.92088 N 
19.26240 E, 28.04.2023.
Annual populations of Tolypella nidifica appearing 
during spring only were spotted in a few plain brackish 
shallow temporary water bodies in Ulcinjska Solana. The 
occurrence in only temporary ponds can be evidence of 
the high susceptibility of this species to interannual dif-
ferences in weather and climate change. This species can 
be referred to “Critically Endangered” (IUCN criteria 
B2a,b, C2a, D) in Montenegro. It was recognized as 
“Critically Endangered” in the Red List of Balkan charo-
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New charophytes for Montenegro, Croatia, and Cyprus
phytes (Blaženčić et al., 2006b) and “Least Concern” in 
the provisional European Red List (Stewart, 2024).
In Southeast Europe, Tolypella nidifica is known from 
a few records from Croatia only (Figure 2g; Visiani, 
1842; Lovrić et al., 1989; Lambert & Schubert, 2024b). 
Some doubts are associated with the old record by Vi-
siani (1842) because his specimen from the same location 
was later cited for Nitella opaca (C. Agardh ex Bruzelius) 
C. Agardh (Leonhardi, 1864). Tolypella nidifica occurs in 
coastal regions of Europe, but it has the main distribu-
tion center in the Baltic Sea region (Lambert & Schubert, 
2024b and reference therein). North Africa, Caucasus, 
and Lake Issyk Kul in Kyrgyzstan harbor very few popu-
lations of Tolypella nidifica outside of Europe (Kausik & 
Bhattacharya, 1971; Hollerbach & Krassavina, 1983; Ro-
manov et al., 2023; Romanov, unpubl. data). 
Conclusion
This research targeting all habitat types suitable for cha-
rophytes was implemented mostly in the Montenegrin 
Adriatic coast considering it was the least studied area 
of the country from a charophyte perspective (Blaženčić 
et al., 1998; Biberdžić et al., 2006; Biberdžić & Blaženčič, 
2013; Blaženčič & Stevanović, 2015; Blaženčić et al., 
2018; Biberdžić et al., 2023). It resulted in new national 
records of seven species: Chara corfuensis, Chara galioides, 
Chara squamosa, Sphaerochara intricata, Tolypella hispani-
ca, Tolypella nidifica and “Mediterranean Chara”, mostly 
typical for the Mediterranean and rare outside this region. 
Chara squamosa was found for the first time in Cyprus. 
The checking of herbarium collections revealed one new 
national record for Croatia, Chara galioides, and two old 
neglected sites of Chara galioides and Tolypella hispanica 
for Greece. In Montenegro, all species were found only 
in the southern part in plain temporal and permanent 
small water bodies, except for Chara squamosa, which was 
also found in the mountainous southern and northern re-
gions in small water bodies. The habitats of all species fit 
the environment known to them (Schubert et al., 2024). 
New locations enriched charophyte species distribution 
data for Southeast Europe adding new sites of regionally 
rare species. They were used in the tentative assessment of 
species protection statuses. They will be helpful for future 
evaluation of important charophyte areas. However, the 
territory of the Nature Park “Ulcinjska Solana”, the east-
ernmost vicinities of Velika plaža, and the coastal wetland 
southern of Buljarica can already be recognized as im-
portant charophyte areas in Montenegro and Southeast 
Europe. The updated species list of Montenegrin charo-
phytes counts 37 species making Montenegro one of the 
charophyte species-rich countries in Europe.
Acknowledgments 
The curators of the herbaria listed above are kindly ac-
knowledged for their encouragement and guidance, and 
the Natural History Museum of Montenegro – for the 
access to laboratory equipment and materials for charo-
phyte identification. Martin Dobrota is acknowledged 
for specimen collection in Cyprus. Mario Klesczewski is 
acknowledged for his kind help with recognizing nearly 
illegible hand-written labels as well as Vedran Šegota, 
University of Zagreb, for providing some references. The 
suggestions by anonymous reviewers and Urban Šilc are 
really appreciated.
Funding
The investigations were supported by the project Bio-
Mon – Establishing Biodiversity Monitoring in Ulcinj 
Salina, a part of the IKI Small Grants program. IKI Small 
Grants is carried out by Deutsche Gesellschaft für Inter-
nationale Zusammenarbeit (GIZ) GmbH and part of the 
International Climate Initiative (IKI). The project is im-
plemented by the Center for Protection and Research of 
Birds (CZIP), BirdLife partner in Montenegro, (permit 
No. 02-UPI-798/6). The opinions put forward in this pa-
per are the sole responsibility of the authors and do not 
necessarily reflect the views of the Federal Ministry for the 
Environment, Nature Conservation and Nuclear Safety 
(BMU) or of Deutsche Gesellschaft für Internationale 
Zusammenarbeit (GIZ) GmbH.
ORCID iDs
Roman Romanov  https://orcid.org/0000-0002-6137-3586
Snežana Dragićević  https://orcid.org/0000-0002-2788-5211
František Bednár  https://orcid.org/0009-0003-4209-0155
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New charophytes for Montenegro, Croatia, and Cyprus
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