PhytoKeys 103: 61–82 (2018)
A peer-reviewed open-access journal
Checklist of gypsophilous vascular flora in Italy
doi: 10.3897/phytokeys.103.25690
61
RESEARCH ARTICLE
http://phytokeys.pensoft.net
Launched to accelerate biodiversity research
Checklist of gypsophilous vascular flora in Italy
Carmelo Maria Musarella1,*, Antonio Jesús Mendoza-Fernández2,*,
Juan Francisco Mota2,*, Alessandro Alessandrini3, Gianluigi Bacchetta4,5,
Salvatore Brullo6, Orazio Caldarella7, Giampiero Ciaschetti8, Fabio Conti9,
Luciano Di Martino8, Amedeo Falci10, Lorenzo Gianguzzi11, Riccardo Guarino12,
Aurelio Manzi13, Pietro Minissale6, Sergio Montanari14, Salvatore Pasta15,
Lorenzo Peruzzi16, Lina Podda4, Saverio Sciandrello17, Leonardo Scuderi18,
Angelo Troia12, Giovanni Spampinato1,*
1 Department of AGRARIA, Mediterranean University of Reggio Calabria, Loc. Feo di Vito, 89122 Reggio
Calabria, Italy 2 Department of Biology and Geology, CECOUAL, University of Almeria, Desp. 1.101. Edificio
Cientifico Tecnico II – B. Ctra. Sacramento s/n. La Cañada de San Urbano, 04120 Almeria, Spain 3 Regional
Institute for Cultural Heritage. Via Galliera 21 40121 Bologna Italy 4 Centre for the Conservation of Biodiversity
(CCB), Life and Environmental Sciences Department, University of Cagliari,Viale S. Ignazio da Laconi 1113, 09123 – Cagliari, Italy 5 Hortus Botanicus Karalitanus (HBK), University of Cagliari, Viale S. Ignazio
da Laconi 9-11, 09123 Cagliari, Italy 6 Department of Biological, Geological and Environmental Sciences,
University of Catania, Via A. Longo 19, 95125 Catania, Italy 7 Via Maria SS. Mediatrice n° 38, 90129
Palermo, Italy 8 Majella National Park, Via Badia 28 I-67039 Sulmona (AQ), Italy 9 Center of floristic research
of the Apennine – School of Biosciences and Veterinary Medicine, University of Camerino, Italy 10 Via Libertà
n° 200, 93100 Caltanissetta, Italy 11 Department of Agricultural and Forest Sciences. Università degli Studi di
Palermo, Italy 12 Department STEBICEF, Botanical Unit. University of Palermo, Italy 13 Via Peligna n° 228,
66010 Gessopalena (CH), Italy 14 SSNR (Società per gli Studi Naturalistici della Romagna), C.P. 143, 48012
Bagnacavallo (RA), Italy 15 Department of Biology, University of Fribourg, Chemin du Musée 10, CH-1700,
Fribourg, Switzerland 16 Department of Biology, Unit of Botany, University of Pisa, Via Derna 1, 56126, Pisa,
Italy 17 Centre for the Conservation and Management of Nature and Agroecosystems (CUTGANA). University
of Catania, Italy 18 Via Andromaca n° 60, 91100 Trapani, Italy
Corresponding author: Carmelo Maria Musarella (carmelo.musarella@unirc.it)
Academic editor: D. Geltman | Received 10 April 2018 | Accepted 27 June 2018 | Published 18 July 2018
Citation: Musarella CM, Mendoza-Fernández AJ, Mota JF, Alessandrini A, Bacchetta G, Brullo S, Caldarella O,
Ciaschetti G, Conti F, Martino L, Falci A, Gianguzzi L, Guarino R, Manzi A, Minissale P, Montanari S, Pasta S, Peruzzi
L, Podda l, Sciandrello S, Scuderi L, Troia A, Spampinato G (2018) Checklist of gypsophilous vascular flora in Italy.
PhytoKeys 103: 61–82. https://doi.org/10.3897/phytokeys.103.25690
*
These authors contributed equally to the writing of this paper, the project and the first list of plants,
coordinating the group, elaborating the data and editing the text. The other authors contributed by
completing and evaluating the plant list.
Copyright Carmelo Maria Musarella et al. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
Abstract
Our understanding of the richness and uniqueness of the flora growing on gypsum substrates in Italy
has grown significantly since the 19th century and, even today, new plant species are still being discovered. However, the plants and plant communities, growing on gypsum substrates in Italy, are still a
relatively unknown subject.
The main aim of this paper was to elaborate a checklist of the Italian gypsophilous flora, to increase
knowledge about this peculiar flora and for which conservation efforts need to be addressed.
Through a structured group communication process of experts (application of the Delphi technique),
a remarkable number of experienced Italian botanists have joined together to select focal plant species
linked to gypsum substrates. From the results obtained, 31 plant species behave as absolute or preferent
taxa (gypsophytes and gypsoclines) and form the ‘core’ Italian gypsophilous flora. The most abundant life
forms were chamaephytes and hemicryptophytes, belonging to Poaceae and Brassicaceae; as for chorotypes,
the most represented are Mediterranean and narrow endemics. By improving on previously available information about the flora with a clear preference for gypsum in Italy, this undertaking represents an important
contribution to the knowledge of a habitat which is today considered a priority for conservation.
Keywords
Edaphism, Gypsophyte, Habitats Directive, Plant preservation
Introduction
The relationship between local bedrock types and vegetation cover has long been highlighted. Andrea Cesalpino, in De plantis libri XVI (1583), had already documented
the existence of endemic plant species on the Italian serpentines. The term ‘edaphism’
– interpreted as a ‘geobotanical phenomenon giving rise to particular floras on certain
substrates’ (Font Quer 1977) or ‘those physical and chemical effects induced on living
beings by the soil’ (Sarmiento 2001) – has been used extensively in Europe since the
19th century (Parsons 1976, Kruckeberg 2002). The species and the plant assemblages
growing on gypsum provide a clear example of the strict relationship between soil and
vegetation, as many plant species grow exclusively or preferentially on such peculiar
substrates. This geobotanical pattern occurs in more than 70 countries worldwide (Pérez-García et al. 2017). Nevertheless, the gypsicolous substrates represent a largely underrated or ignored habitat, with serious consequences for both flora and fauna conservation. These habitats host sparse and scattered vegetation, since the gypsum outcrops
often represent geological islands interrupting the uniformity of other surrounding
landscapes. Local evolutionary processes have probably been facilitated by the geographical isolation of outcrops, so that the appearance of several plant species with a
narrow distribution have been favoured (Moore et al. 2014). This kind of geographic
speciation could have been complemented by some selective pressure that might have
favoured the survival of certain plant lineages on nutrient-unbalanced, water-limited
soils which are unfavourable to the establishment of most plants (Merlo et al. 1998,
2001, Bolukbasi et al. 2016).
Checklist of gypsophilous vascular flora in Italy
63
There is an abundance of accurate information concerning gypsophilous plant communities in Spain (Mota et al. 2011), but not for those other European countries where
such substrata also occur. In Cyprus, for instance, it is only the outcrop located at Rizoelia National Park that are known, but unfortunately it seems that both local flora and
vegetation are degraded due to several impact factors (EIONET 2008). The first scholar
to study the gypsophilous flora of Italy was Macchiati (1888). There have been some
studies in mainland Italy and Sicily which have increased our knowledge about this flora,
e.g. Gallo (2014). Such previous studies on both the rate of endemism and risk of extinction amongst these plant communities pointed out that ‘Mediterranean gypsum vegetation’ should be considered a Priority Habitat (*1520) for conservation according to the
92/43 EU Directive (Anonymous 1992). Since no specific national law has yet been enacted in Italy (Fenu et al. 2017, Rossi et al. 2016), this directive theoretically represents
the major instrument for plant conservation. Several Italian gypsum outcrops have been
identified as SCI (Sites of Community Importance) and SCZ (Special Conservation
Zones) e.g. Piano di Gestione Complessi Gessosi M. Conca, SIC ITA050006 (IUCN
UNEP-WCMC 2014, Mento 2008). However, information about gypsum vegetation
in Italy is still deficient: in fact, neither its current area, nor the recent trends in term of
quantity and quality (last 50 years) are known (Loidi 2016). An accurate checklist of
Italian gypsophilous flora has not yet been produced; hence, only published investigations concerning these flora and vegetation could be used as a reference guide. When
taking into account that conservation policies at any level must be based on scientific
assessments about habitats, species conservation status and existing threats (IUCN 2001;
Joppa et al. 2013), the exhaustive knowledge about the flora typical of gypsum outcrops
in Italy would be a crucial step towards the application of any sort of conservation measure (Mota et al. 2004, 2011, Martínez-Hernández et al. 2011, 2015).
Therefore, the aims of this research were (i) to elaborate a checklist of Italian gypsophilous vascular flora through a structured group communication process of experts;
(ii) to expand the knowledge of this flora type to which conservation efforts need to be
addressed; (iii) to examine the spectrum of taxonomical groups, life forms and chorotypes of this flora. Through this approach, the comparison between the gypsophilous
flora of Italy and that of other countries was carried out in order to detect common
phylogenetic, functional and biogeographic patterns that allow a better understanding
of the gypsophily phenomenon at European and global levels.
Methods
Several approaches have been proposed to elucidate which plant species can be considered as best linked to gypsum substrates (Mota et al. 2016). However, coping with an
extremely species-rich flora over a wide territory such as Italy, using the Delphi technique (Hasson et al. 2000) resulted in being the most effective way to build a checklist
of gypsophilous flora by using the so-called ‘expert criterion’ (Mota et al. 2008, 2009).
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
The Delphi technique
The Delphi technique is a structured, anonymous and iterative survey undertaken by a
panel of ‘experts’, which enables a group of individuals to collectively address a complex
problem through a structured group communication process. This method has been
applied in ecology to fill in data gaps (Eycott et al. 2011), through the experience of the
participants (Ochoa-Gaona et al. 2010). The technique can be classified into four categories relevant to ecology and conservation (Hasson and Keeney 2011). One of these
categories is Decision Delphi, which is used to identify focal species for conservation.
Our scheme comprised two rounds of semi-structured questionnaires, each followed by an aggregation of responses and anonymous feedback from the experts. The
number of rounds was limited and adapted according to the time available. An increased number of rounds would make the process more time-consuming.
Preparation of the first round of the questionnaire
A semi-structured survey, drawing from evidence based on published literature, was
designed. The initial listing of taxa included species issuing from bibliographical references, which recorded the presence of these taxa on gypsum substrates (See Appendix 1:
Methodology References for detailed information).
Selection and invitation of a panel of experts
Participants from a great diversity of backgrounds were included (e.g. teachers, scientists, conservationists, non-governmental organisations, policy-makers, environmental managers and technicians) in order to obtain a wide range of perspectives and
minimise bias arising from self-interest or information preferences. The participants
included the co-authors of this article, i.e. experts situated in the Italian peninsula
and Sardinia – hereinafter Italy (9) – and in Sicily (11). The reason why the number
of Sicilian botanists involved is greater than those from mainland Italy is due to the
considerable extension and importance that gypsum outcrops have in Sicily (Fig. 1).
Collection and analysis of the completed questionnaire for the first round
Once the preliminary plant catalogue was elaborated, the complete list was submitted
to the group of experts (Musarella et al. 2016). The experts were clearly asked to base
their gypsophily assessment for each plant species only on their personal field experience in order to avoid any judgement based on bibliographical references or other
sources of information. In addition, the peers were provided with a series of hierarchical criteria according to the Likert (1932) scale, a method where participants were
asked to rank their responses on a scale of ‘one to five’ (Table 1) where ‘one’ indicated
Checklist of gypsophilous vascular flora in Italy
65
Figure 1. Italian gypsum outcrop presences in 10×10 km UTM grids.
Table 1. Likert scale ranking for the gypsophilous character of the taxa.
5
Strictly gypsophile species; that is, species that do not live outside gypsum
substrates (except accidentally). ALWAYS GROW ON GYPSUM
Strict gypsophyte
4
Species with great preference for gypsum and which are found very rarely outside
this substrate. ALMOST ALWAYS GROW ON GYPSUM
Preferential
gypsophyte
3
Species that live on gypsum, but which can also live on other substrates. If they live
on many other different types of soil, they will not fit into this category. For example,
if they live on limestone, marls and gypsum they could fall into this category. At
least, it is as abundant (or almost) on gypsum as it is on other types of substrates.
GYPSUM AND OTHER HIGHLY RELATED SUBSTRATES ARE THEIR
PREFERRED HABITATS
Subgypsophyte
2
Species that may be abundant on gypsum, although they could be even more
frequent on other types of substrates.
CLEARLY MORE COMMON OUTSIDE OF GYPSUM
Gypsovag
1
Very rare species on gypsum or absent on this type of soil.
NEVER (OR ONLY ACCIDENTALLY) ON GYPSUM
Accidental
an ‘absolute absence on gypsum outcrops (plants that avoid gypsum or, at most, may
eventually occur on this substrate)’, while ‘five’ indicated ‘absolute preference (the considered plant species only grows on gypsum)’.
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
Preparation and analysis of second round questionnaire
The collated responses from the first round were used to prepare a second questionnaire.
The experts were requested to add new taxa candidates to be subsequently evaluated by the
panel (Spampinato et al. 2016). The second round questionnaire was administered only
to respondents who participated to the first round. The responses were collated and analysed using quantitative measures. Statistical summaries were generated for the responses,
central tendencies (mean, median) and the levels of dispersion (standard deviation and
interquartile ranges) of each taxon. The results were compiled into a report, which was
used in the next step as feedback and described quantitative details to the participants.
Plant species data
Information about plant species included in the checklist was collected: (i) taxonomic rank (Bartolucci et al. 2018); (ii) family (Peruzzi 2010); (iii) chorology and
(iv) life form (Pignatti 1982); (v) conservation status (Rossi et al. 2014); and (vi)
functional groups (narrow gypsophiles, wide gypsophiles or gypsovags) according
to Bolukbasi et al. (2016) and Palacio et al. (2007). See Suppl. material 1 for complete information.
Checklist data analysis
Considering that the plant species assessment was made with a quantitative but discontinuous scale, median values could be useful criteria for selecting the gypsophilous
species (Mota et al. 2011). According to the proposed Likert scale, species like gypsoclines (subgypsophytes or preferential gypsophytes) with median values 3 and 4 could
be considered; and those with median values >4, such as gypsophytes.
Several statistical t-student and ANOVA tests were performed (SPSS ver. 22.0.0.0.
IBM SPSS Statistics). Only taxa with gypsophily median values ≥2 were assessed in
order to exclude ‘casual occurring taxa’. This analysis examined statistical differences
between gypsum affinity (gypsophily), functional groups (narrow gypsophile, wide
gypsophile and gypsovag) and distribution (Endemic, Mediterranean s.l., European,
Eurasiatic/Widespread).
Results
Decision Delphi technique
The first round of the questionnaire comprised 115 plant taxa. However, experts included more than 69% of other taxa growing on Italian gypsum substrates. This fact
Checklist of gypsophilous vascular flora in Italy
67
implied that, during the second round questionnaire, the panel of experts made assessments of 380 taxa (Suppl. material 1). All these taxa were assessed by at least one
of the experts. The set of 380 taxa received on average 8.95 valuations out of 20 (i.e.
almost half of the specialists, 44.60%, gave an assessment). The average number of
evaluations from the mainland Italian botanists was 2.29, compared to 6.65 from the
Sicilian botanists.
The species in this catalogue belong to 59 different families. As far as the taxonomic spectrum of the 380 taxa is concerned, the most represented families were Asteraceae (14%), Poaceae (9.5%), Fabaceae (8.4%), Lamiaceae (6.3%) and Orchidaceae
(6.1%). Moreover, the percentages of life forms on this preliminary list were as follows:
therophytes (28.7%), hemicryptophytes (24%), geophytes (17.1%), chamaephytes
(16.3%), nanophanerophytes (7.9%) and phanerophytes (6.1%). According to their
distribution, two groups were clearly highlighted. The first one is composed of species
with Mediterranean distribution (49.5%); the second included Italian endemic species
(16.8%). The rest of the species were grouped (in smaller percentages) under Eurimediterranean, Submediterranean, European and Euroasiatic/Widespread species. Considering the conservation status of the species in the preliminary list, only seven taxa had
IUCN extinction risk assessments and five of them were considered to be threatened
according to IUCN categories: CR [Aizoanthemopsis hispanicum (L.) Klak., Limonium
calcarae (Tod. ex Janka) Pignatti and Astragalus raphaelis Ferro]; EN [Allosorus persicus
(Bory) Christenh.]; VU [Tripolium sorrentinoi (Tod.) Raimondo & Greuter].
Checklist of Italian gypsophilous flora
The consensus, established amongst the responses of the panel of experts, produced the
first checklist of Italian gypsophilous flora. A tiny group of 31 species out of the 380
preliminary taxa (8.16%) obtained median values over 3 from the experts’ assessments,
so that they can be considered as gypsophiles or gypsoclines (Table 2).
The 31 Italian gypsophilous taxa, on average, received 11 evaluations by experts (i.e.
more than 55% of specialists evaluated this group): a result which increased the average
number of assessments that the 380 taxa considered as the preliminary list received by
more than 10%. Specifically, only the 9 species group composed of Chaenorhinum rupestre (Guss.) Speta, Sedum gypsicola Boiss. & Reut. subsp. trinacriae Afferni, Brassica villosa
Biv. subsp. tineoi (Lojac.) Raimondo & Mazzola, Diplotaxis harra (Forssk.) Boiss. subsp.
crassifolia (Rafin.) DC., Erysimum metlesicsii Polatschek, Astragalus caprinus L. subsp.
huetii (Bunge) Podlech, Capparis sicula Veill., Gypsophila arrostii Guss. subsp. arrostii and
Matthiola fruticulosa (L.) Maire subsp. fruticulosa, was evaluated by 11 or more specialists.
Within the evaluation of the Italian Checklist of gypsophilous flora, 8 taxa obtained arithmetic-mean values higher than 4, whilst 8 taxa values were greater than
or equal to 3. In the case of the median calculation, 5 species showed values equal to
5, 8 taxa reached values equal to or greater than 4 and 18 were equal to or above the
median value 3.
Species
Festuca gypsophila Hack.
Chaenorhinum exile (Coss. &
Kralik) Lange
Ctenopsis gypsophila (Hack.)
Paunero)
Sedum gypsicola Boiss. & Reuter
subsp. trinacriae Afferni
Petrosedum ochroleucum (Chaix)
Niederle subsp. mediterraneum
(L.Gallo) Niederle
Allosorus persicus (Bory) Christenh.
Artemisia pedemontana Balb.
Stipa austroitalica Martinovský
subsp. frentana Moraldo & Ricceri
Diplotaxis harra (Forssk.) Boiss.
subsp. crassifolia (Raf.) Maire
Brassica villosa Biv. subsp. tineoi
(Lojac.) Raimondo & Mazzola
Erysimum metlesicsii Polatschek
Limonium catanzaroi Brullo
Limonium optimae Raimondo
Reaumuria vermiculata L.
Gypsophila arrostii Guss.
subsp. arrostii
Cheilanthes persica (Bory)
Mett. ex Kuhn, Notholaena
persica Bory
Diplotaxis crassifolia (Raf.) DC.
Family
Life Choro
IUCN NA Score
form
logy
Me
dian
Mean
SD
Mainland Sicily
Median Median
Plantaginaceae
T
S-Medit.
-
11
55
5.00
5.00
0.00
-
5.00
Poaceae
T
Medit.
-
1
5
5.00
5.00
-
-
5.00
Crassulaceae
Ch
Medit.
-
11
51
5.00
4.64
0.67
-
5.00
Crassulaceae
Ch
Endem.
-
9
38
5.00
4.22
1.30
-
5.00
Pteridaceae
H
Medit.
EN
5
21
5.00
4.20
1.10
5.00
3.00
Asteraceae
Ch
Europ.
-
2
9
4.50
4.50
0.71
5.00
4.00
Poaceae
H
Endem.
LC
5
21
4.00
4.20
0.84
4.50
4.00
Brassicaceae
Ch
S-Medit.
-
11
45
4.00
4.09
0.94
-
4.00
Brassicaceae
Ch
Endem.
-
11
43
4.00
3.91
1.30
-
4.00
Brassicaceae
Plumbaginaceae
Plumbaginaceae
Tamaricaceae
Caryophyllaceae
H
H
H
NP
Endem.
Endem.
Endem.
S-Medit.
-
11
5
5
4
40
18
18
13
4.00
4.00
4.00
4.00
3.64
3.60
3.60
3.25
0.92
1.67
1.67
1.50
-
4.00
4.00
4.00
4.00
Ch
Endem.
-
11
36
3.00
3.27
0.90
-
3.00
Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
Chaenorhinum rupestre (Guss.)
Speta
Synonym
68
Table 2. Checklist of Italian gypsophilous flora. Species are listed in decreasing order of Median. Life-form: Therophyte (T), Chamaephyte (Ch), Hemicryptophyte (H), Nanophanerophyte (NP), Geophyte (G). Distribution: Endemic (Endem.), Mediterranean (Medit.), Sub-Mediterranean (S-Medit.), European (Europ.),
Widespread (Wide.). IUCN category: Endangered (EN), Least Concern (LC) species. Number of assessment (NA). Median (Median) and average (Mean) values of
experts’ assessments. Standard deviation (SD). Median values from mainland Italy and Sicilian experts separately (Mainland Med/Sicily Med).
Species
Capparis sicula Veill.
Teucrium luteum (Mill.) Degen
Parapholis incurva (L.) C.E.Hubb.
Me
dian
Mean
SD
Mainland Sicily
Median Median
Ch
Endem.
-
1
3
3.00
3.00
-
-
3.00
Amaryllidaceae
G
Europ.
-
1
3
3.00
3.00
-
3.00
-
Poaceae
G
Europ.
-
1
3
3.00
3.00
-
-
3.00
Apiaceae
H
Medit.
-
4
11
3.00
2.75
0.50
-
3.00
Matthiola tristis (L.) R.Br.
Brassicaceae
Ch
Europ.
-
11
30
3.00
2.73
0.90
2.00
3.00
Ammi crinitum Guss.
Apiaceae
T
Endem.
-
5
13
3.00
2.60
0.89
-
3.00
Poaceae
Linaceae
H
T
Medit.
Medit.
-
7
9
18
23
3.00
3.00
2.57
2.56
0.79
0.53
-
3.00
3.00
Asteraceae
Ch
Medit.
-
9
23
3.00
2.56
0.88
3.00
2.50
Fabaceae
H
Endem.
-
11
28
3.00
2.55
1.13
-
3.00
Capparaceae
NP
Medit.
-
11
28
3.00
2.55
0.82
1.00
3.00
Lamiaceae
Ch
Medit.
-
7
16
3.00
2.29
0.95
-
3.00
Poaceae
H
S-Medit.
-
9
20
3.00
2.22
0.97
3.00
2.50
Apiaceae
H
Medit.
-
5
11
3.00
2.20
1.10
-
3.00
Poaceae
T
Medit.
-
5
11
3.00
2.20
1.10
1.00
3.00
Amaranthaceae
NP
Wide.
-
5
11
3.00
2.20
1.10
1.00
3.00
Poaceae
T
Medit.
-
7
15
3.00
2.14
1.07
1.00
3.00
Elymus obtusiflorus (DC.)
Conert
Capparis spinosa L. subsp. spinosa var. canescens Coss.
Teucrium polium L. subsp. aureum (Schreb.) Arcang.
Hippomarathrum siculum (L.)
Hoffm. & Link
Parapholis strigosa (Dumort.)
C.E.Hubb.
Suaeda vera J.F.Gmel..
Life Choro
IUCN NA Score
form
logy
Brassicaceae
Lygeum spartum L.
Cachrys sicula L.
Family
Suaeda fruticosa (L.) Forssk.
(auct. Fl. Ital.)
Lepturus incurvus (L.) Druce
Checklist of gypsophilous vascular flora in Italy
Matthiola fruticulosa (L.)
Maire subsp. coronopifolia (Sm.)
Giardina & Raimondo
Allium moschatum L.
Elymus elongatus (Host)
Runemark subsp. elongatus
Thapsia meoides (Desf.) Guss.
Matthiola fruticulosa (L.)
Maire subsp. fruticulosa
Visnaga crinita (Guss.)
Giardina & Raimondo
Stipa barbata Desf. subsp. barbata
Linum decumbens Desf.
Phagnalon rupestre (L.) DC.
subsp. illyricum (H.Lindb.) Ginzb.
Astragalus caprinus L.
subsp. huetii (Bunge) Podlech
Synonym
69
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
The plant species on the Italian Checklist are present in 16 families (Figure 2
and Table 3), amongst which the most abundant are Poaceae (22.6%), Brassicaceae
(16.1%), Apiaceae (9.7%), Asteraceae, Crassulaceae and Plumbaginaceae (6.5%). Interestingly, these 6 families alone represented almost 70% of the total gypsophilous
species. As regards life forms, the spectrum is as follows: chamaephytes (32.26%),
hemicryptophytes (32.26%), therophytes (19.35%), nanophanerophytes (9.67%) and
geophytes (6.45%). In terms of the distribution analysis, the Italian Checklist highlights exactly the same two groups of the initial list, with those species with Mediterranean distribution predominating (38.7%), followed by the group composed of Italian
endemic species. In the case of this last group, the species percentage was double that
in the same analysis on the preliminary list (32.3%) (Figure 2).
When the evaluations of the Sicilian experts are compared with those of experts
from the peninsula (Table 2), the latter consider 23 species with gypsophily values
higher than 3, although only 6 of them were included in the final checklist (19.4%).
These species are Artemisia pedemontana Balb., Allosorus persicus (Bory) Christenh., Stipa austroitalica Martinovský subsp. frentana Moraldo & Ricceri, Allium moschatum L.,
Lygeum spartum L. and Phagnalon rupestre (L.) DC. subsp. illyricum (H.Lindb.) Ginzb.
Table 3. Percentage of gypsophile taxa grouping by taxonomic families and a comparison between Italian
and Spanish Checklists (Mota et al. 2011).
Family
Amaryllidaceae
Apiaceae
Asteraceae
Brassicaceae
Campanulaceae
Capparaceae
Caryophyllaceae
Amaranthaceae
Cistaceae
Crassulaceae
Euphorbiaceae
Fabaceae
Frankeniaceae
Gentianaceae
Lamiaceae
Linaceae
Orobanchaceae
Plantaginaceae
Plumbaginaceae
Poaceae
Primulaceae
Pteridaceae
Resedaceae
Tamaricaceae
Italian Checklist
3.23
9.68
6.45
16.13
–
3.23
3.23
3.23
–
6.45
–
3.23
–
–
3.23
3.23
–
3.23
6.45
22.58
–
3.23
–
3.23
Spanish Checklist
1.41
1.41
14.08
12.68
1.41
–
8.45
–
4.23
1.41
1.41
9.86
1.41
1.41
11.27
–
1.41
5.63
12.68
4.23
1.41
–
4.23
–
Checklist of gypsophilous vascular flora in Italy
71
Figure 2. Percentage of taxa grouping by taxonomic families, life-forms and distribution and comparison
between gypsophilous flora and preliminary data.
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
The Sicilian experts considered 32 taxa with gypsophily values equal to or greater than
3 and 28 of these species are present in the final checklist (90.3%).
Finally, in the case of Sardinia, where few gypsum outcrops are located in the
northwest and in southeast of the island, local experts have highlighted the presence of
three taxa (Euphorbia pithyusa L. subsp. pithyusa, Helichrysum italicum (Roth) G.Don
subsp. tyrrhenicum (Bacch., Brullo & Giusso) Herrando, J.M.Blanco, L.Sáez & Galbany and Teucrium marum L. subsp. occidentale Mus, Mayol & Rossellò) which reached
gypsophily values between 1.86 and 2.33 in the experts’ assessments.
Statistical analysis
Statistical tests (t-student) showed significant differences between the groups of Italian
endemic plants versus species with a wider distribution; the Italian endemics obtained a
higher average value of gypsophily (Table 4). However, when comparing the gypsophily
values and groups of taxonomic family or life form, no significant difference was found.
In addition, when considering the clustering performed by distribution, the
ANOVA analysis showed the existence of significant differences in gypsophily values
between the Italian endemics and those showing both Mediterranean and European
distribution. This is not so for species with a wider distribution range, such as Eurasian
and widespread taxa. However, this result could be an artefact due to the small size of
this sample, since its average gypsophily value is the one that most differed from the
endemic species group (Table 5a).
Finally, according to the grouping variables narrow gypsophile, wide gypsophile
and gypsovag, the species considered as narrow gypsophile, showed on average the
highest gypsophily values and reached maximum values. Both species regarded as narrow gypsophile and wide gypsophile showed statistically significant results with higher
gypsophily values than those considered as gypsovags. Nevertheless, there were no significant differences between the groups of narrow gypsophile and wide gypsophile, so
that this separation was not supported (Table 5b).
For further information about the statistical analyses performed see Suppl. material 2.
Discussion
Ad hoc investigations on gypsophily have been performed in only 12 countries and only
five of these studies approached a functional perspective (Mota et al. 2016). However,
documented gypsophilous flora can be found in at least 75 countries (Pérez-García et
al. 2017, 2018).
This work provides the first Checklist of Italian gypsophytes, including 31 taxa
showing a great affinity for this substrate, 12 of which can be unequivocally considered
as strictly gypsophytes. In addition, a number of further species often found on these
substrates is detailed. As mentioned before, although the studies on gypsophilous flora
Checklist of gypsophilous vascular flora in Italy
73
Table 4. t-student analyses by gypsophily level, taxa grouping by endemic and non-endemic species.
Number of species (N). Average (AV). Standard deviation (SD). Standard error (SE).
Endemism
Rest
Total
N
AV
SD
SE
p-value
46
293
339
2.4000
2.1460
2.1810
0.7731
0.4744
0.5321
0.1153
0.0285
0.0296
0.0030
95% confidence interval
Min
Max
2.1680
2.6320
2.0900
2.2020
2.1230
2.2390
* p-value < 0.05
Table 5. ANOVA analysis by gypsophily level. Average (AV). Standard deviation (SD). 5a) Grouping by
distribution: Italian endemic, Mediterranean, European and Wide distribution. 5b) Grouping by functional group: narrow gypsophile, wide gypsophile and gypsovag.
a) Chorotype
Endemic
Mediterranean
European
Eurasiat/Widespread
Mediterranean
European
Eurasiat/Widespread
Endemic
European
Eurasiat/Widespread
Endemic
Mediterranean
Eurasiat/Widespread
Endemic
Mediterranean
European
b) Functional group
Narrow gypsophile
Wide gypsophile
Gypsovag
Wide gypsophile
Gypsovag
Narrow gypsophile
Gypsovag
Narrow gypsophile
Wide gypsophile
AV
SD
p-value
0.2358
0.2917
0.3412
0.2358
0.0558
0.1054
0.2917
-0.0558
0.0495
-0.3412
-0.1054
-0.0495
0.0868
0.1038
0.1498
0.0868
0.0774
0.1329
0.1038
0.0774
0.1446
0.1498
0.1329
0.1446
0.0350
0.0270
0.1060
0.0350
0.8890
0.8580
0.0270
0.8890
0.9860
0.1060
0.8580
0.9860
AV
SD
p-value
0.1524
1.6692
-0.1524
1.5168
1.6692
1.5168
0.1014
0.0848
0.1014
0.0596
0.0848
0.0596
0.2910
0.0000
0.2910
0.0000
0.0000
0.0000
95% confidence
interval
Min
Max
0.0120
0.4600
0.0240
0.5600
-0.0460 0.7280
-0.4600 -0.0120
-0.1440 0.2560
-0.2380 0.4490
-0.5600 -0.0240
-0.2560 0.1440
-0.3240 0.4230
-0.7280 0.0460
-0.4490 0.2380
-0.4230 0.3240
95% confidence
interval
Min
Max
-0.0860 0.3910
1.4690
1.8690
-0.3910 0.0860
1.3760
1.6570
-1.8690 -1.4690
-1.6570 -1.3760
* p-value < 0.05
in Italy date as far back as the 19th century (Macchiati 1888), there is no knowledge of
the existence of detailed ecological, functional or phylogeographic studies. After this
first analysis of Italian flora, it would not be possible to discard a second evaluation
based on the information presented in this work and any new information generated
in the future (e.g. Montanari et al. 2016). A new revision should not only take into
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
account the vascular flora, but also the plant assemblages associated with these outcrops. It must be noted that Italy presents a complex natural scenario, with a strong
North-South environmental gradient conditioning the composition and dynamics of
plant communities. In the case of vegetation associated with gypsum outcrops, this
gradient is remarkable and it is impossible to overlook the far greater aridity of the
southernmost regions, which exaggerates the gypsophily phenomenon (Merlo et al.
1998, 2001). This probably explains why the largest contingent of gypsophytes in the
Italian territory is concentrated in Sicily.
This research reinforces the idea that, provided there is no definitive criterion for
establishing whether a species is a gypsophyte or not, the inductive approach based on
‘expert criterion’ is not only plausible, but perhaps the only one possible to establish
the groundwork for future research on gypsophily. To further complicate this scenario,
the same species may have different affinity levels for gypsum substrates in isolated
territories: e.g. Sedum gypsicola subsp. trinacriae shows gypsophily median value of 5
in Italy, but in Spain, the nominal subspecies (S. gypsicola subsp. gypsicola) reached a
median value of 4 in a previous study (Mota et al. 2011).
Both the taxonomical and life form spectra concerning the 31 gypsophytes on the
Italian Checklist are largely in agreement with the data recorded in other areas of the
Mediterranean Basin for this type of substrates (EIONET 2008, Bolukbasi et al. 2016).
As far as the most represented families are concerned, the taxonomic spectrum of the
strictly gypsophilous flora in Italy is similar. Brassicaceae and Poaceae are amongst the
families with a higher number of species, although the latter is slightly over-represented
in Italy. Poaceae occur frequently in very stressful environments (Baskin and Baskin
2000) and, consequently, it is easy to understand why there is a high number of them,
which can be considered as peculiar to gypsum substrates. Other conspicuous families
in the taxonomic spectrum of the Italian gypsophytes are Apiaceae, Asteraceae, Plumbaginaceae and Crassulaceae. All of them are found in arid Mediterranean environments and, therefore, it is not surprising to find them so well represented in Italy.
Although there were variations in the abundance percentages, the most common
families with the greatest match with the grouping occurred for the Spanish gypsophilous flora (Mota et al. 2011), where Asteraceae (14.1%), Brassicaceae and Plumbaginaceae (12.68%) correspond to the larger families. Nevertheless, Poaceae and Apiaceae
obtained greater representation in the Italian gypsophilous flora. On the contrary, Lamiaceae, Fabaceae and Caryophyllaceae families appear to be less common on Italian
gypsum outcrops with respect to other countries (Table 3).
Plant formations linked to gypsum substrates are usually dominated by small
plant species such as chamaephytes or hemicryptophytes, similarly to those which
occur in other Mediterranean areas, although there are exceptions of woody plant
formations growing on gypsum (Pérez-García et al. 2017). The intense exposure to
sunshine for these environments, the shortage of water and nutrient imbalance are
probably responsible for this scenery. In addition, the abundance of therophytes may
be explained by water shortage. Thus, some annual plants can be favoured on gypsum
by drought (Merlo et al. 2001).
Checklist of gypsophilous vascular flora in Italy
75
Conclusions
Efforts to ensure the conservation of Mediterranean gypsophilous vegetation, considered as a Priority Habitat, should be focused on endangered, rare or endemic species,
according to the premises established by the EU. All these efforts cannot be easily undertaken unless it is previously determined which species, out of many hundreds, are
to be given top priority. The approach adopted in this work may help both to focus on
certain species and to detect research and conservation priorities. The high proportion
of Italian endemic species and the geographic rarity component of the flora associated
with Italian gypsum outcrops is an aspect that makes these outcrops very interesting
habitats. However, the degree of threat to the gypsophilous flora in Italy could have
been insufficiently assessed. This fact is confirmed since less than 7% of the gypsophytes of the Italian Checklist have been evaluated under IUCN protocol. As there has
been a prior effort to establish a network of sites for the conservation of nature (Natura
2000 Network), it would be worthwhile to extend this initiative to areas supporting
Italian gypsum outcrops that have biodiversity values worthy of consideration. In order
to achieve this purpose, an in-depth review of the conservation status for both the gypsophilous flora and the natural areas where these substrata occur in Italy is crucial. Data
generated by experts in conservation, for which threat categories of red-listed species
are based, should be incorporated into nature protection Acts (Mendoza-Fernández
and Mota 2016) to ensure the preservation of these sites in Italy.
There are numerous SCI and SCZ including gypsum outcrops and their associated
flora. The question that remains to be clarified is whether they are sufficient to ensure
the conservation of this flora as well as the vegetation linked to this peculiar substrate.
In this examination of gaps in conservation, fauna and other plant groups, such as
lichens and bryophytes, should be integrated (Mota et al. 2011).
At this moment, the existence of endemic and rare flora with remarkable eco-morphological adaptations and the description of new taxa growing on gypsum outcrops
fully justifies the conservation of these outcrops (Gallo 2014). Some Italian gypsum
outcrops are currently under protection, but other areas have not yet been included in
the network of nature reserves. As Panuccio et al. (2017) and Spampinato et al. (2018)
pointed out for Calabria (South-Italy) and Mendoza-Fernández et al. (2014) for Andalusia (southern Spain), arid or semi-arid territories are often under-represented in
protected area networks. This is the case for most Italian gypsum sites. Furthermore,
the checklist of Italian gypsophytes should contribute to a better understanding of the
autoecology and synecology of rare and endemic species and, therefore, to better conservation of the biodiversity associated with gypsum areas in Italy. For example, scrub
communities, typical of Italian gypsum substrates, represent one of the major gaps in
conservation habitats in the European Union. In order to fill this gap, the peculiar
Italian gypsum habitats could be considered by modifying the meaning that habitat
1520* currently has in the manual of habitat interpretation and through the addition
of Italian gypsophytes in order to include the gypsum habitat in Italy, amongst those
of European interest.
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Carmelo Maria Musarella et al. / PhytoKeys 103: 61–82 (2018)
Data provided in this paper denote an important advance in this sense, because
only five plant species in Italy have been recognised as characteristic taxa for this habitat on the European Red List of Habitats (Loidi 2016). These are: Brassica villosa
subsp. tineoi, Chaenorhinum rupestre, Festuca gypsophila (Ctenopsis gypsophila), Erysimum metlesicsii and Sedum gypsicola subsp. trinacriae. However, according to the information provided in this paper, at least twelve species, more than double, could be
indicative plants for this Priority Habitat. Thus, the following should be added to those
already mentioned: Petrosedum ochroleucum subsp. mediterraneum, Allosorus persicus,
Artemisia pedemontana, Diplotaxis harra subsp. crassifolia and Stipa austroitalica subsp.
frentana. In addition, some species typically related to saline soils belonging to the
“Serie gessoso-solfifera” of Sicily, such as Reaumuria vermiculata, Limonium catanzaroi
and Limonium optimae, may also occur.
Acknowledgements
The study results from the international collaboration between the “Mediterranea”
University of Reggio Calabria (Italy) and the University of Almeria (Spain). The authors Antonio Jesús Mendoza-Fernández and Carmelo Maria Musarella were sponsored
by the University of Almeria through the Programme ‘Plan Propio de Investigación y
Transferencia’, with a postdoctoral grant (Contrato Puente 2015) and financial assistance for their stay abroad (Estancias en otros Centros de Investigación 2016); and
with support for guest teachers (Programa de Profesores Invitados 2017).
This study was carried out under the GYPWORLD Project, funded by the European Commission under the European Union’s Horizon 2020 Programme/MSCARISE-2017 Grant Agreement n. 777803.
We thank Beatrice Antolin and Edward Parker for reviewing the English version
of the text.
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Supplementary material 1
Plant species data
Authors: Carmelo Maria Musarella, Antonio Jesús Mendoza-Fernández, Juan Francisco Mota, Alessandro Alessandrini, Gianluigi Bacchetta, Salvatore Brullo, Orazio
Caldarella, Giampiero Ciaschetti, Fabio Conti, Luciano Di Martino, Amedeo Falci,
Lorenzo Gianguzzi, Riccardo Guarino, Aurelio Manzi, Pietro Minissale, Sergio Montanari, Salvatore Pasta, Lorenzo Peruzzi, Lina Podda, Saverio Sciandrello, Leonardo
Scuderi, Angelo Troia, Giovanni Spampinato
Data type: species data
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/phytokeys.103.25690.suppl1
Supplementary material 2
ANOVA analysis by gypsophily level
Authors: Carmelo Maria Musarella, Antonio Jesús Mendoza-Fernández, Juan Francisco Mota, Alessandro Alessandrini, Gianluigi Bacchetta, Salvatore Brullo, Orazio
Caldarella, Giampiero Ciaschetti, Fabio Conti, Luciano Di Martino, Amedeo Falci,
Lorenzo Gianguzzi, Riccardo Guarino, Aurelio Manzi, Pietro Minissale, Sergio Montanari, Salvatore Pasta, Lorenzo Peruzzi, Lina Podda, Saverio Sciandrello, Leonardo
Scuderi, Angelo Troia, Giovanni Spampinato
Data type: statistical data
Explanation note: ANOVA analysis by gypsophily level.
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Link: https://doi.org/10.3897/phytokeys.103.25690.suppl2