The genus Odontarrhena in Albania: disentangling the
systematics of a critical group of Ni-hyperaccumulators in a
European serpentine hot-spot
F. Selvi, L. Cecchi, I. Colzi, A. Coppi, G. Echevarria and C. Gonnelli
The genus Odontarrhena
C.A. Mey., 1831• A monophyletic group of
Alysseae, not closely related to
Alyssum !
• 87 sp. (Alybase, Španiel et al.,
Pl. Syst. Evol. 301, 2015).
• Major synapomorphies: ovule solitary in each locule of the ovary, branched inflorescence
Range, edaphism and metal accumulation ability
Introduction
• Europe and Mediterranean, W Asia
• Ultramafic substrates (serpentinite and others), limestone, dolomite, schist…
Excluders Hyperaccumulators
Indicators
• Physio-ecological synapomorphy: Ni-hyperaccumulation ability. All taxa/populations that grow on heavy-metal rich soils have > 1000 μg of Ni g-1 leaf dry weight)
Introduction
Taxonomy, Systematics, Phylogenetic relationships:
is everything clear ?
No, main causes:
•population phenotypic plasticity
•weak cohesion between morphological, ecological or
geographical variation
•relatively homogeneous karyotype features
•low phylogenetic divergence
uncertain species limits, lack of taxonomic consensus between
authors
Introduction
Critical species complexes in Europe
• O. serpyllifolia (Desf.) Jord. & Fourr.: W Mediterranean, Iberian peninsula, facultative serpentinophyte
• O. muralis (Waldst. & Kit.) Endl.: Balkan Peninsula, facultative serp.
Albania:
a poorly known diversity «hot spot»
O. serpyllifolia s.l.
non-serpentine pop. serpentine pop.
Research questions (Agronickel)
Introduction
Q1: how many and which taxa (species and subspecies) do
occur in Albania?
Q2: what are their phylogenetic relationships and patterns
of population genetic structure and diversity?
Q3: Ni-hyperaccumulation capacity: Which are the most
promising accessions for phytoremediation and/or
Approach
Q1. Search and study of taxonomic literature and herbarium
materials - especially types
Q1. Macromorphology and morphometrics
Q1. Micromorphological analyses (SEM)
Q1. Karyological analyses
Q2. Molecular phylogenetic analyses
Q2. Population genetic analyses
Q3. Metal concentration analyses in plant and soil samples
Q3. Experiments in controlled conditions (hydroponics)
Field sampling:
•Geographic: as many populations as possible across Albania (all major
ultramafic areas)
•Taxonomic: type localities of all taxa (species and infraspecific units)
described from Albania:
18
taxa
Methods E.J. Nyárády 1927 1928 1929-1930 1949 F. K. Meyer, 2011 F. Markgraf 1926 1927 a,b 1932
Work in progress
23 populations sampled
in 2016 for:
•Herbarium specimens
•Leaf tissue (12 ind.)
•Seeds (when available)
•Whole plants (metal, 5 x pop.)
•Soil samples (5 reps. x site)
Map of sampled populations July 2016 Results - Q1
bal6 bal4 ba5 bal23 ja1 mkg11-1 mkg11-2 mkg12-1 mkg12-2 mo19 mo20 mu mu15 mu17 mu18 mu21 mu24 mu3 mu7 mu8 mu9 muD1 muD2 muD3 rig10 ri13 ri14 sc2-1 sc2-2 sm 22-1 sm16 sm22 -40 -32 -24 -16 -8 8 16 24 32 Component 1 - 54.5% -20 -16 -12 -8 -4 4 8 12 16 C o m p o ne nt 2 2 1 .9 %
Explorative morphometry
PCA based on 20 characters (qualitative and quantitative)
Morphotype 1: O. muralis sensu lato.
Close to typical O. muralis from type locality (Romania), but: less densely
pubescent stems and leaves; glabrous to nearly glabrous silicles (variable shape)
O. muralis, type locality
= O. chalcidica
(Janka) Španiel, Al-Shehbaz, D.A.German & Marhold
Pop. from N Albania
No appreciable differentiation (continuous variation) from populations of type
localities of:
bertolonii ssp. scutarina (Renc)
markgrafii (Guri Pishkashit)
SEM: stellate hairs on mid part of flowering stem
jancheni (Renc)
markgrafii murale Romania
chalcidica Albania
Hypothesis:
Taxa to be reduced as synonyms of
O. chalcidica
Morphotype 2: O. muralis sensu latissimo.
Mirdita region, East of Kruja, Lure. Tall and robust; sterile basal shoots abundant, large, spathulate, rounded leaves; late flowering.
Alyssum balkanicum
Nyár., Repert. Spec. Nov. Regni Veg. 27. 1930. Described from Shtamës pass east of Kruja. Type destroyed.
Hypothesis:
Taxon to be recognized at species level: O . balkanica, comb. nov.
= A. lurense F.K. Mey. !
Haussknechtia, 2011
Type (JE) Results - Q1
Morphotype 3: O. smolikana: race with glabrous silicles.
Alyssum smolikanum f. glabrum Nyár.
Repert. Spec. Nov. Regni Veg. 27: 1930. Type locality:
mountains of Lure region.
= A. serpentinicola F.K. Mey. !
Haussknechtia, 2011
Type from Lure (JE) Results - Q1
Morphotype 4. Region of Korçë. Dwarf shrublet with pulvinate habit; narrow basal leaves; simple racemes, no lvs. on flw. stems; silicles with undulate margin; early flw.
A. smolikanum subsp. moravense F.K. Mey.
Haussknechtia Beih. 15: 65. 2011.
Hypothesis:
Taxon to be recognized at species level:
O. moravensis, comb. & stat. nov.
Morphotype 5. Elbasan-Prrenjas region.
Low shrublet with narrowly oblong basal leaves, lvs. greenish; glabrous silicles, early flw.
Alyssum rigidum (Nyár.) Nyár., Feddes Repert. 27: 393. 1930. Type locality: Mt. Shpat (SE Elbasan). Type destroyed.
O. smolikana and O. rigida side by side on Mt. Shebenik, 1300 m
Hypothesis:
Taxon to be recognized at species level:
O. rigida, comb. nov.
Results - Q1
O. balkanica, Gjegjan O. «markgrafii», type loc.
Tetraploids, 2n = 4 x = 32: O. chalcidica, O. balkanica, O. smolikana
O. rigida, Mt. Shpat O. moravensis, Voskopoje
Karyology
seeds with broad wing (0.5 mm)
Diploids, 2n = 2x = 16: O. rigida, O. moravensis
ba23 ba4 ba5 ba6 ja1 mkg11-1 mkg11-2 mkg12-1 mkg12-2 mo19 mo20 mu mu15 mu17 mu18 mu21 mu24 mu3 mu7 mu8 mu9 muD1 muD2 muD3 ri10 ri13 ri14 sc2-1 sc2-2 sm 22-1 sm16 sm22 -40 -32 -24 -16 -8 8 16 24 32 Component 1 -20 -16 -12 -8 -4 4 8 12 16 C o m p o ne nt 2 O. balkanica 2n=32 O. smolikana ssp. glabra 2n=32 O. rigida 2n=16 O. moravensis 2n=16 O. chalcidica 2n=32
Morphometric analysis with first taxonomic interpretation
Results - Q1Results - Q2 alpe borz nebr smol held bert robe sibi troo orbe arge2 chal8 chal9 chal10 chal11 arge1 chal2 chal5 chal13 chal18 chal15 chal17 chal16 chal6 chal7 chal3 chal12 mura6 chal4 mura7 6 Rev 23 Rev 16 Rev 22 Rev 12 Rev teni 10 Rev 19 Rev bald2 bald3 bald1 bald4 tort serp cory1 cory2 eubo frag out dens1 out dens2 alpestris borzaeana nebrodensis
smolikana subsp. smolikana (Greece) heldreichii bertolonii tavolarae sibirica troodi orbelica - Bulgaria argentea - Alps argentea - Alps
chalcidica 18 – Greece («chlorocarpum») muralis - Bulgaria muralis - Greece tenium - Greece chalcidica 2 chalcidica 5 chalcidica 13 chalcidica 9
chalcidica «bertolonii ssp. scutarina» (Renc)
chalcidica 10 chalcidica 11 chalcidica 6 chalcidica 7 chalcidica 3 chalcidica 12 chalcidica 6 chalcidica 4 chalcidica 7 balkanica 1 (Valbona) balkanica 2 (Kruja)
smolikana subsp. glabra 1 (Shebenik) smolikana subsp. glabra 2 (Shtames)
chalcidica «markgrafii» (Guri Pishkashit)
chalcidica 8 rigida (Mt. Shpat) moravense baldaccii 2 baldaccii 3 baldaccii 1 baldaccii 4 tortuosa serpyllifolia corymbosoides 1 corymbosoides 2 euboea fragillima Alyssum densistellatum 1
Alyssum densistellatum 2 outgroup
polyploids 87 75 95 56 83 84 O2 Clade O3 68 Consensus tree from MP analysis of ITS sequences (700 bp)
Phylogeny
O1 Poor resolution ! 65 51 diploids Albanian accessionsResults – Q2
No better results with chloroplast trnL-trnF sequences (645 bp)
Phylogeny
Recent radiation processes in Southern Europe ?
O. troodi
Alyssum lenense (outgroup)
O. cr
O. baldaccii 1
O. «markgrafii» type locality O. baldaccii 2
O. sibirica
O. balkanica Valbona
O. smolikana ssp. glabra Shebenik O. heldreichii
O. euboea
O. moravensis
O. rigida type locality
O. smolikana ssp. smolikana Greece 1 O.smolikanassp. glabra Shtames
O. smolikana ssp. smolikana Greece 2
Albanian accessions
Nickel concentration (X 103 µg g-1 d.w.)
0
0 11 22 33 44 55 O. balkanica, Gjegian
O. balkanica, Fierzë O. balkanica, Kruje, Shtames pass O. chalcidica, Krume-Kukes O. chalcidica, Muhurr-Arras
O. chalcidica, Shupenze
O. chalcidica, Pogradec O. chalcidica, Erseke to Barmash O. chalcidica (bertolonii ssp. scutarina), Renc O. chalcidica, Mt. Shebenik O. chalcidica «markgrafi», Elbasan- Mt. Shpat
O. chalcidica «markgrafi», Librazhd, Pishkash
O. moravensis, Voskopoje O. rigida, Elbasan- Mt. Shpat O. rigida, Librazhd, Pishkash O. rigida ,Mt. Shebenik O. smolikana, Mt. Shebenik O. smolikana, Kruje, Shtames pass
6 6 77 88 99 1010 1111 1212 1313 1414 1515 1616 1717 1818 1919 2020 Results – Q3
Metal content
soil plant (shoots)Next steps
• Complete population sampling and analyses.
• Trace and check type specimens of taxa also from nearby Macedonia
• Resolving the two following names («enigmatic» taxa only known from type collection in E Albania):
A. elatius F.K. Mey.
Haussknechtia Beih. 15: 64. 2011
A. albiflorum F.K. Mey.
Thank you !
The Albania 2016 team
Systematics and biodiversity:
active research fields at the base of other
related research topics (metal physiology,
community ecology, conservation biology …).
Need for more data and information.
Whiting et al., Rest. Ecol. 2004
“It is essential that geobotanical reconnaissance be initiated on
metal-rich soils. Good taxonomic skills are crucial, as are the use
of robust ecological techniques. There is an urgent need for more
precise geobotanical explorations, notably to elucidate the
specific links between the plants and their native substrates”.
Three contributions
•Plant diversity on serpentines of Ussangoda (Iqbal et al., also lichens)
•Serpentine Flora of Turkey (Kurt et al.). IUCN categories of the endemic and rare taxa: EX 1; CR 43; EN 35; VU 33
•Endemism in the flora of the Griqualand manganese fields, SA (Siebert et al.).
Important centre of plant endemism. Considerable pressure from mining activities and population growth. Protection needed.
