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Sonchus parathalassius J.G.Costa ex R.Jardim & M.Seq., a new species from Porto Santo Island (Madeira Archipelago)

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Silva Lusitana 19(2): 233 - 245, 2011
© UISPF, L-INIA, Oeiras. Portugal 233
Editores: jorge.capelo@gmail.com; ana.almeida@inrb.pt
Notas do Herbário Florestal
do INRB (LISFA): Fasc.
XXXIII
∫1. Novarum Flora Lusitana Commentarii
In memoriam A.R. Pinto da Silva
(1912 – 1992)
Sonchus parathalassius J.G. Costa ex R.
Jardim & M. Seq., a new species from
Porto Santo Island (Madeira
Archipelago)
The woody pachycaulous species of
Sonchus L. (Asteraceae) are included in
subgenus Dendrosonchus Sch. Bip. ex
Boulos and are endemic to the
macaronesian archipelagos of Canaries
(14 species), Madeira (3 species) and
Cape Verde (1 species), except for one
species common to Canary Islands and
western cost of Morocco (BOULOS, 1972,
1974; LEE et al., 2005). In Madeira
Archipelago this subgenus was so far
thought to be represented by 4 endemic
taxa, Sonchus fruticosus L. f., S. pinnatus
Aiton, S. ustulatus Lowe subsp. ustulatus
and S. ustulatus subsp. maderensis
Aldridge (PRESS, 1994; JARDIM &
MENEZES DE SEQUEIRA, 2008).
The Madeiran botanist José Gonçalves
da Costa (1899-1967) started to publish in
1946 a commented list of plants of Porto
Santo Island (COSTA 1946, 1948). J. G.
Costa added 54 taxa, including the
description of new species, subspecies
and varieties, based on the observations
and collecting during a 3 years stay on
Porto Santo, between October 1938 and
August 1941. On the introduction of the
first publication COSTA (1946) mentioned
a new species of perennial Sonchus, as
"Sonchus parathalassia sp. n." from Ilhéu
de Baixo, the largest Porto Santo islet,
also named Ilhéu da Cal, but never
published a diagnosis or description of
this new species.
COSTA (1946, 1948) listed species from
20 families (Ranunculaceae to
Leguminosae) and described new taxa
according to the plant list order, Fumaria
muralis Sond. ex W.D.J. Koch var.
lacostrum J.G. Costa in 1946 and Vicia
sativa L. subsp. devia J.G. Costa and Vicia
atlantica J.G. Costa (=V. costae A. Hansen)
in 1948. The same year J.G. Costa left
Madeira to Mozambique, which is
probably the reason why the description
of the perennial Sonchus of Porto Santo
was never published.
J. G. Costa was the first author to refer
a perennial Sonchus for Porto Santo
Island. In fact, LOWE (1868) and
MENEZES (1914) only cited for Porto
Santo S. oleraceus L.
PICKERING (1962) published a check-
list of the flowering plants and ferns of
the Island of Porto Santo and referred
that the name "Sonchus parathalassa"
without description in Costa's list should
be disregarded. Curiously, C.H.
Pickering referred to the finding of an
endemic Sonchus sp., from the Ilhéu das
Cenouras, another Porto Santo islet, that
"await determination". However, this
supposed new species of Sonchus was,
also, never published.
HANSEN (1969) cited for Porto Santo
only Sonchus fruticosus (as S. squarrosus
DC.), whereas BOULOS (1974) referred
solely to Sonchus ustulatus, based on the
specimens collected by C.H. Pickering in
1961, on Ilhéu das Cenouras, deposited
at Kew. ERIKSSON et al. (1974) cited
Sonchus ustulatus and S. fruticosus for
234
Porto Santo Island. Latter ALDRIDGE
(1976) included Porto Santo on the
distribution of Sonchus ustulatus subsp.
maderensis.
HANSEN & SUNDING (1979, 1985,
1993) referred S. fruticosus and S.
ustulatus subsp. maderensis for Porto
Santo synonymizing "Sonchus
parathalassicus Costa" as Sonchus
ustulatus. VIEIRA (1992) cited the same
taxa for Porto Santo but synonymized
"Sonchus parathalassicus Costa" under S.
ustulatus subsp. ustulatus (that he did not
mention for Porto Santo). PRESS (1994)
also assigned the same two taxa to Porto
Santo, not referring the name proposed
by J. G. Costa.
FONTINHA & JARDIM (1999)
confirmed the observations by
PICKERING (1962) about the occurrence
of a perennial branched Sonchus on Ilhéu
das Cenouras, different from those cited
for Madeira Archipelago. JARDIM &
MENEZES DE SEQUEIRA (2008)
considered questionable the existence of
S. fruticosus in Porto Santo.
Field observations in Porto Santo
Island and islets, Madeira and Desertas
and the examination of collected material
as well as herbarium specimens (MADJ,
MADM, MADS, K and BM) allowed the
recognition of a new species of Sonchus
subgen. Dendrosonchus endemic to Porto
Santo distinct from S. ustulatus, S.
fruticosus and S. pinnatus. The putative
Sonchus ustulatus subsp. maderensis and S.
fruticosus from Porto Santo correspond to
this new species. The specimens collected
by J.G. Costa on Ilhéu de Baixo and by C.
Pickering on Ilhéu das Cenouras belong
to the same new taxon. Since J.G. Costa
was the first author to observe this new
Dendrosonchus species and due to the fact
that he named it, although without a
description, the validation of the name
and the description of this new species
from Porto Santo is proposed here as:
Sonchus parathalassius J.G. Costa ex R.
Jardim & M. Seq., sp. nov.
Sonchus parathalassia J.G. Costa in Bol.
Mus. Munic. Funchal, 2: 66 (1946), nom.
nud.
= Sonchus ustulatus sensu auct. fl. Portus
Sancti.
Nanophanerophyto ad usque 90 cm alto a
Soncho ustulato affinis sed differt caule
lignoso robusto superne valde ramoso foliis
ad apicem caule rosulato; inflorescentia erecta
ad usque 60 cm alta ultima tertiam
irregulater ramificans, capitulis plusquam
sexaginta; foliis pennatipartitis anguste
triangularis longe acuminatis lobis
lateralibus 5-9, lobo terminali deltoidis longe
acuminatis vel acuminati-caudatis 30-82
mm longo.
Description: Suffruticose perennial to
perennial shrubby plant (Figure 1A), up
to 90cm. STEM thick, branched, woody,
up to 4 cm diameter. LEAVES (Figure 1B)
28-35 x 8-14cm, fleshy, glabrous, sessile
with a persistent sheathing base,
disposed in rosettes at the tips of the
stems, elliptic-lanceolate to oblong-
lanceolate, pinnatipartite, attenuate to
the base, midrib 5-8 mm attenuate to the
apex; lobes 5-9 pairs, ± opposite to
alternate, 3.5-5.1 x 2.1-3.5cm, narrowly
triangular, long acuminate, entire to
scarcely irregularly dentate; terminal
lobe triangular, acuminate to acuminate-
caudate, 3-8 cm, larger than the upper
pair of lateral lobes; interlobular space
round; caulinar leaves smaller and less
divided. INFLORESCENCE (Figure 1C) a
terminal, upright, branched corymbose
cyme, main axis up to 60cm, branching
usually at the terminal 1/3, 2-3 branched,
Notas do Herbário 235
last order branches with up to 2-4
pedunculate capitula; peduncles green
becoming purplish and broader towards
the obconic apex; inflorescence bracts
triangular, long acuminate, small, 1.6-5.2
mm; CAPITULA (Figure 1D) numerous,
60-90; ovoid-conical, up to 18 x 15 mm;
involucral bracts purplish to yellow-
greenish with yellow-greenish apex;
outer bracts ca. 9, 1.1-2.7 x 1.5-2.0 mm,
small, triangular, base thick, adpressed,
purplish with yellow-greenish margins,
slightly scarious, apex ciliate;
intermediate bracts ca. 6, 5.3-7.6 x 1.3-1.9
mm, oblong, apex yellow-greenish,
ciliate, slightly retroflexed; inner bracts
ca. 20, 10.2-12.7 x 1.1-1.8 mm, linear-
oblong, purplish-yellow to yellow, apex
retroflexed and ciliate. FLORETS 12.6-15.3
mm; ligules 7.0-9.2 x 1.5-2.3 mm, narrow-
oblong, yellow; corolla-tube 4.4-5.0 mm,
white, pubescent. CYPSELA 2.8-2.9 x 0.8-1
mm, oblong-elliptical, compressed
brown. PAPPUS white; bristles 7.7-8.5
mm, scabrid; flexuous hairs 5.0-5.3 mm,
less scabrid, fasciculate.
Types: PORTUGAL, MADEIRA, Porto
Santo: Ilhéu da Cal, Costa, MADM
(holotype, Figure 2); Porto Santo: Ilhéu
de Baixo, VI-1939, Costa, MADS 815
(isotype);
Other specimens: PORTUGAL, MADEIRA,
Porto Santo: Rocha das Cenouras, on
steep volcanic rock facing south, 100 ft,
20-IV-1961, C. Pickering 324, K; Ilhéu das
Cenouras, 9-V-1979, R. Santos, MADJ
5987; Ilhéu da Cal, vertente E no topo N,
14-IV-2004, R. Jardim, MADJ 11014;
Miradouro no Pico Castelo. 6-XII-2008,
M. Sequeira, 5701, UMad; Pico do
Concelho, vertente N, alt.270m, 18-IV-
2011, R. Jardim, 1091, UMad; Pico
Castelo, vertente NW, alt. 240m, 10-VII-
2011, R. Jardim 1215, MA 844022 (UMad);
Pico Branco, Lombo dos Amarelos,
vertente NE, alt. 150m, 13-VII-2011, R.
Jardim 1241, UMad.
Etymology: The specific epithet
"parathalassius" derives from the Greek
Para near and Thalassa sea, presumably
used by J.G. Costa as a reference to the
habitat in sea cliffs. The name Sonchus
parathalassius is given by J.G. Costa in the
specimen here elected as holotype,
although the name invalidly published
(COSTA, 1946) as "S. parathalassia", does
not agree with the gender of the genus.
Distribution: Sonchus parathalassius is
endemic from Porto Santo, occurring on
the main island, in Pico Castelo, Pico
Branco, Pico Concelho and also on the
surrounding islets, namely in Ilhéu da
Cal (or Ilhéu de Baixo), Ilhéu das
Cenouras, Ilhéu de Cima (or Ilhéu do
Farol), Ilhéu de Fora and Ilhéu da Fonte
da Areia.
Habitat: Chasmo-comophyte in rocky
slopes, usually, on north exposed sea
cliffs, in the islets up to 100m (Figure 3),
inland in Porto Santo Island, up to 320m.
Phenology: Flowers from April to July.
Conservation status: This new species
should be considered as Critically
Endangered (CR, C2a(i)), due to the
scarce number of populations, reduced
occupancy and occurrence area as
defined by IUCN (2001), and heavy
grazing by rabbits.
Taxonomic remarks: Sonchus
parathalassius closely resembles S.
ustulatus s.s. but differs by the shrubby
habit up to 90 cm, penatipartite leaves
with almost entire lobes, long
236
acuminated terminal lobe and
inflorescence characters (insertion,
branching, number of capitula). The
shrubby habit mainly observed in the
islets could be related with older plants
less grazed by rabbits. Heavy grazing by
goats in Porto Santo Island until recently
(1995) added to rabbit herbivory, could
explain the very late discovery of this
species. Table 1 summarizes the
diagnostic characters of Sonchus
parathalassius and compares them with
the other Madeiran Dendrosonchus.
Nomenclature: The revision of the
herbarium specimens and of
nomenclatural aspects of the Madeiran
Dendrosonchus taxa, further led to
propose the recognition of former
Sonchus ustulatus var. latifolia Lowe at a
specific rank and the recognition of
Sonchus ustulatus var. imbricatus Lowe at
a subspecific level.
1. Sonchus latifolius (Lowe) R. Jardim &
M. Seq., comb. nov.
Sonchus ustulatus Lowe var. latifolia
Lowe in Trans. Cambridge Philos. Soc., 4:
23 (1831). [bas.]
Sonchus ustulatus Lowe subsp.
maderensis Aldridge in Bot. Macaronés.,
2: 91 (1977) [1976 publ. 1977], pro parte.
2. Sonchus ustulatus Lowe subsp.
imbricatus (Lowe) R. Jardim & M. Seq.,
comb. nov.
Sonchus ustulatus Lowe var. imbricata
Lowe, Man. Fl. Mad., 1(5): 548 (1868).
[bas.]
= Sonchus ustulatus Lowe subsp.
maderensis Aldridge in Bot. Macaronés.,
2: 91 (1977) [1976 publ. 1977], pro parte.
Key to the Sonchus subgen.
Dendrosonchus species in Madeira
Archipelago:
1. Leaves petiolate …………… S. pinnatus
- Leaves sessile, the bases sheathing the
stem. …………………………………….... 2
2. Caulirosulate shrub up to 4 m high;
inflorescence with large leafy bracts and
capitula up to 30 x 15 mm: ….S. fruticosus
- Suffruticose or caulirosulate shrub up
to 90 cm high; inflorescence with very
small bracts and capitula up to up to 18 x
15 mm ……………………………....…….3
3. Suffruticose to caulirosulate shrub up
to 90 cm high; capitula numerous, more
than 60, disposed in a terminal
irregularly branched, inflorescence
branching from the upper 1/3.
……………………………..S. parathalassius
- Suffruticose with short woody caudex
up to 30 cm high; capitula fewer, up to
30, disposed in a regularly branched
(usually dichotomous), inflorescence
branching from the lower 1/3 - 1/2. …..4
4. Leaf-lobes linear-lanceolate, more than
2x as long as wide, toothed; bracts
without appendices………..…S. ustulatus
- Leaf-lobes ovate, rarely rhombic,
not more than 2x as long as wide,
subentire; bracts with conspicuous
appendices. …………...…...…. S. latifolius
Concluding remarks: LEE et al. (2005)
molecular studies indicate that Madeiran
Dendrosonchus species derive from a
Tenerife ancestor. Moreover the coastal
S. ustulatus and S. latifolia (as S. ustulatus
subsp. maderensis) were the basal taxa
from which a radiation process initiated
to other ecologies. However, the
recognition of S. parathalassius as Porto
Santo endemic and the fact that this
island is older than Madeira (14 My vs.
Notas do Herbário 237
5.6 My) suggests a possible basal
position of this endemism in the
Madeiran Dendrosonchus taxa. Therefore,
possibly Sonchus parathalassius derived
from the Canarian ancestor, dispersed to
Madeira Island and rapidly radiated in
the other Madeiran Dendrosonchus
species, in accordance with the proposed
patterns by LEE et al. (2005). In fact, S.
parathalassius has characters similar to S.
ustulatus (e.g. leaves), S. pinnatus (e.g.
inflorescence ramification) and to S.
fruticosus (e.g. habit). Further studies,
namely molecular, should contribute to
clarify the phylogenetic position of
Sonchus parathalassius in subgen.
Dendrosonchus.
Acknowledgments
The authors acknowledge J. Capelo
for the Latin translation of the diagnosis
and A. Pupo and Z. Ferreira for revising
the English revision. The study of K and
BM specimens (by M. Sequeira) was
supported by SYNTHESIS GB-TAF-1053,
we are also grateful to the herbarium
curators of both herbaria and to the M.
Carine (NHM).
A
B
C
D
Figure 1Sonchus parathalassius. A, Habit. Ilhéu da Cal, 14-IV-2004; B, Leaf rosette, Porto Santo,
Pico Castelo, 20-V-2011; C, Inflorescence, Porto Santo, Pico Castelo, 10-VII-2011; D, Capitula,
Porto Santo, Pico Castelo, 10-VII-2011; all photos R.Jardim
Notas do Herbário 238
Table 1 - Diagnostic and comparative characters of Sonchus parathalassius
Sonchus ustulatus
Sonchus latifolius
Sonchus pinnatus
Sonchus fruticosus
Habit Suffruticose to shrub, up to 90
cm
Suffruticose up to 30 cm
high
Suffruticose up to 30 cm
high
Caulirosulate shrub up to
2m
Caulirosulate shrub up
to 4 m high
Leaves 28-35 x 8-13.6 cm 14-40 x 5-12 cm 13-40 x 5-13 cm 13-40 x 4-15 cm 28-67 x 7-18 cm
Incision Pinnatipartite pinnatisect Pinnatipartite to pinnatisect Pinnatipartite to pinnatisect Sinuate to pinnatifid
Insertion Sessile with sheathing base Sessile with sheathing
base
Sessile with sheathing base Petiolate Sessile with sheathing
base
Lateral lobes 5-9 9-15 3-10 10-16 2-6
Shape and size Narrowly triangular, long
acuminate , 35-51 x 21-35 mm
Linear-lanceolate, to
rhombic, 16-40 x 12-27
mm
Ovate, to obovate, 19-40 x
12-28 mm
Linear-lanceolate to
triangular, 25-80 x 15-32
mm
Rounded to triangular,
18-36 x 38-43 mm
Terminal lobe 30-82 mm, triangular,
acuminate to acuminate-
caudate
18-37 mm, narrow
acuminate-lanceolate or
subcaudate
19-30 mm, lanceolate to
ovate
20-65 mm, triangular-
caudate
100-150 mm, broadly
triangular
Margin Entire to irregularly scarcely
dentate
Dentate Subentire Irregularly dentate to
subentire
Denticulate
Inflorescence Branched, spreading
corymbose cyme
Regularly branched
corymbose cyme
Regularly branched
corymbose cyme
Branched, spreading
corymbose cyme
Branched corymbose
cyme
Axis 60 cm, branching in the
terminal 1/3
15-50 cm, branching
from lower 1/3-1/2
15-50 cm, branching from
lower 1/3-1/2
35 – 90 cm, branching from
lower 1/3-1/2
30-60, branching from
lower 1/3
Capitula 60-90 8-24(-30) (4-) 8-14 Up to 100 or more Up to 100 or more
Diameter 2.5-3.5 cm 4 cm 5.5-6.5 cm 3-5 cm 4-7 cm
Capitula bracts 35, in 5 series 28-29, in 4 series 30, 5 series 28-29, 4 series 67-70, 7 series
Outer bracts 1.2-2.7 x 1.5-2.0 mm,
triangular, adpressed
1.6-4.0 x 1.8-2.6 mm,
triangular, adpressed
3-6.3 x 3.1-3.9 mm,
triangular, adpressed,
appendix ca. 4 mm (truncate
to lobate)
1.0-1.7x1.7-2.0 mm,
triangular, adpressed
11.5-14.2 x 2.5-4.6 mm,
long acuminate,
retroflexed
Intermediate 5.3-7.6 x 1.3-1.9 mm, oblong,
slightly retroflexed
7.9 x 2.3 mm, lanceolate 8.1x4.2 mm, triangular
acuminate
9.1x2.6 mm, lanceolate 11.8-14.8 x 3.4-5.4 mm,
obovate to spatulate,
retroflexed
Inner 10.2-11.7 x 1.1-1.8 mm, linear-
oblong, retroflexed
8.2 x 1.5 linear-
lanceolate, retroflexed
12.4-13.3 x 2.8-4.4 mm,
linear-lanceolate,
retroflexed
9.6-9.8 x 2.0-2.6 mm, linear-
lanceolate, retroflexed
19.1-19.3 x 1.2-1.6 mm,
linear-lanceolate,
retroflexed
Corolla-tube 4.4-5.0 mm 4.8-5.6 mm 7.4-9.2 mm 4.9-7 mm 12.7-13.5 mm
Ligule 7.0-9.2 x 1.5-2.3 mm 5.8-8 x 1.9-2.8 mm 10.6-20 x 3.4-4.3 mm 5.9-6.8 x 1.4-1.9 mm 13.9-18.6 x 2.7-2.9 mm
Cypsela 2.8-2.9 x 0.8-1 mm 2.6-2.8 x 0.8 mm 3-3.8 x 1-1.3 mm 2.1-2.7 x 0.6-1 mm 3.3-4.5 x 1-1.2 mm
Notas do Herbário 239
Figure 2 Sonchus parathalassius J.G. Costa
ex R.Jardim & M.Seq., holotype. MADM
Figure 3 Sonchus parathalassius, habitat.
Ilhéu de Cima, 14-IV-2004, photo: R. Jardim
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A complete and up-to-date checklist of the vascular plants of Macaronesia (the Azores, the Madeira archipelago, the Salvage Islands, the Canary Islands, and the Cape Verde Islands) is given. 3.125 species belonging to 1.041 genera are listed, as are also a number of intraspecific taxa down to variety level. New combinations are proposed within the genera Cheilanthes, Pericallis and Pulicaria. A second section lists 2.250 synonyms and their presumed identity.
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Determining accurate phylogenetic relationships among the members of the woody Sonchus alliance presents challenges because of an insufficient level of molecular variation and the convergent evolution of similar morphological traits in island settings. To obtain a better resolved phylogeny and to test the potential role of hybridization and introgression, we sequenced all members of the alliance with multiple populations for the ITS of nrDNA and over 4000 base pairs of coding and noncoding regions of cpDNA. The cpDNA phylogeny is not well resolved in the core members of the alliance (i.e., subg. Dendrosonchus and genus Taeckholmia), but like the ITS tree, it has identified basal lineages of monotypic genera. The cpDNA data set was not significantly different from that of ITS, and subsequent combined analysis provided a better resolved and supported phylogeny within the alliance. The combined ML tree identified the same basal lineages, suggested nonmonophyly of Dendrosonchus and Taeckholmia, and did not support either Boulos' or Aldridge's infrasubgeneric classification system. Assessment of the role of hybridization and introgression was limited due to poor resolution in the cpDNA phylogeny. The combined analysis supports a Gran Canaria origin for the alliance and two subsequent long distance dispersal events to Madeira and Cape Verde islands.
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