Smilax canariensis, S. azorica (Smilacaceae) and the genus Smilax in Europe

Abstract

A morphological and molecular systematic analysis of the European species of Smilax
results in the recognition of a species endemic to the Azores, Smilax azorica H. Schaef.
& P. Schoenfelder, nom. nov. (= S. divaricata Sol. ex H. C. Wats., nom. illegit.). Its closest
relative is S. canariensis Brouss. ex Willd. from the Canaries and Madeira. Together with the
Eastern European S. excelsa L. they are nested in a clade of North American species, which
seem to belong to an Asian lineage.

Full text

Homenaje al Prof. Dr.
WOLFREDO WILDPRET
DE LA TORRE
Smilax canariensis, S. azorica (Smilacaceae) and the genus Smilax
in Europe
Ha n n o Sc H a e f e r & Pe t e r Sc H o e n f e l d e r
INSTITUTO DE ESTUDIOS CANARIOS
LA LAGUNA - TENERIFE
2009

Homenaje al Prof. Dr.
WOLFREDO WILDPRET
DE LA TORRE
Esperanza Beltrán Tejera, Julio Afonso-Carrillo,
Antonio García Gallo & Octavio Rodríguez Delgado
(Editores)
INSTITUTO DE ESTUDIOS CANARIOS
LA LAGUNA - TENERIFE
2009

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Smilax canariensis, S. azorica (Smilacaceae) and the genus Smilax
in Europe
Ha n n o Sc H a e f e r
1
& Pe t e r Sc H o e n f e l d e r
2*
1
Imperial College London, Ecology & Evol. Biology. Silwood Park Campus. Ascot SL5 7PY. United Kingdom.
hanno.schaefer@imperial.ac.uk
2
Institut fuer Botanik. Universitaet Regensburg. 93040 Regensburg. Germany. peter.schoenfelder@gmx.com
*
Author for correspondence
Resumen: Un análisis morfológico y de sistemática molecular de las especies Europeas de
Smilax dio como resultado el reconocimiento de una especie endémica de las Azores, Smilax
azorica H. Schaef. & P. Schoenfelder, nom. nov. (= S. divaricata Sol. ex H. C. Wats., nom.
illegit.). Su pariente mas cercano es S. canariensis Brouss. ex Willd. de Canarias y Madeira.
Junto con la especie del este de Europa S. excelsa L., todas ellas forman un grupo dentro del
clado de las especies norteamericanas, las cuales parecen pertenecer a un linaje asiático.
Palabras claves: Azores, biogeografía, Islas Canarias, refugio glacial, Smilax azorica, Smilax
canariensis, Smilax divaricata.
Abstract: A morphological and molecular systematic analysis of the European species of Smi-
lax results in the recognition of a species endemic to the Azores, Smilax azorica H. Schaef.
& P. Schoenfelder, nom. nov. (= S. divaricata Sol. ex H. C. Wats., nom. illegit.). Its closest
relative is S. canariensis Brouss. ex Willd. from the Canaries and Madeira. Together with the
Eastern European S. excelsa L. they are nested in a clade of North American species, which
seem to belong to an Asian lineage.
Key words: Azores, biogeography, Canary Islands, glacial refugia, Smilax azorica, Smilax
canariensis, Smilax divaricata.
I
n t r o d u c t I o n
The genus Smilax comprises about 200 species distributed mainly in the Northern he-
misphere from the temperate regions to the Subtropics (CAMERON & FU, 2006). Diversity
centres of the genus are located in Northern and Central America and East Asia, while the
European region harbours only four species and Africa and Australia only two each (CAME-
RON & FU, 2006). The comparatively reduced diversity in European Smilax is presumably
a result of the ice ages and is well known from other genera like Quercus, Acer, and Cornus
(e.g. SVENNING et al., 2008; XIANG et al., 2006).
Of the four European species, one, Smilax aspera L., is widespread and often common
throughout the Mediterranean region (Fig. 1). The three remaining species are restricted to
glacial refugia: (i) Smilax canariensis Brouss. ex Willd. is endemic to the Canary Islands and
Madeira (SCHOENFELDER & SCHOENFELDER, 2005), (ii) Smilax azorica H. Schaef. &
Schoenfelder, nom. nov. (= S. divaricata Sol. ex H. C. Wats., nom. illegit., see d
I S c u S S I o n )

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Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
is known only from the Azores, and (iii) Smilax excelsa L. is found mainly in the Black Sea
and Caspian Sea region from Greece and Bulgaria to Iran (BROWICZ, 1988).
M
at e r I a l a n d M e t H o d S
Morphology
The authors studied herbarium material at AZU, BM, K, LISU, M, and REG. Both authors
spent many months of fieldwork in the Mediterranean region and the middle-Atlantic Islands
and studied morphology and ecology of Smilax in many different countries and habitats.
Sampling and DNA extraction
Total genomic DNA was isolated from herbarium specimens or, more rarely, silica-dried
material following the standard CTAB method of DOYLE & DOYLE (1987). We ampli-
fied the rbcL and matK genes, the trnL intron and the trnL-F intergenic spacer. Polymerase
chain reactions (PCR) were performed with the standard protocol and primers described
in SCHAEFER et al. (2008), and products were purified with the Wizard SV PCR clean-
up kit (PROMEGA GmbH, Mannheim, Germany). Cycle sequencing was performed with
BigDye Terminator cycle sequencing kits on an ABI Prism 3100 Avant automated sequencer
(Applied Biosystems, Foster City, California, USA).
In addition to these plastid regions, we sequenced the nuclear internal transcribed spacer
region using the ITS primers of CAMERON & FU (2006). Direct PCR amplification of ITS
yielded single bands and unambiguous base calls. Twenty-seven sequences were generated
for this study. Table 1 lists the relevant taxonomic names with authors and plant sources. All
new sequences have been deposited in GenBank (http://www.ncbi.nlm.nih.gov/). Additional
sequences for Asian and American species (mostly generated by CAMERON & FU, 2006)
were downloaded from GenBank.
Sequence alignment and phylogenetic analyses
Sequences were edited with Sequencher (4.6; Gene Codes, Ann Arbor, Michigan, USA)
and aligned by eye, using MacClade 4.06 (MADDISON & MADDISON, 2003). The aligned
plastid matrix comprised 3517 nucleotides. The aligned ITS matrix comprised 867 nucleo-
tides. Maximum likelihood (ML) tree searches and ML bootstrap searches were performed
using RAxML 7.0.3 (STAMATAKIS et al. 2008, available at http://phylobench.vital-it.ch/
raxml-bb/). RAxML searches relied on the GTR + G + I model (six general time-reversible
substitution rates, assuming gamma rate heterogeneity and a proportion of invariable sites),
with model parameters estimated over the duration of specified runs. Analyses in RAxML
were run both with the combined un-partitioned data and with a model that partitioned the
plastid regions from the ITS region. Trees were rooted on Philesia magellanica (sequences
from GenBank). The data matrix and trees have been deposited in TreeBASE (http://www.
treebase.org/).

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The genus Smilax in Europe
Figure 1. European distribution of Smilax species (modified after BOLÓS & VIGO, 2001; BROWICZ, 1988).
Table 1. Species sampled and their origin.
SPECIES ORIGIN
Smilax aspera L. subsp. aspera France, Camargue
Smilax aspera L. subsp. balearica (Willk.) Romo Spain, Balearic Islands, Mallorca
Smilax aspera L. subsp. mauritanica (Desf.) Malag. Spain, Canary Islands, Tenerife
Smilax canariensis Brouss. ex Willd. Portugal, Madeira
Smilax canariensis Brouss. ex Willd. Spain, Canary Islands, Tenerife
Smilax azorica H. Schaef. & P. Schoenfelder Portugal, Azores, São Miguel
Smilax excelsa L. Georgian Republic
Smilax hispida Muhl. ex Torr.
United States, seeds bought from
“BT World Seeds”
Smilax lasioneura Hook.
United States, seeds bought from
“BT World Seeds”

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Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
re S u l t S
Morphology
The mediterranean Smilax aspera differs from the remaining European species above all
in its inflorescences, which are composed of several umbel-like sub-inflorescences and often
more than 20 cm long (fig. 2a). All other European taxa have their flowers in simple umbels
(SCHOENFELDER & SCHOENFELDER, 2005; our Fig. 2 c-d, Fig. 3).
The leaves of S. aspera are coriaceous and very variable, usually with 7-9 main nerves
and a ± cordate base. The leaves of the remaining species are laurophyllous (S. canariensis, S.
azorica) or deciduous (S. excelsa) with usually 3-5 main nerves and a cuneate to rounded or
shallowly cordate base (BROWICZ, 1988; S. Arndt, Jena Botanical Gardens, pers. comm.).
Leaf shape is variable, especially on young shoots. Leaves on second-year or older shoots are
more uniform in shape and in general broadly ovate in S. azorica and more narrowly ovate in
S. canariensis. The older stems of S. excelsa carry considerable thorns (Fig. 2d), while thorns
on stems of S. canariensis and S. azorica are small or absent.
A taxon with extremly narrow leaves that has been described as S. aspera subsp. bale-
arica (Willk.) Romo is apparently restricted to the Balearic Islands and accepted as an en-
demic variety in BOLÒS & VIGO (2001), but not accepted as a separate taxon by AEDO
(2005). Forms with broadly cordate leaves that lack thorns almost completely are known as
S. aspera subsp. mauritanica (Desf.) Malag. (= S. altissima Roxb.). They are found in the
Western Mediterranean region and in the Canary Island’s laurel forest but also in dry lowland
areas and cliffs on Madeira (PRESS & SHORT, 1994), and the Azores (Terceira Island). The
Madeiran plants have been described as endemic species S. pendulina Lowe but they do not
differ considerably from S. aspera subsp. mauritanica.
Phylogenetic analyses
The topologies of the best likelihood tree for the plastid and ITS datasets (not shown)
were not contradicting in any well-supported node. We therefore combined the data and in
the following focus on the result of the combined data (Fig. 4). Resolution and bootstrap
support was in general low, a problem already reported in previous studies (CAMERON &
FU, 2006). However, the placement of S. aspera as sister to all other analysed ingroup taxa
is moderately supported. Furthermore, we found support for a clade consisting of the North
American S. herbacea and other American species, a clade of Asian species, and a clade con-
sisting of S. china, two North American species, S. excelsa, and the middle-Atlantic island
species. The Azorean plants are cleary different from S. canariensis, while the sample from
Madeira seems to be genetically very close to the Canary Island plants.

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The genus Smilax in Europe
Figure 2. Inflorescences and leaves of European Smilax: a) S. aspera subsp. mauritanica (La Palma, Canary Islands,
10-10-1993); b) S. canariensis (La Palma, Canary Islands, 7-10-1993); c) female inflorescence of S. azorica (Faial
Island, Azores, 17-7-1999); d) male inflorescence of S. excelsa (Botanical Garden, Jena, 31-5-2003).

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Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
Figure 3. Illustration of S. canariensis reproduced from WEBB & BERTHELOT (1847).

303
The genus Smilax in Europe
Ke y t o t H e eu r o P e a n S P e c I e S o f Sm i l a x
1 Leaves usually with 7-9 main nerves. Male and female inflorescences
compound of several umbel-like sub-inflorescences. Flowering time VIII-XI. Fruit ripe-
ning blackish-red ............................................................................................... S. aspera
1* Leaves usually with 3-5 main nerves. Male and female inflorescence a simple
umbel .............................................................................................................................. 2
2 Plant deciduous, older stems with strong thorns. Flowering time V-VI. Black sea and
Eastern Mediterranean region ...........................................................................S. excelsa
2* Leaves wintergreen, thorns on older stems weak or absent. Flowering time V-VIII.
Middle-Atlantic Islands ................................................................................................. 3
3 Leaf blades on older branches broadly cordate-ovate, almost as broad as long (relation
length:width c. 1:0.9). Fruit ripening red. Endemic to the Azores ................... S. azorica
3* Leaf blades on older branches narrower (relation c. 1: 0.6). Fruit ripening
black (fide WEBB & BERTHELOT, 1847). Endemic to the Canary Islands and
Madeira ......................................................................................................S. canariensis
d
I S c u S S I o n
Taxonomy
Our results support the separation of a species endemic to the Azores from S. canarien-
sis, endemic to the Canaries and probably Madeira. The species from the Azores was first
collected by Francis Masson in 1777 on São Miguel (specimens in BM) and later described
as Smilax divaricata Sol. ex H. C. Watson (WATSON, 1844), a name that had already been
Figure 4. Maximum likelihood phylogram of Smilax plastid and ITS sequences produced with RAxML 7.0.3 (STA-
MATAKIS et al. 2008). Likelihood bootstrap support values > 60% are given at the nodes.

304
Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
given to a species from the Philippines seven years earlier (BLANCO, 1837). Therefore the
Solander name is a later homonym and illegitimate. A new epithet is required, which we
propose as follows:
Smilax azorica H. Schaef. & P. Schoenfelder, nom. nov.
replaced synonym: Smilax divaricata Sol. ex H. C. Wats. in London J. Bot. 3: 608. 1844. –
Type: F. Masson s.n. (holo BM!), Portugal, Azores, Sao Miguel 1777. – (non Smilax divari-
cata Blanco, Fl. Filip. 795. 1837).
HANSEN & SUNDING (1993) listed both species Smilax canariensis and S. divaricata
(= S. azorica H. Schaef. & P. Schoenfelder, nom. nov.) in their Azores checklist and also
added the East European S. excelsa, certainly a mistake. SCHAEFER (2003, 2005) based
on morphology only, treated S. divaricata as a synonym of S. canariensis but with our new
genetic data, this view is no longer supported. SEUBERT (1844) lists S. tetragona L.f. in his
“Flora Azorica”, a synonym for S. aspera subsp. mauritanica, but his description of plants
from Pico Island (Azores) matches S. azorica. The specimen C. Hochstetter 121, cited by
Seubert, was studied at BM and identified as S. azorica.
Biogeography
The European Smilax species clearly belong to two long separated lineages: the wide-
spread S. aspera is sister to all other Smilax species (see also CAMERON & FU, 2006),
Figure 5. Distribution of S. canariensis on Tenerife, Canary Islands.

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The genus Smilax in Europe
Figure 6. Distribution of S. azorica in the Azores archipelago.
while the remaining species S. excelsa, S. canariensis, and S. azorica form a monophyletic
group. Their closest relatives seem to be North American species but these belong to an Asian
lineage. All possible biogeographic scenarios locate the ancestors of the S. canariensis group
in Asia. From there, in the most parsimonious scenario, the lineage spread via Beringia into
North America and from there across the then narrow North Atlantic back into the European
continent. During the glacial periods, this ancestral lineage was split into an eastern popula-
tion in the Black sea region and a western population in the middle-Atlantic Islands. The lack
of genetic exchange between the Azores and the Madeira/Canary islands population finally
resulted in the evolution of two endemic species. An alternative, less parsimonious scenario
would require at least two independent dispersal events from Asia into North America and
one dispersal/range expansion of the S. canariensis lineage from Asia directly into the Me-
diterranean and the middle-Atlantic islands. A broader genetic analysis of Smilax samples
from the islands, from North America, Asia, and especially from Africa (one or two endemic
species) combined with molecular clock dating techniques will be required to confirm one
of these scenarios.
Conservation
Today, S. canariensis is very rare in the Canaries and restricted to the central laurel forest
regions on Tenerife (Fig. 5), La Palma, and La Gomera. On Madeira, it is a poorly known
species that was collected only a few times (PRESS & SHORT, 1994). At least in the Canary
Islands, it seems to be highly threatened. Despite years of search, the senior author could find

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Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
only one fruiting individual on La Palma and was unable to find a single flowering specimen
in the archipelago. Notably, there is not a single picture of S. canariensis inflorescences in
the contemporary literature on the Canary island flora and the flora of Madeira. An excellent
description and a plate with flowers in all details, however, can be found in WEBB & BER-
THELOT (1847) (our Fig. 3). Apparently, the species was reproducing more frequently in
those days. Maybe the more intense use of the laurel forest in the 19
th
century produced more
open spaces and clearings and favoured less competitive species like S. canariensis.
Smilax azorica, is known from six islands of the Azores archipelago (Fig. 6). While ab-
sent in the western group and rare in the central group, it is locally common on São Miguel
and Santa Maria in the eastern group. Flowers and fruits are regularly found and the species
seems to be not threatened. It is protected by the Berne convention on the conservation of
European wildlife and natural habitats.
a
c K n o w l e d g e M e n t S
The authors are grateful to C. Heibl (Munich), H. J. Esser (M), M. Carine (BM), S. Arndt
(Jena), R. Jahn (Großschirma) and K. Tan (Copenhagen) for material and information, to W.
Lang for drawing figure 1 after the authors’ draft, and to O. Fiz-Palacios (London) for the
Spanish abstract. HS is grateful to the Azorean Direcção Regional do Ambiente for a permit
to collect DNA samples in the islands.
r
e f e r e n c e S
AEDO, C., 2005. Smilax L. in Aedo, C. & Herrero, A. (Eds.). Flora iberica 21: 3-7.
BLANCO, F.M. 1837. Flora de Filipinas. 795 pp.
BOLÓS, O. DE & J. VIGO, 2001. Flora dels Països Catalans 4: 168-169.
BROWICZ, K., 1988. Chorology of trees and scrubs in South-West Asia and adjacent regions. Vol. 6:
1-86.
CAMERON, K.M. & C. FU, 2006. A nuclear rDNA phylogeny of Smilax (Smilacaceae). Aliso 22:
598-605.
DOYLE, J.J. & J.L. DOYLE, 1987. A rapid DNA isolation procedure for small quantities of fresh leaf
tissue. Phytochem. Bull. 19: 11-15.
HANSEN, A. & P. SUNDING, 1993. Flora of Macaronesia. Checklist of vascular plants. 4. rev. ed.
Sommerfeltia 17: 1-295.
MADDISON, W.P. & D.K. MADDISON, 2003. MacClade: analysis of phylogeny and character evo-
lution, version 4.06. Sunderland, Massachusetts: Sinauer Associates.
PRESS, J.R. & M.J. SHORT, 1994. Flora of Madeira. London. 574 pp.
SCHAEFER, H., 2003. Chorology and Diversity of the Azorean Flora. Diss. Bot. 374: 1-670.
SCHAEFER, H., 2005. Flora of the Azores. A Fieldguide. 2nd ed. Margraf Publishers/Backhuys Pub-
lishers, Weikersheim. 346 pp.
SCHAEFER, H., I.R.H. TELFORD & S.S. RENNER, 2008. Austrobryonia (Cucurbitaceae), a new
Australian endemic genus, is the closest living relative to the Eurasian and Mediterranean Bryonia
and Ecballium. Syst. Bot. 33: 125-132.
SCHOENFELDER, I. & P. SCHOENFELDER, 2002. Kosmos Atlas Mittelmeer- und Kanarenflora.
2nd ed. Franckh-Kosmos, Stuttgart. 303 pp.
SCHOENFELDER, P. & I. SCHOENFELDER, 2005. Die Kosmos-Kanarenflora. 2nd ed. Franckh-
Kosmos, Stuttgart. 319 pp.
SEUBERT, M., 1844. Flora Azorica. Bonn. 50 pp.

307
The genus Smilax in Europe
STAMATAKIS, A., P. HOOVER & J. ROUGEMONT, 2008. A rapid bootstrap algorithm for the
RAxML web-servers. Syst. Biol. 75(5): 758-771.
SVENNING, J.-C., S. NORMAND & M. KAGEYAMA, 2008. Glacial refugia of temperate trees in
Europe: insights from species distribution modelling. J. Ecol. 96(6): 1117-1127.
WATSON, H.C., 1844. Notes on the botany of the Azores. London J. Bot. 3: 608.
WEBB, P.B. & S. BERTHELOT, 1847. Histoire naturelle des Îles Canaries [...] Tome troisième.
Deuxième partie. Phytographia canariensis. Sectio III, Smilax: 322-324, tab. 225.
XIANG, Q.-Y., D.T. THOMAS, W. ZHANG, S.R. MANCHESTER & Z. MURELL, 2006. Species
level phylogeny of the Dogwood genus Cornus (Cornaceae) based on molecular and morphological
evidence– implication in taxonomy and Tertiary intercontinental migration. Taxon 55(1): 9-30.

ÍNDICE
TABVLA GRATVLATORIA .......................................................................................
Presentación
Milagros Luis Brito ............................................................................................
Antonio Alarcó Hernández .................................................................................
Eduardo Doménech Martínez .............................................................................
Esperanza Beltrán Tejera: Semblanza de un botánico comprometido con su
tiempo. Profesor Wolfredo Wildpret de la Torre ................................................
Esperanza Beltrán Tejera: Producción bibliográfica de la Unidad de Botánica
de la Universidad de La Laguna. Etapa wildpretiana (1969-2008). I ...............
Jorge Alfredo Reyes-Betancort & María Catalina León Arencibia: Helichrysum x
wildpretii nothosp. nov., un nuevo híbrido natural de las Islas Canarias ..........
Marcelino José del Arco Aguilar, Octavio Rodríguez Delgado, Juan Ramón Ace-
bes Ginovés, Marcos Salas Pascual & Víctor Garzón Machado: Los retamares
de Retama rhodorrhizoides Webb & Berth. en las Islas Canarias: Retamation
rhodorhizoidis all. nov. ...........................................................................................
Arnoldo Santos Guerra & Jorge Alfredo Reyes-Betancort: Contribución al co-
nocimiento de las comunidades comofíticas de la Clase Greenovio-Aeonietea
Santos 1976. Aichryso laxi-Monanthetalia laxiflorae ord. nov............................
Octavio Rodríguez Delgado: El Barranco del Agua de Güímar, un espacio na-
tural de gran interés botánico, turístico y etnográfico .......................................
Pedro L. Pérez de Paz, Vicente L. Lucía Sauquillo & Ricardo González González:
Las Charcas de Erjos: enclave antrópico de singular naturaleza .....................
Werner Nezadal & Walter Welss: Aportaciones al conocimiento del bosque ter-
mófilo en el noroeste de Tenerife (Islas Canarias) ..............................................
13
21
23
25
27
71
159
163
173
181
213
229

Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
Marcos Salas Pascual, Emilio Fernández Negrín & Gregorio Quintana Vega: Salvio
canariensis-Pterocephaletum dumetori ass. nov. (Artemisio thusculae-Rumicion
lunariae; Forsskaoleo angustifoliae-Rumicetalia lunariae; Pegano-Salsoletea)
nueva asociación para la Isla de Gran Canaria (Islas Canarias-España) ........
Salvador Rivas-Martínez: Ensayo geobotánico global sobre la Macaronesia .....
Hanno Schaefer & Peter Schoenfelder: Smilax canariensis, S. azorica (Smilaca-
ceae) and the genus Smilax in Europe ..................................................................
Julia Pérez de Paz, Olga Fernández-Palacios & Rosa Febles: Polimorfismos y
series polínicas en el género canario Parolinia y parientes continentales Dice-
ratella y Morettia (Matthioleae-Brassicaceae). Significado biológico y filoge-
nético .......................................................................................................................
Irene E. La Serna Ramos: Parkinsonia aculeata L.: un ejemplo del interés de la
flora ornamental en la caracterización geográfica de las mieles canarias ........
Victoria Eugenia Martín Osorio: Jardines Sostenibles ..........................................
Beatriz Hernández Bolaños & Victoria Eugenia Martín Osorio: El Jardín Botá-
nico del Parque Nacional del Teide (Tenerife, Islas Canarias), a través de un
Sistema de Información Geobotánica ..................................................................
Antonio García Gallo, Israel Pérez Vargas & Francesco Salomone Suárez: Los
olmos de La Laguna ...............................................................................................
Richard Pott & Joachim Hüppe: Canary Islands: A Botanical Paradise in the
Atlantic Ocean ........................................................................................................
María Candelaria Gil-Rodríguez, Myrian Rodríguez García del Castillo, Óscar
Monterroso Hoyos & Rodrigo Riera Elena: Perturbaciones en ecosistemas ma-
rinos canarios. Un modelo: Guayonje-Tacoronte, Islas Canarias .....................
Julio Afonso-Carrillo & Marta Sansón: Aún lejos de un completo conocimiento
de la biota canaria: el ejemplo de la flora de algas rojas gelatinosas efímeras
del sublitoral ...........................................................................................................
Esperanza Beltrán Tejera, J. Laura Rodríguez-Armas, Luis Quijada, Janira Gu-
tiérrez Peraza, Jonathan Díaz & Ángel Bañares: Contribución al estudio de la
micobiota de los castaños del Norte de Tenerife (Islas Canarias. España). II ..
María Carmen Jaizme-Vega: Las micorrizas, una simbiosis de interés en agri-
cultura .....................................................................................................................
245
255
297
309
329
345
371
383
395
421
433
453
479

Índice
Consuelo Hernández, Israel Pérez-Vargas, Dessire Sicilia & Pedro L. Pérez de
Paz: Los líquenes de la alta montaña canaria ......................................................
Ana Losada-Lima, Sofía Rodríguez-Núñez & Arnoldo Santos Guerra: Referen-
cias a briófitos de las Islas Canarias anteriores al siglo XIX: Dillenius y Leu-
codon canariensis ....................................................................................................
Mari Carmen Alfayate, Eugenia Ron, Agustín Fernández, Belén Estébanez, David
Gómez, Miguel Ángel Pérez-Batista & Benjamín Fernández: Biontes entrometi-
dos en cápsulas de musgos Canarios ....................................................................
Juana María González-Mancebo, Jairo Patiño, Julio Leal Pérez, Stephan Scholz &
Ángel Fernández-López: Amenazas sobre la flora briofítica de la Isla de Fuer-
teventura. SOS para los últimos supervivientes del extinto bosque de Jandía
Marie-Luise Schnetter, Andreas Opitz & Reinhard Schnetter: Estructura y
función de las glándulas submarginales del mangle Laguncularia racemosa
(Combretaceae) ......................................................................................................
Domingo Morales & Mª Soledad Jiménez: Ecofisiología de algunos tipos de ve-
getación de las Islas Canarias ...............................................................................
Juan Felipe Pérez Francés, Isabel Santana López, Emma Suárez Toste, Raquel Mar-
tín Pérez, Miguel Cabrera Pérez, Juan Cristo Luis Jorge & Francisco Valdés: Apli-
caciones del cultivo in vitro a la conservación de plantas canarias en pe ligro ....
Germán Santana Henríquez: Una farmacopea un tanto singular. Sobre los re-
medios para el dolor de cabeza en Galeno ...........................................................
José N. Boada, Eduardo Navarro & C. Marina Álvarez: Nuestras aportaciones
al conocimiento de las propiedades farmacológicas de productos obtenidos de
plantas de Canarias................................................................................................
José Juan Jiménez González: Etnohistoria y arqueología de las plantas entre
los antiguos canarios ..............................................................................................
Fernando Lozano Soldevilla, Ignacio J. Lozano, José Mª. Landeira & Fátima Her-
nández: Antecedentes históricos de la taxonomía zooplanctónica en aguas de
la región Canaria ....................................................................................................
Lázaro Sánchez-Pinto, Francisco García-Talavera, José López Rondón & Merce-
des Martín Oval: Sobre la presencia del icnofósil Dactyloidites ottoi (Geinitz,
1849) en sedimentos neógenos de la costa occidental de Fuerteventura (Islas
Canarias) .................................................................................................................
489
501
509
517
539
555
567
581
591
603
613
625

Homenaje al Prof. Dr. Wolfredo Wildpret de la Torre
Juan José Bacallado, José Espinosa, Jesús Ortea, Lázaro Márquez, Leopoldo
Moro, Osmani Borrego & Manuel Caballero: La península de Guanahacabibes
y su Parque Nacional (Cuba): biodiversidad marina y terrestre ......................
Marisa Tejedor, Jonay Neris, María Ascención Dorta & Concepción Jiménez:
Evaluación del recurso suelo con alta potencialidad agrológica en la isla de
Tenerife. 1981-2008 ................................................................................................
Juan Luis Mora Hernández, Carmen Dolores Arbelo Rodríguez & Antonio Rodrí-
guez Rodríguez: Características de los suelos de las Islas Canarias en relación
a la vegetación natural ...........................................................................................
Constantino Criado, Carmen Machado & José Afonso: Geomorfología eólica en
el Parque Nacional del Teide (Tenerife) ...............................................................
Sara del Río, Luis Herrero & Ángel Penas: Tendencias recientes en la precipi-
tación de las Islas Canarias occidentales y su relación con la oscilación del
Atlántico Norte (NAO) ..........................................................................................
Sebastián Delgado Díaz: Las nuevas aguas en Canarias ......................................
Gonzalo Lozano Soldevilla: Miscelánea académica del quinquenio 1983-1988
en la Facultad de Biología de la Universidad de La Laguna..............................
Nácere Hayek: Un ensayo histórico sobre la aportación matemática a la Biolo-
gía durante períodos anteriores a su creación .....................................................
Andrés Sánchez Robayna: Viene del mar la integridad de más allá del mar .....
Juan Hernández Bravo de Laguna: La Teoría del Estado fallido: Estados débi-
les, Estados aparenciales y otras formas fallidas de Estado ...............................
Matilde Arnay de la Rosa & Emilio González Reimers: La ocupación humana de
Las Cañadas del Teide a partir del siglo XV .......................................................
Conrado Rodríguez Martín, Rafael González Antón & María del Carmen del Arco
Aguilar: La colonización humana de islas en la prehistoria. Un modelo teórico
para el estudio de poblamientos insulares ...........................................................
Cristóbal Corrales Zumbado & Dolores Corbella Díaz: Creación y adaptación
del término malpaís ................................................................................................
Josefa Dorta Luis & María del Carmen Muñiz Cachón: La entonación de las
interrogativas en el español de Canarias y en asturiano ....................................
633
651
665
685
705
723
731
739
753
755
767
785
797
809

Índice
Juan Antonio Frago Gracia: El español de Canarias en la historia de la lengua
española ...................................................................................................................
Javier Medina López: La gramática olvidada de D. Ireneo González y Hernán-
dez: el Compendio de gramática castellana (1895) ...............................................
Francisco Salas Salgado: Influencia clásica en los poemas a Filis de Juan Bau-
tista Poggio Monteverde ........................................................................................
Teodoro Ravelo Mesa, María Carmen Moreno Perdigón & Moulaye Ahmed Ould
Ahmed Deoula: Un análisis multicriterio de la capacidad de atracción de los
destinos turísticos en la Isla de Tenerife ...............................................................
823
837
849
861