Anales del Jardín Botánico de Madrid
ISSN: 0211-1322
anales@ma-rjb.csic.es
Consejo Superior de Investigaciones
Científicas
España
Galán Cela, Pablo; Gamarra, Roberto
Check list of the iberian and balearic orchids. 1. Aceras R. Br. - Nigritella Rich
Anales del Jardín Botánico de Madrid, vol. 59, núm. 2, 2002, pp. 187-208
Consejo Superior de Investigaciones Científicas
Madrid, España
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CHECK LIST OF THE IBERIAN AND BALEARIC ORCHIDS
1. ACERAS R. BR. – NIGRITELLA RICH.
by
PABLO GALÁN CELA 1 & ROBERTO GAMARRA 2
1
Departamento de Producción Vegetal: Botánica y Protección Vegetal, E.U.I.T. Forestal,
Universidad Politécnica. E-28040 Madrid <pgalan@forestales.upm.es>
2
Departamento de Biología, Facultad de Ciencias, Universidad Autónoma de Madrid. E-28049 Madrid
<roberto.gamarra@uam.es>
Resumen
GALÁN CELA, P. & R. GAMARRA (2002). Catálogo de las orquídeas ibéricas y baleares. 1.
Aceras R. Br. – Nigritella Rich. Anales Jard. Bot. Madrid 59(2): 187-208 (en inglés).
Se presenta un catálogo, ordenado alfabéticamente, de los táxones de la familia Orchidaceae
incluidos en los géneros Aceras a Nigritella, presentes en la Península Ibérica e Islas Baleares,
excluidos los híbridos. Se recogen todos los nombres, hasta el rango varietal, mencionados de
dicho ámbito geográfico, con su correspondiente lugar de publicación; por supuesto, los
considerados correctos, sus sinónimos e indicatio locotypica. Se incluyen, cuando se estiman
oportunas, observaciones de índole nomenclatural y taxonómico.
Palabras clave: Orchidaceae, nomenclatura, taxonomía, Península Ibérica, Islas Baleares.
Abstract
GALÁN CELA, P. & R. GAMARRA (2002). Check list of the iberian and balearic orchids. 1.
Aceras R. Br. – Nigritella Rich. Anales Jard. Bot. Madrid 59(2): 187-208.
A check list of the taxa of the family Orchidaceae found in the Iberian Peninsula and the
Balearic Islands is presented in alphabetical order. This section includes the genera from
Aceras to Nigritella, down to the rank of variety, but excluding hybrids. We have gathered
together the correct names, with their corresponding place of publication, synonyms and
indicatio locotypica. Also included are some observations in those genera and species that
present nomenclatural and taxonomic problems.
Key words: Orchidaceae, nomenclature, taxonomy, Iberian Peninsula, Balearic Islands.
INTRODUCTION
The family Orchidaceae is widely
represented throughout the entire world, with
maximum diversity in tropical and subtropical zones. In Europe, traditionally 35
genera are recognized –cf. MOORE (1980),
BUTTLER (1991), DELFORGE (1994), etc.–, of
which, 25 appear in the Iberian Peninsula and
the Balearics, where the representation of this
family is one of the broadest, at least by
extratropical standards, equivalent or only
slightly less than other countries of the
Mediterranean region. But, in the last five
years, in the cladistic studies published by
BATEMAN & al. (1997) and BATEMAN (2001),
the number has decreased, because some
genera are included in others. Therefore,
BUTTLER (2001) on the basis of traditional
characters, only accepts the inclusion of
188
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Aceras in Orchis. D ELFORGE (2001) only
considers 28 genera. In this paper, we prefer
to maintain the current status presented in
recent floras, such as STACE (1997).
At present, it is difficult to determine the
number of recognized taxa, due to the
confusion resulting from the taxonomic
treatments in highly difficult groups and the
notable grade of variation, such as found in
representatives of the genera Dactylorhiza,
Epipactis, Ophrys and Orchis. Taxonomic
studies of this family in the Iberian Peninsula
and the Balearic Islands have been slowly
increasing, but in a fragmented manner
through time. Although a diverse group of
Spanish, Portuguese and foreign botanists have
described new taxa for the Peninsula and
Balearics, we would say that the first author
completing the most in-depth study was José
d’Ascenção Guimarães, who published a
treatment of the Portuguese orchids in 1887.
Since the 1960s, numerous French, Belgian,
German and English orchidologists have
visited our peninsula and have contributed,
frequently with the description of new taxa, to
the taxonomic and chorologic knowledge of
this family. From 1970 onward, botanizing by
numerous Spanish and Portuguese scientists
has proliferated many studies, resulting in
development of abundant local, provincial, and
regional catalogues of the Iberian and Balearic
territories, which contain new information
concerning representatives of this family, e.g.
RIVERA & LÓPEZ VÉLEZ (1987), DELFORGE
(1989), PÉREZ RAYA & MOLERO MESA (1990),
PÉREZ CHISCANO & al. (1991), BOUILLIE & al.
(1992), H ERMOSILLA & S ABANDO (1993,
1996a, 1996b, 1997, 1998), LOWE (1995),
MARÍN & GALÁN CELA (1994), SANZ & NUET
(1995), TYTECA (1997), LOWE (1998), BENITO
AYUSO & al. (1999), CORTIZO & SAHUQUILLO
(1999), PALLARÉS (1999), LIZAUR (2001). Also
in many floras, such as M ERINO (1909),
CADEVALL (1933), BONAFÉ (1978), VALDÉS &
al. (1987), MATEO (1991), AIZPURU & al.
(1999), BOLÒS & VIGO (2001), and many
others.
Additional proof of awaking interest in this
family is the publication of various guides of
European and Mediterranean scopes, such as
those of SUNDERMANN (1980), BAUMANN &
KÜNKELE (1988), BUTTLER (1991), DELFORGE
(1994, 2001).
All of this investigative labor has generated
an extensive proliferation of names given for
our country. From this arises the necessity of
this present work, which has two goals. First,
and of highest priority, it aims to compile of
the nomenclatural information within this
family for the defined geographic range,
without taking into account ranks lower than
variety, nor hybrids, although several taxa are
present in the list which were originally
treated as such. Secondly, the order within
this list of names follows the criteria of
morphological similarity and taxonomic
affinity, a task which was easily accomplished in some cases, but in others,
particularly in the aforementioned genera,
one that forced us to group names that
belonged to related taxa. We do not intend
this to constitute a systematic study in any
manner; rather, it should be treated as nothing
more than a thoughtful opinion derived from a
thorough review of the current status of
knowledge.
MATERIAL AND METHODS
The search for names of Orchidaceae has
entailed the review of numerous bibliographic sources, as much books as periodical
publications, principally about the Iberian
flora, but also others that cited the occurrence
of taxa in the Iberian Peninsula and the
Balearic Islands. For this, we utilized resources held in the libraries of the Real Jardín
Botánico de Madrid, the Facultad de Farmacia in the Universidad Complutense de Madrid and Kew Gardens of London.
The review of the names using the different
volumes of Index Kewensis uncovered
numerous homonyms of taxa named in the
Iberian Peninsula. In relation to this task, we
detected and brought to light several citation errors and omission of taxa in this
distinguished work. In the same manner,
redundant combinations were also discovered.
The catalogue is organized alphabetically
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
189
by accepted names of the taxa included into
the genera from Aceras to Nigritella. The
valid names are designated in bold type and
each one is followed by the accepted
authority, the place and date of publication,
indicating the exact pages and figures. Below,
in italics, are collected all the homotypic
synonyms, with the corresponding indicatio
locotypica, and occasionally including more
details, with a view to facilitating possible
typifications. In some cases access to the
publication of the original description was not
possible, and it was transcribed from other
work of the same author, or collected from
other authors. Each of these cases is specified
as such in the observations.
In addition, the heterotypic synonyms and
invalid names are listed in chronological
order, with the information indicated previously. The abbreviation “n. v.” is used to
indicate cases in which it was not possible to
review the corresponding publication.
In several taxa we added an observation
section, which included explanations about
nomenclatural problems that resulted in the
adoption of one or the other name.
The taxa which have been described for
Iberian Peninsula and Balearic Islands are
preceded by an asterisk (*).
Lectotypus: “Vaill., Bot. paris: tab. 31, fig.
19-20. 1727 sub nom. Orchis” –cf. BAUMANN
& al. (1989: 437).
Observations.–In Index Kewensis and
in numerous works about orchids, the
authorship of this taxon is cited in the
following form: “A. anthropophorum R. Br.
in W. T. Aiton, Hortus Kew. ed. 2, 5: 191
(1813)” The aforementioned index describes
the genus Aceras and makes reference to
Ophrys anthropophora Willd., Sp. Pl. 4(1):
63 n.º 3 (1805), which actually described
Ophrys anthropomorpha. To this one adds
the fact that in the publication of Aiton of
1813, this author writes in the margin the
epithet of the species treated, but in this case it
was lacking for the genus Aceras, and the
name Aceras anthropophorum was used for
the first time in 1814.
CHECK LIST
Ind. loc.: “Centro littoral: Appellação (J.P.
Gomes!).- Baixo Alemtejo littoral: Arrabida
(W.!)”.
Observations.–GUIMARÃES (1887: 69-71)
adjudged this taxon to have a hybrid origin
between Aceras anthropophorum and Orchis
longicruris, and indicated that it more closely
approaches the former.
Aceras R. Br. in W.T. Aiton, Hortus Kew. ed.
2, 5: 191 (1813)
Aceras anthropophorum (L.) W.T. Aiton,
Epitome ed. 2 Hortus Kew.: 281 (1814)
⬅ *Ophrys anthropophora L., Sp. Pl. 2: 948
n.º 14 (1753); Orchis anthropophora (L.)
All., Fl. Pedem. 2: 148 n.º 1835 (1785);
Arachnites anthropophora (L.) F.W.
Schmidt, Fl. Boëm. 1: 77 n.º LXXXIX, tab.
103 (1793); Loroglossum anthropophorum
(L.) Rich., Orchid. Eur. Annot.: 32 (1817);
in Mém. Mus. Hist. Nat. 4: 54 (1818);
Himantoglossum anthropophorum (L.)
Spreng., Syst. Veg. 3: 694 n.º 3 (1826)
Ind. loc.: “Habitat in Italia, Lusitania,
Gallia”.
= *Ophrys anthropomorpha Willd., Sp. Pl.
4(1): 63-64 n.º 4 (1805); Aceras anthropomorphum (Willd.) Sm. in Rees, Cycl.
39(1), Aceras n.º 2 (1818); Aceras anthropophora subsp. anthropomorpha
(Willd.) Gand., Nov. Consp. Fl. Eur.: 464
(1910)
Ind. loc.: “Habitat in collibus Lusitaniae”.
= *Orchis henriquesea J.A. Guim. in Bol.
Soc. Brot. 5: 71 (1887)
– Aceras anthropophora var. balearica
R. Chodat in Bull. Trav. Soc. Bot. Genève
11: 88 (1905), nom. nud.
Ind. loc.: “colline de San Salvador”.
Anacamptis Rich., Orchid. Eur. Annot.: 25
(1817); in Mém. Mus. Hist. Nat. 4: 41, 47
(1818)
In B ATEMAN & al. (1997: 120), some
species belonging to genus Orchis are
190
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
combined under Anacamptis, but we maintain
the traditional systematic treatment for this
genus including only the following species.
Anacamptis pyramidalis (L.) Rich., Orchid.
Eur. Annot.: 33 (1817); in Mém. Mus. Hist.
Nat. 4: 55 (1818)
⬅ Orchis pyramidalis L., Sp. Pl. 2: 940 n.º 5
(1753); Aceras pyramidalis (L.) Rchb. f.,
Icon. Fl. Germ. Helv. 13/14: 6 n.º 6, tab. 9
figs. I (1850)
Ind. loc.: “Habitat in Helvetiae, Belgii,
Angliae, Galliae arenosis, cretaceis”.
Lectotypus: “Ray, Syn. meth. stirp. brit. ed.
3: tab. 18. 1724, ca. 1723, leg. Johann Jacob
Dillenius” –cf. BAUMANN & al. (1989: 439).
= Orchis condensata Desf., Fl. Atlant. 2:
316-317 (1799); Anacamptis condensata
(Desf.) K. Koch in Linnaea 22: 285 (1849),
in syn. Anacamptis pyramidalis var.
asiatica
Ind. loc.: “Habitat ad flumen Schelif
Algeriae”.
= Aceras pyramidalis var. brachystachys
Rchb. f., Icon. Fl. Germ. Helv. 13/14: 7,
tab. 9 figs. II (1850)
Ind. loc.: “Da: Lesina, Botteri! Graec.:
Parnassus Sartori! Metana Friedrichsthal!
Corinthi herb. Zuccarin! Kythaeron v.
Spruner! Hymettes Fraas! Ott. as.: Brussa
Thirke! (in herb. C. Koch)”.
Observations.–G UIMARÃES (1887: 58)
indicated this variety for Portugal. Accepting
the criteria of DELFORGE (2001: 262), we
consider that it forms a part of the wide
variability of this species in the Iberian
territory.
= Anacamptis pyramidalis subsp. algarbiensis Gand., Nov. Consp. Fl. Eur.: 463
(1910)
Ind. loc.: “Lusit. mer., Algarve Lagos
(Bourgeau N.º 2037 ex parte) [sec. Gand., Fl.
Eur. 23: 110 (1890) sub Orchis algarbiensis
Gand.]”.
Barlia Parl., Due Nuov. Gen. Monocot.: 5
(1858); Fl. Ital. 3: 445 (1858)
Barlia robertiana (Loisel.) Greuter in
Boissiera 13: 192 n.º 553 (1967)
⬅ Orchis robertiana Loisel., Fl. Gall. 2: 606
n.º 4, tab. 21 (1807); Himantoglossum
robertianum (Loisel.) P. Delforge in
Naturalistes Belges 80 (Orchid. 12): 401
(1999)
Ind. loc.: “In collibus siccis petrosis circa
Telonem”.
= Orchis longibracteata Biv., Sicul. Pl.,
Cent. 1: 57-59 n.º 66 (1806), nom. illeg.,
non F.W. Schmidt in Samml. Phys.
Aufsätze Böhm. Naturgesch. 1: 233, tab. 2
(1791); Aceras longibracteata Biv. ex
Rchb. f., Icon. Fl. Germ. Helv. 13/14: 3-4
n.º 3, tab. 27 (1850); Barlia longibracteata
(Biv. ex Rchb. f.) Parl., Due Nuov. Gen.
Monocot.: 6 (1858); Fl. Ital. 3: 447 n.º 901
(1858); Loroglossum longibracteatum
(Biv. ex Rchb. f.) Moris ex Ardoino, Fl.
Anal. Alpes-Mar.: 351 (1867); Himantoglossum longibracteatum (Biv. ex Rchb.
f.) Schltr., Orchideen 1: 52 (1914)
Ind. loc.: “In campestribus succulentis.
Panormi dietro S. Francesco di Paola fra li
ficu d’Innia di Cannatedda”.
– Ophrys aromatica Pourr. ex Colmeiro,
Enum. Pl. Penins. Hispano-Lusit. 5: 22
(1889), nom. inval., pro syn. Aceras
longibracteata
– Ophrys lusitanica Pourr. ex Colmeiro,
Enum. Pl. Penins. Hispano-Lusit. 5: 22
(1889), nom. inval., pro syn. Aceras
longibracteata
Cephalanthera Rich., Orchid. Eur. Annot.:
29 (1817); in Mém. Mus. Hist. Nat. 4: 43,
51 (1818)
Cephalanthera damasonium (Mill.) Druce
in Ann. Scott. Nat. Hist. 60: 225 (1906)
⬅ Serapias damasonium Mill., Gard. Dict.
ed. 8, Serapias n.º 2 (1768)
Ind. loc.: “Stoken Church woods in
Oxfordshire, and in several parts of
Westmoreland and Lancashire”.
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
= Epipactis alba Crantz, Stirp. Austr. Fasc.
ed. 2, 2(6): 460-462 n.º 2 (1769);
Cephalanthera alba (Crantz) Simonk.,
Enum. Fl. Transsilv.: 504 n.º 1809 (1886);
(Crantz) Fritsch in Österr. Bot. Z. 38: 81
n.º 1 (1888), comb. superfl.
Ind. loc.: “In glareosis sylvae Dornbach,
cum Epipactis purpurea, abunde in Caltenberg”.
= Serapias lonchophyllum L.f., Suppl. pl.:
405 (1781); Cephalanthera lonchophyllum
(L.f.) Rchb. f., Icon. Fl. Germ. Helv. 13-14:
136, tab. 119, in syn. Cephalanthera
grandiflora Bab.
Ind. loc.: “Habitat in Germania, Helvetia,
Ehrhart”.
= Epipactis pallens Sw. in Neues J. Bot. 1(1):
65 (1806); Cephalanthera pallens (Sw.)
Rich., Orchid. Eur. Annot.: 38 (1817); in
Mém. Mus. Hist. Nat. 4: 60 (1818)
Ind. loc.: not indicated expressly in the
protologue [Hall. helv. t. 41. Sowerb. C. Bot.
t. 271].
= Cephalanthera ochroleuca Rchb., Fl.
Germ. Excurs. 1: 140 n. 864 (1830)
191
Cephalanthera longifolia (L.) Fritsch in
Österr. Bot. Z. 38: 81 (1888)
⬅ Serapias helleborine var. longifolia L., Sp.
Pl. 2: 950 (1753)
Ind. loc.: “Habitat in Europae asperis” [sec.
L., Sp. Pl. 2: 949 n.º 1 (1753) for the species].
Neotypus: “Oeder, Fl. Dan. 3,9: 4. 1770:
Danemark: im Walde bey Dronninggaard”
–cf. BAUMANN & al. (1989: 443).
= Serapias ensifolia Murray, Syst. Veg. ed.
14: 815 n.º 3 (1784); Cephalanthera
ensifolia (Murray) Rich., Orchid. Eur.
Annot.: 38 (1817); in Mém. Mus. Hist. Nat.
4: 60 (1818)
Ind. loc.: “Habitat in Germania, Helvetia”
[sec. L.f., Suppl. pl.: 405 (1781)].
= Serapias xiphophyllum L.f., Suppl. pl.: 404
(1781); Cephalanthera xiphophyllum
(L.f.) Rchb. f., Icon. Fl. Germ. Helv. 13/14:
135 n.º 2, tab. 118 (1851)
Ind. loc.: “Habitat in Germania, Helvetia,
Ehrhart”.
Ind. loc.: “In shattigen Bergwälden in
Siebenbürgen bei Segesvár mi Reisselt nach
B m g.”.
= Cephalanthera xiphophyllum var. latifolia
Maire in Bull. Soc. Hist. Nat. Afrique N.
5(6): 239 (1914); Cephalanthera longifolia
var. latifolia (Maire) D. Rivera & López
Vélez, Orquíd. Prov. Albacete: 55 (1987)
– Serapias grandiflora L., Syst. Nat. ed. 12,
2: 594 n.º 6 (1767), p.p.; Cephalanthera
grandiflora (L.) Gray, Nat. Arr. Brit. Pl. 2:
210-211 n.º 1 (1821), p.p.
Ind. loc.: “Forêts de Quercus ilex audessous du lac de Mouzaïa; forêts de Cèdres
de l’Atlas de Blida; forêts de Quercus suber à
la Réghaïa; El-Aftroun (Brichi)”.
Ind. loc.: “in a Beachwood [sic], just below
the duke of Bedford’s seat at Cheyneis,
Buckinghamshire” [sec. Huds., Fl. Angl.: 342
(1762)].
Observations.–In the description of
Serapias grandiflora L., Syst. Nat. ed. 12, 2:
594 (1767) some species are included
–cf. MANSFELD (1938: 238)–, like S. longifolia Huds., Fl. Angl.: 341 (1762), not based
on Serapias helleborine var. longifolia L., Sp.
Pl. 2: 950 (1753), which is the basionym
of Cephalanthera longifolia (L.) Fritsch.
Therefore, Gray’s combination belongs to a
group of taxa.
= *Cephalanthera ensifolia var. cesarigonzalezii [“cesar-gonzalezii”] Rivas Goday & Bellot in Anales Jard. Bot. Madrid
6(2): 191-192 n.º 692, fig. 31 (1948)
Ind. loc.: “Habitat in sylvaticis montanis,
solo siliceo prope Santa Elena, Montium
Marianorum. 30-III-1940”.
Cephalanthera rubra (L.) Rich., Orchid.
Eur. Annot.: 38 (1817); in Mém. Mus. Hist.
Nat. 4: 60 (1818)
⬅ Serapias rubra L., Syst. Nat. ed. 12, 2: 594
n.º 7 (1767)
192
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Ind. loc.: not indicated expressly in the
protologue.
Lectotypus: “LINN 1057.5, ca. 1760, leg.
Friedrich Wilhelm von Leysser” –cf. BAUMANN & al. (1989: 444).
Observations.–W ILLKOMM (1861: 175)
includes the binomial “Epipactis rubra Asso”
among the synonyms of this species;
nevertheless, in reviewing the publications of
this author, we have only found the name
Serapias rubra L. in ASSO (1779: 131).
Coeloglossum Hartm., Handb. Skand. Fl.:
329 (1820)
BATEMAN & al. (1997: 127) subordinates
this genus under Dactylorhiza, but we
maintain the traditional systematic treatment.
Coeloglossum viride (L.) Hartm., Handb.
Skand. Fl.: 329 n.º 398 (1820)
⬅ Satyrium viride L., Sp. Pl. 2: 944 n.º 2
(1753); Orchis viridis (L.) Crantz, Stirp.
Austr. Fasc. ed. 2, 2(6): 491 n.º 10 (1769);
Habenaria viridis (L.) R. Br. in W.T.
Aiton, Hortus Kew. ed. 2, 5: 192 (1813);
Gymnadenia viridis (L.) Rich., Orchid.
Eur. Annot.: 35 (1817); in Mém. Mus.
Hist. Nat. 4: 57 (1818); Platanthera viridis
(L.) Lindl., Syn. Brit. Fl.: 261 (1829);
Himantoglossum viride (L.) Rchb., Fl.
Germ. Excurs. 1: 119 n.º 813 (1830);
Peristylus viridis (L.) Lindl., Syn. Brit. Fl.
ed. 2: 261 (1835); Dactylorhiza viridis (L.)
R.M. Bateman, Pridgeon & M.W. Chase in
Lindleyana 12(3): 129 (1997)
Ind. loc.: “Habitat in Europae frigidioris
asperis”.
Lectotypus: “Loesel, Fl. pruss.: fig. 59 dextra.
1703, sub nom. Orchis palmata flore galericulato dilute viridi, ca. 1650, leg. Johannes Loesel”
–cf. BAUMANN & al. (1989: 447).
= *Orchis viridis var. labellifida Costa ex
Vayr. in Anales Soc. Esp. Hist. Nat. 9: 97
(1880); Coeloglossum viride var. labellifidum (Costa ex Vayr.) E.G. Camus, Monogr.
Orchid.: 320 (1908); [Orchis viridis var.
labellifida Costa, Suppl. Cat. Pl. Cataluña:
78 n.º 2077 (1877), nom. nud. –Ind. loc.:
“Monseny c. de Viladrau, Trem!”]
Ind. loc.: “Bosch de las Féxas, Prat de
Campdedeu, c. Olot. Junio, 1801 (Bolós hb.),
Viladrau (Trem.)”.
Corallorhiza Châtel., Sp. Inaug. Corall.: 5
(1760), n. v.
Corallorhiza trifida Châtel., Sp. Inaug.
Corall.: 8 (1760), n. v.
⬅ Ophrys corallorhiza L., Sp. Pl. 2: 945 n.º 2
(1753), syn. subst.
Ind. loc.: “Habitat in Europae borealis
desertis”.
Lectotypus: “LINN 1056.5, pl. media
06/07. 1732, leg. Carl Linnaeus” –cf. BAUMANN & al. (1989: 449).
= Corallorhiza innata R. Br. in W.T. Aiton,
Hortus Kew. ed. 2, 5: 209 (1813)
Ind. loc.: “Nat. of Scotland”.
Cypripedium L., Sp. Pl. 2: 951 (1753)
Cypripedium calceolus L., Sp. Pl. 2: 951
n.º 1 (1753)
Ind. loc.: “Habitat in Europae, Asiae,
Americae septentrionalibus”.
Lectotypus: “Dodoens, Fl. coronar. hist.:
77. 1568 sub nom. Damasonium nothum,
ca. 1564, leg. Adolph Occo III & Hieronymus
Harder” –cf. BAUMANN & al. (1989: 452).
Dactylorhiza Neck. ex Nevski in Trudy Bot.
Inst. Akad. Nauk S.S.S.R., ser. 1, Fl. Sist.
Vyssh. Rast. 4: 332 (1937)
For the systematic treatment of this genus,
we accept the subdivision in groups assigned
in TYTECA (2001).
Dactylorhiza gr. incarnata
Dactylorhiza incarnata (L.) Soó, Nom. Nov.
Gen. Dactylorhiza: 3 (1962)
⬅ Orchis incarnata L., Fl. Suec. ed. 2: 312
n.º 802 (1755); Orchis latifolia subsp. incarnata (L.) Hook. f., Student Fl. Brit. Isl.:
353 (1870); Dactylorchis incarnatus (L.)
Verm., Stud. Dactylorch.: 65, 101 (1947)
Ind. loc.: “Habitat in pratis rarius”
[Sweden].
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
Lectotypus: “Linnaeus’s herbal in the
herbal of the Linnean Society in London sub
“incarnata” in Linnaeus’s own handwriting”,
VERMEULEN (1947: 89, 101); “LINN 1054.33,
ca. 1754, leg. Carl Linnaeus”, cf. BAUMANN &
al. (1989: 458).
= *Orchis orientalis subsp. africana Klinge
in Trudy Imp. S.-Peterburgsk. Bot. Sada
17: 186-187 n.º 3 (1898); Dactylorhiza
incarnata subsp. africana (Klinge)
H. Sund., Europ. Medit. Orchid. ed. 3: 40
(1980)
Ind. loc.: “Africa borealis: Tunesia, Algeria, Marocco; Europa: Hispania, Lusitania
(et Gallia australis?) –crescit in pratis et paludosis litoralibus et montanis et ad montium
aquas”.
= *Orchis incarnata var. maxima Cuatrec. in
Treb. Inst. Catalana Hist. Nat. 12: 253-254
(1929)
Ind. loc.: “Acequias del Gargantón, bajo el
Boquetón a unos 1000 m, 24-VI-25; Barranco
de la Tejeda, 1200 m, 19-VI-26; Cambil,
800 m, 13-VI-26; Fuenmayor, 27-VI-26”.
= *Dactylorhiza incarnata var. brevibracteata Landwehr in Orchideen 37(3): 78
(1975); Wilde Orchid. Eur. 2: 556, 55,
fig. 1 (1977)
Ind. loc.: “Teruel, Spanje. Typus: in colle.
J. Landwehr, plaat nr. 53-1, 27 mei 1963”.
= Dactylorhiza incarnata subsp. pyrenaica
Balayer in Bull. Soc. Bot. France, Lett. Bot.
133(3): 280 (1986)
Ind. loc.: “Col de Jau. Typus: Balayer,
juillet 1981 (Herbier du Laboratoire de Biologie, Université de Perpignan)”.
– Orchis latifolia L., Sp. Pl. 2: 941-942 n.º 11
(1753), p.p.; Orchis fistulosa Moench,
Meth.: 713 (1794), nom. illeg., p.p.
Ind. loc.: “Habitat in Europae pratis”.
Observations.–For comments on this last
name, see the observations in D. majalis.
Dactylorhiza incarnata shows a great
variability through the range of its distribution, and many infraspecific taxa have
193
been described. We consider them as synonyms of the species.
Dactylorhiza gr. maculata
The different taxa integrated in this group
have been considered indistinct at the specific
level or as subspecies of D. maculata. In this
paper, we accept the first proposal but they
need more detailed taxonomic studies.
Dactylorhiza caramulensis (Verm.)
D. Tyteca in Orchidophile (Asnières) 88:
155, figs. 1-2 (1989)
⬅ *Dactylorhiza maculata subsp. caramulensis Verm. in Bol. Soc. Brot. ser. 2, 44:
94-95, figs. 6-8 (1970)
Ind. loc.: “Serra do Caramulo, S. João do
Monte. Typus: in Herb. Vermeulen n.º 6089”.
Dactylorhiza ericetorum (E.F. Linton)
Aver. in Bot. Zhurn. (Moscow & St.
Petersburg) 67(3): 309 n.º 7 (1982)
⬅ Orchis maculata subsp. ericetorum E.F.
Linton, Fl. Bournemouth: 209 n.º 7
(1900); Dactylorchis maculata subsp.
ericetorum (E.F. Linton) Verm., Stud.
Dactylorch.: 69 (1947); Dactylorhiza
maculata subsp. ericetorum (E.F. Linton)
P.F. Hunt & Summerh. in Watsonia 6(2):
132 (1965); (E.F. Linton) P.F. Hunt &
Summerh. ex E.C. Nelson, Monogr.
Ikonogr. Orchid. Gattung Dactylorhiza:
101 n.º 35, Taf. 71a-c, 84e (1976), comb.
superfl.
Ind. loc.: “from Caithness and Sutherland
to the S. Coast and Guernsey, and also from
Co. Wicklow”.
Dactylorhiza fuchsii (Druce) Soó, Nom.
Nov. Gen. Dactylorhiza: 8 (1962)
⬅ Orchis fuchsii Druce in Bot. Exch. Club
Brit. Isles Rep. 4(1): 105-106 (1915);
Dactylorchis fuchsii (Druce) Verm., Stud.
Dactylorch. : 69, 142, 147 (1947)
Ind. loc.: “From Kent to Cornwall
northwards to Caithness, and in Ireland from
Cork to Antrim”.
Lectotypus: “Dr. Druce’s herbal in Oxford.
194
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Sheet with 5 specimens, no. 3.” –sec.
VERMEULEN (1947: 144, 146, 147).
= Orchis maculata var. meyeri Rchb. f., Icon.
Fl. Germ. Helv. 13/14: 67, tab. 164 (1851);
Dactylorhiza meyeri (Rchb. f.) Aver. in
Bot. Zhurn. (Moscow & St. Petersburg)
67(3): 307 (1982); Orchis maculata subsp.
meyeri (Rchb. f.) O. Bolòs & Vigo, Fl.
Països Catalans 4: 629 (2001)
Ind. loc.: “Ga: Colombelles, près de Caen
Vict. de Villaine! Carabillon près Falaise De
Brébisson! Bat. Zuid Beveland et Hulst Van
den Bosch!, Ross. Petersburg Weinmann!
Moskau W. Stolle!”.
Dactylorhiza maculata (L.) Soó, Nom. Nov.
Gen. Dactylorhiza: 7 (1962)
⬅ Orchis maculata L., Sp. Pl. 2: 942 n.º 12
(1753); Dactylorchis maculata (L.)
Verm., Stud. Dactylorch.: 68, 130 (1947)
Ind. loc.: “Habitat in Europae pratis succulentis”.
Lectotypus: “Linnaeus’s herbarium at
London or elsewhere; «12» and «maculata»
in Linnaeus’s own handwriting on the sheet”,
–sec. VERMEULEN (1947: 128-130); “LINN
1054.36, ca. 1750, leg. Carl Linnaeus”,
cf. BAUMANN & al. (1989: 467).
= *Orchis maculata var. lusitanica J.A.
Guim. in Bol. Soc. Brot. 5: 79 (1887)
Ind. loc.: “Alemdouro littoral: Serra do
Gerez (Brot., Link, Hoffmannsegg, W. fide
Reich., M. Ferreira!), Borrageiro, Ponte Feia,
Manga de Maceira, Agua da Adega, Rol da
Fonte (Moller!); Caldas do Gerez (D. Maria
L. Henriques!), Sabugueira pr. ao Covão do
Urso (Dr. J. Henriques!); Miranda (Brot.).
Beira transmontana: Alcaide (R. da Cunha!)”.
= Orchis maculata var. palustris E.G.
Camus, Monogr. Orchid.: 50 (1892)
Ind. loc.: “Frome des marais tourbeux”
[France].
– Orchis maculata var. angustifolia Lázaro
Ibiza ex Rivas Goday, Rev. Orquíd. España: 18 (1930), nom. nud.
Dactylorhiza gr. majalis
Dactylorhiza alpestris (Pugsley) Aver. in
Bot. Zhurn. (Moscow & St. Petersburg)
68(9): 1164 (1983)
⬅ Orchis alpestris Pugsley in J. Linn. Soc.,
Bot. 49: 587-588 n.º 7 (1935); Dactylorchis alpestris (Pugsley) Verm., Stud.
Dactylorch.: 67 (1947); Dactylorhiza latifolia subsp. alpestris (Pugsley) Soó, Nom.
Nov. Gen Dactylorhiza: 4 (1962); Dactylorhiza fistulosa subsp. alpestris (Pugsley)
H. Baumann & Künkele in Mitt. Arbeitskr.
Beob. Heimischer Orchideen 20(3): 628
(1988); Dactylorhiza majalis subsp.
alpestris (Pugsley) Senghas in Jahresb.
Naturw. Ver. Wuppertal 21-22: 126 (1968)
Ind. loc.: “Exsicc. Pugsley no. 517 (Zurs,
Vorarlberg) (type); Reverchon, Lautaret,
1869, in Hb. Manchester, and Gap, 1871, in
Hb. Kew., as Orchis latifolia; Krättli, Bevers,
1868, in Herb. Mus. Brit., as Orchis latifolia”.
Dactylorhiza elata (Poir.) Soó, Nom. Nov.
Gen. Dactylorhiza: 7 (1962)
⬅ Orchis elata Poir., Voy. Barbarie 2: 248
(1789); Dactylorchis elata (Poir.) Verm.,
Stud. Dactylorch.: 68 (1947); Dactylorhiza incarnata subsp. elata (Poir.) H. Sund.,
Europ. Medit. Orchid.: 45 (1975)
Ind. loc.: “Cette plante est abondante dans
les plaines de la Mazoule”.
= Orchis vestita Lag. & Rodr. in Anales Ci.
Nat. 6(16): 142-143 n.º 170 (1803); Orchis
elata var. vestita (Lag. & Rodr.) Losa & Rivas Goday in Arch. Inst. Aclim. Almería
13(1): 79 n.º 318 (1968); Dactylorhiza vestita (Lag. & Rodr.) Aver. in Bot. Zhurn.
(Moscow & St. Petersburg) 71(1): 92 (1986)
Ind. loc.: “Se cría en Tánger”.
Observations.–Index Kewensis attributes
the authority of Orchis vestita to Cavanilles.
= *Orchis sesquipedalis Willd., Sp. Pl. 4(1):
30 n.º 48 (1805); Orchis incarnata var.
sesquipedalis (Willd.) Rchb. f., Icon. Fl.
Germ. Helv. 13/14: 53, tab. 48 (1851);
Orchis incarnata subsp. sesquipedalis
(Willd.) Gand., Nov. Consp. Fl. Eur.: 461
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
(1910); Orchis elata subsp. sesquipedalis
(Willd.) Soó in Repert. Spec. Nov. Regni
Veg. 24: 31 n.º 28(2) (1928); Dactylorchis
sesquipedalis (Willd.) Verm., Stud.
Dactylorch.: 68 (1947); Dactylorhiza elata
subsp. sesquipedalis (Willd.) Soó, Nom.
Nov. Gen. Dactylorhiza: 7 (1962);
Dactylorhiza sesquipedalis (Willd.) Verm.
in Bol. Soc. Brot. sér. 2, 44: 89 (1970),
comb. inval.; (Willd.) Laínz, Aport. Fl.
Gallega VII: 31 (1971); (Willd.) Aver. in
Bot. Zhurn. (Moscow & St. Petersburg)
68(7): 895 (1983), comb. superfl.; Dactylorhiza elata var. sesquipedalis (Willd.)
Landwehr, Wilde Orchid. Eur. 1: 204
(1977)
Ind. loc.: “Habitat in Lusitania”.
Observations.–The combination of Dactylorhiza sesquipedalis (Willd.) Vermeulen is
not found in Index Kewensis.
= *Orchis latifolia var. labrovaria Brot.,
Phytogr. Lusit. Select. 2: 25-26, tab. 92
(1827); Orchis lusitanica Steud., Nom. ed.
2, 2: 224 (1841); Orchis latifolia subsp.
lusitanica (Steud.) Gand., Nov. Consp. Fl.
Eur.: 461 (1910)
Ind. loc.: “Habitat in uliginosis, et pratis
humidis prope Conimbricam, circa Setubal, et
alibi in Beira et trans Tagum”.
= *Orchis durandii Boiss. & Reut., Pugill. Pl.
Afr. Bot. Hispan.: 111-112 (1852); Orchis
incarnata var. durandii (Boiss. & Reut.)
Willk. in Willk. & Lange, Prodr. Fl. Hispan.
1(1): 170 n.º 744 (1861); Orchis latifolia
var. durandii (Boiss. & Reut.) Ball in
J. Linn. Soc., Bot. 16: 672 (1878); Orchis
latifolia subsp. durandii (Boiss. & Reut.)
Trab. in Batt. & Trab., Fl. Algérie
(Monocot.): 30 (1895); Malag. in Acta
Phytotax. Barcinon. 1: 64 (1968), comb.
superfl.; Orchis incarnata subsp. durandii
(Boiss. & Reut.) Gand., Nov. Consp. Fl.
Eur.: 461 (1910); Orchis elata subsp.
durandii (Boiss. & Reut.) Soó in Repert.
Spec. Nov. Regni Veg. 24: 32 n.º 3 (1928);
Dactylorchis durandii (Boiss. & Reut.)
Verm., Stud. Dactylorch.: 68 (1947);
Dactylorhiza elata subsp. durandii (Boiss.
195
& Reut.) Soó, Nom. Nov. Gen. Dactylorhiza: 7 (1962); (Boiss. & Reut.) Molero Mesa & Pérez Raya, Fl. Sierra Nevada:
365 (1987), comb. superfl.; Dactylorhiza
durandii (Boiss. & Reut.) Verm. in Bol.
Soc. Brot. ser. 2, 44: 89 (1970), comb.
inval.; (Boiss. & Reut.) Laínz, Aport. Fl.
Gallega VII: 31 (1971); Dactylorhiza elata
var. durandii (Boiss. & Reut.) Landwehr,
Wilde Orchid. Eur. 1: 557 (1977)
Ind. loc.: “Hab. in humidis propè Tingidem
(Durand in herb. Pavon), ad aquas Sierra
Nevada in valle Dylar (Reuter) Sierra de
Alcaraz (Bourg.!)”.
= Orchis munbyana Boiss. & Reut., Pugill.
Pl. Afr. Bot. Hispan.: 112 (1852); Orchis
elata subsp. munbyana (Boiss. & Reut.)
E.G. Camus in E.G. Camus & A. Camus,
Iconogr. Orchid. Europe 1: 221 (1928);
Dactylorhiza munbyana (Boiss. & Reut.)
Holub in Folia Geobot. Phytotax. 19(2):
214 (V-1984); (Boiss. & Reut.) Aver., Bot.
Zhurn. (Moscow & St. Petersburg) 69(6):
875 n.º 17 (VI-1984), comb. superfl.
Ind. loc.: “Hab. in paludosis Algeriae prope
La Maison Carrée (Reut.)”.
= *Orchis incarnata var. ambigua J.A. Guim.
in Bol. Soc. Brot. 5: 77 (1887); Orchis
sesquipedalis var. iberica T. Stephenson in
Bull. Soc. Bot. France 75: 492 (1928), non
Orchis sesquipedalis var. ambigua Rouy,
Fl. Fr. 13: 152 (1912) [“ambiguus”]; Dactylorhiza elata var. iberica (T. Stephenson)
Soó in Ann. Univ. Sci. Budapest. Rolando
Eötvös, Sect. Biol., 3: 347 (1960), comb.
inval.
Ind. loc.: “Beira littoral: Valladares (E. Johnston! Felgueiras!), Granja (E. Johnston!
C. Barbosa), entre a Granja e Gulpilhares
(E. Johnston!)”.
Dactylorhiza majalis (Rchb.) P.F. Hunt &
Summerh. in Watsonia 6(2): 130 n.º 16
(1965); (Rchb.) Rauschert apud Hudziok in
Wiss. Z. Martin-Luther-Univ. HalleWittemberg, Math.-Naturwiss. Reihe 14:
492 (1965), n. v.
⬅ Orchis majalis Rchb., Iconogr. Bot. Pl.
196
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Crit. 6, Decas Tab. 57: 7 n.º 770, tab.
DLXV (1828); Orchis latifolia subsp.
majalis (Rchb.) Gand., Nov. Consp. Fl.
Eur.: 461 (1910); Dactylorchis majalis
(Rchb.) Verm., Stud. Dactylorch.: 67
(1947)
Ind. loc.: “nach dem Leben, aus der Flora
von Dresden”.
= Orchis latifolia L., Sp. Pl. 2: 941-942 n.º 11
(1753), p.p.; Orchis fistulosa Moench,
Meth.: 713 (1794), nom. illeg., p.p.; Orchis
maculata subsp. latifolia (L.) Bonnier &
Layens, Tabl. Syn. Pl. Vasc. France: 311
(1894); Dactylorhiza latifolia (L.) Soó,
Nom. Nov. Gen. Dactylorhiza: 4 (1962);
Dactylorhiza fistulosa Moench ex
H. Baumann & Künkele in Mitt. Arbeitskr.
Beob. Heimischer Orchideen 15: 419
(1983), nom. illeg., p.p.
Ind. loc.: “Habitat in Europae pratis”.
Observations.–In the description of Orchis
latifolia L., Sp. Pl. 2: 941-942 (1753), three
species are included. Two of them are
described subsequently in L., Fl. Suec. ed. 2:
312 (1755) with the names Orchis sambucina
and Orchis incarnata, and the third one is the
same as Reichenbach describes as O. majalis.
Recently, PEDERSEN (2000a: 299) has proposed the reject of the first linnaean taxa, and later,
the same author has published the synonymy of
these taxa, cf. PEDERSEN (2000b: 540).
– Orchis fistulosa sensu Moench, Meth.: 713
(1794); Dactylorhiza fistulosa sensu
H. Baumann & Künkele in Mitt. Arbeitskr.
Beob. Heimischer Orchideen 15: 419 (1983)
Dactylorhiza gr. sambucina
A very complex group, requiring a detailed
taxonomic and biogeographic study. Assorted
names are given for the Peninsula and
Balearics, but we maintain the latest tendency
to accept the following species.
Dactylorhiza insularis (Sommier) Landwehr in Orchidee (Hamburg) 20(3): 128
(1969); (Sommier) E.C. Nelson, Monogr.
Ikonogr. Orchid. Gattung Dactylorhiza:
104 n.º 39 (1976), comb. superfl.
⬅ Orchis insularis Sommier in Boll. Soc.
Bot. Ital., ser. 2, 1895: 247 (1895); Orchis
sambucina subsp. insularis (Sommier)
Gand., Nov. Consp. Fl. Eur.: 462 (1910);
Dactylorhiza iberica subsp. insularis
(Sommier) Soó, Nom. Nov. Gen. Dactylorhiza: 3 (1962); Dactylorhiza sambucina subsp. insularis (Sommier) Soó,
Nom. Nov. Gen. Dactylorhiza: 3 (1962)
Ind. loc.: “Isola di Giglio”.
Observations.–Index Kewensis adjudges
the authority to Sommier ex Martelli, Monocot. Sardoae fasc. 1: 58 (1896).
= *Orchis pseudosambucina subsp. castellana Rivas Goday in Farmacognosia 6:
193, 196, 197 (1945); Orchis romana var.
castellana Rivas Goday in Farmacognosia
6: 193, 197 (1945), nom. altern.; Orchis
sambucina var. castellana Rivas Goday in
Farmacognosia 6: 193, 197 (1945), nom.
altern.; Orchis sulphurea subsp. castellana
(Rivas Goday) Rivas Goday in Anales
Jard. Bot. Madrid 21(2): 306 (1963);
Dactylorhiza insularis subsp. castellana
(Rivas Goday) D. Rivera & López Vélez,
Orquíd. Prov. Albacete: 82 (1987)
Ind. loc.: “In monte S. de Alfacar regni
Granat. (Lge.)” [sec. Willk. in Willk. &
Lange, Prodr. Fl. Hispan. 1(1): 169 n.º 742
(1861), sub Orchis pseudosambucina Ten.].
= *Dactylorhiza romana subsp. bartonii
Huxley & P.F. Hunt in J. Roy. Hort. Soc.
92: 309, figs. 165-166 (1967); Dactylorhiza insularis var. bartonii (Huxley
& P.F. Hunt) Landwehr in Orchidee
(Hamburg) 20(3): 128 (1969); Dactylorhiza bartonii (Huxley & P.F. Hunt)
Aver. in Bot. Zhurn. (Moscow & St. Petersburg) 69(6): 876 n.º 20 (1984); Dactylorhiza insularis var. bartonii (Huxley
& P.F. Hunt) D. Rivera & López Vélez,
Orquíd. Prov. Albacete: 82 (1987)
Ind. loc.: “Hispania, Serranía de Cuenca,
Montes Universales, prope Tragacete. Habitat in pinetis humidis umbrosis, solo humoso.
Coll. A. J. Huxley, R. Gorer, I. Barton, A.H.
Venison, 3.VI.1966, no. 115. Holotypus: K,
s/n”.
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
– Orchis pseudosambucina sensu Willk. in
Willk. & Lange, Prodr. Fl. Hispan. 1(1):
169 n.º 742 (1861), non Ten., Cat. Hort.
Neapol. 1815: 72-73 (1815)
Dactylorhiza markusii (Tineo) H. Baumann
& Künkele in Mitt. Arbeitskr. Beob.
Heimischer Orchideen 13(4): 461 (1981)
⬅ Orchis markusii Tineo, Pl. Rar. Sicil. 1: 9
n.º 5 (1846); Orchis pseudosambucina
subsp. markusii (Tineo) Gand., Nov.
Consp. Fl. Eur.: 462 (1910); Orchis
sulphurea var. markusii (Tineo) Maire in
Jahand. & Maire, Cat. Pl. Maroc 1: 151
(1931); Orchis romana var. markusii
(Tineo) Rivas Goday in Farmacognosia
4(6): 194-195 (1945); Dactylorhiza romana subsp. markusii (Tineo) Holub in Folia
Geobot. Phytotax. 19(2): 214 (1984)
Ind. loc.: “Palermo a Gibilrossa, nel boschetto de’sugheri presso il Convento”.
= *Orchis mediterranea subsp. siciliensis
Klinge in Trudy Imp. S.-Peterburgsk. Bot.
Sada 17: 165-166 (1898); Dactylorhiza
sulphurea subsp. siciliensis (Klinge)
Franco in Bot. J. Linn. Soc. 76(4): 367
(1978); Dactylorhiza sambucina subsp.
siciliensis (Klinge) H. Sund., Europ. Medit.
Orchid. ed. 3: 40 (1980)
Ind. loc.: “Sicilia, Sardinia (Corsica?),
Algeria: an in Hispania et Lusitania?”.
= *Orchis romana var. guimaraesii E.G.
Camus in E.G. Camus & A. Camus,
Iconogr. Orchid. Europe 1: 216, pl. 33 figs.
14-15 (1928); Orchis guimaraesii (E.G.
Camus) Rivas Goday in Farmacognosia
4(6): 193, 197 (1945)
Ind. loc.: “Portugal, montagnes arides”.
– Orchis sambucina sensu Brot., Fl. Lusit. 1:
21 n.º 9 (1804), non L., Fl. Suec. ed. 2, 2:
312-313 (1755)
– Orchis sulphurea Link in Neues J. Bot.
1(3): 132 (1806), nom. inval.; Orchis
sulphurea Link in Steud., Nomencl. Bot. 1:
225 (1821), nom. inval.; Orchis pseudosambucina subsp. sulphurea Link in
Gand., Nov. Consp. Fl. Eur.: 462 (1910),
nom. inval.
197
– Orchis sulphurea sensu Samp., Lista Esp.
Herb. Portug.: 35 (1913), non Link ex
Samp., Lista Esp. Herb. Portug.: 35 (1913),
nom. illeg., ⬅ O. pseudosambucina Ten.
– Dactylorhiza sulphurea sensu Franco in
Bot. J. Linn. Soc. 76(4): 366 (1978), non
Link ex Franco in Bot. J. Linn. Soc. 76(4):
366 (1978), nom. illeg., ⬅ Orchis romana
Sebast.
Observations.–Link published a new name,
Orchis suplhurea, without description, for
O. sambucina sensu Brotero, non L., Fl. Suec.
ed. 2, 2: 312-313 (1755). Brotero, Fl. Lusit. 1:
21 n.º 9 (1804), did not publish a new
description for the portuguese plants –”Hab. in
montosis humidis circa Torre de Moncorvo et
alibi in Transmontana”–, he just transcribed
the linnaean description with a small change:
“labio ovato, trilobo” instead of “labio ovato,
subtrilobo”. Therefore, the name from Link is
nomen nudum. Later, other authors (Steudel,
Gandoger) cited O. sulphurea Link without
description. As far as we know, the first valid
publication for this name appears in Sampaio,
who mentioned O. pseudosambucina Ten. as
synonym. Finally, in FRANCO (1978: 366) the
earliest legitimate name included in Dactylorhiza sulphurea is Orchis romana Sebast.
Dactylorhiza sambucina (L.) Soó, Nom.
Nov. Gen. Dactylorhiza: 3 (1962)
⬅ Orchis sambucina L., Fl. Suec. ed. 2: 312313 n.º 803 (1755); Orchis maculata
subsp. sambucina (L.) Bonnier & Layens,
Tabl. Syn. Pl. Vasc. France: 311 (1894);
Dactylorchis sambucina (L.) Verm., Stud.
Dactylorch.: 65 (1947)
Ind. loc.: “Holmiae ad praedium Mart.
Triewald Mariaeberg, in planitie montium humentium; eandem aliis in locis juxta Holmian
letam misit Eric. Tuwen”.
Lectotypus: “LINN 1054.34, 1755,
vermutlich leg. Eric Tuwén” –cf. BAUMANN
& al. (1989: 468).
= *Orchis laurentina R. Bolós ex Vayr., Pl.
Not.: 160-161, lám. V (1879); Anales Soc.
Esp. Hist. Nat. 9: 95-96, lám. I (1880);
Orchis sambucina var. laurentina
(R. Bolós ex Vayr.) Willk., Suppl. Prodr.
198
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Fl. Hispan.: 42, 356 (1893); Orchis
sambucina subsp. laurentina (R. Bolós ex
Vayr.) Malag. in Acta Phytotax. Barcinon.
1: 64 (1968)
Ind. loc.: “Inmediaciones del lago de
Laurenti, Julio, 1801; Platraver (Bolós hb.)”.
= *Orchis sambucina subsp. asturica Gand.,
Nov. Consp. Fl. Eur.: 462 (1910)
Ind. loc.: “Hisp. bor., Burgos Soncillo
(Estebanez)” [sec. Gand., Fl. Eur. 23: 95
(1890) sub Orchis asturica Gand.].
= Orchis latifolia L., Sp. Pl. 2: 941-942 n.º 11
(1753), p.p.; Orchis fistulosa Moench,
Meth.: 713 (1794), nom. illeg., p.p.
Ind. loc.: “Habitat in Europae pratis”.
Observations.–About this last name, see
the observations in D. majalis.
= Orchis incarnata Willd., Sp. Pl. 4(1): 30
(1805), nom. illeg., non L., Fl. Suec. ed. 2:
312 n.º 802 (1755); Orchis sambucina var.
incarnata (Willd.) Amo, Fl. Faner. Penins.
Iberica 1: 455 (1871), comb. illeg.
Taxa whose presence requires confirmation:
Dactylorhiza elodes (Griseb.) Aver. in Bot.
Zhurn. (Moscow & St. Petersburg) 67(3):
309 n.º 6 (1982)
⬅ Orchis elodes Griseb., Über Bild. Torfs: 25
(1845), n. v.; Gesam. Abh.: 67 (1880); Orchis maculata subsp. elodes (Griseb.)
Gand., Nov. Consp. Fl. Eur.: 462 (1910);
Dactylorchis maculata subsp. elodes
(Griseb.) Verm., Stud. Dactylorch.: 68
(1947); Dactylorchis elodes (Griseb.)
Verm., Stud. Dactylorch.: 137 (1947);
Dactylorhiza maculata subsp. elodes
(Griseb.) Soó, Nom. Nov. Gen. Dactylorhiza: 7 (1962)
Ind. loc.: “in ericetis turfosis totius paludis
Bourtangensis sparsim” [sec. Griseb., Gesam.
Abh.: 67 (1880)].
Lectotypus: “Herb. Vermeulen Rev. Dact.
0480, 2, from the locus classicus Bourtanger
Moor”, VERMEULEN (1947: 137).
Observations.–Under the name of D. ma-
culata subsp. elodes, this plant is cited for
Portugal and central and northern Spain in SOÓ
(1960: 349). We have more indications of this
species in the Peninsula and Balearics, and all
of them probably correspond to D. ericetorum
or other species of Dactylorhiza gr. maculata.
Dactylorhiza romana (Sebast.) Soó, Nom.
Nov. Gen. Dactylorhiza: 3 (1962)
⬅ O. romana Sebast., Roman. Pl. Fasc.
Prim.: 12, tab. III (1813); Dactylorchis
romanus (Sebast.) Verm., Stud.
Dactylorch.: 65 (1947), n. v.; Dactylorhiza
sulphurea Link ex Franco in Bot. J. Linn.
Soc. 76(4): 366 (1978), nom. illeg.
Ind. loc.: “in Ericeto vulgo dicto Pigneto di
Sacchetti”.
= Orchis bracteata Ten., Prodr. Fl. Neapol.:
LII (1812), nom. illeg., non Muhl. ex
Willd., Sp. Pl. 4(1): 34 n.º 56 (1805);
Orchis pseudosambucina Ten., Cat. Hort.
Neapol. 1815: 72-73 (1815); Orchis
mediterranea subsp. pseudosambucina
(Ten.) Klinge in Trudy Imp. S.–Peterburgsk. Bot. Sada 17: 164 (1898); Orchis
pseudosambucina subsp. romana (Sebast.)
Gand., Nov. Consp. Fl. Eur.: 462 (1910),
nom. illeg.; Orchis sulphurea Link ex
Samp., Lista Esp. Herb. Portug.: 35 (1913),
nom. illeg.; Dactylorhiza sulphurea subsp.
pseudosambucina (Ten.) Franco in Bot. J.
Linn. Soc. 76(4): 366 (1978); Dactylorhiza
sambucina subsp. pseudosambucina (Ten.)
H. Sund., Europ. Medit. Orchid. ed. 3: 40
(1980)
Ind. loc.: “Nasce in gran copia al Monte
Nuovo presso Pozzuoli”.
Observations.–E.G. CAMUS & A. CAMUS
(1928: 126) indicates the presence of Orchis
romana in the Iberian territory. SOÓ (1960:
337) admits the occurrence of D. romana
subsp. romana in Spain and Portugal. In later
publications he did not cite this taxon in the
geographic area of our study. These citations
probably correspond to D. insularis or
D. markusii.
Dactylorhiza saccifera (Brongn.) Soó,
Nom. Nov. Gen. Dactylorhiza: 8 (1962)
⬅ Orchis saccifera Brongn. in Bory, Exp.
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
Sci. Morée, Bot.: 259, tab. 30 fig. 1 (1832);
Orchis maculata var. saccifera (Brongn.)
Rchb. f., Icon. Fl. Germ. Helv. 13/14: 67
n.º 5, tab. 57 (1851); Orchis maculata
subsp. saccifera (Brongn.) E.G. Camus in
E.G. Camus & A. Camus, Iconogr. Orchid.
Europe 1: 242 (1928); Dactylorchis
maculata subsp. saccifera (Brongn.)
Verm., Stud. Dactylorch.: 68 (1947);
Dactylorhiza maculata subsp. saccifera
(Brongn.) Diklić, Fl. SR Srbije 8: 77
(1976); (Brongn.) H. Sund., Europ. Medit.
Orchid. ed. 3: 40 (1980), comb. superfl.
Ind. loc.: “Lieux montueux, particulièremente dans le vallon de Messène et en
montant au Taygète”.
Observations.–SOÓ (1960: 350) cited the
type subspecies from Spain and Portugal, but
D ELFORGE (2001: 211) indicated that the
species is typical of the central and eastern
Mediterranean. Because testimonies referring
to this taxon in our geographical region are
few and quite uncertain, we doubt its presence
in the Iberian Peninsula.
Dactylorhiza traunsteineri (Saut. ex Rchb.)
Soó, Nom. Nov. Gen. Dactylorhiza: 5
(1962)
⬅ Orchis traunsteineri Saut. ex Rchb., Fl.
Germ. Excurs. 1: 14018 n.º 853 (1830);
Dactylorchis traunsteineri (Saut. ex
Rchb.) Verm., Stud. Dactylorch.: 66
(1947)
Ind. loc.: “Gegend von Kitzbühl in Tyrol:
Traunsteiner”.
Observations.–LÖVE & KJELLQVIST (1973:
176s) published a chromosome count of
material from Sierra de Cazorla (Jaen) that
they attributed to this taxon, admitting that
it is also found in other Mediterranean
countries, although they recognized the possibility that it could have been D. incarnata.
SOÓ (1960: 345) and DELFORGE (2001: 186)
considered it to be a plant of central Europe.
Epipactis Zinn, Cat. Pl. Gott.: 85 (1757)
In the last ten years, the number of taxa has
increased in the Iberian Peninsula and
Balearic Islands with the description of new
species and new mentions, and we agree with
199
BENITO AYUSO & C. HERMOSILLA (1998: 104)
when they indicate that this genus is specially
complex. The most recent studies defend the
existence of phenomena such as autogamy
which lead to considering it as a composite
of a high number of microspecies, whose
taxonomic relationships and relative ranks
require detailed analysis, including karyologic and molecular investigations.
Epipactis gr. atrorubens
Epipactis atrorubens Hoffm. ex Besser,
Prim. Fl. Galiciae Austriac. 2: 220 n.º 1091
(1809)
⬅ Epipactis latifolia subsp. atrorubens
(Hoffm. ex Besser) Bonnier & Layens,
Tabl. Syn. Pl. Vasc. France: 309 (1894)
Ind. loc.: “In collibus pagi Kleparow prope
Leopolim. In sylva prope Stracz”.
Observations.–In many publications, this
taxa is mentioned as E. atrorubens (Hoffm.)
Besser, based in Serapias atrorubens Hoffm.,
Deutschl. Fl. ed. 2, 2: 182 (1804), but this
author doesn’t want to publishe this taxon, and
he include it into the variability of S. latifolia,
cf. BAUMANN & KÜNKELE (1999: 625).
= Epipactis helleborine var. rubiginosa
Crantz, Stirp. Austr. Fasc. ed. 2, 2(6): 467468 (1769); Epipactis latifolia var. rubiginosa (Crantz) Gaudin, Fl. Helv. 5: 465 n.º
2082 (1829); Epipactis rubiginosa (Crantz)
W.D.J. Koch, Syn. Fl. Germ. Helv. ed. 2, 2:
801 n.º 2 (1844); Epipactis latifolia subsp.
rubiginosa (Crantz) Hook. f., Student Fl.
Brit. Isl.: 358 (1870); Epipactis latifolia
raça rubiginosa (Crantz) Samp., Man. Fl.
Port.: 116 (1910); Helleborine rubiginosa
(Crantz) Samp., Lista Esp. Herb. Portug.:
36 (1913); Helleborine viridans raça
rubiginosa (Crantz) Samp., Fl. Port. ed. 2:
139 (1947)
Ind. loc.: “in Baden”.
Observations.–According to Index Kewensis, Epipactis rubiginosa was described
by Crantz, but in the work of this author we
only found the variety that W.D.J. Koch later
combined.
= Epipactis helleborine var. viridans Crantz,
200
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Stirp. Austr. Fasc. ed. 2, 2(6): 467-468
(1769); Helleborine viridans (Crantz)
Samp., Lista Esp. Herb. Portug.: 36 (1913)
Ind. loc.: “in Baden, locis apricis”.
= Epipactis atropurpurea Raf., Caratt. Nuov.
Gen.: 87 n.º 208 (1810); Helleborine
atropurpurea (Raf.) Schinz & Thell. in
Vierteljahrsschr. Naturf. Ges. Zürich
53(4): 588 (1909); (Raf.) Druce in Bull.
Torrey Bot. Club 36: 547 (1909)
Ind. loc.: “L’ho osservata nei boschi
dell’Etna sopra Nicolasi e Linguagrossa”.
*Epipactis cardina Benito Ayuso & C. Hermosilla in Estud. Mus. Ci. Nat. Álava 13:
108-109, figs. 3-4, 6 (1998)
Ind. loc.: “Cantavieja (Teruel), puerto de
Cuarto Pelado, YK1489, 1640 m. Holotypus:
leg. J. Benito Ayuso & C. E. Hermosilla,
SALA n.º 95211”.
Observations.–Species recently described
from the southern part of the Sistema Ibérico mountain range. The authors defend
the specific rank of the taxon, in spite of
presenting features that could indicate a
hybrid between E. kleinii and E. tremolsii.
Epipactis kleinii M.B. Crespo, M.R. Lowe &
Piera in Taxon 50(3): 854 (2001)
= *Epipactis atrorubens subsp. parviflora
A. Niesch. & C. Niesch. in Philippia 1(2):
59-60, abb. 1-2 (1971); Epipactis parviflora (A. Niesch. & C. Niesch.) E. Klein
in Orchidee (Hamburg) 30(2): 46 (1979),
comb. illeg., non (Blume) A.A. Eaton in
Proc. Biol. Soc. Wash. 21: 65 (1908)
Ind. loc.: “Sierra de Alcaraz, südlich
von Vianos bei Alcaraz, Provinz Albacete,
Spanien”.
Epipactis microphylla (Ehrh.) Sw. in Kongl.
Vetensk. Acad. Nya Handl. ser. 2, 21: 232
(1800)
⬅ Serapias microphylla Ehrh., Beitr. Naturk.
4: 42 (1789); Epipactis latifolia var. microphylla (Ehrh.) DC. in Lam. & DC., Fl.
Franç. ed. 3, 6: 334 n.º 2039 (1815);
Epipactis helleborine var. microphylla
(Ehrh.) Rchb. f., Icon. Fl. Germ. Helv.
13/14: 141 n.º 1, tab. 132 figs. I-II (1851);
Epipactis latifolia subsp. microphylla
(Ehrh.) Bonnier & Layens, Tabl. Syn. Pl.
Vasc. France: 309 (1894); (Ehrh.) Rivas
Goday & Borja in Anales Inst. Bot. Cavanilles 19: 537 (1961), comb. inval.;
Helleborine microphylla (Ehrh.) Schinz &
Thell. in Vierteljahrsschr. Naturf. Ges.
Zürich 53(4): 589 (1909); Epipactis helleborine subsp. microphylla (Ehrh.) Rivas
Goday & Borja in Anales Inst. Bot.
Cavanilles 19: 537 (1961), comb. inval.
Ind. loc.: “Habitat in sylvis montanis
Electoratus Brunsvico-Luneburgici”.
– Epipactis confusa auct., non D.P. Young in
Bot. Not. 1953: 263 (1953)
Epipactis gr. helleborine
We maintain the criteria of other authors
who have conducted diverse investigations in
this genus (BENITO AYUSO & HERMOSILLA,
1998: 104) considering the components of
this group at the rank of species.
*Epipactis campeadorii P. Delforge in
Naturalistes Belges 76 (Orchid. 8): 90
(1995)
Ind. loc.: “Hispania, regio Castilla-León,
provincia Bravum Burgi (Burgos), prope
Manzanedo (UTM: 30TVN4549), alt. s. m.
600 m, in populeto juxta flumen Hibericum.
Holotypus: In herb. Pierre Delforge sub
n.º 9221”.
= *Epipactis hispanica Benito Ayuso &
C. Hermosilla in Estud. Mus. Ci. Nat.
Álava 13: 106, figs. 1-2, 5 (1998)
Ind. loc.: “Munilla (la Rioja), 30TWM5571,
820 m, en chopera. Holotypus: leg. J. Benito
Ayuso, SALA n.º 95208”.
= *Epipactis hispanica var. viridis Benito
Ayuso & C. Hermosilla in Estud. Mus. Ci.
Nat. Álava 13: 107, fig. 5d (1998).
Ind. loc.: “(la Rioja), Daroca, 30T
WM3491, 720 m, en chopera. Holotypus: leg.
J. Benito Ayuso, SALA n.º 95209”.
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
Epipactis helleborine (L.) Crantz, Stirp.
Austr. Fasc. ed. 2, 2(6): 467 n.º 5 (1769)
⬅ Serapias helleborine L., Sp. Pl. 2: 949
n.º 1 (1753)
Ind. loc.: “Habitat in Europae asperis”.
Lectotypus: “J. Bauhin, Hisp. Pl. 3,2: 516.
1651, fig. excl. descr. sub nom. Elleborine Dodonaei (= E. helleborine), ca. 1550, leg. Leonhart
Fuchs” –cf. BAUMANN & al. (1989: 470).
*Epipactis lusitanica D. Tyteca in Orchidophile (Asnières) 84: 218-219, figs. 2-6
(1988)
Ind. loc.: “Lusitania, Algarve, ad Monchique, alt. 550 m. Holotypus: Leg. D. Tyteca
(BR) sub n.º 1 1988/4”.
Epipactis muelleri Godfery in J. Bot. 59: 106
(1921)
⬅ Epipactis helleborine subsp. muelleri
(Godfery) O. Bolòs, Masalles & Vigo in
Collect. Bot. (Barcelona) 17(1): 96 (1988)
Ind. loc.: “Thorenc, above Grasse”.
201
on Serapias viridiflora Hoffm., Deutschl. Fl.
ed. 2, 2: 182 (1804), but this author did not
want to publish this taxon, and he included it
under the variability of S. latifolia, cf. BAUMANN & KÜNKELE (1999: 625s).
= Epipactis helleborine var. varians Crantz,
Stirp. Austr. Fasc. ed. 2, 2(6): 478 (1769);
Epipactis helleborine subsp. varians
(Crantz) H. Sund., Europ. Medit. Orchid.
ed. 3: 41 (1980)
Ind. loc.: “Vero in sylvis umbrosis.
Dornbach”.
= Epipactis purpurata Sm., Engl. Fl. 4: 4142 n.º 2 (1828); Epipactis latifolia subsp.
purpurata (Sm.) Gand., Nov. Consp. Fl.
Eur.: 459 (1910)
Ind. loc.: “In shady woods. Parasitical on
the stump of a maple or hazle [sic], in a wood
near the Noris farm, at Leigh, Worcestershire,
in 1807. Rev. Dr. Abbot”.
Epipactis gr. palustris
*Epipactis tremolsii Pau in Bol. Soc.
Aragonesa Ci. Nat. 13(2): 43 (1914)
Epipactis palustris (L.) Crantz, Stirp. Austr.
Fasc. ed. 2, 2(6): 462 (1769)
⬅ Epipactis helleborine subsp. tremolsii
(Pau) E. Klein in Orchidee (Hamburg)
30(2): 49 (1979)
⬅ Serapias helleborine var. palustris L., Sp.
Pl. 2: 950 (1753); Helleborine palustris
(L.) Schrank, Fl. Monac. 2: 190 (1814),
n. v., non Hill, Brit. Herb., fasc. 47: 478
n.º 5 (1756), nom. inval.
Ind. loc.: “Habitat in Europae asperis” [sec.
L., Sp. Pl. 2: 949 n.º 1 (1753) for the species].
Lectotypus: “Dodoens, Purgant. Hist.: 186.
1574 sub nom. Helleborine, ca. 1570, leg.
Rembert Dodoens” –cf. B AUMANN & al.
(1989: 474).
Ind. loc.: “Moncada versus Cerdañola
(Tremols: V. 1875 – Pau: V. 1882)”.
= *Epipactis tremolsii var. viridiflora Benito
Ayuso in Estud. Mus. Ci. Nat. Álava 14:
30, fig. 1.6 (1999)
Ind. loc.: “La Rioja: WM6165, Enciso.
Holotypus: Herb. J. Benito 593/98”.
Epipactis viridiflora Hoffm. ex Krock., Fl.
Siles. 3: 41 n.º 1536 (1814)
Ind. loc.: “cum Epipactis atrorubens in
pratis, ac sylvis locis humidis, turfosis, inveni
ben Starsine, item auf dem Zobtenberge”.
Neotypus: “Rchb. f., Icon. Fl. Germ. Helv.
13/14, tab. 134 figs. II (1851), sub ‘Epipactis
latifolia brevifolia IRM’ ” –cf. BAUMANN &
KÜNKELE (1999: 628).
Observations.–In some publications, this
taxon is mentioned as E. viridiflora (Hoffm.)
Rchb., Fl. Germ. Excurs. 1: 134 (1830), based
= *Serapias longiflora Asso, Syn. Stirp.
Aragon.: 131 n.º 899 (1779)
Ind. loc.: “Oritur in udis en la Palomita”.
Epipactis gr. phyllanthes
Epipactis phyllanthes G.E. Sm. in Gard.
Chron. 1852: 660 (1852)
⬅ Epipactis helleborine subsp. phyllanthes
(G.E. Sm.) H. Sund., Europ. Medit.
Orchid. ed. 3: 41 (1980)
202
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Ind. loc.: “This pretty plant was gathered
from a single locality, upon the eleveted part
of Phillis Wood, near Westdean, Sussex, on a
gravelly soil, where it occurred with Galium
verum, Fragaria vesca, &c., under the shade
od dwarf Sallows and Birches, first in seed in
Sept. 1838, and the second time in blossom
July, 1839, ...”.
= *Epipactis phyllanthes var. fageticola
C. Hermosilla in Estud. Mus. Ci. Nat. Álava 13: 136-138, fig. 9 (1998)
Ind. loc.: “Ezcaray (Bonicaparra), provincia de la Rioja, 30TWM0184, 1000 m / Holotypus: Herbario VIT n.º 56.348”.
– Epipactis leptochila auct., non (Godfery)
Godfery in J. Bot. 59: 146-147 (1921)
Taxa whose presence requires confirmation:
Epipactis distans Arv.-Touv., Essai sur
l’espèce et les variétés: 11 (1872)
Ind. loc.: “Cette plante croît sous les bois,
entre Briançon et le col du Mont-Genèvre,
souvent en compagnie de l’atrorubens; ...”.
Observations.–This species is not found in
Index Kewensis.
BENITO AYUSO & al. (1998: 55s) cite this
species, described from the Alps, for the Sistema Ibérico mountain range, that traditionally
has been reported under the name of
E. helleborine. According to DELFORGE (2001:
72), it only lives in France and Italy, and the
Iberian individuals belong to atypical forms of
E. tremolsii and E. cardina.
Epipogium S.G. Gmel. ex Borkh., Tent.
Displ. Pl. German.: 139 (1792)
Epipogium aphyllum Sw., Summa Veg.
Scand.: 32 (1814), syn. subst.
⬅ Satyrium epipogium L., Sp. Pl. 2: 945 n.º 5
(1753)
Ind. loc.: “Habitat in Sibiriae montibus
arenosis”.
Lectotypus: “J.G. Gmelin, Fl. Sibir. 1:
tab. 2, fig. II (1747)” –cf. R ENZ & T AUBENHEIM (1984: 470).
Gennaria Parl., Fl. Ital. 3: 404 (1858)
Gennaria diphylla (Link) Parl., Fl. Ital. 3:
405 n.º 888 (1858)
⬅ *Satyrium diphyllum Link in J. Bot.
(Schrader) 1799(2): 323 (1800); Orchis
cordata Willd., Sp. Pl. 4(1): 27-28 n.º 44
(1805), nom. illeg.; Habenaria cordata
Willd. ex R. Br., Prodr. 1: 312 (1810), nom.
illeg.; Gymnadenia diphylla (Link) Link,
Handbuch I: 243 n.º 7 (1829); Herminium
cordatum Willd. ex Lindl., Edward’s Bot.
Reg. 18, tab. 1499 (1832), nom. illeg.;
Peristylus cordatus Willd. ex Lindl., Gen.
Sp. Orchid. Pl.: 298 n.º 8 (1835), nom.
illeg.; Platanthera diphylla (Link) Rchb. f.,
Icon. Fl. Germ. Helv. 13/14: 128 n.º 11, tab.
84 (1851); Orchis diphylla (Link) Willk.,
Suppl. Prodr. Fl. Hispan.: 43 (1893); (Link)
Samp., Lista Esp. Herb. Portug.: 34 (1913),
comb. superfl.; Habenaria diphylla (Link)
T. Durand & Schinz, Consp. Fl. Afr. 5: 7677 n.º 40 (1894); Coeloglossum diphyllum
(Link) Fiori in Fiori & Paol., Fl. Italia 1(1):
248 (1896)
Ind. loc.: “in den Hainen an der Serra
da Arrabida, und den Fichtenwäldern um
Setuval”.
Goodyera R. Br. in W.T. Aiton, Hortus Kew.
ed. 2, 5: 197 (1813)
Goodyera repens (L.) R. Br. in W.T. Aiton,
Hortus Kew. ed. 2, 5: 198 n.º 1 (1813)
⬅ Satyrium repens L., Sp. Pl. 2: 945 n.º 6
(1753); Neottia repens (L.) Sw. in Kongl.
Vetensk. Acad. Nya Handl. 21: 226
(1800), n. v.; Gonogona repens (L.) Link,
Enum. Hort. Berol. Alt. 2: 369 n.º 4612
(1822)
Ind. loc.: “Habitat in Sueciae, Angliae,
Sibiriae, Helvetiae sylvis”.
Lectotypus: “J. Camerar. fil., Hort. Med.:
tab. 35. 1588 sub nom. Orchis radice repente,
ca. 1584, leg. Joachim Camerarius fil.”
–cf. BAUMANN & al. (1989: 479).
Gymnadenia R. Br. in W.T. Aiton, Hortus
Kew. ed. 2, 5: 191 (1813)
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
Gymnadenia conopsea (L.) R. Br. in W.T.
Aiton, Hortus Kew. ed. 2, 5: 191 n.º 1 (1813)
⬅ Orchis conopsea L., Sp. Pl. 2: 942 n.º 13
(1753)
Ind. loc.: “Habitat in Europae pratis montosis”.
Lectotypus: “L. Fuchs, Hist. Stirp.: 712.
1542 sub nom. Satyrium basilicum mas, ca.
1535-1538, leg. Leonhart Fuchs” –cf. BAUMANN & al. (1989: 449).
= Orchis conopsea var. densiflora Wahlenb.
in Kongl. Vetensk. Acad. Nya Handl. 27: 68
n.º 34 (1806); Gymnadenia densiflora
(Wahlenb.) A. Dietr. in Allg. Gartenzeitung
7(22): 170 (1839); Gymnadenia conopsea
subsp. densiflora (Wahlenb.) Gand., Nov.
Consp. Fl. Eur.: 463 (1910)
Ind. loc.: “Den ar ej funnen forr i Sverige”.
= Gymnadenia comigera Rchb., Fl. Germ.
Excurs. 1: 121 n.º 823 (1830); Gymnadenia
conopsea var. comigera (Rchb.) Rchb. f.,
Icon. Fl. Germ. Helv. 13/14: 115, tab. 71
figs. II (1851); Gymnadenia conopsea
subsp. comigera (Rchb.) Gand., Nov.
Consp. Fl. Eur.: 463 (1910)
Ind. loc.: “Auf feuchten sonnigen Wiesen
bei Suchl im Thüringer Walde, neben
F. Kletts Rohrhammer: D. Hofmann”.
= Orchis pyrenaica Philippe, Fl. Pyrénées 2:
354 (1859); Gymnadenia conopsea var.
pyrenaica (Philippe) Nym., Consp. Fl. Eur.
4: 695 (1882); Gymnadenia conopsea
subsp. pyrenaica (Philippe) Gand., Nov.
Consp. Fl. Eur.: 463 (1910); Gymnadenia
odoratissima var. pyrenaica (Philippe)
P. Delforge in Naturalistes Belges 81
(Orchid. 13): 397 (2000)
Ind. loc.: “Pyrénées centrales, pâturages et
bois subalpins; Lhéris; Pla des Pouts; Barèges
(Lap.)”.
= Gymnadenia conopsea var. intermedia
Gren., Fl. Jurass. 2: 751 (1869)
Ind. loc.: “habite les mêmes lieux [“Toute
la châine jurassique depuis la plaine jusque
sur les sommités”], et surtout la région
montagneuse”.
203
Gymnadenia odoratissima (L.) Rich.,
Orchid. Eur. Annot.: 35 (1817); in Mém.
Mus. Hist. Nat. 4: 57 (1818)
⬅ Orchis odoratissima L., Syst. Nat. ed. 10,
2: 1243 (1759); Orchis conopsea subsp.
odoratissima (L.) Bonnier & Layens,
Tabl. Syn. Pl. Vasc. France: 311 (1894)
Ind. loc.: “Habitat in Italia, Gallia, Germania, rarissima in Svecia” [sec. L., Sp. Pl. ed. 2,
2: 1335 n.º 25 (1763)].
Lectotypus: “C. Bauhin, Prodr. ed. 1: 30,
fig. dextra. 1620 sub nom. Orchis palmata
angustifolia minor, flore odoratissima,
ca. 1595, leg. Caspar Bauhin” –cf. BAUMANN
& al. (1989: 485).
= Orchis suaveolens Vill., Hist. Pl. Dauph.
2: 38-39 n.º 18, tab. 1 (1787); Gymnadenia suaveolens (Vill.) Rchb. f. in Bonplandia 4: 321 (1856); Gymnadenia odoratissima subsp. suaveolens (Vill.) Gand., Nov.
Consp. Fl. Eur.: 463 (1910)
Ind. loc.: “Il vient sur les Alpes, dans les
prairies herbeuses, à Palanfré, sous la Moucherolle, près de Grenoble”.
– Orchis conopsea sensu Asso, Syn. Stirp.
Aragon.: 130 n.º 893 (1779)
*Gymnadenia odoratissima subsp. longicalcarata C. Hermosilla & Sabando in
Estud. Mus. Ci. Nat. Álava 10-11: 123128, figs. 4c, 5, 7 (1996)
Ind. loc.: “Burgos, Valpuesta, VN8947. Holotypus: C. E. Hermosilla & J. Sabando. In
herb. VIT 25920”.
Himantoglossum Spreng., Syst. Veg. 3: 675,
694 (1826)
Himantoglossum hircinum (L.) Spreng.,
Syst. Veg. 3: 694 (1826)
⬅ Satyrium hircinum L., Sp. Pl. 2: 944 n.º 1
(1753); Orchis hircina (L.) Crantz, Stirp.
Austr. Fasc. ed. 2, 2(6): 484 n.º 5 (1769);
Loroglossum hircinum (L.) Rich., Orchid.
Eur. Annot.: 32 (1817); in Mém. Mus. Hist.
Nat. 4: 54 n.º 2(I) (1818); Aceras hircina
(L.) Lindl., Gen. Sp. Orchid. Pl.: 282 n.º 1
(1835)
204
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Ind. loc.: “Habitat in Galliae, Cantii campestribus”.
Lectotypus: “Vaill., Bot. Paris.: tab. 30,
fig. 6. 1727 sub nom. Orchis barbata foetida
J.B., ca. 1720, leg. Sebastian Vaillant”
–cf. BAUMANN & al. (1989: 491).
Limodorum Boehm. in Ludw., Defin. Gen.
Pl.: 358 (1760)
Limodorum abortivum (L.) Sw. in Nova
Acta Regiae Soc. Sci. Upsal. 6: 80 (1799)
⬅ Orchis abortiva L., Sp. Pl. 2: 943 n.º 17
(1753); Centrosis abortiva (L.) Sw., Summa Veg. Scand.: 32 (1814); Ionorchis
abortiva (L.) Beck, Fl. Nieder-Österreich
1: 215 n.º 1 (1890)
Ind. loc.: “Habitat in Gallia, Helvetia,
Anglia, Italia”.
Lectotypus: “LINN 1054.43, pl. media,
vermutlich 1738, leg. Carl Linnaeus”
–cf. BAUMANN & al. (1989: 493).
Limodorum trabutianum Batt. in Bull. Soc.
Bot. France 33: 297-298 (1886); Batt. &
Trabut, Atlas Fl. Alger. 1: 15, pl. 10 figs. 15 (1886)
⬅ Limodorum abortivum subsp. trabutianum
(Batt.) Rouy, Fl. France 13: 208 (1912);
Centrosis trabutiana (Batt.) Samp., Lista
Esp. Herb. Portug.: 36 (1913); Limodorum abortivum var. trabutianum (Batt.)
Schlecht., Monogr. Icon. Orchid. Eur. 1:
285 (1928); (Batt.) Raynaud, Orchid.
Maroc: 105 (1985), comb. superfl.
Ind. loc.: “Floret junio in fruticetis Quercus Ballotae Desf. montis Zaccar prope
Milianah”.
Syntypus: “leg. J.A. Battandier, 07-061886 (P!, herbar Cosson)” –cf. BAUMANN &
HOFFMANN (1985: 166)
= *Limodorum lusitanicum J.A. Guim. in Polytechnia 3(6): 13-14[in separatum] (1907)
Ind. loc.: “na azinhaga da Quinta do Bispo,
nos arredores de Alemquer”.
Observations.–This binomial is not reported in Index Kewensis.
= Limodorum abortivum subsp. occidentale
Rouy, Fl. France 13: 208 (1912); Limodorum abortivum var. occidentale (Rouy)
G. Keller & Soó, Monogr. Icon. Orchid.
Eur. 2(10): 361 (1938)
Ind. loc.: “Hab. – Charente-Inférieure: bois
secs à Doeuil et environs”.
Listera R. Br. in W.T. Aiton, Hortus Kew.
ed. 2, 5: 201 (1813)
Listera cordata (L.) R. Br. in W.T. Aiton,
Hortus Kew. ed. 2, 5: 201 n.º 2 (1813)
⬅ Ophrys cordata L., Sp. Pl. 2: 946 n.º 6
(1753); Neottia cordata (L.) Rich.,
Orchid. Eur. Annot.: 37 (1817); in Mém.
Mus. Hist. Nat. 4: 59 (1818)
Ind. loc.: “Habitat in Europae frigidae
sylvis humentibus”.
Lectotypus: “LINN 1056.11, vermutlich
30.08.1741, leg. Carl Linnaeus” –cf. BAUMANN & al. (1989: 497).
Listera ovata (L.) R. Br. in W.T. Aiton, Hortus Kew. ed. 2, 5: 201 n.º 1 (1813)
⬅ Ophrys ovata L., Sp. Pl. 2: 946 n.º 5
(1753); Epipactis ovata (L.) Crantz, Stirp.
Austr. Fasc. ed. 2, 2(6): 473 (1769)
Ind. loc.: “Habitat in Europa subhumidis
pratis”.
Lectotypus: “Fuchs, Hist. Stirp.: 566. 1542
sub nom. Ophris, ca. 1540, leg. Leonhart
Fuchs” –cf. BAUMANN & al. (1989: 499).
Neotinea Rchb. f., Pollin. Orchid.: 18, 29
(1852)
Neotinea maculata (Desf.) Stearn in Ann.
Mus. Goulandris 2: 79 (1975) [“1974”]
⬅ Satyrium maculatum Desf., Fl. Atlant. 2:
319-320 (1799); Peristylus maculatus
(Desf.) Rchb. f., Icon. Fl. Germ. Helv.
13/14: 2 (1850), comb. inval., pro syn.
Aceras intacta (Link) Rchb. f.; Aceras
maculata (Desf.) Gren. in Gren. & Godr.,
Fl. France 3(1): 282 (1855), comb. inval.,
in observ. Aceras densiflora (Brot.) Boiss.
Ind. loc.: “Habitat in Atlante prope Belide”.
Observations.–R EICHENBACH (1850: 2)
attributes the binomial Peristylus maculatus to
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
Lindley, Gen. Sp. Orchid. Pl.: 300 (1838). This
name does not appear either on this page nor
on others, but Peristylus atlanticus (Willd.)
Lindl. does, based on Orchis atlantica Willd.
WILLDENOW (1805: 42) wanted to combine
Satyrium maculatum Desf. under the genus
Orchis, but with the previously existing Orchis
maculata L. (1753), he gave it the name of
Orchis atlantica.
= *Orchis intacta Link in J. Bot. (Schrader)
1799(2): 322 (1800); Aceras intacta (Link)
Rchb. f., Icon. Fl. Germ. Helv. 13/14: 2 n.º 2,
tab. 148 (1850); Neotinea intacta (Link)
Rchb. f., Pollin. Orchid.: 29 (1852); Tinea intacta (Link) Boiss., Fl. Orient. 5(1): 58 (1882)
205
Ind. loc.: “cerca de Olot, Bolós” [sec. Vayr.
in Anales Soc. Esp. Hist. Nat. 11: 137-138
(1882) sub Aceras anthropophora-simia
Vayr.].
Observations.–This name was published at
first as Aceras anthropophora-simia Vayr. in
Anales Soc. Esp. Hist. Nat 11: 137-138 (1882),
and later, K. Richter proposed a new hybrid
based in the plant described previously.
ROUY (1891: 141) ran the specific epithet
as “Vayredae”, already that hybrid was
dedicated to the Catalan botanist Estanislao
Vayreda y Vila (1848-1901).
= *Neotinea maculata var. stricta Landwehr,
Wilde Orchid. Eur. 2: 556, figs. 1-4 (1977)
Ind. loc.: “an der Serra da Arrabida in der
dortigen achattigen Lorbeerhainen”.
Ind. loc.: “Villaviciosa, Spanje. Typus: in
collectie Landwehr, plaat 11, april 1963”.
= *Satyrium densiflorum Brot., Fl. Lusit. 1: 22
n.º 2 (1804); Peristylus densiflorus (Brot.)
Lindl., Gen. Sp. Orchid. Pl.: 298 n.º 7 (1835);
Aceras densiflorum (Brot.) Boiss., Voy. Bot.
Espagne 2(19): 595 n.º 1597 (1842); Coeloglossum densiflorum (Brot.) Hartm. ex
Willk. & Lange, Prodr. Fl. Hispan. 1(1): 164
n.º 719 (1861), comb. inval., pro syn. Aceras
densiflora (Brot.) Boiss.
Neottia Guett. in Hist. Acad. Roy. Sci. (Paris)
1750, Mém.: 374 (1754)
Ind. loc.: “Hab. in Arrabida (...) Commun.
a Cl P. Link”.
= Orchis secundiflora Bertol., Rar. Ital. Pl. 2:
42 n.º 8 (1806); Himantoglossum secundiflorum (Bertol.) Rchb., Fl. Germ. Excurs.
1: 120 n.º 815 (1830); Aceras secundiflora
(Bertol.) Lindl. in Edward’s Bot. Reg. 18,
n.º & tab. 1525 (1833)
Ind. loc.: “Reperitur in editiori collium
parte circum Sarzanam”.
= *Orchis ecalcarata Costa & Vayr. in Costa,
Suppl. Cat. Pl. Cataluña: 92-93 n.º 2074bis
(1877); in Anales Soc. Esp. Hist. Nat. 9: 97
(1880).
Ind. loc.: “Crescir raro c. Olot, Fontfreda!,
Bolós; Roca de Brugués supra Gabá Vayr!;
versus Reixach propè Moncada, Csta.”.
= *Aceras ⫻ vayredae K. Richt., Pl. Eur. 1:
276 n.º 3 (1890) [“Vayrae”]
Neottia nidus-avis (L.) Rich., Orchid. Eur.
Annot.: 37 n.º 1 (1817); in Mém. Mus. Hist.
Nat. 4: 59 (1818)
⬅ Ophrys nidus-avis L., Sp. Pl. 2: 945 n.º 1
(1753); Epipactis nidus-avis (L.) Crantz,
Stirp. Austr. Fasc. ed. 2, 2(6): 475 (1769);
Listera nidus-avis (L.) W. Hook., Fl. Lond.,
tab. 58 (1819); Neottidium nidus-avis (L.)
Schltdl., Fl. Berol. 1: 454 n.º 904 (1823)
Ind. loc.: “Habitat in Sueciae, Germaniae,
Galliae nemoribus”.
Lectotypus: “Lobel, Pl. Icon.: 195, fig. sin.
1581 sub nom. Nidus avis, ca. 1576-1580,
leg. Matthias de Lobel” –cf. BAUMANN & al.
(1989: 504).
Nigritella Rich., Orchid. Eur. Annot.: 26
(1817); in Mém. Mus. Hist. Nat. 4: 41, 48
(1818)
In their taxonomic and karyological study,
TEPPNER & KLEIN (1998) included the species
of Nigritella in the genus Gymnadenia, basing
that on the phylogenetic works of BATEMAN
& al. (1997) and PRIDGEON & al. (1997). In
this check list, we maintain the traditional
separation of the two genera, although large
differences do not exist between the two.
206
ANALES JARDÍN BOTÁNICO DE MADRID, 59(2) 2002
Traditionally, it was thought that all
material from Spain belonged to N. nigra s.l.
and frequently, N. nigra subsp. nigra, in the
manner that the diverse synonyms utilized by
authors in the distant past can not be ascribed
with certainty to either of the two taxa now
under consideration. Actually, two taxa are
accepted for our flora, N. gabasiana among
the diploid taxa, and N. nigra subsp. gallica
among the polyploid and apomictic taxa.
*Nigritella gabasiana Teppner & E. Klein in
Phyton (Horn) 33(2): 182-192 n.º 3 (1994)
⬅ Gymnadenia gabasiana (Teppner &
E. Klein) Teppner & E. Klein in Phyton
(Horn) 38(1): 221 (1998)
Ind. loc.: “Zentral-Pyrenäen, Prov. Huesca,
oberstes Rio Gallego-Tal, S-Hänge der Peña
Foradada oberhalb El Formigal, ca. 1740 m,
Kalk, Rasen. Holotypus: leg. H. Teppner &
E. Klein (P).-Isotypen: GZU, MA”.
Observations.–DELFORGE (1994: 123) cites
N. rhellicani Teppner & E. Klein for the
Spanish Pyrenees and, with a question mark,
includes N. gabasiana as a possible synonym
of this species. TEPPNER & KLEIN (1998: 221)
accepted the treatment of different taxa that
are subordinate within the genus Gymnadenia,
both diploids but with separate distributions;
G. rhellicani grows from the Alps to northern
Greece, while G. gabasiana is distributed
between the Pyrenees and the Cantabrian
mountains.
Nigritella nigra (L.) Kirschl., Prodr. Fl. Alsace: 159 (1836); (L.) Rchb. f., Icon. Fl.
Germ. Helv. 13/14: 102 n.º 1, tab. 115
(1851), comb. superfl.
⬅ Satyrium nigrum L., Sp. Pl. 2: 944 n.º 3
(1753); Orchis nigra (L.) Scop., Fl. Carniol.
ed. 2, 2: 200 n.º 1123 (1771); Habenaria
nigra (L.) R. Br. in W.T. Aiton, Hortus
Kew. ed. 2, 5: 192 n.º 1 (1813); Gymnadenia
nigra (L.) Rchb. f. in Bonplandia 4: 321
(1856); (L.) Wettst. in Ber. Deutsch. Bot.
Ges. 7: 308 (1889), comb. superfl.
Ind. loc.: “Habitat in Alpibus Helveticis,
Lapponicis”.
Lectotypus: “LINN 1055.4, planta dextra,
ca. 1745, leg. Johan Otto Hagström” –cf. BAUMANN & al. (1989: 507).
= Nigritella angustifolia Rich., Orchid. Eur.
Annot.: 34 (1817); in Mém. Mus. Hist. Nat.
4: 56 (1818)
Ind. loc.: “Habitat in alpibus summis Europae borealis et australis”.
Nigritella nigra subsp. gallica E. Breiner &
R. Breiner in Mitt. Arbeitskr. Beob.
Heimischer Orchideen 25(4): 471 (1993);
Gymnadenia austriaca var. gallica (E. Breiner & R. Breiner) P. Delforge in Naturalistes
Belges 79 (Orchid. 11): 255 (1998)
Ind. loc.: “l’Alpe d’Huez, 1880 m, Gallia.
Holotypus: 26.06.1993, leg. E. u. R. Breiner,
Naturmuseum Augsburg, 93-351/1075”.
= Nigritella nigra var. pyrenaica Schltr. in
Repert. Spec. Nov. Regni Veg. 16: 271
(1919)
Ind. loc.: “Pyrenäen”.
Observations.–SCHLECHTER (1919: 271)
doesn’t specify if this variety is found on both
sides of the Pyrenees or not.
= *Nigritella nigra subsp. iberica Teppner &
E. Klein in Phyton (Horn) 33(2): 192-204
n.º 4 (1994); Gymnadenia nigra subsp.
iberica (Teppner & E. Klein) Teppner &
E. Klein in Phyton (Horn) 38(1): 222
(1998); Gymnadenia austriaca var. iberica
(Teppner & Klein) G. Foelsche & W. Foelsche in J. Eur. Orch. 31(4): 803 (1999)
Ind. loc.: “Zentral-Pyrenäen, Prov. Huesca,
Esera-Tal, Massiv von Castanesa, Umgebung
von Cerler, Llanos del Ampriu, ca. 1900 m,
flach gereigter, NNW exponierter Hang.
Holotypus: leg. H. Teppner & E. Klein
(MA).-Isotypen: GZU, P”.
Observations.–SANZ & NUET (1995: 112)
cite a sample for the Vall de Ribes (Gerona)
under the name of Nigritella nigra subsp.
corneliana Beauverd. However, TEPPNER &
KLEIN (1998: 221) consider this taxon as a
synonym of Gymnadenia corneliana (Beauverd) Teppner & E. Klein, that is distributed
exclusively in the western Alps.
P. GALÁN CELA & R. GAMARRA: THE IBERIAN AND BALEARIC ORCHIDS. 1
ACKNOWLEDGEMENTS
The authors wish to express their gratitude to the
personnel of the libraries of the Real Jardín
Botánico de Madrid, the Facultad de Farmacia of
the Universidad Complutense de Madrid, the
Conservatoire et Jardin Botaniques de la Ville de
Genève and of the Kew Gardens. To Drs. Félix
Muñoz Garmendia and Gonzalo Nieto Feliner, of
the Real Jardín Botánico de Madrid, for their
nomenclatural commentaries and diverse appraisals of the work. To Dr. Daniel Sánchez Mata,
who gave us very valuable bibliographic material.
To the orchidologist Carlos Hermosilla who
contributed bibliographic documentation and
important information. And, with special affection,
to Dr. Lulú Rico, of Kew Gardens, for her
imponderable help and collaboration during our
stay in London, and Dr. Cindy Roché, for the
translation to english.
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Editado por Félix Muñoz Garmendia
Aceptado para publicación: 20-XI-2001