55663202

Anales del Jardín Botánico de Madrid

Consejo Superior de Investigaciones Científicas “Real Jardín Botánico"

anales@ma-rjb.csic.es

ISSN (Versión impresa): 0211-1322

ESPAÑA

2006

Caspar Venhuis / Pepijn Venhuis / Albertine C. Ellis Adam

A NEW TONGUE-ORCHID (ORCHIDACEAE) IN SOUTHWEST SPAIN: SERAPIAS

OCCIDENTALIS

Anales del Jardín Botánico de Madrid,

july-december, año/vol. 63, número 002

Consejo Superior de Investigaciones Científicas “Real Jardín Botánico"

Madrid, España

pp. 131-143

Red de Revistas Científicas de América Latina y el Caribe, España y Portugal

Universidad Autónoma del Estado de México

http://redalyc.uaemex.mx

A new Tongue-orchid (Orchidaceae) in southwest Spain:

Serapias occidentalis

Caspar Venhuis, Pepijn Venhuis & Albertine C. Ellis-Adam

Institute for Biodiversity and Ecosystem Dynamics (IBED), Universiteit van Amsterdam, Kruislaan 118, 1098 SM,

Amsterdam, The Netherlands

casvenhuis@gmail.com

Abstract

Serapias occidentalis is described from several populations

(Campo Lugar, Obando and Aljucén) scattered over the Gua-

diana river basin in Extremadura, Spain. Morphological charac-

ters defining the new species are analysed, and differences with

related Serapias taxa from the Iberian Peninsula and other Euro-

pean countries are established. In addition, distribution, ecology

and reproduction are discussed.

Keywords: Orchidaceae, Serapias, taxonomy, Spain, Ex-

tremadura.

Resumen

Se describe Serapias occidentalis a partir de diferentes poblacio-

nes (Campo Lugar, Obando y Aljucén) situadas a lo largo de la

cuenca del río Guadiana en Extremadura, España. Se analizan

los caracteres morfológicos que definen la nueva especie y sus

diferencias con otras Serapias de la Península Ibérica y de Euro-

pa. Además, se discuten diversos aspectos de su distribución,

hábitat y reproducción.

Palabras clave: Orchidaceae, Serapias, taxonomía, España, Ex-

tremadura.

Introduction

The genus Serapias L. (Tongue-orchids) comprises

26 species (Delforge, 2002), with a predominantly

Mediterranean distribution. Its range extends from

the Azores and the Canaries in the west to the Cauca-

sus in the east, and north as far as Brittany (France)

(Gölz & Reinhard, 1980; Pérez Chiscano & al., 1991;

Delforge, 1995b, 2002).

Venhuis & al. (in prep) performed multivariate sta-

tistical analyses on all Serapias species occurring in the

Iberian Peninsula and France and distinguished two

main groups: a S. parviflora group and a S. vomeracea

group. These two groups were separated predomi-

nantly on the basis of the epichile width, which varied

between 3-8(9) mm and (7)8-28 mm, respectively. Se-

rapias vomeracea (Burm. fil.) Briq. has been reported

for nearly the entire European Mediterranean

zone (Richter, 1890; Koch, 1907; Schlechter, 1923;

Hermann, 1956; Meusel & al., 1965; Nelson,

1968; Landwehr, 1977; Gölz & Reinhard, 1980;

Moore, 1980; Meikle, 1985; Pérez Chiscano & al.,

1991; Delforge, 1995b, 2002). Most recent studies,

however, report several other Serapias species cover-

ing large parts of this distribution area and suggest

that at least parts of the populations previously con-

sidered as S. vomeracea would, in fact, be representa-

tives of those species. The taxa concerned are S. stric-

tiflora Welwitsch ex Veiga and S. elsae P. Delforge (S.

parviflora group) and S. bergonii E.G. Camus, S. ori-

entalis (Greuter) H. Baumann & Künkele, S. levanti-

na H. Baumann & Künkele and S. feldwegiana H.

Baumann & Künkele (S. vomeracea group).

According to Kreutz (2004), S. vomeracea consists

of three subspecies: S. vomeracea subsp. vomeracea

(Figs. 3c, d), S. vomeracea subsp. longipetala (Ten.) H.

Baumann & Künkele (Fig. 4a) and S. vomeracea

subsp. istriaca (Perko) Kreutz (Fig. 4b). S. vomeracea

subsp. longipetala occurs in Italy and Greece (Bau-

mann & Künkele, 1991) and S. vomeracea subsp. is-

triaca is endemic for Istria (Delforge, 2004). The dis-

tribution of S. vomeracea subsp. vomeracea probably

extends from the northeastern part of Spain (Benito

Ayuso & Tabuenca Marraco, 2001), Basque country

Anales del Jardín Botánico de Madrid

Vol. 63(2): 131-143

July-December 2006

ISSN: 0211-1322

(Lizaur & Lazare, 2004), the lower Pyrenees, south-

ern France (Gölz & Reinhard, 1980) and southern

Switzerland (Gölz & Reinhard, 1980; Moser & al.,

1999) to northern Italy (Gölz & Reinhard, 1980; Bau-

mann & Künkele, 1991; Kropf, 2002).

In the last 150 years, S. vomeracea and its synonyms

–S. pseudocordigera (Sebastiani) Moricand and S.

longipetala (Tenore) Pollini– have often been report-

ed from the Iberian Peninsula. Intensive studies dur-

ing the past 20 years have improved insight into the

Serapias group. On the basis of these studies, we sup-

posed that earlier presentations of S. vomeracea (or

equivalent names) from predominantly southwestern

coastal regions of the Iberian Peninsula by Pérez Lara

(1886), Coutinho (1913), Martínez Gámez (1921),

Camus & Camus (1928), Vicioso (1948), Bodegom

(1972), Landwehr (1977), Rivera & Cabezudo (1985),

Valdés & al. (1987), Buján & al. (1990) and Sáez & al.

(2005) probably represent S. strictiflora (Fig. 4c) or

S. elsae. Recent studies indeed confirm that S. vome-

racea does not occur in northwestern Spain (Cor-

tizo & Sahuquillo, 1999), Portugal (Tyteca, 1997;

Delforge, 2002; Venhuis & al., 2004, Sáez & al., 2005),

the province of Málaga (Lowe, 1998) and other south-

ern and central parts of Spain up to Madrid (Benito

Ayuso & Tabuenca Marraco, 2001). However, reports

of S. vomeracea by Willkomm (1861), Gandoger

(1890), Montserrat (1962), Van der Sluys & González

Artabe (1980), Romero & Rico (1989), Cebolla & Ri-

vas (1994), Delforge (1995a) and Sáez & al. (2005) of

the northeastern part of Spain presumably involve

S. vomeracea subsp. vomeracea (Table 4; Figure 8).

This is supported by the map of distribution of

S. vomeracea presented by Benito Ayuso & Tabuenca

Marraco (2001), which is based on observations to the

north of Madrid and from more northeastern locali-

ties.

In Extremadura, S. vomeracea was first reported by

Rivas Mateos (1931) as S. pseudocordigera. Subse-

quently, Rivas Goday (1964) used the name S. lon-

gipetala for plants of the same region. More recently,

Pérez Chiscano & al. (1991) mentioned S. vomeracea

subsp. vomeracea for Extremadura. However, a

meticulous morphological study by us of this taxon

from that region revealed significant differences from

data on populations collected in southern France and

western Italy, northern Italy by Gölz & Reinhard

(1980) and Baumann & Künkele (1991) and Madrid

by Benito Ayuso (pers. comm., 2005) for S. vomeracea

subsp. vomeracea. We also compared our measure-

ments with data reported by Baumann & Künkele

(1991) and data provided by Pellegrino (pers. comm.,

2005) of S. vomeracea subsp. longipetala from

132

C. Venhuis & al.

northern, central and southern Italy and Greece.

Flowers from specimens of this subspecies are more

slender than S. vomeracea subsp. vomeracea and defi-

nitely different from the plants from Extremadura.

Pérez Chiscano (1977) describes Extremaduran

populations that probably evolved by hybridisation

between S. vomeracea and S. lingua (= S. × intermedia

Forest.), but in 1991 Pérez Chiscano & al. mentioned

no hybrids with any Serapias species. Devesa Alcaraz

(1995) notes that Extremaduran S. vomeracea and

S. cordigera often form transitions towards S. lingua.

Recently, Benito Ayuso & Tabuenca Marraco (2001)

suggested that individuals on photographs of S.

vomeracea subsp. vomeracea, taken of populations in

Extremadura by Pérez Chiscano & al. (1991) differ to

some extent from S. vomeracea from the northeastern

part of Spain. Confusion is further augmented by the

fact that Tyteca (1997), Kreutz (pers. comm., 2004),

Benito Ayuso (pers. comm., 2004) and Venhuis & al.

(2004) postulated that the individuals shown in these

photos display morphological similarities with S.

cordigera L, which occurs throughout Iberian Penin-

sula (Willkomm, 1861; Nieschalk & Nieschalk, 1973;

Landwehr, 1977; Pérez Chiscano & al. 1991, Sáez &

al., 2005).

On the basis of the morphological data presented

in this paper, the putative S. vomeracea from Ex-

tremadura probably originated from hybridisation

between S. vomeracea subsp. vomeracea and S. cordi-

gera. Consequently, these plants should be considered

a new taxon, which we propose to name S. occidenta-

lis.

Methods

In the spring of 2004 we compared all the Serapias

species that occur at the western part of the Mediter-

ranean zone (mainland of Spain, Portugal, France,

and western Italy) we sought stable and uniform

characteristics that could distinguish between popu-

lations of different regions. To get a representative

view, we studied three populations per species (Table

1) and measured 25 randomly selected specimens of

each population, for a total of 75 specimens per

species. Vouchers are kept at the AMD herbarium

(Table 1). For each population, we measured fifteen

quantitative (Table 2) and eight qualitative charac-

ters; the hair density, distribution, curvation, shape

and position of the epichile, the shape and lamellae

position, hood position, the petal shape and the bract

versus hood ratio. The three populations for each

species were all chosen with a maximum possible dis-

tance (min. 50 km) between, to avoid spatial and ge-

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

netic autocorrelation as much as possible. The flow-

ers of S. × kelleri that were used in the flower analy-

ses, were collected by O. Gerbaud in April 1997 from

the Var, southern France. Endogenic habitat charac-

ters were measured in the form of the soil character-

istics acidity (pH) and electric conductivity (EC).

Grain size analyses were classified with a triangular

texture system for non-aeolian sediments after de

Bakker & Schelling (1966). Discriminant analyses

were carried out in SPSS 11.5 to determine whether

morphological characters could differentiate the

taxa.

Related species

Characters shared with S. cordigera and S. vome-

racea subsp. vomeracea suggest that S. occidentalis

originated from hybridisation between those two

species. Hybrids between S. vomeracea subsp. vome-

racea and S. cordigera were already described by Ca-

mus (1926) under the name S. × kelleri (Fig. 4d).

These hybrids occur sporadically and only where

S. vomeracea subsp. vomeracea and S. cordigera grow

abundantly together at a site. S. × kelleri has been

found near Pisa (Italy), the Var (France) and Kythera

(Greece) (Nelson, 1968).

Pérez Chiscano & al. (1991) and Delforge

(1995b, 2002) propose that S. perez-chiscanoi Acedo

is closely related to S. vomeracea subsp. vomeracea.

Discriminant analyses (Fig. 1) on morphological

data of S. vomeracea subsp. vomeracea, S. cordigera,

S. perez-chiscanoi and S. occidentalis reveal four

nicely clustered groups. The first two functions,

each with an eigenvalue above one, explained

90.8% of the total morphological variance (Table

3). Function 1 distinguishes the species by length

of the hypochile (correlation = 0.543), the width

A new Tongue-orchid (Orchidaceae) in southwest Spain: Serapias occidentalis

133

(0.211) and the length of the bracts (0.144) and

petal length (0.183) (Table 2). Function 2 distin-

guishes the species by the width of the epichile

(0.674), the width of the hypochile (0.525) and the

length of the sepals (0.214) (Table 2). Our analysis

reveals that the lamellae (stigmatic surface) in all

four species present very constant features. The

lamellae of S. occidentalis have a stable and uniform

appearance and differ from S. vomeracea subsp.

vomeracea, S. perez-chiscanoi and from S. cordigera

as well. The lamellae in S. vomeracea subsp. vomer-

acea and S. perez-chiscanoi are parallel but widely

positioned, in S. cordigera they are divergent and in

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Table 1. Sampled populations of the studied Serapias species belonging to the S. vomeracea group, including vouchers.

Species

Location

Region

Country

Voucher

S. cordigera

Cotifo

Algarve

Portugal

S. cordigera

Badajoz

Extremadura Spain

S. cordigera

Frejus

Var

France

AMD122572-122573

S. perez-chiscanoi

Badajoz

Extremadura Spain

S. perez-chiscanoi

Aljucén

Extremadura Spain

S. perez-chiscanoi

Trujillanos

Extremadura Spain

S. occidentalis

Campo Lugar

Extremadura Spain

AMD122200-122202

S. occidentalis

Obando

Extremadura Spain

AMD123300-123302

S. occidentalis

Aljucén

Extremadura Spain

S. vomeracea subsp. vomeracea Arquettes-en-Val

Aude

France

S. vomeracea subsp. vomeracea Pierrefeu-du-Var

Var

France

S. vomeracea subsp. vomeracea

Taggia

Liguria

Italy

AMD122570

Fig. 1. Simple scatterplot of discriminant scores for the first two

functions based on sixteen quantitative morphological charac-

ters.

S. occidentalis

S. cordigera

S. vomeracea

subsp. vomeracea

S. perez-chiscanoi

S. occidentalis the base is parallel with divergent

tips. The lamellae indeed show a slight resemblance

to the deeply grooved swellings of species that prob-

ably evolved from hybrid origin with S. lingua L. (S.

strictiflora, S. elsae, S. gregaria Godfery and S. olbia

Verguin), but no characters of S. lingua are present

in S. occidentalis. The combined characteristics of

the lamellae of S. vomeracea subsp. vomeracea and S.

cordigera also show similarities with those of pre-

sumed hybrids of S. lingua.

Following identification keys from Nelson

(1968), Baumann & Künkele (1991) and Delforge

(1995b, 2002), we ended up at S. orientalis. This

species, however, has an east Mediterranean distrib-

ution extending from Greece to Turkey (Nelson,

1968; Baumann & Künkele, 1991; Delforge, 1995b,

2002) so S. occidentalis cannot be considered the

same taxon.

Characters shared with S. cordigera

The dark red (red-purple) colour of the flowers is

a constant character in both taxa. The hairs on the

labellum are pinkish to reddish, growing predomi-

nantly in a central strip on both epichile and hypo-

chile, most densely at the middle of the transition be-

tween those two parts. In both taxa the length of the

sepals and petals is almost identical. The position of

the epichile is the same as in S. cordigera: both are

normally reflexed in a position parallel to the stem,

occasionally pointing slightly forwards or back-

wards. The hood is (sub-) horizontally positioned,

contrary to S. vomeracea, in which it has an erect po-

sition. When the labellum is in a flat position, the lat-

eral lobes generally slightly cover the epichile (Fig.

2). S. perez-chiscanoi (Fig. 7c, d) seems closely relat-

ed to S. cordigera (Figs. 3a, b) instead of to S. vome-

134

C. Venhuis & al.

racea (Tyteca, 1997; Venhuis & al., 2004; Sáez & al.,

2005) and S. occidentalis.

Characters shared with S. vomeracea

subsp. vomeracea

The two species have inflorescences with a similar

density of sublaxiflorous flowers: both S. occidentalis

and S. vomeracea subsp. vomeracea have on average

1.4 flowers per cm whereas the inflorescence of S.

cordigera is dense, with on average 2.1 flowers per cm.

The shape of the petals is identical. The shape and

curves of the epichile can be almost identical, al-

though the epichile of S. occidentalis can be much

wider, up to 19 mm, in contrast with a maximum of

only 14 mm in S. vomeracea subsp. vomeracea (Fig.

5a). S. vomeracea is well-known for its bracts, which

are taller than the hood (Moricand, 1820; Pollini,

1824; Willkomm, 1861; Boissier, 1881; Arcangeli,

1882; Briquet, 1910; Camus, 1928; Nelson, 1968; Pol-

unin, 1980; Moore, 1980; Pignatti, 1982; Davis, 1984;

Meikle, 1985; Feinbrun-Dothan, 1986; Delforge,

1995b, 2002, Sáez & al., 2005), a character that gener-

ally does not apply to S. occidentalis.

Differences between S. occidentalis

and the other two species

The inflorescence of S. occidentalis is rather pauci-

florous, having an average of only 4 flowers (Fig. 5d),

compared to S. vomeracea subsp. vomeracea and S.

cordigera, which both contain an average of 6 flowers

per inflorescence. Some dimensions of S. occidentalis

are larger than in both other species, which is not an

unusual phenomenon in orchid specimens of hybrid

origin. The average length of both the hypochile (Fig.

5c) and ovary is greater than in the other two species.

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Fig. 2. Labellum and lamellae shapes of Serapias perez-chiscanoi, S. cordigera, S. x kelleri, S. occidentalis and S. vomeracea subsp.

vomeracea based on averaged measurements.

1 cm

A new Tongue-orchid (Orchidaceae) in southwest Spain: Serapias occidentalis

135

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Fig. 3. a, Serapias cordigera. Badajoz, Extremadura, Spain. 16-IV-2004 (Photo: C. Venhuis); b, S. cordigera (semi-hypochromatic). Bada-

joz, Extremadura, Spain. 16-IV-2004 (Photo: C. Venhuis); c, S. vomeracea subsp. vomeracea. Biot, Alpes-Maritimes, France. 11-V-2004

(Photo: C. Venhuis); d, S. vomeracea subsp. vomeracea. Miraflores de la Sierra, Madrid, Spain. 4-VI-2000 (Photo: J. Benito Ayuso).

136

C. Venhuis & al.

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Fig. 4. a, Serapias vomeracea subsp. longipetala. Mazzarino, Sicilia, Italy. 21-IV-2004 (Photo: C.A.J. Kreutz); b, S. vomeracea subsp. is-

triaca. Premantura, Istria, Croatia. 13-V-2005 (Photo: C. Venhuis); c, S. strictiflora (semi-hypochromatic). La Línea de la Concepción,

Andalucia, Spain. 11-IV-2004 (Photo: C. Venhuis); d, S. × kelleri. Palayson, Var, France. 11-V-2003 (Photo: O. Gerbaud).

The hypochile width (Fig. 5b) is intermediate be-

tween the two proposed parental species.

Differences between S. occidentalis

and S. x kelleri

We found several S. × kelleri specimens near les

Mayons (Var, France), which at first sight showed

strong similarities with S. occidentalis. A closer exami-

A new Tongue-orchid (Orchidaceae) in southwest Spain: Serapias occidentalis

137

nation of S. × kelleri’s flowers disclosed in several mor-

phological differences. In S. × kelleri, the bracts ex-

tended well above the hood (Fig. 4d) (like S. vomeracea

subsp. vomeracea), whereas the bracts of S. occidental-

is were generally shorter (like S. cordigera). S. × kelleri’s

hypochile shape was like S. vomeracea subsp. vome-

racea and its epichile shape looked like S. cordigera. In

contrast to S. × kelleri, S. occidentalis’ hypochile shape

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Fig. 5. Boxplots of Serapias vomeracea subsp. vomeracea, S. occidentalis, S. cordigera and S. perez-chiscanoi; outliers and extremes

were removed a, Epichile width; b, Hypochile width; c, Hypochile length; d, Number of flowers per inflorescence.

S. vom. subsp. vom. S. cordigera

S. occidentalis S. perez-chiscanoi

S. vom. subsp. vom. S. cordigera

S. occidentalis S. perez-chiscanoi

S. vom. subsp. vom. S. cordigera

S. occidentalis S. perez-chiscanoi

S. vom. subsp. vom. S. cordigera

S. occidentalis S. perez-chiscanoi

resembled S. cordigera and its epichile shape was simi-

lar to S. vomeracea subsp. vomeracea, though wider.

Furthermore, S. occidentalis’ labellum colour matched

S. cordigera’s (deep red/purple), while S. × kelleri’s la-

bellum had a colour intermediate between the two

parental species. It thus seems that S. × kelleri and S.

occidentalis contain reversed characters.

Descriptio

Serapias occidentalis C. Venhuis & P. Venhuis, sp. nov.

Holotypus: HISPANIA. Extremadura: Cáceres, Cam-

po Lugar, 30STJ54, 320 m, 23-IV-2005, C. Venhuis

& P. Venhuis (AMD 122200) (isotypi, AMD

122201-122202).

Inter S. cordigeram et S. vomeraceam subsp.

vomeraceam intermediam –etsi notas discrepantes

quoque ferens–, verisimiliter ex hybridatione harum

specierum orta: differt ab ambobus hypochilo saepis-

sime longiore atque inflorescentia comparate pauciflora

(medio numero florum 4 in S. occidentali, 6 in S. cordi-

gera et in S. vomeracea subsp. vomeracea). Intermedia

certe inter eas quoad mediam latitudinem epichili et

hypochili. Orta ut supra dicitur –probabiliter– ab iis-

dem parentibus quam rara S. × kelleri; sed ab hac dis-

clusa notis compluribus –de quibus antea dictum est.

Species nova occidentalis appellatur cum proveniat

in parte occidentali dispersionis generis.

Description

Plants with 2 sessile, subglobose or ovoid pseudotu-

bercules. Stem straight and cylindrical, 10-37 cm high,

green with red-spotted base. Leaves linear-lanceolate,

4-8, i.e. 3-5 basal-leaves 4-16 cm long and 0.6-1.8 cm

wide, green with red-spotted base and 1-3 bract-like

cauline leaves. Inflorescence sub-laxiflorous and pau-

ciflorous comprising (1)2-6(9) large flowers. Bracts

ovate-lanceolate, grey-lilac with reddish or green veins

and a reddish to greenish base, 3.0-5.2(6.0) cm long

and 1.2-2.2 cm wide, normally shorter than the hood.

Sepals and petals form a pointed hood, which is

normally sub-horizontally positioned. Sepals ovate-

lanceolate, 1.8-3.2 cm long, abaxial side grey-lilac

coloured with dark-lilac to reddish veins, adaxial side

red-purple with purple veins. Petals usually 0.3 cm

shorter than sepals, from a deep-purple, orbicular

base abruptly acuminate into a slender, tapering, red

apical part. Labellum divided by a constriction into

hypochile and epichile. Hypochile with a bright red

coloured centre and covered with pinkish hairs, 1.1-

1.6 cm long and 1.7-2.5 cm wide, while the lateral

138

C. Venhuis & al.

lobes, hidden inside or slightly emerging from the

hood, are deep red to purple. Base with 2 subparallel,

closely positioned lamellae, deep purple or almost

blackish. Epichile ovate, 1.5-2.8 cm long and 0.8-1.9

cm wide, bright red, with dark red veins and short

pinkish to reddish hairs. Ovary cylindrical, sessile,

1.0-2.3 cm long, green. Pollinia yellow-green.

Distribution

The main area of distribution of S. occidentalis prob-

ably is the Guadiana river basin in central eastern Ex-

tremadura, with scattered populations throughout the

autonomic region (Fig. 8). Pérez Chiscano & al. (1991)

described 18 localities as S. vomeracea subsp. vome-

racea in Extremadura, some of them very west- and

eastwards. This distribution suggests that this taxon

can probably also can be found also in adjacent Portu-

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Table 2. Pooled within-group correlations between the discrim-

inating quantitative characters and standardized canonical dis-

criminant functions. Characters are ordered by absolute size of

correlation within function.

Function

Hypochile length

-,543* ,484 -,102

}

No of stem leaves

,241* -,086

}

,094

Bract width

-,210* ,003 ,094

Petal length

-,183* ,178 -,092

}

Bract length

-,144* -,003

}

-,066

}

Width of rosette leaves -,116* -,029

}

-,076

}

Epichile width

,101 ,674* -,302

}

Hypochile width

-,098

}

,525* -,272

}

Length of rosette leaves

,017 ,251* ,015

Sepal length

-,160

}

,214* -,062

}

Plant height

-,001

}

,143* ,009

No of flowers/inflorescence ,133 ,012 -,424*

Epichile length

-,190

}

,301 -,412*

Ovary length

-,036

}

,067 ,190*

No of rosette leaves

,147 ,129 ,158*

* Largest absolute correlation between each variable and any discrimi-

nant function.

Table 3. Eigenvalues, percentage of variance and the correla-

tion for the first three canonical functions that were used in the

discriminant analyses.

% of Cumulative Canonical

Function Eigenvalue Variance % Correlation

1 5,221

59,3 59,3 ,916

2 2,766

31,4 90,8 ,857

,810

9,2 100,0 ,669

The first 3 canonical discriminant functions were used in the analysis.

A new Tongue-orchid (Orchidaceae) in southwest Spain: Serapias occidentalis

139

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Fig. 6. Serapias occidentalis. Obando, Extremadura, Spain. 7-IV-2003 (Photo: C. Venhuis).

140

C. Venhuis & al.

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Fig. 7. a, Serapias occidentalis: Obando, Extremadura, Spain, 19-IV-2004 (Photo: C. Venhuis); b, S. occidentalis: Campo Lugar, Ex-

tremadura, Spain, 6-IV-2004 (Photo: C. Venhuis); c, S. perez-chiscanoi: Badajoz, Extremadura, Spain, 26-IV-2006 (Photo: C. Venhuis);

d, S. perez-chiscanoi: Trujillanos, Extremadura, Spain, 24-IV-2006 (Photo: C. Venhuis).

S. strictiflora

S. occidentalis

S. vomeracea

subsp.

vomeracea

gal and in the regions Castilla y León, Castilla la Man-

cha and maybe even in Andalucía. With the help of Dr.

José Luis Pérez Chiscano, we tried to retrace six locali-

ties of S. occidentalis out of the 18 known from Ex-

tremadura. At least five of them have disappeared,

probably due to vegetation succession, altered land-use

and/or livestock grazing. Nowadays, populations of S.

occidentalis have become very localized and contain

only a few plants per population, contrary to observa-

tions made by Pérez Chiscano & al. (1991) and Devesa

(1995), who described S. vomeracea subsp. vomeracea

as (relatively) common. By the end of our investigation,

we were able to find three populations in Extremadura;

the first (type locality), near Campo Lugar, contained

one hundred and ten flowering plants; the second, near

Obando, presented ninety flowering plants; and the

third population, near Aljucén, contained up to sixty

A new Tongue-orchid (Orchidaceae) in southwest Spain: Serapias occidentalis

141

flowering plants. We also found a single plant in an ex-

tended dehesa near Trujillanos. In the spring of 2005,

we found the population near Aljucén destroyed due to

ploughing for the cultivation of oats.

Habitat

S. occidentalis prefers siliceous to muddy soils,

which are humid during spring. These soils are ex-

tremely poor in nutrients (average Electric Conduc-

tivity (EC) 17.3 µS/cm) and slightly acidic (pH 4.6 -

5.2). Grain size analyses from the samples from Cam-

po Lugar and Obando reveal a very similar grain size,

so they can be classified as sandy loam, while the soil

of the third population near Aljucén may be labelled

as clay.

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Fig. 8. Probable distributions of Serapias strictiflora (Benito Ayuso & Tabuenca Marraco, 2001), S. occidentalis and S. vomeracea

subsp. vomeracea. The dots with numbers represent known localities of S. vomeracea subsp. vomeracea (Table 4) and the stars rep-

resent our study populations of S. occidentalis (A = Aljucén, B = Campo Lugar, C = Obando). S. cordigera probably occurs throughout

the entire Iberian Peninsula.

Serapias occidentalis can be found in grasslands

(dehesas) in plant communities of several alliances:

the Agrostidion castellanae-alliance and the Molinio-

Holoschoenion-alliance as well as, though less fre-

quently, the Tuberarion guttatae-alliance (Pérez

Chiscano & al., 1991). Although it prefers the same

habitat as S. perez-chiscanoi (Fig. 7c, d) (Pérez Chis-

cano & al., 1991), we only found them growing to-

gether in an extended dehesa near Obando, where

almost all Serapias-species of Extremadura are pre-

sent. The population near Campo Lugar also con-

tained 2 plants of S. lingua at the edge of the area; we

did not find any other Serapias species in the popula-

tion of Aljucén.

DENTIFICATION KEY FOR THE GENUS

ERAPIAS IN THE

BERIAN

ENINSULA

1. One blackish-purple swelling at the base of hypochile,

whole, deeply grooved or channelled, sometimes divided

into 2 joined lumps ........................................................... 2

1. Two parallel or divergent lamellae at the base of hypochile ..

......................................................................................... 3

2. Swelling at base of hypochile whole or very slightly grooved

and emarginated .................................................. S. lingua

2. Swelling at base of hypochile deeply grooved or channelled

...................................................................... S. strictiflora

3. Epichile width 3-6 mm; petals wide 2-4 mm .... S. parviflora

3. Epichile width 7-28 mm; petals wide 4-9 mm ................... 4

4. Base of hypochile with 2 clearly diverging lamellae ...............

....................................................................... S. cordigera

4. Base of hypochile with 2 +/- parallel lamellae .................... 5

5. Ratio hypochile width / hypochile length = 2.0–2.6; entire

epichile greenish ................................... S. perez-chiscanoi

5. Ratio hypochile width / hypochile length = 1.0–2.0; epichile

pink to red ........................................................................ 6

6. Hypochile 8-12(13) mm long; lateral lobes do not cover

epichile when labellum is flat positioned; bracts generally

taller than the hood; epichile pointing generally backwards..

...................................... S. vomeracea subsp. vomeracea

6. Hypochile (10)12-16 mm long; lateral lobes slightly cover

epichile when labellum is flat positioned; bracts generally

shorter than the hood; epichile generally parallel to stem ....

................................................................... S. occidentalis

142

C. Venhuis & al.

Reproduction

S. occidentalis flowers from the end of March until the

beginning of June, depending on weather and altitude.

Like most Serapias species, S. occidentalis is an obligate

outcrosser and is pollinated by small bees that use the

hoods for shelter during bad weather, spending the

night and by inexperienced bees seeking nectar (Ven-

huis & al., 2004). Pérez Chiscano & al. (1991) studied a

population near Madrigalejo. Of 150 open flowers, 110

(66.7%) had been visited (pollinia removed and pollen

on stigma surface); 51 were occupied by male Euceras

longicornus bees. Of these 51 male bees, 19 (37.2%) had

pollinia on their heads, some with 2 as well as 4, 6, 8, 10

and even 12 pollinia due to 4, 6, 8, 10 and even 12

pollinia, due to visits to multiple flowers.

Acknowledgements

We thank Dr. José Luis Pérez Chiscano for his kind help with pro-

viding localities of the species. Dr. Francisco María Vázquez Pardo is

gratefully acknowledged for his help in providing literature about the

genus Serapias in Spain from the past 150 years. We thank Javier Ben-

ito Ayuso for photographs of Serapias vomeracea. We thank Maurice

Hooyen (Vrije Universiteit Amsterdam) for performing the grain size

analyses. Dr. Guiseppe Pellegrino (University of Calabria), Olivier

Gerbaud, Javier Benito Ayuso are gratefully acknowledged for flower

measurements of respectively Italian, French and Spanish Serapias

species. We thank Karel Kreutz, Noel Kerremans, Kees Jager, Rien

Schot, Helmut Presser, Rob Poot and Frank Verhart for providing lo-

calities of several Serapias species. Dr. Gerard Oostermeijer (Univer-

siteit van Amsterdam) is kindly acknowledged for suggestions for im-

provement on earlier versions of this paper.

References

Arcangeli, G. 1882. Compendio della Flora Italiana. Loescher,

Torino.

Bakker, H. de. & Schelling, J. 1966. Systeem van Bodemclassificatie

voor Nederland. Centrum voor Landbouwpublicaties en Land-

bouwdocumentatie, Wageningen.

Baumann, H. & Künkele, S. 1991. Die Gattung Serapias L.- eine

taxonomische Übersicht. Mitteilungsblatt Arbeitskreis Heimis-

che Orchideen, Baden-Württemberg 21: 701-946.

Anales del Jardín Botánico de Madrid 63(2): 131-143. July-December 2006. ISSN: 0211-1322

Number

Town

Region

Author(s)

Year

El Escorial

Madrid

Gandoger

1890

Guadarrama

Madrid

Willkomm

1861

Miraflores de la Sierra

Madrid

Benito Ayuso & Tabuenca Marraco

2001

Los Montes de Orduntes

Burgos

Delforge

1995

Cizur Mayor

Navarra

van der Sluys & González Artabe

1980

Mataró

Catalonia

Montserrat

1962

La Roca

Catalonia

Montserrat

1962

Blanes

Catalonia

Montserrat

1962

Table 4. Localities of Serapias vomeracea subsp. vomeracea reported in literature on the Iberian Peninsula. Numbers refer to the dis-

tribution map in Figure 8.

Benito Ayuso, J. & Tabuenca Marraco, J.M. 2001. Apuntes sobre

orquídeas Ibéricas. Estudios del Museo de Ciencias Naturales de

Álava 16: 67-87.

Bodegom, J. van. 1972. Orchideeën in Spanje. Orchideeën 34(5):

114-116.

Boissier, E. 1884. Flora Orientalis. Vol. 5. Georg, Basel.

Briquet, J.I. 1910. Prodrome de la Flore Corse. Vol. 1. Georg,

Geneva.

Buján, M., Romero, M.I. & Cremades, J. 1990. Fragmenta Choro-

logica Occidentalia, 2370-2380. Anales del Jardín Botánico de

Madrid 47(1): 223-224.

Camus, A. 1926. S. × kelleri. Riviéra Scientific 3(1): 70.

Camus, E.G. & Camus, A. 1928. Iconographie des Orchidées D’Eu-

rope et du Bassin Méditerranéen. Vol. 1. Lechevalier, Paris.

Cebolla, C. & Rivas, M.A. 1994. Atlas florae matritensis (Amarylli-

daceae, Iridaceae, Liliaceae, Orchidaceae). Fontqueria 41: 1-206.

Cortizo, C. & Sahuquillo, E. 1999. Aportaciones a la familia Or-

chidaceae en Galicia. Anales del Jardín Botánico de Madrid

57(1): 180-184.

Davis, P.H. 1984. Flora of Turkey and the East Aegean Islands. Vol.

8. University Press, Edinburgh.

Delforge, P. 1995a. Contribution à la connaissance des orchidées

de la province de Burgos (Vieille Castille, Espagne). Natura-

listes Belges 76 (Orchid. 8): 232-276.

Delforge, P. 1995b. Orchids of Britain & Europe. Harper Collins

Publishers, London.

Delforge, P. 2002. Guía de las Orquídeas de España y Europa, Norte

de África y Próximo Oriente. Lynx Edicions, Barcelona.

Delforge, P. 2004. Le Sérapias d’Elsa. Naturalistes Belges 85 (Or-

chid. 17): 103-109.

Devesa Alcaraz, J.A. 1995. Vegetación y flora de Extremadura. Uni-

vérsitas Editorial. Badajoz.

Feinbrun-Dothan, N. 1986. Flora Palaestina. Part 4. The Israel

Academy of Sciences and Humanities, Jerusalem.

Gandoger, M. 1890. Flora Europae. Vol. 22. Savy, Paris.

Gölz, P. & Reinhard, H.R. 1980. Serapias (Orchidaceae) Ergeb-

nisse statistischer und chorologischer Untersuchungen. AHO

Mitteilungsblatt, Beitrage zur Erhaltung und Erforschung

Heimischer Orchideen 12(3): 123-189.

Hermann, F. 1956. Flora von Nord- und Mitteleuropa. Fischer,

Stuttgart.

Koch, W.D.J. 1907. Synopsis der Deutschen und Schweizer Flora.

Vol. 3. Reisland, Leipzig.

Kreutz, C.A.J. 2004. Catalogue of European Orchids. Kreutz Pub-

lishers, Landgraaf.

Kropf, M. 2002. Serapias vomeracea (Burm. Fil) Briquet ssp.

vomeracea - Beobachtungen zur Bestäubungsbiologie. Die Or-

chidee 53(4): 442-446.

Landwehr, J. 1977. Wilde Orchideeën van Europa. Vol. 2. Vereniging

tot Behoud van Natuurmonumenten in Nederland, ’s Graveland.

Lizaur, X. & Lazare, J.J. 2004. Adiciones y precisions a:

“Orquídeas de Euskal Herria". Le Journal de Botanique de la

Société Botanique de France 27: 21-25.

Lowe, M.R. 1998. The Orchids of the Province of Malaga, Spain.

Journal Europäischer Orchideen 30(3): 501-570.

Martínez Gámez, V. 1921. El Paraíso de las Orquídeas ofrídeas en Es-

paña. Real Sociedad Española de Historia Natural, Extra: 471-472.

Meikle, R.D. 1985. Flora of Cyprus. Vol. 2. Bentham-Moxon Trust,

Royal Botanic Gardens, Kew.

Meusel, H., Jäger, E. & Weinert, E. 1965. Vergleichende Chorolo-

gie der Zentralleuropäischen Flora. Vol. 1. Fischer, Jena.

Montserrat, P. 1962. Flora de la cordillera litoral catalana (porción

comprendida entre los ríos Besòs y Tordera). Collectanea

Botanica (Barcelona) 6: 434-439.

A new Tongue-orchid (Orchidaceae) in southwest Spain: Serapias occidentalis

143

Moore, D.M. 1980. Serapias. In: Tutin, T.G., Heywood, V.H.,

Burges, N.A., Moore, D.M., Valentine D.H., Walters, S.M. &

Webb, D.A. (eds.). Flora Europaea. Vol. 5. Cambridge Univer-

sity Press. Cambridge.

Moricand, S. 1820. Flora Veneta. Vol. 1. Paschoud, Geneva.

Moser, D.M, Palese, R, Bäumler, B; Gygax, A, Wyler, N. 1999.

Fortschritte in der Floristik der Schweizer Flora (Gefäß-

pflanzen). Botanica Helvetica 109(2): 229-252.

Nelson, E. 1968. Monographie und Ikonographie der Orchidaceen-

Gattungen Serapias, Aceras, Loroglossum, Barlia. Nelson, Cher-

nex-Montreux.

Nieschalk, A. & Nieschalk, C. 1973. Beiträge zur Orchideenflora

Spaniens. Die Orchidee 24: 163-168.

Coutinho, A.X.P. 1913. Flora de Portugal (Plantas Vasculares). Ail-

laud, Alves & Cia, Lisboa.

Pérez Chiscano, J.L. 1977. Aportación al estudio de las orquídeas

de Extremadura. Anales del Instituto Botánico A.J. Cavanilles

34(1): 175-181.

Pérez Chiscano, J.L., Gil Llano, J.R. & Durán Oliva, F. 1991.

Orquídeas de Extremadura. Fonda Natural, Madrid.

Pérez Lara, J.M. 1886. Florula gaditana. Anales de la Sociedad Es-

pañola de Historia Natural 15: 349-475.

Pignatti, S. 1982. Flora D’Italia. Vol. 3. Edagricole, Bologna.

Pollini, C. 1824. Flora Veronensis. Vol. 3. Societas Typographica,

Verona.

Richter, K. 1890. Plantae Europeae. Vol. 1. Engelmann, Leipzig.

Polunin, O. 1980. Flowers of Greece and the Balkans. Oxford Uni-

versity Press, Oxford.

Rivas Goday, S. 1964. Vegetación y flórula de la cuenca extremeña

del Guadiana (Vegetación y flórula de la Provincia de Badajoz).

Diputación Provincia de Badajoz, Madrid.

Rivas Mateos, M. 1931. Flora de la provincia de Cáceres. Sánchez

Rodrigo, Serradilla.

Rivera, J. & Cabezudo, B. 1985. Aportaciones al conocimiento

florístico de la Sierra de Aracena (Huelva, España). Acta

Botánica Malacitana 10: 61-78.

Romero, T. & Rico, E. 1989. Flora de la cuenca del río Duratón.

Ruizia 8: 1-438.

Sáez, L., Quijada, M.P., Alarcón, M.L. & Aldasoro, J.J. 2005. Se-

rapias. In: Aedo, C. & Herrero, A. (eds.), Flora iberica. Vol. 21.

Real Jardín Botánico de Madrid (CSIC). Madrid.

Schlechter, R. 1923. Mitteilungen über europäische und mediter-

rane Orchideen. IV. Repertorium specierum novarum regni ve-

getabilis 19(1): 33-48.

Tyteca, D. 1997. The orchid flora of Portugal. Journal Europäisch-

er Orchideen 29(2/3): 185-581.

Valdés, B., Talavera, S. & Fernández Galiano, E. 1987. Flora Vas-

cular de Andalucía Occidental. Vol. 3. Ketres Editora, Barce-

lona.

Van der Sluys, M. & González Artabe, J. 1982. Orquídeas de

Navarra. Institución Príncipe de Viana, Pamplona.

Venhuis, C., Oostermeijer, J.G.B. & Tonk, J.Th.P. 2004. Conser-

vation biology of Serapias perez-chiscanoi Acedo in the Gua-

diana river basin in Extremadura (Spain). Eurorchis 16: 49-63.

Vicioso, C. 1948. Notas sobre la flora española. Anales del Jardín

Botánico de Madrid 6(2): 5-92.

Willkomm, M. 1861. Orchidaceae. In: Willkomm, M. & Lange, J.

(eds.), Prodromus Florae Hispanicae 1: 161-177. Schweizerbart,

Stuttgart. (reprint 1972. Schweizerbart, Stuttgart).

Associate Editor: C. Aedo

Received: 19-IV-2005

Accepted: 28-VI-2006

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