Phytotaxa 403 (4): 293–300 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2019 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.403.4.3

TWO NEW HYBRID DAFFODILS (NARCISSUS, AMARYLLIDACEAE)

FROM NORTHERN EXTREMADURA (CÁCERES, SPAIN)
PEDRO ESCOBAR GARCÍA
Department of Botany and Biodiversity Research, University of Vienna, Rennweg 14, 1030 Vienna, Austria,
pedro.escobar.garcia@univie.ac.at

Hybridization is since long time considered a major force shaping the diversity of daffodils (Fernandes 1967), where it
is well-documented at the homoploid (Sánchez & Castro 2011, Barra & Ureña 2014, Hervás et al. 2018, López Tirado
2018, 2019, and many others) and polyploid levels (Díaz Lifante et al. 2009, Marques et al. 2016, Marques et al. 2017).
Hybrid zones and introgression have also been studied in Narcissus (Marques et al. 2011), as well as the occurrence of
orphan hybrid populations (Marques et al. 2010), and within-species polyploid and aneuploid series (Barra & López
1984). To our knowledge, and despite morphological trait inheritance in Narcissus has not been specifically addressed
(but see Marques et al. 2011), reciprocal crosses between two given co-occurring species produce F1 offspring more
closely resembling one of the parents (heterogamy, Correns 1909) and display a marked maternal effect (Roach &
Wulff 1987). While overall uncommon among angiosperms (de Vries 1911) heterogamy has been repeatedly reported
in Asparagales (for instance in orchids, Bateman et al. 2008, Bateman et al. 2017).
Following the description of N. vitekii P. Escobar (2018: 144), the presence of its hybrids with other surrounding
species at its remarkably diverse typus locality was expected, as the Sierra de Gata (Extremadura, Spain) sustains large
populations of 7 sympatric species of daffodils growing in immediate vicinity across the ecological gradient of the
mountain: Narcissus minor subsp. asturiensis (Barra & López 1984: 376), N. bulbocodium Linnaeus (1753: 289), N.
cantabricus De Candolle in Redouté (1816: 126), N. pseudonarcissus subsp. portensis (Pugsley 1933: 61) Fernandes
(1951: 183), N. rupicola Dufour ex Schult. & Schult. fil. in Roemer & Schultes (1830: 958), N. triandrus subsp.
pallidulus (Graells 1854: 9–10) Rivas Goday (1964: 710) and N. vitekii. Many of these species hybridize, especially
N. triandrus. Here, we focus on two hitherto unreported hybrids found at the aforementioned site during the spring of
2018 (Figs. 1, 2).

MATERIALS AND METHODS

The type locality of N. vitekii in San Martín de Trevejo (Extremadura, Spain) was visited at intervals of one week during
daffodil flowering time (March–May) and screened for the presence of potentially hybrid individuals. Putative hybrid
flowering individuals were photographed in situ. Inflorescences and up to 2 leaves were collected for desiccation using
a heat press, leaving the bulbs undisturbed in the soil and avoiding to collect leaves from adjacent young individuals
originating from vegetative reproduction. Herbarium vouchers were deposited at the herbarium of the Natural History
Museum of Vienna (W) and duplicates sent to the herbaria of La Orden-Valdesequera (HSS), Real Jardín Botánico de
Madrid (MA) and University of Salamanca (SALA) when possible. Specimens representing the parental species were
photographed at 24 megapixel using a Canon EOS 80D SLR camera and a Canon macro 100 mm 2.8 L objective;
images were measured in Photoshop CS6 (Adobe Systems Incorporated 2000).
Plant morphology was recorded following Escobar (2018). Quantitative characters were recorded in a Past
spreadsheet (Hammer et al. 2001) for 122 flowers belonging to 73 individuals of N. rupicola (6), N. vitekii (20) and
N. triandrus subsp. pallidulus (47) (see Annex). Studied characters included plant size, leaf dimensions, scape length,
number of flowers per inflorescence, flower pedicel length (FPL), ovary length (OL) and width (OW), spathe length
(SL), corolla tube length (TuL), mean tepal (MTeL, MTeW) and corona (CL, CW) length and width; ratios of OL/OW,
TuL/CL, TuL/MTeL, CW/CL, SL/FPL and MTeL/MTeW. Tepal and corona shape and colour were also recorded. The
obtained dataset was merged with the matrix of Escobar (2018).
Linear Discriminant Analysis (LDA) of FPL, SL, TuL and the ratios OL/OW, TuL/CL, TuL/MTeL, CW/CL, SL/
FPL and MTeL/MTeW was performed in Past (Hammer et al. 2001), using default settings (Fig. 3).
Accepted by Mario Martínez-Azorín: 8 May 2019; published: 15 May 2019 293
FIGURE 1. A. Typical colony of short-scaped N. vitekii on an open pasture at the locus classicus in Sierra de Gata (Extremadura, Spain).
B. N. rupicola in a shady quartzite crevice near Hornachos (Badajoz). C. N. triandrus subsp. pallidulus in the immediate vicinity of N.
vitekii in a forest clearing. Note the more elevated habit of the latter in a partially shady habitat. D. N. × richardianus in a neighbouring
scrub clearing. Pale flowers and widely open coronas are the typical outcome of N. triandrus subsp. pallidulus acting as the maternal
parent (see also Fig. 2D). E. N. × kirchnerianus in an open pasture close to granite outcrops. Note the intense yellow and the narrowed
corona mouth, typical outcome of a crossing where N. vitekii acted as the maternal parent (compare with Fig. 2E). All photographs by the
author.

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FIGURE 2. Comparative plate depicting A. N. triandrus subsp. pallidulus, B. N. vitekii, and C. N. rupicola. The two first species hybridize
to form N. × richardianus (D1. Offspring of N. triandrus subsp. pallidulus ♀ × N. vitekii ♂; D2. Offspring of N. triandrus subsp. pallidulus
♂ × N. vitekii ♀; E. N. × kirchnerianus: offspring of N. rupicola ♀ × N. vitekii ♂, note the widely open corona in contrast with Fig. 1E.

TAXONOMIC TREATMENT

Narcissus × richardianus P. Escobar, sp. hybr. nov. (Figs. 1D; 2D 1, 2)

=Narcissus triandrus subsp. pallidulus (Graells) Rivas Goday × N. vitekii P. Escobar
Narcissus foliis glaucescentibus, fistulosis; scapo unifloro atque bifloro; flore pedunculato, sulphureo; laciniis acutis, corona urceolata.
Habitat in dumosis ad Sierra de Gata prope pagum San Martin de Trevejo, Extremadura borealis.
Type:—SPAIN. Extremadura: San Martín de Trevejo, monte Jálama, 26 April 2018, P. Escobar García 111/2018 (holotype W; isotypes
HSS, MA, SALA).

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Perennial bulbose herbs (8)10–20 cm tall. Leaves 2, 80–220 × 1.5–2.5 mm, erect, keeled, glaucous, glabrous,
trapezoidal in cross-section. Scapes (60)80–150 mm long, cylindrical, hollow, glaucous. Spathe (14)16–20 mm long,
membranous. Flowers solitary or in pairs, sometimes unpleasingly scented; pedicels (9)10–15(18) mm long. Ovary
(4)5–7(8) × 3–3.5(4) mm, elliptic, dark green, slightly curved; ratio ovary length/width 1.5–2 at anthesis. Perianth tube
(14)15–16.5 mm long, narrowly conical, widest at tepal insertion point, slightly curved, yellowish green. Tepals 6,
(6.5)7–9.5 × 3–4.5 mm, the inner slightly narrower, pale yellow to citrine, elliptical, mucronulate, reflexed but never
twisted; 0.4–0.6 times as long as the perianth tube. Corona bowl-shaped, sometimes narrower at mouth, 4–6 × 8–12
mm, 1.5–2.5 times as wide as long, pale yellow to citrine, with a smooth margin. Stamens unequal. Style included in
the perianth tube. Fructification not observed.
Examined material (paratypes):—SPAIN. Cáceres: San Martín de Trevejo, monte Jálama, 18 April 2018, fl., P.
Escobar García 98/2018 (W); ibidem, 18 April 2018, fl., P. Escobar García 99/2018 (W); ibidem, 26 April 2018, fl.,
P. Escobar García 108/2018 (W).
Etymology:—The species is named after the Vienna-based Austrian botanophile and benefactor Richard Grasl
(1958–).

Narcissus × kirchnerianus P. Escobar, sp. hybr. nov. (Figs. 1E, 2E)

= Narcissus rupicola Dufour ex Schult. & Schult. fil. in Roemer & Schultes × N. vitekii P. Escobar
Narcissus foliis linearibus, glaucescentibus, fistulosis; scapo unifloro; flore subsessili, aureo; laciniis acutis, corona plicata marginibus
inaequaliter crenatis atque serrulatis. Habitat in rupibus Extremadurae borealis, prope oppidulum San Martin de Trevejo.
Type:—SPAIN. Extremadura: San Martín de Trevejo, monte Jálama, 18 April 2018, P. Escobar García 100/2018 (holotype W).

Perennial bulbose herbs 10–15 cm tall. Leaves 2, 80–150 × 1–2 mm, erect, keeled, glaucous and glabrous, trapezoidal
in cross-section. Scapes 80–120 mm long, terete. Spathe 24–27 mm long, membranous. Flowers solitary, unscented;
pedicels 2–3 mm long. Ovary 5–9 × 2.5–4 mm, elliptic, green; ratio ovary length/width approximately 2. Perianth
tube 14–16 mm long, narrowly conical, yellowish green. Tepals 6, 7–8 × 4–5.5 mm, the inner slightly narrower,
deep yellow, broadly ovate with maximum width towards their central region, mucronulate, erect; approximately 0.5
times as long as the perianth tube. Corona widely conical, 3.5–4.5 × 9.5–10.5 mm, 2–3 times as wide as long, deep
yellow, with an irregularly dentate or dentate-crenated margin, sometimes narrowed at mouth. Stamens unequal. Style
included in the perianth tube. Fructification not observed.
Examined material (paratype):—SPAIN. Cáceres: San Martín de Trevejo, monte Jálama, 26 April 2018, fl., P.
Escobar García 115/2018 (W).
Etymology:—The species is named after the Austrian archivist and benefactor Otto Kirchner (1961–).

DISCUSSION

Narcissus vitekii is an ecologically versatile species that thrives in a variety of habitats, from open dry pastures to
scrub and forest clearings, tolerating partial shade from the surrounding oaks. It is on these spots where its populations
contact the abundant N. triandrus subsp. pallidulus, apparently one of the most promiscuous species of the genus (from
the number of times it is reported to participate in hybrid formation, Aedo 2003, Marques et al. 2017). At these contact
sites, N. × richardianus is frequent, and even though this hybrid taxon is biometrically well characterized and clearly
intermediate between the two parental species (Fig. 3), some qualitative characters suggest heterogamy with maternal
effect. While the offspring of the putative crossing N. triandrus subsp. pallidulus ♀ × N. vitekii ♂ (Fig. 2D1) displays
pale yellow tepals and a typically straigh corona with maximum aperture at mouth, the offspring of the putative N.
triandrus subsp. pallidulus ♂ × N. vitekii ♀ (Fig. 2D2) is more intense in colour and presents a characteristic corona
narrowed at mouth. Both variants are equally frequent but slightly different in ecological requirements and occupy
segregated spaces; the first morph grows in forest clearings among N. triandrus while the second thrives in the pastures
among N. vitekii. While overall resembling N. triandrus due to the completely reflexed tepals, N. × richardianus is
readily distinguishable from it by its broader, completely flat tepals. Even though genetic data were not gathered,
qualitative traits along with population spatial structure strongly support that N. triandrus is the maternal parent of the
forest clearing hybrids while N. vitekii is the maternal parent of the hybrids recorded in the pasture.
Where the open pastures inhabited by N. vitekii contact rocky outcrops, this species becomes a close neighbour of
N. rupicola, producing N. × kirchnerianus (Figs. 1E, 2E), a much rarer hybrid than N. × richardianus that thrives close

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to the rocks. Similarly to the latter, and in the absence of genetic data, both morphology and population spatial structure
are strongly suggestive of maternal effect. However, contrary to the latter, N. × kirchnerianus is morphologically
unstable and resembles closely the putative maternal parent both biometrically (Fig. 3) and in qualitative characters
(Figs. 1E, 2E). The offspring of N. rupicola ♀ × N. vitekii ♂ (Fig. 2E) is very similar to N. rupicola in ecology (rocky
outcrops) and morphology but smaller in plant size, with shorter tepals, a bright yellow corolla (pure N. rupicola
individuals from northern Extremadura are normally citrine), but is deprived of the intense fragrance of the alleged
maternal parent. On the other hand, the offspring of N. rupicola ♂ × N. vitekii ♀ (Fig. 2E) grows in pastures close to
the rocks and is readily distinguishable by its erect, wider tepals than N. vitekii, and its typical irregularly denticulate
corona closed at mouth.
Alternative explanations for offspring similarity to one parent in crosses do exist. Backcrossing could be responsible
for the bimodal character distribution, but lack of fructification and absence of perfect intermediate hybrid individuals
do not support this scenario. Chartier et al. (2016) documented consistent hybrid similarity to the parental species with
higher ploidy level in heteroploid crosses in Arum. However, the retrieved bimodal morphological distribution of traits
in the hybrids and the spatial structure of populations, with hybrid plants resembling more closely the parent in their
immediate vicinity, strongly suggest pervasive maternal effect.

FIGURE 3. Linear Discriminant Analysis (LDA) scatter plots of morphometrical traits with 95% confidence ellipses. 3A. Comparison of
N. × richardianus with its parental species. 3B. Same for N. × kirchnerianus.

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ACKNOWLEDGEMENTS

The retired teacher of Latin and Greek at Colegio Claret de Don Benito (Badajoz, Spain) Manuel Vicente is thankfully
acknowledged for his kind revision of the Latin diagnoses. The author would also like to thank Ernst Vitek and
Anton Igersheim (Department of Botany and Herbarium, Natural History Museum of Vienna) for generously allowing
unrestricted access to herbarium and library facilities. And last but not least, I would like to mention Richard Grasl
and Otto Kirchner, to whom these hybrids are dedicated, because without their support this contribution would have
never been possible.

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ANNEX. MATERIAL STUDIED

The following herbarium materials were added to the matrix of Escobar (2018).
Narcissus rupicola Dufour ex Schult. & Schult. fil. in Roemer & Schultes (1830: 958)
—SPAIN. Cáceres: San Martín de Trevejo, 26 April 2018, P. Escobar 117/2018 (W).
Narcissus vitekii Escobar (2018: 144)
—SPAIN. Cáceres: San Martín de Trevejo, 18 April 2018, P. Escobar 104/2018 (W); ibidem, 26 April 2018, P. Escobar
112/2018 (W).
Narcissus triandrus subsp. pallidulus (Graells 1854: 9–10) Rivas Goday (1964: 710)
—SPAIN. Badajoz: Arroyo de San Serván, 4 March 2016, P. Escobar 128/2016 (W); La Codosera, 21 February 2017,
P. Escobar 28/2017 (W); Cáceres: Cañamero, 16 March 2016, P. Escobar 199/2016 (W); San Martín de Trevejo, 18
March 2017, P. Escobar 510/2017 (W); ibidem, 26 April 2018, P. Escobar 110/2018 (W); Ciudad Real: Fuencaliente,
10 March 2017, P. Escobar 400/2017 (W); ibidem, 10 March 2017, P. Escobar 408/2017 (W).
—PORTUGAL. Alto Alentejo: Mortinho, 21 February 2017, P. Escobar 19/2017 (W); ibidem, 21 February 2017, P.
Escobar 20/2017 (W); Cabroeira de Baixo, 21 February 2017, P. Escobar 21/2017 (W); Rabaça, 21 February 2017,
P. Escobar 26/2017 (W); ibidem, 21 February 2017, P. Escobar 27/2017 (W); Beira Alta: Carregal do Sal, 14 March
2017, P. Escobar 466/2017 (W); Ervedal, 14 March 2017, P. Escobar 468/2017 (W); ibidem, 14 March 2017, P.
Escobar 469/2017 (W); Beira Interior: Mangualde, 14 March 2017, P. Escobar 470/2017 (W).

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