An Updated Taxonomic Appraisal of Narcissus (Amaryllidaceae) in the Maltese Islands
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EcoGozo Directorate, Ministry for Gozo and Planning, St Francis Square, VCT1335 Victoria, Malta
2
Department of Biology, University of Malta, MSD2080 Msida, Malta
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Author to whom correspondence should be addressed.
Diversity 2024, 16(7), 397; https://doi.org/10.3390/d16070397
Submission received: 30 May 2024
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Revised: 3 July 2024
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Accepted: 4 July 2024
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Published: 11 July 2024
(This article belongs to the Special Issue Plant Diversity on Islands)
Abstract
:A previous study of Narcissus species on the Maltese Islands had suggested the existence of a natural hybridogenous species. A fresh study on Narcissus in Malta was conducted in 2021 and 2023 with the primary aim of studying the previously reported populations of intermediate forms and to update the taxonomy of Narcissus in Malta. Four main taxa of native species were identified: N. deficiens, two distinct morphotypes of N. tazetta, and a new hybrid species. The species identified as N. deficiens replaces records of N. serotinus and N. obsoletus. One of the two morphotypes (“autumn-flowering”) is now combined as subsp. aequilimbus (previously described from Maltese material as Hermione aequilimba) and the other one (“spring-flowering”) corresponds to subsp. tazetta. The new hybrid is derived from N. deficiens and N. tazetta and is named N × briffae. This hybrid was previously incorrectly reported as N. elegans. Detailed morphometric analysis and ecological data justify these new taxonomic appraisals. A dichotomous identification key for eight Narcissus taxa (including non-native ones) is provided to facilitate the identification of the occurring and doubtful species recorded on the Maltese Islands.
1. Introduction
Narcissus is a monocot genus that has weak interspecific hybridisation barriers, and that consequently has numerous hybrids described [1], including several natural ones [2]. A few examples described from the Mediterranean region in the last 20 years, mainly by Spanish authors, include N. × alleniae Donnison-Morgan [3], N. × perangustus Fern.Casas [4], N. × alentejanus Fern.Casas and N. × perezlarae Font Quer [5], N. × egabrensis J.López-Tirado, N. × repulloi J.López-Tirado and N. × trianoi J.López-Tirado [6], and N. × hispalensis J.López-Tirado [7], amongst others. A comprehensive list of Narcissus hybrids can be found in [2].
The insularity and small size of the Maltese archipelago, its comparatively rich flora of some 800 native species and the high density of pollinators make these islands an ideal geographic niche for processes of plant hybridisation. In the last decade, several hybridogenous species have been reported from Malta, including Ophrys melitensis × tenthredinifera [8], Romulea columnae × variicolor [9], Allium commutatum × polyanthum [10], and Bolboschoenus glaucus × maritimus [11]. A natural hybridogenous species of Narcissus from the Maltese Islands was already anticipated in a preliminary study of the Narcissus [12], where two main taxa with various stages of intermediate morphotypes were identified, leading the authors to postulate a range of morphological transitions from N. serotinus L. at one end of the spectrum flowering in October and N. tazetta L. at the other end reaching its flowering peak in February–March. Within this range of morphotypes, a distinct wild daffodil has been identified and, according to the available keys used for that study [13,14], this daffodil species morphologically matched best with N. elegans Spach, owing to its narrow caniculate leaves formed during anthesis, scapes with 3–6 flowers, and tufted habitat. Introgression and hybridisation might be the explanation for these plants, as their morphology seems to be intermediate between the putative parent species [12]. No formal description of a new hybrid or declaration of an already described hybrid was reported [12].
A study on Narcissus occurring in the Maltese Islands is much needed, primarily to investigate those intermediate morphotypes [12], for possible undescribed hybrids, and to update the taxonomy of Narcissus spp. This is necessary since various works on Mediterranean Narcissus during recent years [3,5,15,16,17,18] have presented new concepts and drastic changes in the taxonomy and species delimitation of several wild daffodils in the region.
2. Materials and Method
A total of 29 populations were studied, each representing a different sample collection at least 50 m apart. These were labelled and were labelled NRC01 to NRC29 (Table 1). For each population, at least 15 individuals were examined. Selected samples were either a number of individuals scattered within a population, as in “N. serotinus”, or individuals within a tuft or cluster of plants for species such N. tazetta s.l. In any case, at least fifteen scapes were examined per population, either as individuals, or from a single tuft, or, in widespread populations, three scapes from at least five different tufts.
Morphological characters selected to compare and demarcate species were those traditionally used by various authors to discriminate between Narcissus species [2,13,14,15,19], as shown in Table 2. Character states were recorded between October 2021 and December 2023 for known populations of “N. serotinus”, “N. elegans”, N. tazetta s.l., and assumed intermediates, some of which were reported in [12]. Species demarcation was, however, measured with a new approach based on recent studies and knowledge, as discussed below.
Bulbs were not examined in order not to harm the plants and because they are not particularly diagnostic except in N. gr. bulbocodium. Whole plants were only collected for typification and subsequently deposited in an herbarium. Photographs were taken both in situ and on dissected flower parts in the laboratory. When necessary, these structures (e.g., small coronae, anthers) were observed through a stereomicroscope equipped with a calibrated micrometer (SM-4TZ-144A, AmScope, Irvine, CA, USA). Palynological analysis was carried out on fresh material as it may provide new data for the genus and give an important indication of sterility (hybridisation). Pollen from different anthers of the same flower was admixed on a droplet of 1% cotton blue solution (to increase contrast) on a microscope slide, covered with a 32 × 22 mm coverslip and pressed gently with the thumb. The slide was mounted and observed using a compound microscope (Axiolab 5, Carl-Zeiss, Oberkochen, Germany), and pollen viability, shape, and size were determined.
A leaf sample was taken and dried in silica gel at 30 °C for possible cytological or molecular investigations. The habitat and dominant vegetation present near the studied populations were also recorded.
Numerical data, namely, measurements or flower counts, were presented in a very simplified format of (minimum–maximum), hence the average followed by the range. Extreme or exceptional measurements are further indicated in parenthesis prior to or behind the minimum or maximum value.
A more meticulous analysis using more material was carried out on intermediate/introgressed morphotypes that did not match described species, and that therefore represent potential new hybrids or taxa. Further remarks on how some specific character states were observed, measured, obtained, or reported are given below.
Habitat: a general label the dominant vegetation structure and landscape in the vicinity of the studied plants.
Flowering period: Month, and period within month, subdivided into three month-periods as “beg.” (1–10 days), “mid.” (11–20 days), and “end” (21–30/31 days) of the month. Flowering dates recorded during autumn 2023 were later than expected as the first consistent rains of that season started approximately 6–8 weeks later than the climatic norm.
Plant (scape) height: Length from the ground to the tip of the scape below the pedicels. The inflorescence not included due to variable orientation affecting the observed height.
Habit: plants growing as isolated individuals, in small open clumps, in dense tufts, etc.
Leaf formation at anthesis: presence of leaves when mature flowers are present and an approximate ratio of leaf length to scape length.
Width: Maximum width (usually at the lower third of the leaf length), given as a range and average if more than one leaf was present. Length is not diagnostic and was not taken into account; however, its proportion with the length of the flowering scape was recorded.
Leaf cross-section: the mature leaf (when present) was dissected along its midpoint, and its cross-section was observed.
Cross-section of scape: observed at the upper half of the scape.
Pedicel length: Length from its attachment with the scape (base of pedicel) up to the base of the flower (apex of pedicel). The pedicels elongate with age, and hence this character has limited diagnostic value.
Flower tube: Length from the top of the ovary to its base or insertion of the lower whorl of tepals. The ovary is distinct by always being darker than the perianth tube. The shape (lateral view) of the tube was also observed, especially where it widens, and whether it was gently tapering or widening abruptly,
Tepal size: Maximum length and width of the upper and lower whorl was measured. For scapes with three or more flowers, an average was taken on three flowers per scape. The mucro was not included in the length of the tepals.
Corona colour: greenish-yellow, bright yellow (lemon-like), golden yellow, amber (orange hue).
Corona general shape: a lateral view of the corona was observed to assess which of the following shapes it possessed: parallel sides (cup-shaped), divergent sides from the base (bowl-shaped), convex sides (urn-shaped), or very shallow and divergent sides (saucer-shaped).
Corona outline shape: Observation of the outline shape observed from the frontal (top) view. Details of the margin (entire, subentire, crenulate, crenate, and lobed) were also observed. If lobed or somewhat discontinuous, a further detailed description is provided as indicated below.
Corona wall: examination of incisions or lobes in the corona wall and rim was carried out, generally using the following categories: pleated or pinched (irregular projection), notched (minute cut), cleft (and the degree of split down the corona wall), or completely incised to the base, hence forming a free segment or lobe
Corona size: given as the diameter of the longest width, either if this is the rim in divergent coronas or the walls in urn-shaped forms, followed by the height from the rim to the basal attachment with the underlying tepal.
Fruit ‘sterility’: Indicated by the presence of fruit and the presence of fertile (hard) seeds within the fruit. Failure of seed production in at least 80% of the sample size (15 plants) was considered a sign of “sterility”. This threshold was established arbitrarily but was deliberately high so as to account for the relatively small (in statistical terms) sample size.
The following palynological characters were studied:
Pollen viability: Pollen was viewed at a magnification of ×100, and its viability estimated by a simple proportion of viable turgid grains and non-viable ones. Shrunken, deformed, or empty pollen grains were assumed to be dysfunctional and inviable. This assumption was based on the authors’ past experience and was considered reasonable. This was repeated for five individual plants within a population to confirm viability, hence ensuring that sterility was not due to a localised disorder of a single aberrant plant. An approximate percentage of viable pollen was then estimated.
Pollen shape: Only the general shape of the pollen grains was described. It was beyond the scope of this study to give a rigorous description of pollen morphology.
Pollen size: The mean size and range of the pollen grains was given after measuring the longest width and breadth of 30 pollen grains using a calibrated ocular micrometer and using Pixemetre V5.10 to extrapolate the statistical measurements. The ratio of the length to the width is denoted as the Q ratio.
Morphological analysis: Morphological traits of the Narcissus specimens were analysed to determine variations among four morphs: NRC-A, NRC-B, NRC-C1, and NRC-C2. Non-metric multidimensional scaling (nMDS) was performed using Euclidean distance to visualise morphological differentiation. Missing values in the dataset were imputed with the mean of the corresponding trait. The nMDS plots were generated using the seaborn and matplotlib libraries in Python 3.11.8.
3. Results
The morphological results in this study are consistent with those reported in an earlier study [12] but are interpreted using a recently updated taxonomic approach (see Introduction). After examining all populations and samples, three distinct morphotypes were recognised, here referred to as Narc-A, Narc-B, and Narc-C. The latter was further and marginally demarcated into two slightly different entities, primarily based on differences in their phenology, habit, habitat, and some morphological traits. These were labelled as Narc-C1 and Narc-C2. The morphological features of these Narcissus morphotypes are provided in a detailed account below, tabulated and compared in Table 2 and Table 3, illustrated in Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5, while their taxonomic interpretation is analysed below.
3.1. Morphotype Narc-A (Figure 1)
This is an early-flowering Narcissus, traditionally referred to as N. serotinus, which flowers soon after the first substantial autumn rains, typically blooming as early as late September but also as late as the start of December (as was the case in autumn 2023, when the first substantial precipitation took place at the end of November 2023). These are small plants with one to (rarely) few flowers, with the leaves appearing almost always after anthesis and fruiting.
Material examined: NRC01-NRC05, NRC10: Ħondoq ir-Rummien (Qala, Gozo), 29 October 2022, 5 November 2022; NRC08: Dingli Cliffs, (Dingli, Malta), 4 November 2022; NRC09: Qortin tal-Mistra, (Mellieħa, Malta), 4 November 2022; NRC11: Ta’ Ċenċ, (Sannat, Gozo), 12 November 2022; NRC12: il-Fekruna area (Munxar, Gozo), 12 November 2022; NRC13: il-Pergla (Xaghra, Gozo) 12 November 2023; these samples shared the same habitat, which were low garigues or steppes on coralline limestone bedrock and with (at the time of anthesis) few potential competitors.
Typical accompanying vegetation include the following species: Allium polyanthum Schult. and Schult.f., Arisarum vulgare O.Targ.Tozz., Asphodelus ramosus L., Brachypodium hybridum Catalán, Joch.Müll., Hasterok and G.Jenkins, Bromus fasciculatus C.Presl, Bromus madritensis L., Chiliadenus bocconei Brullo, Erica multiflora L., Ferula melitensis Brullo, C.Brullo, Cambria, Giusso, Salmeri and Bacch., Galactites tomentosus Moench., Hyparrhenia hirta (L.) Stapf., Leontodon autumnalis L. (=Scorzoneroides autumnalis (L.) Moench), Muscari parviflorum Desf., Plantago lagopus L., Prospero cf. autumnale (L.) Speta, Ranunculus bullatus L., and Drimia pancration (Steinh.) J.C. Manning and Goldblatt.
Description: Habit, solitary, sometimes in pairs, rarely more. Leaves, absent during anthesis, formed after fruiting, linear, erect and ached, up to 10 cm long. Scape, terete, 14 ((5–20)(–30)) cm long (flower excluded), 1.0–2.0 mm diameter. Basal sheath indistinct, when occasionally emerging above ground, not more than 3 mm high, hyaline with very faint veins. Bract, membranous, frail, lanceolate, beige, 22 ((18–25(–34)) × 6 (5–7) mm. Flowers 1, occasionally 2–3 (Figure 1b), very rarely more, 17–35 mm in diameter (remarkably variable in size, Figure 1a), pure white, with a very light flushing of pale green or yellow adjacent to the corona, delicately fragrant. Pedicel(s), 5–15 mm long. Perianth tube, 14 (12–17) mm long, tubular with a linear outline, gently diverging below the corolla, yellowish green (Figure 1c). Tepals, free or less often touching to slightly imbricated at the basal region; lower tepals, 12.5 (8.0–16.0(–19.0)) × 5.0 (4.0–6.5(–8.0)) mm, tip cuspidate (0.5–2.0 mm long); upper tepals, 11.0 (7.0–13.5(–18.0) × 4.0 (3.0–5.0(–6.5)) mm, tip acute to blunt, rarely shortly mucronate. Corona, (Figure 1d,e) 2.2 (1.8–3.0) × 1.5 (0.8–2.2) mm, bright yellow to greenish yellow, mostly subtrigonous to subcircular, vertically incised to the base or partly so at three equidistant locations (occasionally with only one or two incisions), incisions either reaching the base forming three separated flaps, partially incised halfway the corona or only clefted (notched) minutely at the top, yet forming three distinct flaps or lobes; edge irregularly subentire, shallowly crenulate or notched, sometimes bifid giving the allusion of six lobes. Anthers, of upper stamens pale yellow 1.0 (0.8–1.2) mm long; those of lower stamens 1.4 (1.2–1.7) mm long and covered with more pollen but not abundantly so. Fruit, a cylindrical, broadly rugose capsule opening from the apex into three valves, 8–22 mm long. Seeds, numerous, black, polished, pip-shaped with irregular edges, 1–2 mm long.
Pollen, (Figure 1f) copious on the anthers, >90% fertile (very few deflated pollen grains observed), turgid and looking viable. Grains broadly ellipsoid, compressed or flattened at one side. NRC01: 42.1 (38.0–45.9) × 31.9 (30.8–33.2) µm; Q = 1.39 (1.3–1.4) (n = 31). NRC02: 43.7 (41.2–45.2) × 32.7 (30.5–34.9) µm; Q = 1.34 (1.3–1.4) (n = 31). NRC05: 42.2 (40.5–44.2) × 31.6 (30.5–32.4) µm; Q = 1.35 (1.3–1.4) (n = 31). NRC12: 42.0 (39.1–43.9) × 31.1 (28.5–33.1) µm; Q = 1.40 (1.3–1.5) (n = 29). Testa featuring some small bump-like rounded projections about 3–5 µm wide.
3.2. Morphotype Narc-B (Figure 2)
Rather early-flowering plants, typically flowering from mid-October (more or less 3–4 weeks after anthesis of morphotype Narc-A) characterised by forming small tufts of few to 70 small plants each with 2–5 flowers per scape and narrow leaves present during anthesis. In the year 2023, flowering was delayed to the beginning of December due to the absence of substantial rain in most of the autumn—the first significant precipitation occurred during the last week of November.
Material examined: NRC06, NRC07: limits of Ħondoq ir-Rummien (Qala, Gozo), 29 October 2022; NRC12, NRC18: Il-Fekruna area (Munxar, Gozo), 12 November 2022; NRC15, Rdum il-Bies, limits of Selmun area, (Mellieħa, Malta), 15 November 2022, 12 December 2023; NRC27: il-Ħarrieq limits of Selmun area, (Mellieħa, Malta), 12 December 2023. These sampled specimens shared a common habitat represented by a low and sloped or banked garrigue of coralline limestone, located close to cliffs, rocky escarpments, or valley sides, and often accompanied by xeric grasses and steppic flora, indicative of some disturbance or degradation.
Typical accompanying vegetation includes the following species: Asperula aristata subsp. scabra Nyman, Arisarum vulgare O.Targ.Tozz, Asparagus aphyllus L., Asphodelus ramosus L., Bituminaria bituminosa (L.) C.H.Stirt., Brachypodium hybridum Catalán, Joch.Müll., Hasterok and G.Jenkins, Bromus fasciculatus C.Presl, Bromus madritensis L., Chiliadenus bocconei Brullo, Erica multiflora L., Euphorbia dendroides L., Ferula melitensis Brullo, C.Brullo, Cambria, Giusso, Salmeri and Bacch., Galactites tomentosus Moench., Hyparrhenia hirta (L.) Stapf, Periploca angustifolia Labill., Prasium majus L., Ranunculus bullatus L., Teucrium fruticans L., Thymbra capitata (L.) Cav., Drimia pancration (Steinh.) J.C.Manning and Goldblatt.
Description: Habit, dense tufts of (10–)25–70 plants (Figure 2a). Leaves, present during anthesis, usually one per plant/bulb (Figure 2b), rarely two, variably long but usually about half the scape’s length, narrow and not twisted, 2–5 mm wide, cross-section crescent with rounded ends, moderately succulent or fleshy (Figure 2f). Basal sheath, present, about 5–12 mm above ground level, single-layer, semi-membranous, pale green with 10–16 distinct longitudinal bright green veins, margin smooth. Scape, terete, 25 (22–35) cm long (excluding flowers), ca. 3 mm in diameter. Bract, membranous, frail, lanceolate, beige, 24 (28–45) × 8 (7–10) mm. Flowers, 2–4(–6) per scape (Figure 2c), 22–35 mm in diameter, whitish-cream, slightly more colourful adjacent to the corona, very fragrant. Pedicels, of variable length, the longest 30–42 mm long at anthesis (longer in fruit). Perianth tube, 15 (14–16) mm long, tubular (Figure 2d), sometimes with an irregular (angled) outline, narrow and widening slightly below the tepals, sometimes slightly ribbed above, fading to slightly light green at the base and contrasting with the dark green ovary below. Tepals, imbricated up to their basal third, very rarely free or slightly touching each other; lower tepals, 10.5 (9.5–12.5(–15.0)) × 5.5 (5.0–6.5(–8.5)) mm, tip obtuse and distinctly cuspidate (1.5–3 mm long); upper tepals, 9.5 (8.5–11.0(–13.0)) × 5.0 (4.5–6.5) mm, tip acute mucronate (ca. 0.5 mm long) rarely blunt and without an appendage. Corona, 5.0 (3.5–6.5) × 2.0 (1.6–2.4) mm, butter to golden yellow, seldom with a hint of green in young flowers, shallow bowl-shaped with somewhat divergent profile, margin always irregular, outline shape subcircular to subtrigonous occasionally subdeltate (in very young flowers), wall continuous and often with three (sometimes two or one), small, equidistant pleats or minute notches, rarely with one vertical shallow cleft (incision) reaching down not more than half the corona’s height; edge subentire to crenulate. Anthers of upper stamens bright yellow, 1.0–3.0 mm long often smooth without copious pollen; those of the lower stamens 3.0–4.0 mm long with little pollen cover. Bulbs dark brown, often tightly grouped together in three to five clumps. Fruit capsules undeveloped and fall prematurely (Figure 2e). Seeds seldomly formed, mostly shrivelled. Plants are sterile.
Pollen (Figure 2g,h) scant or not seen on the anthers, mostly sterile, shrivelled or empty (less than 25 µm wide), only about 5–15% with a normal turgid shape. Viable pollen grains dimorphic, one form having a broadly ellipsoid ovate shape with one side laterally compressed and the other form are markedly larger with a subspherical, potato-like shape, and stain stronger in cotton blue. Measurements of the small and large forms as follows: NRC06: small form—42.0 (34.4–46.5) × 32.9 (26.3–38.1) µm; L/W ratio 1.3 (1.2–1.4) (n = 21); large form—57.7 (52.2–70.4) × 50.3 (43.2–57.0) µm; L/W ratio 1.15 (1.1–1.2) (n = 13). NRC07: small form—39.9 (35.6–43.1) × 28.1 (24.6–33.7) µm; L/W ratio 1.3 (1.2–1.4) (n = 17); large form—51.3 (48.5–55.5) × 40.3 (37.4–45.8) µm; L/W ratio 1.2 (1.1–1.3) (n = 15); NRC15: small form—38.5 (35.6–40.4) × 30.7 (28.7–33.6) µm; L/W ratio 1.29 (1.2–1.4) (n = 17); large form—49.4 (43.4–58.2) × 41.4 (37.4–57.4) µm; L/W ratio 1.19 (1.1–1.3) (n = 13). Yellow, bump-shaped or hemispherical lipid projections, up to 6.0 µm wide observed on the testa of the small-type pollen grains but often absent on the larger type.
3.3. Morphotype Narc-C (Figure 3, Figure 4 and Figure 5)
This morphotype has a longer flowering period, starting from November up to March, forming plants grouped in small (5–12 plants) to crowded tufts (up to 80 plants) with 4–9 flowers per scape and well-developed leaves before and during anthesis. The morphology of the flowers is not very variable, with minor differences in sizes and flower numbers, but because of apparent divergences in phenology, habitat, and habit, this group can be further divided into the early-flowering “autumnal” rock-dwelling morphotype Narc-C1, and the later winter–spring morphotype Narc-C2 growing more profusely in comparatively open areas such as abandoned fields, clay slopes and open ground with deep, usually clayey soil.
3.4. Morphotype Narc-C2 (Figure 5)
Material examined: NRC22: limits of Verdala close to Buskett Woodland (Siġġiewi, Malta), 11 January 2023; NRC23: Selmun area (Mellieħa, Malta), 4 February 2023; NRC24: Il-Muxi area, (Qala, Gozo), 5 February 2023; NRC25: Tar-Rehba area (Dingli, Malta), 23 February; NRC26: l-Għolja tal-Gelmus, (Rabat, Gozo), 28 February 2023. These plants grew on clayey soil, namely, open abandoned fields, except for NRC22, which was found in a habitat with stony loam soil in shaded patches of an open woodland area.
Typical accompanying vegetation includes the following species: Asparagus aphyllus L., Avena barbata Brot., Avena sterilis L., Bituminaria bituminosa (L.) C. H.Stirt., Cynara cardunculus L., Dittrichia viscosa (L.) Greuter, Foeniculum vulgare Mill., Galactites tomentosa Moench., Hordeum leporinum Link., Medicago polymorpha L., Oxalis pes-caprae L., and Drimia pancration (Steinh.) J.C.Manning and Goldblatt.
Description: Habit, in tufts of (20–)30–80 ca. plants, mostly crowded with bulbs closely packed (Figure 5a). Leaves present before and during anthesis, 3–4 per plant (bulb), almost of equal length and a few cm shorter than the flowering scape, strap-shaped, twisted by one revolution, (18–)25–40 cm × (8–)15–21 mm wide, glaucous green, cross-section gently canaliculate, thin (not fleshy) with acute to obtuse ends, margin smooth. Basal sheath up to 52 mm above ground level, 2–3 concentric layers exposed above ground level (1–2 below ground level), semi-membranous, pale green, uppermost sheath with >15 distinct longitudinal, parallel, bright-green veins, margin smooth (Figure 5d). Scape terete and slightly compressed laterally (broadly ellipse in cross-section) (20–)27–55 cm long, ca. 6–7 mm diameter, minutely ribbed longitudinally. Bract membranous, frail, lanceolate, beige, 30–45 × 10–16 mm. Flowers, 6–10(–12) per scape (Figure 5a), rarely less in unmature plants; 24–36 mm in diameter, whitish-cream, very fragrant. Pedicels of variable length, the longest 45–58 mm long at anthesis (longer in fruit). Bracteoles, four usually present, linear, 25–35 mm long × 1 mm wide, unequal in length, pale green, very thin and hyaline. Perianth tube, 17–19 mm long, subtrigonous, slightly expanding gradually towards the perianth, light green throughout. Tepals broadly ovate/lanceolate, imbricated up to the basal half of the tepal length, sometimes slightly more; lower tepals 16.0 (14.0–17.5) × 11.5 (9.5–13.5), broadly ovate (widest part below the centre); tip subacute and mucronate (1 mm long), white and often with a pale green midrib; upper tepals subequal in size and shape to the upper, but with a reduced mucro. Corona, 11.0 (9.0–13.5) × 5.0–7.0 mm, yolk-yellow, cup-shaped (sides straight) to bowl-shaped (sides slightly convex sides), shallow with a slightly irregular or smooth circular margin, sometimes with 1(–2) shallow and tiny notches or angular pleats; edge entire to subentire to shallowly undulate (Figure 5c). Stamens, in two rows attached to the corolla by short filaments, one attached at the base of the corona, the other about 5 mm below at the neck of the tube. Anthers, subequal length, 3.5–5.5 mm long; moderately abundant in pollen, included but occasionally slightly exerting the corona. Seeds, fertile and numerous, black, pip-shaped, about 3 mm wide
Pollen, (Figure 5e) copious, about 80% fertile (some deflated pollen observed), turgid, and looking viable. Grains broadly ellipsoid, compressed or flattened on one side. NRC24: 47.7 (45.2–50.7) × 35.6 (33.1–38.8) µm; L: W ratio 1.3 [1.2–1.4] (n = 33). NRC25: Lipid projections large, hemispherical or cushion-shaped, up to 6.7 µm across, numerous, and prominent.
3.5. Morphotype Narc-C1 (Figure 3 and Figure 4)
Material examined: NRC14: Ta’ Ċenċ (Sannat, Gozo), 14 November 2022; NRC16, NRC17: Wied Mġarr ix-Xini (Sannat, Gozo), 16 November 2022; NRC19, NRC20, NRC21: Wied tal-Munxar, (Munxar, Gozo), 25 November 2022; NRC28: Wied Babu, (Żurrieq, Malta), 29 December 2023; NRC29: Wied il-Għasel, (Mosta, Malta), 29 December 2023. These specimens were found specifically on rocky valley banks, close to the valley bed (up to 50 m above), but seldom at the valley bed, even if it is not inundated. Hence, this morphotype shows a rupestral preference, often growing in loamy soil present in deep pockets within karstic rock and rocky shelves of valley sides. Interestingly, they seem to prefer a semi-shaded, moist, south-facing side and seldom occur on the south-facing valley sides exposed to sun.
Typical accompanying vegetation includes the following species (those underlined are the most abundant): Arisarum vulgare O.Targ.Tozz.; Asphodelus ramosus L.; Erica multiflora L.; Euphorbia dendroides L.; Galium murale M.Bieb.; Geranium dissectum L.; Geranium molle L.; Hypericum aegyptium Blanco; Oxalis pes-caprae L.; Prasium majus L.; Rhamnus oleoides L.; Rubus ulmifolius J.Presl and C.Presl.; Teucrium flavum L.; Teucrium fruticans L.; Drimia pancration (Steinh.) J.C.Manning and Goldblatt; Bryophytes.
Description: Habit, isolated plants, small groups of 4–10 plants (Figure 4b) or, rarely, loose tufts of groups of plants close to each other merging together, occasionally a cluster of 15–25 plants. Leaves present before anthesis, (3–)4 per plant slightly shorter to longer than the scape, strap-shaped, twisted by not more than one revolution, 20–35 (–60) cm × 7–14(–19) mm wide, can grow much longer and surpass the inflorescence when occurring beside higher shrubby vegetation; cross-section canaliculate to flattened V-shaped, thin, not fleshy, with subacute to obtuse ends, margin smooth, glaucous, non-bluish green with a hint of yellowish hue. Basal sheath, 15–35(–100) mm above ground level, 2–3 concentric layers (1–2 more below ground, total of 4), semi-membranous, pale green with >15 distinct longitudinal, parallel, bright-green veins, margin smooth. Scape, terete and slightly compressed laterally (broadly ellipse in cross-section) above, 20–45 (–70) cm long, 5–7 mm diameter, minutely ribbed longitudinally. Bract membranous, frail, lanceolate, beige, 45 (35–52) × 9 (10–15) mm. Flowers, (3–)4–7(–8) per scape, 18–30 mm in diameter, whitish-cream, sometimes with a pale green midrib, very fragrant, somewhat nauseating. Pedicels of variable length, the longest 45–68 mm long at anthesis (longer in fruit). Bracteoles, four when present, filiform, 20–40 mm long × <1 mm wide, unequal in length, pale green or beige, membranous. Tepals broadly ovate/lanceolate, imbricated up to the basal half, sometimes slightly more, tinged in pale yellow at the base, close to the corona, often seen deflexed backwards; lower tepals, 13.4 (11.5–15.0) × 10 (8.5–11.0), broadly elliptical (widest part at the centre) to oblanceolate (widest part slightly above the centre); tip obtuse-rounded and shortly mucronate (1.0 mm long), white often with a pale green midrib; upper tepals, subequal to the lower tepals, same shape but tip truncate to emarginate, without a mucro. Perianth tube, 19 (16–24) mm long, pale to light green throughout, tubular, and expanding slightly below the tepals, trigonous in cross-section at the distal part. Corona very variable, 9.0 (7.0–11.5) × 5.0 (3.5–7.0) mm, bright lemon to golden yellow, never orangish or amber, cup-shaped (with vertical parallel sides), bowl-shaped (with long diverging sides) to sometimes saucer-shaped (with wide, shallow diverging sides) (Figure 3e), generally rim with a circular outline or rarely three shallow-lobed (Figure 3d) and having a margin the varies from entire and smooth (especially in cup-shaped forms), crenulate or crenated, pleated (especially in saucer-shaped forms), or rarely notched, with angular folds or tiny clefts. Stamens in two rows attached to the corolla by short filaments, one attached at the base of the corona, the other about 5 mm below at the neck of the tube. Anthers, subequal length, 4.0–7.5 mm long; covered with abundant pollen, included but occasionally slightly exerting the corona. Seeds, fertile and numerous, black, pip-shaped, about 3–4 mm wide. N.B.: when plants are found within thick vegetation or in very shaded pockets, they etiolate, forming abnormally longer leaves and scapes reach almost 1 m in height.
Pollen (Figure 3h), copious on the anthers, about 85–90% fertile (only very few deflated pollen grains observed), generally turgid and viable. Grains broadly ellipsoid, flattened at one side. NRC14: 39.8 (36.3–42.3) × 30.7 (29.1–32.2) µm; L/W ratio 1.3 [1.2–1.4] (n = 31). NRC16: 41.5 (39.9–43.7) × 31.7 (30.8–32.6) µm; L/W ratio 1.3 [1.2–1.4] (n = 37). NRC19: 39.3 (37.6–41.7) × 31.0 (29.3–32.2) µm; L/W ratio 1.3 (1.2–1.4) (n = 31). Lipid projections numerous, large, hemispherical or cushion-shaped, up to 7.2 µm across.
3.6. Morphological Analysis
The separation between the four morphotypes emerged clearly in the results of the morphological analysis (Figure 6). There is a clear separation between NRC-C1 and NRC-C2 morphs on axis 1 of the nMDS plot, whilst morphs NRC-A and NRC-B were separated on axis 2. The results of the analysis lend support to the field observations by the authors.
4. Discussion
Morphotype Narc-A represents the early-flowering species first mentioned by Zerapha [20], which in the literature has long been referred to as Narcissus serotinus. However, the description given in [20] as a multi-flowered spathe with canaliculate leaves does not match perfectly with this species, which usually has one–two flowers and thin subcylindrical leaves. A later flora [21] assumed that this species was N. cupanianus Guss., but Parlatore confirmed a voucher specimen sent to him by Grech Delicata (not the same one that Zerapha described) from Malta as N. serotinus [22], leaving an open question regarding what Zerapha had actually seen (see discussion on morphotype Narc-C, below). N. serotinus was reported to occur in the Maltese Islands repeatedly in subsequent works [12,22,23,24,25,26,27,28] until taxonomic questions and inquiries started to emerge sometime later following broad taxonomic studies and monographs on autumnal Narcissus [5,15,16]. This species was also reported differently as N. obsoletus (Haw.) Spach [29], stating that the closely related N. serotinus is confined to the extreme west of the Mediterranean Region. This taxon was also mentioned in another work on the Maltese flora [30].
However, morphotype Narc-A, characterised by a three-cut or segmented corona and linear perianth tube, neither matches N. serotinus having a six-cut corona and diverging (expanding) perianth tube nor N. obsoletus having an entire, unsegmented, urn-shaped corona [31].
Donnisan-Morgan et al. [3] examined plants—at that time, all referred to as “N. serotinus”—from Spain, Portugal, and Morocco. Their results obtained from flow cytometry revealed that there are two morphotypes of N. serotinus: the sensu stricto with a DNA content of 20 picograms and another one with 50 picograms. The latter showed a few important morphological differences from N. serotinus s. str., namely, in a more linear perianth tube, and, according to the authors, a corona with stronger reddish or orange tones. They described the new species as N. miniatus Donn.-Morg., Koop. and Zonn. and postulated it to have originated from a long-established allotetraploid hybrid between N. obsoletus (or N. elegans) and N. serotinus.
Soon after, Spanish authorities resurrected a forgotten taxon—N. deficiens Herb., described by Herbert (1847) from Santa Maura on the Greek Island of Lefkada. For a long time, this taxon was used as a synonym for N. obsoletus and neglected by everyone—also by Donnisan-Morgan et al. [3]. This has been described as having a distinct three-flapped (deeply dissected) or three-lobed corona, typically trigonous in outline, which differs from N. serotinus having a six-lobed or flapped corona with a subtrigonous to almost circular outline and from N. obsoletus, which have an undivided, bowl-shaped corona with a circular in outline (refer to details in [4]). These scholars suggested that N. deficiens corresponds with the same Narcissus previously described as N. miniatus, which is common in Spain. Moreover, Fernández Casas [4] put N. miniatus in synonymy with N. deficiens but supported its allotetraploid hybrid origin as suggested by Donnisan-Morgan et al. [3] and later by other authors [15].
Some taxonomic disagreements still exist between the different schools of thought on whether N. miniatus and N. deficiens are different or conspecific. The controversy originates from N. deficiens being described and depicted from an aberrant form with a much-reduced corona (hence the epithet “deficiens”). Hence, some suggested that N. deficiens should refer specifically to corona-reduced forms, and the “typical forms” with a developed three-parted corona should be treated as N. miniatus; while others prefer a more conservative approach and include all forms within a single species and accept N. deficiens for being described first. This misperception was settled when specimens from a population of an autumnal daffodil collected from the locus classicus of N. deficiens (Alexander’s Castle, now known as Agia Mavra Castle in Lefkas, Greece) had variable, well-defined coronae dissected in three parts or three lobes with a trigonous outline (Soc. Med. Comm: Carles Jiménez, Narcissus FB group, 27 October 2023), hence supporting further that Herbert’s N. deficiens should be attributed to a more diverse species rather than restricted to a corona-deficient form as the name implies. This ambiguity may have led many researchers in the past to assume N. deficiens to be a daffodil without or only a remnant corona and became neglected.
With regards to the Maltese Islands, the early-flowering Narcissus (morphotype Narc-A) characterised by having no leaves during anthesis and forming one to few flowers with a subtrigonous corona composed of three lobes or flaps should be attributed to N. deficiens.
However, N. serotinus was recorded from the Maltese Islands precisely from Triq il-Batterija, Marsalforn, Gozo, close to a hillock known as il-Qolla l-Bajda [5]. According to these investigations, their specimen labelled S.MLT genetically corresponded to Narcissus serotinus. Surveys carried out by the present author at il-Qolla l-Bajda in 2020 and 2022 did not confirm N. serotinus, but only a fragmented population of N. deficiens. Narcissus serotinus is distinct in having an inflated (abruptly widening) perianth tube and six distinct and fully incised corona segments. An example found by the author on 30 October 2004 with a corona divided into six distinct free segments (Figure 1g) might be additional evidence of the presence of N. serotinus in Malta, but photos showing the lateral view to fully confirm the species were not taken on that occasion. Since N. serotinus is regarded as a species restricted to eastern Spain, western Portugal, and northern Morocco [32], then the presence of N. serotinus in the Maltese Islands is biogeographically improbable, as it is out of the currently reported natural range. Notwithstanding this, the occurrence of N. serotinus in Malta should be considered as tentative, at best, until further studies are conducted.
Morphotype Narc-C (Figure 3, Figure 4 and Figure 5) represents two forms, of Narcissus tazetta. The typical and locally frequent form, morphotype Narc-C2 (Figure 5), flowers between mid-January and March and grows on clay slopes or in clayey soil of abandoned fields or field margins, along pathways, grassland, and sometimes in rocky ground with deep soil. The other form, morphotype Narc-C1 (Figure 3 and Figure 4) flowers from the end of October and occurs on rocky banks, shelves, and ledges in semi-shaded valley sides amongst shrubby, garigue, phrygana, or scrub-type sclerophyllous vegetation.
Narcissus tazetta has been reported since the first modern listing of the Maltese flora [20], and like N. serotinus, in all subsequent floras, guidebooks, or accounts of the Maltese flora [12]. The reported flowering time of five to six months is relatively too long for a single monocot species, as these, including other member species of Narcissus, typically have specific and short flowering times. Hence, this raises the question of whether Narc-C1 is distinct from the typical N. tazetta. When compared (Table 2), there are no significant morphological differences other than that Narc-C2 is a winter–spring-flowering species that is more robust, has larger leaves, denser and larger tufts, more flowers per scape, and slightly larger and deeper coronas.
Interestingly, the distinction of the autumnal variety of N. tazetta had already been identified [33] from Maltese material sent to Herbert by Zerapha. Applying the same arguments above with the early flowering time and habitat, Herbert could not assign it to N. tazetta and described the early-flowering Maltese daffodil as Hermione aequilimba Herb. [34]. He gave a very detailed account and an excellent illustration (Figure 7), which matches with morphotype Narc-C1. All accounts synonymising H. aequilimba with N. obsoletus (e.g., [35]) are therefore not correct because this species is noticeably different from N. obsoletus and resembles the stockier N. tazetta. Similarly, Herbert [34] mentions explicitly that Zerapha made a mistake in assigning this plant to N. serotinus: “Zerapha has certainly made an error in referring this African plant [H. aequilimba] to the little 1-flowered (rarely 2-flowered) N. serotinus of Clusius, which grows in Spain, Naples, and Sardinia”.
According to Errol Vela (pers. comm. To S. Mifsud, November 2020), similar early-flowering tazetta exist in the Middle East and he suggested comparing this with the description of N. tazetta subsp. syriacus (Boiss. and Gaill.) Boiss. This description of this subspecies was checked from the protologue [36] and it was found that while no reference is made to the early flowering character, the main distinction from N. tazetta s. str. seems to be based on differences in the corolla, namely, its narrower tepals: “Perigonii laciniae oblongae longiores et angustiores magis discretae…” [36]. This minor difference might not justify the distinction from N. tazetta s. str., and, more importantly, does not match with the Maltese “autumnal N. tazetta”, which have broad tepals and hence cannot be ascribed to subsp. syriacus in the sense as described in the protologue. Further specimens should however be examined to understand what N. tazetta subsp. syriacus really refers to and whether it merits distinction at subspecies level.
Herbert’s description from Maltese material ventures into whether to assign the autumn-flowering morphotype of N. tazetta (Narc-C1) to H. aequilimba (=Narcissus aequilimbus (Herb.) Nyman) based on different chorology, habitat, stature, the number of flowers per scape, and minor differences in the size of some flower parts. In accordance with Herbert’s approach, the findings in this study, and the opinions of scholars of this genus (e.g., pers. comm. Errol Vela and Oron Peri), the autumnal N. tazetta merits taxonomic distinction from the spring-flowering N. tazetta. It must be noted that a bulb that blooms in autumn would not flower again in spring; thus, this situation does not implicate flowering of the same bulb at two different seasons. Notwithstanding that the distinction between the two morphotypes is not pronounced, N. aequilimbus is proposed to be ranked at the infraspecific level in line with other infraspecific taxa of N. tazetta with slight differences from N. tazetta s. str. such as N. tazetta subsp. aureus (Haw.) Baker, N. tazetta subsp. canariensis (Burb.) Baker, and N. tazetta subsp. italicus (Ker Gawl.) Baker.
For taxonomic procedures, Hermione aequilimba is further epitypified on collected material from Malta, including entire plants with bulb, where currently the type is only based on a partial illustration (Figure 7) of a plant [34]. It is here combined into a subspecies ranking as follows:
Narcissus tazetta L. subsp. aequilimbus (Herb.) Mifsud comb. nov.
=Hermione aequilimba Herb. (basionym) in Amaryllidaceae: 404 (1837)
=Narcissus aequilimbus (Herb.) Nyman in Syll. Fl. Eur.: 365 (1855)
Type—coastal areas in Malta, but locality and exact date (only “flowering in October”) are not given; epitype is based on illustration Pl. 48 in the protologue [34].
Epitype—Designated here from Ta’ Ċenċ (ta’ fuq il-Gruwa), Sannat, Gozo, Malta. Collected 12 November 2022, leg. S. Mifsud. Deposited at ARG! (collection SM140; Neoisotype SM141).
Isoepitype—Second epitype is designated here from Ħondoq ir-Rummien, Qala, Gozo, Malta. Collected 12 November 2022, leg. S. Mifsud. Deposited at ARG! (collection SM142).
Diagnosis—Same as N. tazetta s. str. but flowers much earlier, about six weeks after the first substantial autumnal rains, typically starting by the end of October to November, plants formed in loose groups, scattered, sometimes singular, very rarely in clusters (Figure 4b) on rocky banks, shelves and ledges of valley sides and mouths over the coast, preferring shaded (hence damper) sides typically in loamy soil besides garigue or scrub shrubby vegetation (Figure 4b). Leaves narrower, mostly between 9–14 mm wide, can reach up to 50 cm in length due to etiolation, somewhat less glaucous and less twisted. Scapes with less flowers, averaging 5–6 per scape. Tepals smaller by 2–3 mm, broadly ovate/lanceolate, creamy-white normally, with pale greenish midribs. Corona variable, cup-shaped (parallel sides) or bowl-shaped (divergent sides) with subentire to crenated rim, diameter smaller, on average measuring 9 × 5 mm. A full description is given in above (see Section 3 Results, morphotype Narc-C1).
Specimens examined were Ta’ Ċenċ (Sannat, Gozo); Wied Mġarr ix-Xini (Sannat, Gozo); Wied tal-Munxar, (Munxar, Gozo); Wied Babu (Żurrieq, Malta); Wied il-Għasel, (Mosta, Malta). Refer to Table 1 for more details.
The autumnal N. tazetta (morphotype Narc-C1) should be referred to subsp. aequilimbus, whereas the typical winter/spring flowering form (Narc-C2) should be referred to N. tazetta s. str. It is already assumed that this subspecies is not endemic to Malta because similar autumnal forms have been observed in the Eastern Mediterranean Region (pers. comm. Oron Peri, August 2019; Errol Vela, November 2020).
Morphotype Narc-B represents plants that were identified as N. elegans for several matching morphological characteristics with that species [12], the most important being the narrow, 3–5 mm wide caniculate leaves present at anthesis, the early flowering in mid-October, and the formation of tufts and the multi-flowering scapes. These plants reported as N. elegans were also perceived by the authors to be an intermediate species between N. serotinus (=N. deficiens) and N. tazetta, and the authors further suggested that they may have introgressed with either species to form hybrid swarms.
The present study provided further crucial knowledge on this morphotype by indicating new populations in addition to those reported in [12] and, more importantly, detecting sterility in all examined material with reference to the dysfunctional, unviable pollen (Figure 2h) as well the lack of seed formation in poorly developed and often dropping fruit capsules (Figure 2f). All examined plant populations were largely sterile, leading to two fundamental conclusions: these plants are not N. elegans, and they should represent a natural sterile hybrid.
There are only two species that can act as putative parents where their flowering period overlaps: hence N. deficiens and N. tazetta subsp. aequilimbus. Both species are within pollinator reach, for example, at Il-Fekruna/Wied il-Munxar, and at Hondoq ir-Rummien, the parents and preseumed hybrid (morphotypes Narc-B) were in sympatry. Cross-pollination is also feasible because populations of both parents are locally frequent to abundant. The hypothesis that morphotype Narc-B is a natural hybrid between N. deficiens and the autumnal N. tazetta s.l. is also strongly supported by morphological findings. Some character states of the presumed hybrid resemble one of the parents, some are similar to the other parent and others are considered as intermediate forms of both parents. For example, the moderately fleshy leaf with a crescent cross-section ca. 3–5 mm wide of the presumed hybrid is intermediate between the cylindrical 1 mm leaf of N. deficiens and the 7–16 mm flattened leaf of N. tazetta s.l. Table 3 provides a detailed comparison of many morphological characters, indicating which has affinity to either parent and which is intermediate. This new hybrid is described as follows:
N. deficiens Herb. × N. tazetta L. subsp. aequilimbus (Herb.) Mifsud
Etymology—Named after Michael Briffa (1926–2019), a student of the Maltese flora and macrofungi for most of his life, leaving behind several important findings and publications enriching the knowledge and conservation of the biodiversity of the Maltese Islands.
Diagnosis—Plants in tufts of about 25–70 plants. Leaves present during anthesis, 3–5 mm wide, crescent with rounded edges in cross-section, fleshy. Basal sheath present, pale green with bright green veins, ca. 10 mm long above ground level. Perianth tube tubular with parallel sides tapering gently, green, but fading considerably at the lower half. Scape up to 30 cm long, 3 mm in diameter, bearing 2–5(–6) flowers. Tepals cream-white, broadly lanceolate or ovate, imbricated up to the lower half, approx. 10 × 5 mm. Corona bowl-shaped with a subcircular to faintly trigonous outline, with three equidistant small pleats (sometimes shallow clefts) on the rim, lemon to golden yellow, 5 × 2 mm. Pollen not abundant on anthers, mostly shrivelled and empty, hence unviable. Fruit capsules small, shrunk without seeds or seldom a few. (Refer to the description of morphotype Narc-B in Section 3 Results for a full description).
Type—MALTA. Qala: Ħondoq ir-Rummien, elevation 70 m, 36°1′45.42″ N 14°19′15.47″ E, 7 December 2023, Leg. S. Mifsud (holotype ARG!, collection SM150, isotype ARG!, collection SM151).
Studied material in the Maltese Islands—Selmun-Mistra area (various locations), Mellieħa, Malta! (1 November 2008); Dwejra, San Lawrenz, Gozo (17 October 2009); Qortin tal-Magun, Nadur! (Gozo, 14 November 2009) (Figure 4a); Limits of Wied is-Simar, Qala, Gozo (14 November 2010); Ta’ Ċenċ, Sannat, Gozo! (16 November 2014); Ħondoq ir-Rummien, Qala, Gozo! (29 October 2022); Il-Fekruna/Wied tal-Munxar, Munxar, Gozo! (3 November 2022). (! Indicates both parents have been seen together within a distance of 100 m; date given is that of the first observation).
Narcissus × briffae was mistaken for N. elegans in [12], and in historical records [20,21] that reported their “N. elegans” as being frequent in Malta. More recently, the record of N. elegans by H. Spiteri found in the 1980s and cited in the Red Data Book of the Maltese Islands [37] in a similar habitat to N. × briffae is likely a similar misidentification. Unless typical records of N. elegans are confirmed, it is safe to reject it from the flora of Malta and replace it with N. × briffae.
5. Conclusions
Narcissus is a very dynamic genus, with fluid taxonomy (especially considering nothotaxa), changes in distribution or species demarcation, and synonymising of taxa are ongoing regularly. Its taxonomy can be subjective sometimes, given the difficulty in assessing member species with simple and affordable molecular techniques such as ITS sequencing. This study reshaped the inventory of Narcissus occurring in the Maltese Islands, now represented at least by four native taxa: N. deficiens (syn. N. miniatus), previously referred as N. serotinus or N. obsoletus; Narcissus × briffae (N. deficiens × N. tazetta subsp. aequilimbus) a newly described hybrid replacing N. elegans; and N. tazetta s.l., which comprises of N. tazetta subsp. aequilimbus—previously described from Malta [34] but neglected—and N. tazetta s. str. The intriguing records of N. serotinus from Malta ([5]; Figure 1g) are currently considered as doubtful and probably safe to exclude because this species is assumed to have a restricted distribution in western parts of Spain and adjacent regions in Portugal.
This study advocates authors of Mediterranean flora to revise this genus in their territories since, as resulted in this study, it warrants similar taxonomic updating, with special reference to the autumnal species. Other alien species occurring in Malta include N. papyraceus Ker. Gawl. and N. tazetta subsp. italicus (Ker. Gawl.) Baker, sometimes forming localised naturalised populations [38,39]. This study provides a dichotomous identification key of the Narcissus species occurring or previously recorded from the Maltese Islands.
Key to Narcissus spp. in the Maltese Islands.
Characters should be based on observing at least five flowers since due to the variability within the species and occasionally the presence of atypical examples which may not key properly, especially with regard to the number of flowers and sizes.
- 1.
- Scape with 1–2 flowers; leaves ca. 1 mm wide, absent during anthesis. 2
- +
- Scape with 3 or more flowers; leaves > 2 mm wide present during anthesis. 3
- 2.
- Perianth tube widens abruptly halfway its length (bottle-shaped); corona always with six isolated segments. N. serotinus *
- +
- Perianth tube tubular with almost parallel sides and widens slightly (barely detectable) at the tip below the tepals; corona mostly with three segments or continuous a single unit with three shallow lobes or subentire. N. deficiens
- 3.
- Corona distinctly lemon to yolk yellow, similar in colour to the anthers; flower tube light green or yellowish-green. 4
- +
- Corona white, cream or pale yellow, much contrasting with the golden-coloured anthers; flower tube white. 7
- 4.
- Tepals very narrow, linear-oblong, never imbricated. Corona < 4 mm wide. N. elegans +
- +
- Tepals broadly ovate, overlapping at the lower half. Corona > 4 mm wide. 5
- 5.
- Plants with 1–2 leaves; 3–5 flowers per scape; fruit capsules shrivelled, undeveloped and seedless, anthers with little scanty pollen, which is mostly shrivelled or empty when observed under the microscope, dimorphic. N. × briffae
- +
- Plants with 3–4 leaves; five or more flowers per scape; fruit capsules barrel-shaped with viable black seeds; anthers covered with copious pollen, turgid, and broadly ellipsoid with a slightly flattened side, all the same shape. 6 (N. tazetta s.l.)
- 6.
- First flowers blossoming in October–November, average 4–6 flowers per scape, plants in small groups or singular (seldom forming sizeable tufts). subsp. aequilimbus
- +
- First flowers blossoming in January, average of 8–9 flowers per scape, often forming large, dense tufts of plants. subsp. tazetta
- 7.
- Corona white and the same colour as the tepals. N. papyraceus ^
- +
- Corona cream and dissimilar from the white tepals. N. tazetta subsp. italicus ^
- *
- species recorded from Malta, but occurrences are doubtful and require further verification; unlikely to be present in Malta.
- +
- past misidentifications.
- ^
- alien species for the Maltese Islands.
Author Contributions
Conceptualization, S.M.; methodology, S.M. and S.L.; software, S.L.; validation, S.M..; formal analysis, S.M. and S.L.; investigation, S.M.; resources, S.M. and S.L.; data curation, S.M. and S.L.; writing—original draft preparation, S.M.; writing—review and editing, S.L.; visualization, S.M. and S.L.; supervision, S.M. and S.L.; project administration, S.M.; funding acquisition, S.L. All authors have read and agreed to the published version of the manuscript.
Funding
This research received no external funding.
Data Availability Statement
Raw data is available on the University of Malta Data Repository (drum.um.edu.mt).
Acknowledgments
This work benefited from conversations and correspondences with botanists or naturalists who had worked or studied the genus Narcissus, particularly (alphabetical order) Julián Manuel Fuentes, Carles Jiménez, Harold Koopowitz, Oron Peri, Fernando Ureña Plaza, and Errol Vela, many of whom are members of the Facebook group Narcissus. We received considerable assistance from Tony Meli and Christian Borg at the Argotti Botanic Garden and Resource Centre, Malta while depositing the specimens. This is greatly appreciated.
Conflicts of Interest
The authors declare no conflicts of interest.
References
- Royal Horticultural Society. The International Daffodil Register and Classified List (2008) and Its Supplements: Botanical Classification Updated December 2017; Royal Horticultural Society: London, UK, 2017. [Google Scholar]
- Aedo, C. Narcissus L. In Flora Ibérica; Talavera, S., Andrés, C., Arista, M., Fernández Piedra, M.P., Rico, E., Crespo, M.B., Quintanar, A., Herrero, A., Aedo, C., Eds.; The Royal Botanical Garden of Madrid: Madrid, Spain, 2013; pp. 340–397. [Google Scholar]
- Donnison-Morgan, D.; Koopowitz, H.; Zonneveld, B.; Howe, M. Narcissus miniatus Donnison-Morgan, Koopowitz & Zonneveld sp. nov. a new species of Narcissus (Amaryllidaceae) from southern Spain. In Daffodils, Snowdrops and Tulips Yearbook 2005–2006; Bradbury, M.S., Ed.; Royal Horticultural Society: London, UK, 2005; pp. 19–25. [Google Scholar]
- Fernández Casas, F.J. Narcissorum notulæ XI. Fontqueria 2008, 55, 571–572. [Google Scholar]
- Marques, I.; Nieto Feliner, G.; Draper, D.; Martins-Louçao, M.A.; Fuertes Aguilar, J. Unravelling cryptic reticulate relationships and the origin of orphan hybrid disjunct populations in Narcissus. Evolution 2010, 64, 2353–2368. [Google Scholar]
- López-Tirado, J. A natural laboratory in southern Spain: New hybrids of wild daffodils (Narcissus, Amaryllidaceae). Phytotaxa 2019, 394, 161–170. [Google Scholar] [CrossRef]
- López-Tirado, J. A New Contribution to the Wild Daffodils of the Iberian Peninsula: Description of the Autumnal Narcissus × hispalensis (Amaryllidaceae). Rhodora 2022, 123, 11–18. [Google Scholar] [CrossRef]
- Mifsud, S. Ophrys × gaulosana (Orchidaceae)—A new hybrid from Gozo (Maltese Islands). J. Eur. Orchid. 2018, 50, 213–220. [Google Scholar]
- Mifsud, S. Rediscovery and taxonomic analysis of Romulea melitensis (Iridaceae) from the Maltese Islands. Phytotaxa 2021, 483, 229–243. [Google Scholar] [CrossRef]
- Mifsud, S.; Mifsud, O. A revision of Allium subsect. Allium (Amaryllidaceae) for the Maltese Islands. Flora Mediterr. 2018, 28, 27–51. [Google Scholar] [CrossRef]
- Mifsud, S. Bolboschoenus (Cyperaceae) in the Maltese Islands—An enigma of taxonomic interest. Flora Mediterr. 2022, 32, 53–71. [Google Scholar] [CrossRef]
- Mifsud, S.; Caruana, E. Records of Narcissus elegans (Fam. Amaryllidaceae) and notes on the wild Narcissus in the Maltese islands. Cent. Mediterr. Nat. 2010, 5, 19–29. [Google Scholar]
- Pignatti, S. Flora d’Italia; Edagricole: Bologna, Italy, 1982; Volume 13. [Google Scholar]
- Webb, D.A. Narcissus. In Flora Europaea; Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M., Webb, D.A., Eds.; Cambridge University Press: Cambridge, UK, 1980; Volume 5, pp. 78–84. [Google Scholar]
- Díaz Lifante, Z.; Camacho, C.A. Morphological variation of Narcissus serotinus L. s.l. (Amaryllidaceae) in the Iberian Peninsula. Bot. J. Linn. Soc. 2007, 154, 237–257. [Google Scholar] [CrossRef]
- Marques, I.; Feliner, G.; Martins-Louçao, M.A.; Aguilar, J. Genome size and base composition variation in natural and experimental Narcissus (Amaryllidaceae) hybrids. Ann. Bot. 2011, 109, 257–264. [Google Scholar] [CrossRef] [PubMed]
- Marques, I.; Aguilar, J.; Martins-Louçao, M.A.; Moharrek, F.; Feliner, G. A three-genome, five-gene comprehensive phylogeny of the bulbous genus Narcissus (Amaryllidaceae) challenges current classifications and reveals multiple hybridisation events. Taxon 2017, 66, 832–854. [Google Scholar] [CrossRef]
- Boukhebache, N.; Amirouche, N.; Amirouche, R. Cytotaxonomic investigations on species of genus Narcissus (Amaryllidaceae) from Algeria. Comp. Cytogenet. 2022, 16, 55–76. [Google Scholar] [CrossRef] [PubMed]
- Pignatti, S. Flora d’Italia, 2nd ed.; Edagricole: Bologna, Italy, 2019; Volume 4. [Google Scholar]
- Zerapha, S. Florae Melitensis Thesaurus: Sive Plantarum Enumeratio, Quae in Melitae Gaulosque Insulis aut Vulgatissimae; Fasc. 2: Valletta, Malta, 1831. [Google Scholar]
- Grech Delicata, G.C. Flora Melitensis, Sistens Stirpes Phanerogamas in Melita Insulisque Adjacentibus Hucusque Detectas Secundum Systema Candolleanum Digestas; ex typis FW Franz: Valletta, Malta, 1853. [Google Scholar]
- Sommier, S.; Caruana Gatto, A. Flora Melitensis Nova; Stabilimento Pellas: Firenze, Italy, 1915. [Google Scholar]
- Borg, J. Descriptive Flora of the Maltese Islands: Including the Ferns and Flowering Plants; Government Printing Office: Marsa, Malta, 1927. [Google Scholar]
- Lanfranco, G. A Field Guide to the Wild Flowers of Malta; Progress Press: Valletta, Malta, 1969. [Google Scholar]
- Haslam, S.M.; Sell, P.D.; Wolseley, P.A. A Flora of the Maltese Islands; Malta University Press: Msida, Malta, 1977. [Google Scholar]
- Lanfranco, E. Il-Flora. In Flora u Fawna ta’ Malta; Sultana, J., Falzon, V., Eds.; Dipart. ghall-Harsien ta’l-Ambient: Valletta, Malta, 1995; pp. 34–143. [Google Scholar]
- Weber, H.C.; Kendzior, B. Flora of the Maltese Islands: A Field Guide; Margraf Publishers: Weikersheim, Germany, 2006. [Google Scholar]
- Casha, A. Flora of the Maltese Islands; self-published: Valletta, Malta, 2014; Volume 4. [Google Scholar]
- Lanfranco, E.; Bonett, G. Wild Flowers of the Maltese Islands; Book Distributors Ltd.: San Ġwann, Malta, 2015. [Google Scholar]
- Casha, A. Flora of the Maltese Islands, 2nd ed.; self-published: Valletta, Malta, 2017. [Google Scholar]
- Fernández Casas, F.J. Narcissorum notulæ, X. Fontqueria 2008, 55, 547–558. [Google Scholar]
- López-Tirado, J.; González-Armenteros, J. Narcissus serotinus Loefl. ex L. (Amaryllidaceae), confirmed for the province of Cádiz (Andalusia, Spain). Acta Bot. Malacit. 2019, 44, 1–2. [Google Scholar] [CrossRef]
- Herbert, W. Narcissi. In Edward’s Botanical Register V33: Tab. 22; Lindley, J., Ed.; James Midway: London, UK, 1847. [Google Scholar]
- Herbert, W. Amaryllidaceae: Preceded by an Attempt to Arrange the Monocotyledonous Orders, and Followed by a Treatise on Cross-Bred Vegetables, and Supplement; J. Ridgway & Sons: London, UK, 1837. [Google Scholar]
- WFO. World Flora Online. 2024. Available online: http://www.worldfloraonline.org (accessed on 2 January 2024).
- Boisser, E. Flora Orientalis Sive Enumeratio Plantarum in Oriente a Graecia et Aegypto ad Indiae Fines Hucusque Observatarum; H. Georg Publisher: Basel, Switzerland, 1882; Volume 5, pp. 150–151. [Google Scholar]
- Lanfranco, E. 1989: The Flora. In Red Data Book for the Maltese Islands; Schembri, P.J., Sultana, J., Eds.; Department of Information: Valletta, Malta; pp. 5–70.
- Mifsud, S. Narcissus Papyraceus—MaltaWildPlants.com, Online Flora of the Maltese Islands, Datasheet Created on Dec-2007, Last Updated 2022. Available online: https://www.maltawildplants.com/AMRY/Narcissus_papyraceus.php (accessed on 17 January 2024).
- Mifsud, S. Narcissus tazetta subsp. Italicus—MaltaWildPlants.com, Online Flora of the Maltese Islands, Datasheet Created on Mar-2012, Last Updated 2022. 2022. Available online: https://www.maltawildplants.com/AMRY/Narcissus_tazetta_subsp_italicus.php (accessed on 17 January 2024).
Figure 1.
Narcissus deficiens Herb. from the Maltese Islands: (a) flowers from two individuals, very variable in size, generally 1–2 per scape; (b) rare occasions of 3-flowered example; (c) linear floral tube that expands slightly below the tepals (grid = 1 × 1 cm; (d,e) variation of the corona—yellowish green to golden yellow to amber; terete to trigonous outline with subentire to fully incised wall forming three segments, each being emarginate, notched to clefted (scale bar 1 mm); (f) pollen grains at ×100 magnification (scale bar = 10 µm) with an average size of 43 × 32 µm (stained in cotton blue); (g) corona with a circular outline and six fully incised, free segments characteristic of N. serotinus (scale bar 2 mm).
Figure 1.
Narcissus deficiens Herb. from the Maltese Islands: (a) flowers from two individuals, very variable in size, generally 1–2 per scape; (b) rare occasions of 3-flowered example; (c) linear floral tube that expands slightly below the tepals (grid = 1 × 1 cm; (d,e) variation of the corona—yellowish green to golden yellow to amber; terete to trigonous outline with subentire to fully incised wall forming three segments, each being emarginate, notched to clefted (scale bar 1 mm); (f) pollen grains at ×100 magnification (scale bar = 10 µm) with an average size of 43 × 32 µm (stained in cotton blue); (g) corona with a circular outline and six fully incised, free segments characteristic of N. serotinus (scale bar 2 mm).
Figure 2.
Narcissus × briffae Mifsud nothosp. nov. from the Maltese Islands: (a) habit in small dense tufts of about 40 plants; (b) individual plants with one narrow leaf (ca. 5 mm wide) and 3–5 flowers (grid = 5 × 5 mm); (c) detail of flower with slightly imbricated tepals and a shallow corona with a subtrigonous to subterete outline having three pleats or pinched-like folds (scale bar in (c) = 5 mm); (d) lateral view of the corolla showing shallow corona and dimensions (grid = 10 × 10 mm); (e) scapes with fallen or shrivelled seedless fruit due to sterility of the hybrid; (f) narrow leaves with fleshy crescent cross-section (ruler scale = 1 mm/div); (g) pollen grains at ×100 magnification (g) and × 400 magnification (h) (scale bar = 100 µm) mostly shrivelled or empty, with few turgid grains of two different sizes, the small ones ((h) bottom arrow) measuring on average 38 × 29 µm and the larger ones ((h) top arrow) measuring 52 × 40 µm and stain stronger in cotton blue.
Figure 2.
Narcissus × briffae Mifsud nothosp. nov. from the Maltese Islands: (a) habit in small dense tufts of about 40 plants; (b) individual plants with one narrow leaf (ca. 5 mm wide) and 3–5 flowers (grid = 5 × 5 mm); (c) detail of flower with slightly imbricated tepals and a shallow corona with a subtrigonous to subterete outline having three pleats or pinched-like folds (scale bar in (c) = 5 mm); (d) lateral view of the corolla showing shallow corona and dimensions (grid = 10 × 10 mm); (e) scapes with fallen or shrivelled seedless fruit due to sterility of the hybrid; (f) narrow leaves with fleshy crescent cross-section (ruler scale = 1 mm/div); (g) pollen grains at ×100 magnification (g) and × 400 magnification (h) (scale bar = 100 µm) mostly shrivelled or empty, with few turgid grains of two different sizes, the small ones ((h) bottom arrow) measuring on average 38 × 29 µm and the larger ones ((h) top arrow) measuring 52 × 40 µm and stain stronger in cotton blue.
Figure 3.
Narcissus tazetta L. subsp. aequilimbus (Herb.) Mifsud from the Maltese Islands: (a) inflorescence usually made of 5–7 flowers; (b) closeup of flowers showing corona and imbricated tepals; (c) individual plant in its natural habitat with four leaves ((c) 12 mm wide); (d,e) variation in the corona from lemon to gold yellow, cup-shaped ((e) top), bowl-shaped ((e) middle), or saucer-shaped ((e) bottom) with a circular or (three-) shallow-lobed outline usually unsegmented and without clefts or incisions (scale 10 mm); (f) very rare example of tetramerous corolla with 8 tepals; (g) flowers with deflexed petals; (h) pollen grains at ×100 magnification (scale bar = 10 µm) with an average size of 40 × 31 µm and numerous yellow warts (stained in cotton blue).
Figure 3.
Narcissus tazetta L. subsp. aequilimbus (Herb.) Mifsud from the Maltese Islands: (a) inflorescence usually made of 5–7 flowers; (b) closeup of flowers showing corona and imbricated tepals; (c) individual plant in its natural habitat with four leaves ((c) 12 mm wide); (d,e) variation in the corona from lemon to gold yellow, cup-shaped ((e) top), bowl-shaped ((e) middle), or saucer-shaped ((e) bottom) with a circular or (three-) shallow-lobed outline usually unsegmented and without clefts or incisions (scale 10 mm); (f) very rare example of tetramerous corolla with 8 tepals; (g) flowers with deflexed petals; (h) pollen grains at ×100 magnification (scale bar = 10 µm) with an average size of 40 × 31 µm and numerous yellow warts (stained in cotton blue).
Figure 4.
(a) Field comparison of Narcissus × briffae Mifsud (left) and N. tazetta subsp. aequilimbus (Herb.) Mifsud (right) growing together (Qortin tal-Magun, Nadur, Gozo, 14 November 2009): (b) habit and habitat of N. tazetta subsp. aequilimbus scattered as individuals or small groups (rarely well-formed tufts) along shrubby vegetation on rocky shelves and banks (plateaux overlooking Wied Mġarr ix-Xini, Ta’ Ċenċ, Sannat, Gozo, locus of the neotype, 8 January 2024).
Figure 4.
(a) Field comparison of Narcissus × briffae Mifsud (left) and N. tazetta subsp. aequilimbus (Herb.) Mifsud (right) growing together (Qortin tal-Magun, Nadur, Gozo, 14 November 2009): (b) habit and habitat of N. tazetta subsp. aequilimbus scattered as individuals or small groups (rarely well-formed tufts) along shrubby vegetation on rocky shelves and banks (plateaux overlooking Wied Mġarr ix-Xini, Ta’ Ċenċ, Sannat, Gozo, locus of the neotype, 8 January 2024).
Figure 5.
Narcissus tazetta subsp. tazetta L. from the Maltese Islands: (a) habitat, typically steppe and abandoned fields on clay slopes or clayey soil; (b) inflorescence with 9 flowers; (c) detail of flower with imbricated broadly ovate tepals and golden yellow corona with a circular outline, subentire margin and unlobed or undivided wall; (d) stocky plants with well-developed sheath (3 layers) enveloping 4 wide leaves (typically 15–20 mm wide) and a central scape; (e) pollen grains at ×100 magnification (scale bar = 10 µm) with an average size of 47 × 38 µm covered with numerous yellow warts (stained in cotton blue).
Figure 5.
Narcissus tazetta subsp. tazetta L. from the Maltese Islands: (a) habitat, typically steppe and abandoned fields on clay slopes or clayey soil; (b) inflorescence with 9 flowers; (c) detail of flower with imbricated broadly ovate tepals and golden yellow corona with a circular outline, subentire margin and unlobed or undivided wall; (d) stocky plants with well-developed sheath (3 layers) enveloping 4 wide leaves (typically 15–20 mm wide) and a central scape; (e) pollen grains at ×100 magnification (scale bar = 10 µm) with an average size of 47 × 38 µm covered with numerous yellow warts (stained in cotton blue).
Figure 6.
Separation of four morphs based on their morphometry based on non-metric multidimensional scaling (nMDS).
Figure 6.
Separation of four morphs based on their morphometry based on non-metric multidimensional scaling (nMDS).
Figure 7.
Iconotype of Hermione aequilimba Herb. (=Narcissus tazetta subsp. aequilimbus (Herb.) Mifsud) collected from the Maltese Islands, flowering in October [6].
Figure 7.
Iconotype of Hermione aequilimba Herb. (=Narcissus tazetta subsp. aequilimbus (Herb.) Mifsud) collected from the Maltese Islands, flowering in October [6].
Table 1.
Locations of examined populations, including altitude, coordinates and attributed morphotype code.
Table 1.
Locations of examined populations, including altitude, coordinates and attributed morphotype code.
| Sample Code | Examination Date | Island | Locality | Toponym (Valley Name) | Altitude (m) | Geographic Coordinates | Morphotype |
|---|---|---|---|---|---|---|---|
| NRC01 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | 45–55 | 36°1′45.50″–47.88″ N 14°19′12.04″–13.70″ E | Narc-A |
| NRC02 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | Narc-A | ||
| NRC03 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | Narc-A | ||
| NRC04 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | Narc-A | ||
| NRC05 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | Narc-A | ||
| NRC06 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | Narc-B | ||
| NRC07 | 29 October 2022 | Gozo | Qala | Ħondoq ir-Rummien | Narc-B | ||
| NRC08 | 4 November 2022 | Malta | Dingli | Dingli Cliffs (Radar station) | 240 | 35°51′9.36″ N 14°23′5.75″ E | Narc-A |
| NRC09 | 4 November 2022 | Malta | Mellieħa | Qortin tal-Mistra | 35 | 35°57′16.76″ N 14°23′21.31″ E | Narc-A |
| NRC10 | 5 November 2022 | Gozo | Qala | Ħondoq ir-Rummien | 70 | 36°1′55.10″ N 14°19′39.08″ E | Narc-A |
| NRC11 | 12 November 2022 | Gozo | Sannat | Ta′ Ċenċ | 135 | 36°1′18.87″ N 14°15′16.99″ E | Narc-A |
| NRC12 | 12 November 2022 | Gozo | Munxar | Il-Fekruna | 60 | 36°1′57.64″ N 14°13′45.18″ E | Narc-B |
| NRC13 | 12 November 2022 | Gozo | Xagħra | Tal-Pergla | 80 | 36°4′1.55″ N 14°16′12.76″ E | Narc-A |
| NRC14 | 14 November 2022 | Gozo | Sannat | Ta′ Ċenċ (il-gruwa) | 75 | 36°1′14.77″ N 14°15′43.50″ E | Narc-C1 |
| NRC15 | 15 November 2022 | Malta | Mellieħa | Selmun area | 25 | 35°57′49.33″ N 14°23′51.66″ E | Narc-B |
| NRC16 | 16 November 2022 | Gozo | Sannat | Wied Mġarr ix-Xini | 50–55 | 36°1′28.24–28.41″ N 14°15′45.43–45.95″ E | Narc-C1 |
| NRC17 | 16 November 2022 | Gozo | Sannat | Wied Mġarr ix-Xini | Narc-C1 | ||
| NRC18 | 16 November 2022 | Malta | Mellieħa | Selmun area | 40 | 35°57′33.50″ N 14°23′43.16″ E | Narc-B |
| NRC19 | 25 November 2022 | Gozo | Munxar | Wied il-Munxar | 40–60 | 36°1′52.20–52.84″ N 14°13′56.46–57.49″ E | Narc-C1 |
| NRC20 | 25 November 2022 | Gozo | Munxar | Wied il-Munxar | Narc-C1 | ||
| NRC21 | 25 November 2022 | Gozo | Munxar | Wied il-Munxar | Narc-C1 | ||
| NRC22 | 11 January 2023 | Malta | Siġġiewi | Verdala | 220 | 35°51′38.48″ N 14°23′54.45″ E | Narc-C2 |
| NRC23 | 4 February 2023 | Malta | Mellieħa | Selmun-Mistra area | 30 | 35°57′41.89″ N 14°23′38.26″ E | Narc-C2 |
| NRC24 | 5 February 2023 | Gozo | Qala | Tal-Muxi area | 100 | 36°2′20.69″ N 14°19′31.68″ E | Narc-C2 |
| NRC25 | 18 February 2023 | Malta | Dingli | Tar-Rehba area | 230 | 35°51′23.48″ N 14°23′3.20″ E | Narc-C2 |
| NRC26 | 28 February 2023 | Gozo | Rabat | Gelmus Hill | 80 | 36°2′50.13″ N 14°14′7.40″ E | Narc-C2 |
| NRC27 | 12 December 2023 | Malta | Mellieħa | Selmun-Mistra area | 40 | 35°57′39.70″ N 14°23′38.37″ E | Narc-B |
| NRC28 | 29 December 2023 | Malta | Żurrieq | Wied Babu | 60 | 35°49′23.56″ N 14°27′33.47″ E | Narc-C1 |
| NRC29 | 29 December 2023 | Malta | Mosta | Wied il-Għasel | 40 | 35°55′5.87″ N 14°25′38.37″ E | Narc-C1 |
Table 2.
Comparison of examined characters of Narcissus morphotypes occurring in the Maltese Islands.
Table 2.
Comparison of examined characters of Narcissus morphotypes occurring in the Maltese Islands.
| Character | Narc-A | Narc-B | Narc-C1 | Narc-C2 |
|---|---|---|---|---|
| Habitat | Exposed rocky ground, steppe, low garigue | Steppe, low garigue, rocky valley banks | Rocky ground at valley sides or escarpments, especially rocky banks and ledges | Abandoned clayey fields or clay slopes, gardens, derelict agricultural areas |
| Habit | Solitary, rarely in small gregarious groups | Dense tufts of (10–)25–50(–70) plants closely packed to each other | Singular or isolated small groups of 4–10 plants, often close to each other and merging to form larger clusters of plants. | Dense tufts of (10–)25–50(–80) plants are closely packed to each other. Very rarely seen as singular individuals. |
| Flowering time | Autumnal (September–November) | Autumnal (October–November) but later from Narc-A | Autumnal (November–December) much earlier from Narc-C2 | January–March |
| Plant length | 15 (10–30) cm | 25 (22–35) cm | 40 (20–70) cm | 50 (30–120) cm |
| Presence of leaves with flowers | Absent | Present, about half the length of the scape | Present, reaching the same length of the scape | Present, reaching the same length of the scape |
| Leaves per plant | 1–2 | 1(–2) | 3–4 | (3–)4 |
| Basal sheath | Indistinct | Pale green with longitudinal parallel bright green veins. | Pale green with longitudinal parallel bright green veins. | Pale green with longitudinal parallel bright green veins. |
| Basal sheath length | N/a | 5–12 mm long, 1 layer exposed above ground level | 15–35 mm or even up to 100 cm in shaded locations, 3–4 layers exposed above ground level | 10–30 mm 2–3(–4) layers exposed above ground level |
| Leaf length | 5–18 cm | 15–28 cm | 20–40 cm, subequal to the scapes. Can get longer when plants are growing within bushes due to etiolation | 30–40 cm, shorter than the scapes |
| Leaf width | 1 mm | 2–5 mm | 7–14(–19) mm | 10–25 mm |
| Leaf cross-section | Terete | Crescent with rounded ends, fleshy | Canaliculate to flattened V-shaped, thin and not fleshy | Canaliculate to flattened V-shaped, thin and not fleshy |
| Flowers per scape | 1–2(–4) | (2–)3–5(–6) | (3–)4–7(–8) | (4–)6–10(–12) |
| Length of longest pedicels | Up to 15 mm | Up to 42 mm | Up to 68 mm | Up to 58 mm |
| Length of perianth tube | 14 (12–17) | 15 (14–16) | 16–24 mm | 17–19 mm |
| Flower diameter | 17–35 mm | 22–35 mm | 18–30 mm | 24–36 mm |
| Tepal colour | Pure white | Whitish-cream | Whitish-cream | Whitish-cream |
| Length lower tepals | 12.5 (8.0–16.0) mm | 10.5 (9.5–12.5) mm | 13.4 (11.5–15.0) mm | 16.0 (14.0–17.5) mm |
| Width lower tepals | 5.0 (4.0–6.5) mm | 5.5 (5.0–6.5) mm | 10 (8.5–11.0) mm | 11.5 (9.5–13.5) mm |
| Length upper tepals | 11.0 (7.0–13.5) mm | 9.5 (8.5–11.0) mm | Subequal to lower tepals | Subequal to lower tepals |
| Width upper tepals | 4.0 (3.0–5.0) mm | 5.0 (4.5–6.5) mm | Subequal to lower tepals | Subequal to lower tepals |
| Tepal imbrication | Usually free, but sometimes marginally imbricated at the base | Free to slightly imbricated from the basal third, then free | Imbricated from the basal half, free above | Imbricated from the basal half, free above |
| Diameter of corona | 2.2 (1.8–3.0) | 5.0 (3.5–6.5) | 9.0 (7.0–11.5) mm | 11.0 (9.0–13.5) |
| Height of corona | 1.5 (0.8–2.2) | 2.0 (1.6–2.5) | 5.0 (3.7–7.0) mm | 6.0 (5.0–7.5) |
| Corona colour | Greenish yellow to yellow and golden yellow sometimes deeper hues to amber (but not orange or reddish) | Golden yellow | Bright lemon-yellow | Bright lemon-yellow |
| Corona outline shape | Subdeltate, trigonous, or rarely almost circular | Circular to subtrigonous, rarely triangular-like | Circular occasionally trigonous or with three shallow lobes (esp. in examples with shallow coronae) | Circular or subcircular, seldom showing any signs of trigonometry |
| Corona margin | Subentire to crenated, occasionally shallowly lobed | Entire with three equidistant pleats | Subentire, crenated or rarely 3-lobed | Subentire |
| Corona lateral profile | Parallel sides (cup-shaped) sometimes slightly converging at the upper part (urn-shaped) | Parallel or more frequently divergent from the base (bowl-shaped) | Parallel or divergent from the base, comparatively deep (bowl-shaped) or shallow (saucer-shaped) | Mostly parallel or slightly divergent and deep. |
| Corona wall | Variable, notched, clefted or deeply incised to the bottom from (1–)3 equidistant locations forming 3 distinct flaps | Usually pinched (pleated) at the top from 3 locations, or rarely with 1(–3) clefts or full incisions. | Mostly entire but sometimes with 1–3 pleats at the top, rarely with 1(–2) clefts or partial incisions | Entire, sometimes with 1–3 pleats or a pinched area at the top |
| Tepal length/corona diameter ratio | 5–6 | 2–2.5 | ≈1.3 | ≈1.4 |
| Viable seeds | Yes | No or very few | Yes | Yes |
| Viability pollen | Yes (≈95% viable) | ≈10% viable | Yes (≈92% viable) | Yes (≈80% viable) |
| Pollen shape | Broadly ellipsoid with one compressed side | Dimorphic, broadly ellipsoid and sub-spherical | Broadly ellipsoid with one compressed side | Broadly ellipsoid with one compressed side |
| Average pollen size | 42.5 × 32.0 | Broadly elliptic pollen 38.0 × 29.5 µm; Subspherical pollen ca. 52.0 × 45.5 µm | 40.2 × 31.1 | 47.7 × 35.6 |
| Lipid texture on pollen testa | Scanty, small, about 3–5 µm wide | Prominent, sometimes as large semispherical warts, up to 5.9 µm wide | Prominent, numerous sometimes as large semispherical warts, up to 7.2 µm wide | Prominent, sometimes as large semispherical warts, up to 6.7 µm wide |
Table 3.
Comparison of selected characters between N. × briffae and its parents N. deficiens and N. tazetta indicating which characters of the hybrid have an affinity to either parent (yellow for N. deficiens and blue for N. tazetta), which are intermediate (green), and which are unique (grey) due to its hybridogenous nature (e.g., sterility).
Table 3.
Comparison of selected characters between N. × briffae and its parents N. deficiens and N. tazetta indicating which characters of the hybrid have an affinity to either parent (yellow for N. deficiens and blue for N. tazetta), which are intermediate (green), and which are unique (grey) due to its hybridogenous nature (e.g., sterility).
| Character | N. deficiens | N. × briffae | N. tazetta (subsp. aequilimbus) | Character Affinity |
|---|---|---|---|---|
| Habitat | Exposed rocky ground, steppe, low garigue | Steppe, low garigue, rocky valley banks | Rocky ground at valley sides or escarpments, especially rocky banks and ledges | Closer to N. deficiens |
| Habit | Solitary, rarely in small gregarious groups | Dense tufts of (10–)25–50(–70) plants closely packed to each other | Singular or isolated small groups of 4–10 plants, often close to each other and merging to form larger clusters of plants | N. tazetta |
| Flowering time | Autumnal (September–November) | Autumnal (October–November) | Autumnal (November–December) | Intermediate |
| Average plant length | 15 cm | 25 cm | 40 cm | Intermediate |
| Presence of leaves with flowers | Absent | Present, about half length of the scape | Present, reaching same length of the scape | N. tazetta |
| No. of leaves/plant | 1–2 | 1(–2) | 3–4 | N. deficiens |
| Basal sheath | Indistinct or very short (above ground level) | Pale green with longitudinal parallel bright green veins. | Pale green with longitudinal parallel bright green veins. | N. tazetta |
| Basal sheath length | Nil—2 mm | 5–12 mm long, 1 layer exposed above ground level | 15–35 mm or even up to 100 cm in shaded locations, 3(–4) layers exposed above ground level | N. tazetta (reduced) |
| Leaf length | 5–18 cm | 15–28 cm | 20–40 cm (can get longer when plants are growing within bushes due to etiolation) | Intermediate |
| Leaf width | 1 mm | 2–5 mm | 7–19 mm | Intermediate |
| Leaf cross-section | Terete | Crescent with rounded ends, fleshy | Canaliculate to flattened V-shaped, thin and not fleshy | Considered as an Intermediate form between terete and flattened V-shaped |
| No of flowers/scape | 1–2(–4) | 2–5(–6) | (3–)4–7(–8) | Intermediate or reduced N. tazetta |
| Length of longest pedicels | Up to 15 mm | Up to 42 mm | Up to 68 mm | Intermediate |
| Tepal imbrication | Sometimes, usually free tepals | Imbricated from the basal third, then free | Imbricated from the basal half, isolated above | N. tazetta |
| Diameter of corona | 2.2 (1.8–3.0) | 5.0 (3.5–6.5) | 9.0 (7.0–11.5) mm | Intermediate |
| Height of corona | 1.5 (0.8–2.2) | 2.0 (1.6–2.5) | 5.0 (3.7–7.0) mm | N. deficiens |
| Corona outline shape | Subdeltate, trigonous, or rarely almost circular | Circular to subtrigonous, rarely triangular-like | Circular occasionally trigonous or with three shallow lobes (esp. in examples with shallow coronae) | Intermediate or both parents |
| Corona lateral profile | Parallel sides (cup-shaped), sometimes slightly converging at the upper part (urn-shaped) | Parallel or more frequently divergent from the base (bowl-shaped) | Parallel or divergent from the base, comparatively deep (bowl-shaped) or shallow (saucer-shaped) | Both parents |
| Corona wall | Variable, notched, cleft or deeply incised to the bottom from (1–)3 equidistant locations forming 3 distinct flaps | Usually pinched (pleated) at the top from 3 locations, or rarely with 1(–3) clefts or full incisions. | Mostly entire but sometimes with 1–3 pleats at the top, rarely with 1(–2) clefts or partial incisions | Somewhat intermediate (pleats can be assigned as halfway between incised and entire |
| Tepal length/corona diameter | 5–6 | 2–2.5 | ≈1.4 | Intermediate |
| Viable seeds | Yes | No or negligible | Yes | Hybrid character |
| Pollen abundance on the anthers | Copious | Scanty | Copious | Hybrid character |
| Pollen viability | Yes (≈95% viable) | ≈10% viable | Yes (≈90% viable) | Hybrid character |
| Pollen shape | Broadly ellipsoid with one compressed side | Dimorphic, broadly ellipsoid and sub-spherical | Broadly ellipsoid with one compressed side | Unique character possibly due to pollen dysfunction/sterility |
| Average pollen size | 42.5 × 32.0 | Broadly elliptic pollen 38.0 × 29.5 µm; Subspherical pollen ca. 52.0 × 45.5 µm | 40.2 × 31.1 | Both parents |
| Warts on pollen testa | Scanty, small, about 3–5 µm wide | Prominent, sometimes hemispherical up to 5.9 µm wide | Prominent, numerous hemispherical and large, up to 7.2 µm wide | N. tazetta |
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MDPI and ACS Style
Mifsud, S.; Lanfranco, S. An Updated Taxonomic Appraisal of Narcissus (Amaryllidaceae) in the Maltese Islands. Diversity 2024, 16, 397. https://doi.org/10.3390/d16070397
AMA Style
Mifsud S, Lanfranco S. An Updated Taxonomic Appraisal of Narcissus (Amaryllidaceae) in the Maltese Islands. Diversity. 2024; 16(7):397. https://doi.org/10.3390/d16070397
Chicago/Turabian StyleMifsud, Stephen, and Sandro Lanfranco. 2024. "An Updated Taxonomic Appraisal of Narcissus (Amaryllidaceae) in the Maltese Islands" Diversity 16, no. 7: 397. https://doi.org/10.3390/d16070397
APA StyleMifsud, S., & Lanfranco, S. (2024). An Updated Taxonomic Appraisal of Narcissus (Amaryllidaceae) in the Maltese Islands. Diversity, 16(7), 397. https://doi.org/10.3390/d16070397
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