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Daucus carota L. Map Illustration.
Linnaeus, Sp. pl.: 242 (1753). – Type: Linnaean Herbarium 340.1 (LINN) lectotype, sel. by Sáenz Laín, Anales Jard. Bot. Madrid 37: 487 (1981).
D Gulerod.
Therophyte (winter annual) or hemicryptophyte (biennial). Plant bristly to almost glabrous, to 80(–110) cm (sometimes very small), with a tap-root; both root and leaves with a distinct aromatic scent. Stem solid; basal part 2–5(–9) mm thick, sulcate to angular, rarely terete, usually not violet, indistinctly glaucous; upper internodes distinctly to indistinctly sulcate. Leaves 3–7(–10) at the base and 2–6(–9) on the stem (one of the basal ones is usually the largest); sheath rather narrow, usually not violet; petiole 2–11(–20) cm; blade 1–2(–3)-pinnate, (3–)6–17(–26) × 2–5(–11) cm. Primary leaflets (2–)3–6 pairs; angle leaflet/rachis 45–60(–70°); longest petiolules (0–)2–10(–18) mm. Ultimate leaflets 2–3-pinnatifid, with 3–7 pairs of lobes/teeth; petiolule 3–10(–19) mm; blade 10–36 × 8–26(–37) mm, with a length/width ratio of (0.8–)1–1.4(–1.9); margin flat; base shortly attenuate to shallowly cordate (usually truncate). Ultimate lobes (0.8–)1.5–4(–7.8) × 0.6–2.3 mm, with a length/width ratio of (1.1–)1.3–2.7(–4.3); apices acuminate or sometimes acute, hyaline or violet.
Umbels concave to flat (rarely convex to semiglobose) in flower, 2.5–5 cm high, 5.5–7(–12) cm wide, usually nest-like (with incurved rays) in fruit; peduncle 18–31 cm; rays 2.1–4.8(–6.3) cm, in their upper part papillose or sometimes bristly on the adaxial side. Bracts 7–14, persistent, 11–46(–75) mm, 1(–2)-pinnatifid, with 1–4 pairs of linear lobes; basal part 0.7–2.5(–3.5) mm wide, distinctly membrane-bordered, usually ciliate. Umbellules (16–)22–52, sometimes with a central small umbellule with one to few dark violet flowers; pedicels 0.6–1.2 cm, glabrous or upper part indistinctly papillose on the adaxial side. Bractlets 7–10(–11), persistent, 4–13(–20) mm long, pinnatifid or entire, with 0–2 pairs of linear lobes (usually 3-fid), the centrifugal ones distinctly larger; basal part 0.5–1.5 mm wide, distinctly membrane-bordered, usually ciliate (entire bractlets usually membranous and ciliate all around). Flowers usually distinctly zygomorphic (see subspecies), (16–)22–52 per umbellule; sepals 0–0.5 mm; petals cream-white to cream-yellow (in dry specimens turning more intensely yellow on ageing), rarely violet-tinged, 1.3–3.6 × 1.4–4.1 mm, bifid (cut halfway or almost to the base), sometimes with a green patch at the base of the incision; lobes usually uneven (the larger one often constituting the main part of the petal); filaments 0.9–2.2 mm; anthers 0.3–0.5 mm. Fruit not flattened, elliptic to broadly elliptic in outline; carpophore slightly flattened, almost entire. Mericarps 2.1–3.1 × 1–1.8 × 0.6–0.9 mm (length/width ratio 1.5–2.4), excluding spines; secondary ridges consisting of 13–19 whitish, 0.5–1.5 mm long spines; vittae light brown, partly covered by the secondary ridges; stylopodium conical, 0.3–0.5 mm wide; style 0.8–1.5 mm, directed outwards. – Mid-summer to late summer.
2n=18 (S Sk 5). – [2n=18]
Distribution and habitat. An archaeophyte in parts of the south, but also continually introduced with agriculture and gardening; possibly partly native on seashores and dunes (for details see under the subspecies). – D common in most areas, but less so in central Jylland. N fairly rare to rare in the southeastern parts, north to He Elverum and Ringsaker and along the southern coast west to VA Farsund; casual on the west coast and in the fjords (Ho several records); ST Trondheim and Skaun, and Tr Balsfjord 1963 (garden weed) and Sørreisa 1946 (with German troops). S common to scattered (but unevenly distributed) in the southern lowlands north to and including BhG, eastern Dls, Nrk, southeastern Vsm and central Upl; further north rare and mostly casual. F several records from A, apparently resident in some places; otherwise casual north to OP Oulu and Kn Hyrynsalmi. I ÍNo Fnjóskadal 1965.
Europe, N Africa, W Asia; introduced in China, Australia, New Zealand, North and South America.
Biology. The primary umbels consist of only bisexual flowers in subsp. carota and subsp. sativus, while there are also male flowers in those of subsp. gummifer. The percentage of male flowers is successively higher in secondary and tertiary umbels (cf. Heywood 1983). The dark violet, sterile flowers in the central umbellule, which occur in many specimens, have usually been regarded as a pollinator guide (Heywood 1983). The dense umbel with enlarged outer flowers has also been regarded as an insect guide, but Bell & Lindsey (1978) argued that it is more likely a structure that favours multiple pollination.
The contraction of the umbel after fertilization is probably an adaptation to prolong the period of fruit dispersal. When the umbel is drying, the outer umbellules bend outwards and drop their fruits, while the fruits of the central umbellules are released first when the umbel falls and decays. This should increase the probability of seed germination (cf. Heywood 1983). In subsp. carota the primary umbel produces most of the seeds, while c. 50% of the seeds are produced by the secondary umbels in subsp. sativus (Hawthorn et al. 1962).
Variation. Infraspecific variation is complex. A genetic differentiation of natural taxa, followed by domestication and breeding, including hybridization and selection of favourable cultivars, have obscured the limits of some of the taxa. Both phenotypic plasticity and genotypic variation within the taxa is rather large; deviating specimens (e.g., with red flowers, with reduced secondary ridges or dwarf morphotypes) may occur independently in several of the taxa (cf. Reduron 2007).
Daucus carota has been divided into two informal groups, viz. the carota group and the gummifer group (= D. gingidium L.). The carota group is characterized by thin leaves, umbels with rather narrow-lobed bracts and bractlets, and flexible rays forming a birdnest-like infructescence; the plants are often fairly tall with rather few branches, and occur in varying habitats. The group is represented in Norden by subsp. carota, subsp. sativus and subsp. maximus, of which subsp. sativus is regarded as a hybrid derivative (see further comments under subsp. sativus). Among the characters for the gummifer group are thick leaves, umbels with rather broad-lobed bracts and bractlets, and stiff rays not forming a birdnest-like infructescence; the plants are low and distinctly branched, and occur mainly on seashores. The group is only represented in Norden by subsp. gummifer
(see further comments under this subspecies). The two groups have sometimes been treated at specific level (Small 1978), but since intermediates are rather frequent (cf. Heywood 1983, Reduron 2007), infraspecific status is preferable.
Similar taxa. Leaves of Daucus carota are similar to those of Carum carvi, but the latter have sessile leaflets. – Glabrous specimens of Daucus carota may resemble Ammi majus or A. visnaga, but those species have glabrous ovaries/fruits with entire ridges.
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Basal parts densely covered with long bristles; lower primary leaflets inserted to the rachis at an angle of more than 70°; fruiting umbels not nest-like (rays stiff, straight) |
C. subsp. gummifer |
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Basal parts glabrous or sparsely covered with short bristles; lower primary leaflets inserted to the rachis at an angle of less than 70° ; fruiting umbels nest-like (rays flexible, incurved so that the umbellules meet) |
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Root swollen and brittle, usually orange; lower leaf blades broadly triangular, weak and almost glabrous; outer petals in the umbel not distinctly larger than the others |
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Root not swollen, tough, whitish; lower leaf blades narrowly ovate, stiff and usually densely covered with bristles; outer petals in the umbel distinctly larger than the others and radiating |
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Daucus carota ssp. sylvestris (Mill.) Hyl. (1945).
D Vild Gulerod. F rikkaporkkana. N vill gulrot. S vildmorot.
Hemicryptophyte (biennial) or rarely annual. Sparsely to moderately branched herb; taproot not swollen (3–9 mm thick), tough, whitish, often branched. Lower leaves ± patent, narrowly ovate, with a length/width ratio of 2–3.8(–5), greyish green, rather thick, densely to sparsely covered with bristles; angle primary leaflet/rachis 45–65(–70°); leaflets ± narrow-lobed.
Umbels small to medium-sized, usually concave to flat in flower, nest-like in fruit (rays thin, flexible, bent inwards in upper part); outer flowers zygomorphic, usually distinctly larger than the inner ones. Fruit with secondary ridge spines indistinctly to distinctly connate, united part 0.1–0.3 mm wide, and with 1–2 barbs at each tip; vittae equilaterally triangular in cross section.
2n=18 (S Sk 5). – [2n=18]
Distribution. Archaeophytic in D and the southern, calcareous lowlands of S; elsewhere introduced later, e.g. with ley seed. – D distribution mainly as in the species. N resident along the coast from Øf to VA Kristiansand; Ro Haugesund 1898, Ho Bergen 1946 (with German troops) and Stord 1924, SF Leikanger 1918, and ST Trondheim 2001 (with meadow seed). S distribution in the south mainly as in the species; in the north casual, northernmost records Ång Grundsunda 2000 (bark pile), Nb Piteå 1914 (sawmill), 1998 (filling earth). F several records from A, apparently resident in some places; EP Vaasa 1943–47 (railway), Kristiinankaupunki 1943 (with German troops), 1958 (docks), Kaskinen 1946 (railway). – Unconfirmed records from F V Naantali 1998 and Turku 1991, U Vantaa 2005, and OP Oulu 1988.
In most of the range of the species.
Habitat. On dry to mesic, sun-exposed, sandy or clayey, often bare soil; favoured by lime, but also occurring on nutrient-rich soil in non-calcareous areas. Mainly in man-made sites, such as roadsides, field margins, railways, gravel pits and other ruderal places, but in D and southern S also in seminatural sites, such as heathlands, dry meadows and seashores; probably favoured by slight overgrowth.
Variation. Daucus carota subsp. carota is very variable, in size, hairiness, shape of leaf-lobes and colour of flowers. However, the characters are not correlated and intermediate morphotypes are common. In S Gtl dwarf specimens have been collected in alvar vegetation, and several specimens with reduced, red-tinged petals have been collected throughout the distribution area.
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B. subsp. sativus (Hoffm.) Arcang. Map Illustration.
Arcangeli, Comp. fl. ital.: 299 (1882). – Daucus carota var. sativus Hoffm., Deutschl. fl.: 94 (1791). – Described from cultivated material.
D Have-Gulerod. F ruokaporkkana. I gulrót. N dyrka gulrot. S morot.
Hemicryptophyte (biennial). Sparsely to moderately branched herb; taproot in the first year swollen (14–30 mm thick), orange, usually unbranched, brittle. Lower leaves ± erectopatent, broadly ovate to deltate (length/width ratio 1.6–1.9), dark green, rather thin, glabrous to moderately densely covered with bristles; angle primary leaflets/rachis 30–60°; leaflets narrow-lobed to broad-lobed.
Umbels medium-sized to large, usually distinctly convex to semiglobose in flower, nest-like in fruit (rays thin, flexible, bent inwards in upper part); outer flowers like the inner ones (or sometimes slightly zygomorphic and/or slightly larger). Fruit with secondary ridge spines indistinctly connate, united part 0–0.15 mm wide, and with 1–6 barbs at each tip; vittae variable in shape, but mostly equilaterally triangular in cross section.
[2n=18]
Distribution and habitat. Cultivated as a root vegetable and occasionally escaped in abandoned fields (e.g. potato fields), earth fillings, refuse tips, or as a remnant in gardens or throwout; apparently rarely escaped, since the Nordic material is very scarce, but also most probably undercollected. – D records few and scattered, mostly post-1980, from Sjæ (Lammefjord), FyL and Jylland (all provinces); confirmed from Sjæ København 1962 (docks) and VJy Vind 1980. N seen from Ak Oslo 1946 (with German troops), Asker 1950 (railway station), He Elverum 1938, Ringsaker 1982 (field margin), Ro Strand 1927 (mill), Ho Stord 1922 (weed), ST Trondheim 1943 (dump) and Skaun 1945 (mill). S several records in the southern parts, north to Nrk Kumla 1917 (stone quarry), Örebro 1995 (earth heap), and Srm Ösmo 1912 (roadside); Jmt Frösön 1914. F several records in the southern parts, north to Kn Hyrunsalmi 1945 (with German troops). I INo Fnjóskadal 1965 (potato field). – Also reported from S Vrm Karlskoga (Nilsson 2008), but specimens not seen.
Origin uncertain but perhaps SW Asia (Heywood 1983); widely cultivated in temperate regions.
Comments. Subsp. sativus is not known in the wild, but has been cultivated since ancient times. The origin of the cultivars has been disputed, but most authors agree with Thellung (1927) that they originate from hybrid derivatives of subsp. carota and subsp. maximus (cf. Heywood 1983, Small 1978). Subsp. sativus has been subdivided into an ”eastern carrot”, containing both carotenes and anthocyanins (var. atrorubens Alef.), and a ”western carrot”, comprising the cultivars predominant in Europe, containing only carotenes (var. sativus).
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C. subsp. gummifer (Syme) Hook. f. Map.
Hooker, Student fl. Brit. Isl., Ed. 3: 185 (1884). – D. carota var. gummifer Syme in Sm., Engl. Bot. ed. 3 [B], 4: 157 (1865). – Described from the British Isles.
D Strand-Gulerod. S strandmorot.
Literature. Faurholdt & Hinke (2006), Reduron (2007).
Hemicryptophyte (biennial). Robust, richly branched herb, 10–35 cm tall, densely covered with bristles; taproot not swollen, whitish. Lower leaves narrowly ovate to triangular (length/width ratio 2–2.8), greyish green, thick; angle primary leaflets/rachis 65–90°; leaflets narrow-lobed to broad-lobed.
Umbels small to medium-sized, usually distinctly convex in flower, not nest-like in fruit (rays stiff, straight); outer flowers zygomorphic, usually distinctly larger than the other ones. Fruit with secondary ridge spines indistinctly to distinctly connate, and with 1–4 barbs at each tip.
Distribution and habitat. D possibly spontaneous on coastal sand dunes on the eastern islands, known from ØJy Tunø, FyL Korshavn and Langeland (Ristinge), Sjæ Sjællands Odde, Sejerø and (since 2002) Peberholm.
Atlantic coasts of Britain, France and northern Spain.
Comments. Subsp. gummifer was recognized in D by K. Wiinstedt under the name var. libanotifolius, which, however, was never validly published. It is quite distinct: the small, richly branched shoots are densely covered with bristles, the primary leaflets are inserted at a wide angle, and umbels are never contracted, due to the stiff rays.
According to Reduron (2007), subsp. gummifer also has rather broad-lobed bracts and bractlets, fruits with rather short spines on the secondary ridges, and glossy leaves; the Danish material is not fully typical, having bracts and bractlets with rather narrow lobes and fruits with rather long spines (glossiness of leaves could not be assessed in dried specimens). |
D. subsp. maximus (Desf.) Ball 1878. – Lit.: Heywood (1983), Thellung (1926a; also ill.). – Annual, large herb; taproot not swollen. Lower leaves narrowly ovate (length/width ratio 2.3–2.4), greyish green, moderately densely covered with bristles; leaflets rather broad-lobed. Umbels large; outer flowers zygomorphic, usually distinctly larger than the inner ones. Fruit with secondary ridge spines distinctly connate, united part 0.1–0.3 mm wide, and with 1–4 barbs at each tip; vittae elongated triangular in cross section. – [2n=18]. – Illustration.
S BhG Västra Frölunda 1961 (ditch). – The Mediterranean, SW Asia.. – Occurrence data (no map in the book).
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Rare casuals
Daucus glochidiatus (Labill.) Fisch., C.A. Mey. & Avé-Lall. 1843. – Lit.: Thellung (1926b), Saénz Laín (1981, ill.) – Annual, small herb with weak bristles. Umbels sessile or subsessile with few (up to 6) unequally long rays; bractlets 3–6. Fruit c. 0.5 cm with a papillose or smooth surface and minute styles; secondary ridges with 4–13 short and rather narrow spines; vittae small, flattened in cross section; ovary wall thin.
S Sk Lackalänga 1949 (wool alien). – Australia. – Map (not in the book).
Daucus muricatus (L.) L. 1762. S spikmorot. – Lit.: Saénz Laín (1981, ill.). – Annual, small herb with strong bristles. Umbels with pinnatifid bracts and many (usually more than 10) unequally long rays; bractlets entire or pinnatifid. Fruit to c. 1 cm; secondary ridges with 4–8 long, distinctly barbed spines with very broad, connate bases. – [2n=22]
S BhG Göteborg 1955–56 (Ringön; waste from fruit imports); also reported from S BhG Backa (Blom 1961) but no voucher has been seen. – The W Mediterranean. – Map (not in the book).
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