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The Biology of the Citron (Citrus medica L., Rutaceae-Aurantioideae-Aurantieae), its Hybrids and their Allies

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The Citron Compendium

Abstract

The biology of citrus is briefly reviewed, pointing up how surprisingly little is known of pollination and dispersal in wild species. The classification of the citron, Citrus medica, in modern science is set out in a historical context: its taxonomic relationships in the light of the evolution of the genus Citrus and the citron's role in the origin of major citrus crops as well as the importance of the etrog citron in the traditional Jewish Tabernacles festival are outlined. The global threat to the citron and all other citrus from the bacterial disease, huanglongbing, is explained. As an aid to understanding the much-confused citrological literature, the formal taxonomy of the citron is presented in an Appendix, complete with a nomenclatural account of those commercial crops which have citron in their make-up; for example, the Rangpur lime (a rough lemon) is Citrus × otaitensis (syn. C. × volcameriana, C. × jambhiri).

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Notes

  1. 1.

    * = Addition to POWO/IPNI and other databases.

  2. 2.

    * = Addition to POWO/IPNI and other databases.

  3. 3.

    * = Addition to POWO/IPNI and other databases.

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Acknowledgements

Thanks are due to George Beattie (Western Sydney University, Australia), for reviewing the section on huanglongbing, to Valery Malécot (Angers, France) for valuable information on Risso’s Citrus specimens, to John McNeill (Edinburgh) for nomenclatural advice and to Ian Warrender (Palmerston North, New Zealand) for redrawing Fig. 1.10.

Author information

Authors and Affiliations

  1. Wadham College, University of Oxford, Oxford, OX1 3PN, UK

    David J. Mabberley

  2. School of Natural Sciences, Macquarie University and Australian Institute of Botanical Science (Royal Botanic Gardens and Domain Trust), Sydney, NSW, 2000, Australia

    David J. Mabberley

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  1. David J. Mabberley
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Correspondence to David J. Mabberley .

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Editors and Affiliations

  1. Inst of Plant Science & Genetics in Agri, The Hebrew University of Jerusalem, Rehovot, Israel

    Eliezer E. Goldschmidt

  2. The S. Tolkowsky Laboratory, The Volcani Institute, ARO, Rishon LeZion, Israel

    Moshe Bar-Joseph

Appendix—Formal Classification

Appendix—Formal Classification

(Expanded and updated from Mabberley 1997, 2004, 2022; Zhang and Mabberley 2008).

As the names of citrus crops involving the citron are very much confused in the literature, regrettably so even in modern papers, this synopsis should allow the reader to place correctly germane names encountered there.

1.1 Citrus medica—Citrons

Citrus medica L., Sp. Pl. 2: 782 (1753) ≡ C. tuberosa Mill., Gard. Dict. ed. 8: Citrus no. 2 (1768), nom. superfl. ≡ C. fragrans Salisb., Prodr. Stirp. Chapel Allerton: 378 (1796), nom. superfl. ≡ C. medica var. vulgaris Risso & Poit., Hist. Nat. Orang.: 194, tt. 96. 97 (1820), nom. superfl. pro var. medica ≡ C. cedra Link, Handbuch 2: 346 (1829) & Risso, Fl. Nice: 87 (1844), isonym, ?nom. superfl. ≡ C. × limonum var. medica (L.) Risso, Hist. Nat. Prod. Eur. Mérid. 1: 431 (1826) ≡C. × aurantium L. [unranked] medica (L.) Wight & Arn., Prodr. Fl. Ind. Orient. 1: 98 (1834) ≡ C. medica subsp. genuina Engl., Nat. Pflanzenfam. III, 4: 200 (1896), nom. superfl. pro subsp. medica ≡ Aurantium medicum (L.) M. Gómez, Fl. Habanera: 205 (1897); C. medica var. proper Guillaumin in Agric. Prat. Pays Chauds n. s. 14: 118 (1932), nom. superfl. pro var. medica.

Type: “Habitat in Asia, Media, Assyria, Persia”. Type [icon]: ‘Citreum’ in Tournefort, Inst. Rei Herb. 3: t. 396 (1700) lectotype designated by D.M. Porter in C.E. Jarvis et al. (ed.), Regnum Veg. 127: 34 (1993).

[Citreum vulgare Tournef. [Inst. Rei Herb. 1: 621 (1700)] ex Mill., Gard. Dict.: Citreum no. 1 (1754), nom. illeg.]

Citrus spinosa S.G. Gmel., Reise Russland 3: 278 + t. 22 (1774).

Type: Iran [cult.?], S.G. Gmelin (in Herb. Pallas) (BM?).

Citrus medica var. tuberosa Risso in Ann. Mus. Hist. Nat. 20: 200 (1813) ≡ C. medica var. monstrosa C. Presl, Fl. Sicul.: 183 (1826), nom. superfl.

Type: France [cult.], ?not preserved.

? Citrus lumia Risso, Hist. Nat. Prod. Eur. Mérid. 1: 414 (1826) ≡ C. medica var. lumia (Risso) Engl., Nat. Pflanzenfam. III, 4: 200 (1896).

Syntypes: France [cult.], October, Risso ‘No. 37’ (P P06137324) and ‘39’ (P P05471819).

Citrus cedrata Raf., Sylva Tell.: 141 (1838).

Type: lost? (P-DU?).

Citrus papaya Hassk., Cat. Pl. Hort. Bog. Alt.: 218 (1844).

Type: Indonesia [cult.], Java, Bogor, ?Anon. s.n. [Hort. Bog. III-G-82] (?L holo; BO iso].

Citrus crassa Hassk., Cat. Pl. Hort. Bog. Alt.: 217 (1844).

Type: Indonesia [cult.], Java, Bogor, ?Anon. s.n. (?L holo).

Citrus medica var. bajoura Bonavia, Cult. Oranges lemons: 67 (1890) ≡ C. medica subsp. bajoura (Bonavia) Engl., Nat. Pflanzenfam. III, 4: 200 (1896).

Type: Not indicated.

? Citrus kwangsiensis Hu in J. Arnold Arbor. 12: 153 (1931).

Type: China, Kwangsi, 14 June 1928, R.C. Ching [Kwangsi Expedition 6456] (A A00044028 holo; NY NY00388449 iso).

? Citrus medica var. alata Yu. Tanaka in Trans. Nat. Hist. Soc. Taiwan 22: 431 (1932) ≡ C. alata (Yu Tanaka) Tanaka, Syst. Pomol.: 140 (1951).

Type: Philippines, Mindanao, Todaya, Mount Apo, 1924, M.S. Clemens s.n. (UC holo).

[Citrus medica var. yunnanensis S.Q. Ding ex C.C. Huang in Guihaia 11: 8 (1991), nom. inval. (ICN Art. 40.6)].

There are many cultivars (one being ‘Etrog’ [Citrus medica var. conifera Michel, C. medica var. ethrog Engl.], the etrog) and cultivar groups to which some of the above botanical varietal names (and others published by Risso and subsequent authors) may be assignable (but for Florentine citrons, see Citrus × limon). Another cultivar group (Fingered Group) discussed above includes a number of cultivars including:

C. medica var. digitata Lour., Fl. Cochinch. 2: 469 (1791) ≡ C. medica f. digitata (Lour.) Guillaumin in Agric. Prat. Pays Chauds n. s. 14: 119 (1932).

Type: Not preserved?

(N.B. Although Loureiro used the binomial C. digitata, he called it a variety of C. medica).

Sarcodactilis helicteroides Gaertn.f., Suppl. Carp.: 39 + t. 185 [f. 1 ‘Sarcodactylis helicterioides’] (1805).

Type: ‘E Guiana belgica [Surinam]’, Anon. s.n. (TUB holo?).

Citrus sarcodactylis Hoola van Nooten, Fl. Java 1(6): t. 3 (1863) ≡ C. medica L. var. sarcodactylis (Hoola van Nooten) Swingle, Pl. Wilson. (Sargent) 2: 141 (1914).

Type: “Native of East India.“ Probably not preserved, in which case the plate would be a good candidate as lectotype.

Citrus × limonum var. cheirocarpa H. Lév. ex Cavalerie in Bull. Géogr. Bot. 21: 211 (1911).

Type: ? not preserved.

The Buddha-hand or Buddha’s hand citron with separated carpels is best treated as a cultivar group. Also to be referred here is Citrus medica var. fructucornuto Michel (1816). Citrus limonum var. digitata Risso & Poit., Hist. Nat. Orang.: 192 + t. 95 (1820) and C. medica var. muiliensis H. Wang & Y. Ding in Acta Hort. Sin. 10: 181 (1983), nom. nud., are somewhat intermediate between ‘Fingered’ and typical C. medica.

1.2 Commercial Citrus with Citron Parentage and Grown in the Mediterranean

Citrus medica is the male parent of many citrus cultigens (see Fig. 1.10), of which the principal (with commonly used synonyms) are:

  1. 1.

    Citrus × aurantiifolia —(Key) Limes

Citrus × aurantiifolia (Christm.) Swingle in J. Wash. Acad. Sci. 3: 465 (1913), pro sp. ≡ Limonia × acidissima Houtt., Nat. Hist. II, 2: 444 (1774), pro sp., non L. (1753), ≡ L. × aurantiifolia Christm., Vollst. Pflanzensyst. 1: 618 (1777), pro sp. ≡ C. × limonellus Hassk. in Flora 25, Beibl. 2: 43 (1842), pro sp., nom. superfl. ≡ C. medica f. aurantiifolia (Christm.) Hiroe, Forest Pl. Hist. Jap. Islands 1: 219 (1974).

Type [icon]: ‘Limonellus sive Limon Nipis’ Rumpf, Herb. Amb. 2: t. 29 (1741), lectotype designated by B.C. Stone in Dassanayake & Fosberg, Revis. Handb. Fl. Ceylon 5: 424 (1985).

Citrus × nipis Michel, Traité Citronier: 44 (1816) pro sp.

Type: Nor preserved? Michel includes ‘Limonellus sive Limon Nipis’ Rumpf, Herb. Amb. 2: t. 29 (1741), type of C. × aurantiifolia, in his protologue.

Citrus × spinosissima G. Mey., Prim. Fl. Esseq.: 247 (1818), pro sp. ≡ C. medica var. spinosissima (G. Mey.) Mart., Syst. Mat. Med. Bras.: 30 (1843) ≡ C. × aurantium L. var. spinosissima (G. Mey.) Griseb., Fl. Br. West Ind. Is.: 132 (1864) ≡ Aurantium × spinosissima (G. Mey.) M. Gómez, Fl. Habanera: 205 (1897).

Type: Guyana [naturalized], R. Essequibo (=Araunama), ‘Arowabisch’ Is., Rodschied 131 (GOET).

? Citrus × mellarosa Risso, Hist. Nat. Prod. Eur. Mérid. 1: 405 (1826), pro sp.

Type: France [cult.], ‘primavere’, Risso s.n. (P P05308995).

? C. × aurata Risso, Hist. Nat. Prod. Eur. Mérid. 1: 409 (1826), pro sp.

Type: France [cult.], not found.

Citrus × lima Macfad. in Bot. Misc. 1: 300 (1830), pro sp. ≡ C. × aurantium subsp. lima (Macfad.) Guillaumin in Agric. Prat. Pays Chauds n. s. 14: 113 (1932).

Type: Jamaica, not preserved?

C. × acida Roxb., Fl. Ind. 3: 390 (1832), pro sp. ≡ C. medica L. var. acida (Roxb.) Hook.f., Fl. Brit. India 1: 515 (1875) ≡ C. hystrix DC. subsp. acida (Roxb.) Bonavia ex Engl., Nat. Pflanzenfam. III, 4: 200 (1896).

Type [icon]: India, “Cultivated in Bengal”, ‘Citrus acida’, Fig. 1 (Icones Roxburghianae) (K, lectotype designated here).

Citrus × notissima Blanco, Fl. Filip.: 607 (1837), pro sp.

Type: Philippines, not preserved?

Citrus × lima Raf., Sylva Tell.: 143 (1838), pro sp., nom. illeg., non Macfad. (1830).

Type: lost? (P-DU?).

*Footnote 2Citrus × rissoa Risso in Hortic. Univ. 1: 200 + t. 27 (1839) & Herb. Gén. Amat., sér. 2, 2: 15 + t. (1841), pro sp., syn. nov.

Syntypes: France [cult.], Risso s.n. (P P05309085), Alpes-Maritimes, Beaulieu-sur-Mer, Risso s.n. (P P05309084), ‘fleur janvier’, Risso s.n. (P P05309082).

  1. 2.

    Citrus × floridana —Limequats

Citrus  ×  floridana (J. Ingram & H.E. Moore) Mabb. in Telopea 7: 337 (1998) ≡ × Citrofortunella floridana J. Ingram & H.E. Moore in Baileya 19: 170 (1975).

Holotype: [icon] ‘Evstis [sic = ‘Eustis’] limequat (No 48798), grown in the greenhouse at Washington, D.C.‘, J. Agric. Res. 23: [237] t. 4 (1923).

 × Citrofortunella swinglei J. Ingram & H.E. Moore in Baileya 19: 170 (1975), non Citrus swinglei Burkill ex Harms (1931).

Holotype: [icon] ‘Tavares limequat’ (No 48792), J. Agric. Res. 23: [238] t. 5B (1923). 

  1. 3.

    Citrus × latifolia —Persian or Tahitian (Seedless) Limes

Citrus  ×  latifolia (Yu. Tanaka) Tanaka, Syst. Pomol.: 140 (1951), pro sp. ≡ C. × aurantiifolia var. latifolia Yu. Tanaka in Agr. & Hort. 9: 2346 (1934) ≡ C. × aurantiifolia subsp. latifolia (Yu.Tanaka) S. Ríos & al., Varied. Trad. Frut.: 101 (1998).

Type: Not preserved?

  1. 4.

    Citrus × limon —Lemons

Citrus × limon (L.) Osbeck, Reise Ostindien: 250 (1765) as ‘limonia’, pro sp. ≡ C. medica L. var. limon L., Sp. Pl. 2: 782 (1753) ≡ C. medica var. odoratissima Andrews, Bot. Rep. 10: t. 609 (1810), excl. ic. & descr., nom, superfl. pro var. limon ≡ C. medica L. f. limon (L.) Hiroe, Forest Pl. Hist. Jap. Islands 1: 218 (1974).

Type [icon]:’Limon vulgaris’ in Ferrari, Hesperides: 191, 193 (1646) lectotype designated by D.J. Mabberley in Telopea 7: 169 (1997).

Citrus × limonum Risso in Ann. Mus. Natl. Hist. Nat. 20: 201 (1813), pro sp. ≡ C. × limonum var. vulgaris Risso & Poit., Hist. Nat. Orang.: 174 + t. 84 (1820), nom. superfl. pro var. limonum ≡ C. × aurantium [unranked] limonum (Risso) Wight & Arn., Prodr. Fl. Ind. Orient. 1: 98 (1834) ≡ C. medica var. limonum (Risso) Hook. f., Fl. Brit. India 1: 515 (1875) ≡ C. medica subsp. limonum (Risso) Engl., Nat. Pflanzenfam. III, 4: 200 (1896).

Type: “Ce lemonier s’élève jusqu’a quatre metres dans notre department; et quoique il soit moins sensible au froid que les varietés suivantes, il est peu cultivé parce qu’il ne donne qu’une petite quantité de fruits.“ (?not preserved).

Citrus medica var. florentina Risso in Ann. Mus. Hist. Nat. 20: 200 + t. 4 [II] Fig. 1 (1813) ≡ C. cedra var. florentina (Risso) Risso, Fl. Nice: 87 (1844).

Type: France [cult.], ?not preserved, in which case the figure would make an appropriate lectotype.

Citrus × limetta Risso in Ann. Mus. Natl. Hist. Nat. 20: 195 + t. 4 (II) Fig. 2 (1813), pro sp.

C. × lumia var. limetta (Risso) Risso & Poit., Hist. Nat. Orang.: 144 + t. 69 (1819); C. medica var. limetta (Risso) Hook. f., Fl. Brit. India 1: 515 (1875); C. × aurantium var. limetta (Risso) Guillaumin in Agric. Prat. Pays Chauds n. s. 14: 115 (1932).

Syntypes: France [cult.], Risso s.n. (P P05309097, P05309098).

Citrus × limetta var. bergamia Risso in Ann. Mus. Natl. Hist. Nat. 20: 197 (1813) ≡ C. × aurantium var. bergamia Michel, Citronier: 31, t. 26 + Fig. 3 (1816) ≡ C. × bergamia (Risso) Risso, Hist. Nat. Prod. Eur. Mérid. 1: 403 (1826), pro sp. ≡ C. × aurantium [unranked] bergamia (Risso) Wight & Arn., Prodr. Fl. Ind. Orient. 1: 98 (1834) ≡ C. × aurantium subsp. bergamia (Risso) Engl., Nat. Pflanzenfam. III, 4: 198 (1896).

Type: France [cult.] (? not preserved).

?Citrus × aurata var. gordonia Risso, Hist. Nat. Prod. Eur. Mérid. 1: 411 (1826) ≡Footnote 3*C. × gordonia (Risso) Risso in Hortic. Univ. 1: 352 + t. 41 (1839) & Herb. Gén. Amat., sér. 2, 2: 33 + t. (1841), pro sp.

Syntypes: France [cult.] ‘Citrus Gordon’, Risso s.n. (P P05308992); ‘C. Gordon’, Risso s.n. (P P05308993).

Citrus × bergamota Raf., Sylva Tell.: 141 (1838), pro sp.

Type: lost? (P-DU?).

Citrus × pachiderma Risso, Fl. Nice: 87 (1844), pro sp., syn. nov.

Type: France [cult], Risso s.n. (P P05309047 holo).

Citrus × dimorphocarpa Lush. in Ind. Forester 36: 346 (1910).

Type: Not indicated, but probably a cultivated plant at Dehra Dun, India (?not preserved).

Citrus × limonimedica Lush. in Ind. Forester 36: 348 (1910).

Type: Not indicated, but probably a cultivated plant at Dehra Dun, India (?not preserved).

Citrus × medicolimonum Lush. in Ind. Forester 36: 349 (1910).

Type: Not indicated, but probably a cultivated plant at Dehra Dun, India (?not preserved).

Citrus × meyeri Yu. Tanaka, Icon. Jap. Citrus Fruits 1: 91 (1946).

Type: Not found.

Citrus × limodulcis D. Rivera et al., Varied. Trad. Frut.: 125 (1998).

Type: Spain [cult.], Murcia, Churra, 1996, D. Rivera & F. Mendez (MUB 47,654 holo; MUB iso).

  1. 5.

    Citrus × otaitensis —Rough Lemons (including Rangpur Lime)

Citrus  ×  otaitensis (Risso & Poit.) Risso, Fl. Nice: 86 + t. [7] (1844 ‘taitensis’), pro sp. ≡ C. × aurantium var. otaitensis Risso & Poit., Hist. Nat. Orang.: 66 + t. 27 (1819) ≡ C. × lumia var. otaitensis (Risso & Poit.) Lémon in Rev. Bot. 1: 115 (1829) ≡ C. × limon var. otaitensis (Risso & Poit.) Tanaka in Sci. Bull., Facult. Agric. Kyushu Univ. 1: 107 (1925).

Type: France [cultivated], Paris, Noisette’s garden and ?not preserved, so the plate is here designated as lectotype (see Fig.  9). Apparently the pot-plant grown as ‘Otaheite’ for winter decoration in USA etc. Note that the only (undated) specimen in Risso’s herbarium is ‘Citrus otaitense [sic]’, Risso s.n. (P P02441072) and that, as so often in his flora, Risso ‘recycled’ his earlier varietal epithets (see for example C. bergamia above) as species names, this time inadmissibly replacing the by then archaic ‘Otaite’ with ‘Taite’, but in the horticultural literature of the nineteenth century the name was usually rendered C. otaitensis. I am indebted to John McNeill (E) for advice here.

Citrus × bigaradia var. volcameriana Risso & Poit., Hist. Nat. Orang.: 91, t. 40 (1819); Ten., Cat. Orto Bot. Napoli: 20 (1845) [’Wolkameriana R.’] ≡ C. × volcameriana (Risso & Poit.) V. Ten. & Pasq., Comp. Bot.: 414 (1847), pro sp. Syn. nov.

Type [icon]: Risso & Poit., Hist. Nat. Orang.: t. 40 (1819) lectotype designated here (see Fig. 1.11); the illustration and description published by Risso & Poiteau match the Rangpur lime (a rough lemon).

[Citrus × limetta var. sinensis Hort. Angl. ex Lémon, op. cit.: 117, nom. in synon.]

[‘C. × limonia’ auctt., non C. × limon (L.) Osbeck, ‘limonia’].

Citrus × volkameriana Pasq., Cat. Ort. Bot. Napoli: 29 (1867), pro sp., nom. illeg., non C. × volcameriana V. Ten. & Pasq. (1847).

Type: Italy [cult.], Reggio Calabria and Naples (?NAP not seen).

Citrus × jambhiri Lush. in Ind. Forester 36: 342 (1910), pro sp. ≡ C. × aurantium subsp. jambhiri (Lush.) Engl., Nat. Pflanzenfam. III, 4: 200 (1896) as ‘jambiri’. Syn. nov.

Type: Not indicated, but probably a cultivated plant at Dehra Dun, India (?not preserved), fide D. J. Mabberley in Telopea 7: 170 (1997).

? Citrus x rangpuriensis Traub, Plant Life 29: 172 (1973).

Type: USA [cult.], California (n.v.).

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Mabberley, D.J. (2023). The Biology of the Citron (Citrus medica L., Rutaceae-Aurantioideae-Aurantieae), its Hybrids and their Allies. In: Goldschmidt, E.E., Bar-Joseph, M. (eds) The Citron Compendium. Springer, Cham. https://doi.org/10.1007/978-3-031-25775-9_1

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