(3133–3134) Proposals to reject the names <i>Ficus serrata</i> and <i>F</i> . <i>maculata</i> ( <i>Moraceae</i> )

(3133) Ficus serrata L., Syst. Nat., ed. 10, 2: 1315. 7 Jun 1759 [Angiosp.: Mor.], nom. utique rej. prop. Typus: non designatus. (3134) Ficus maculata L., Sp. Pl., ed. 2, 2: 1515. Jul–Aug 1763 [Angiosp.: Mor.], nom. utique rej. prop. Typus: non designatus. Ficus serrata was described by Linnaeus (Syst. Nat., ed. 10, 2: 1315. 1759) with a concise diagnosis “A. F. fol. lanceolatis serratis” and a sole reference to “Plum. ic. t. 131. f. 1”, i.e., Claude Aubriet's copy of Charles Plumier's drawing of an unusual fig from the West Indies (most likely from Haiti) published in the sixth fascicle of Johannes Burman's Plantarum Americanum on 22 August 1757 (Burman, Pl. Amer.: t. 131, fig. 1. 1757; Stafleu & Cowan in Regnum Veg. 110 [TL-2, 2]: 301, 303. 1983). The prefix “A” preceding the nomen specificum legitimum in the protologue indicates that F. serrata was treated by Linnaeus as a new species in addition to the seven already described by him in the first edition of Species plantarum (Linnaeus, Sp. Pl. 2: 1059–1060. 1753; Baum in Taxon 17: 507. 1968). Linnaeus did not cite any additional elements, and the illustration cited is taxonomically ambiguous as it shows a branch with finely serrated leaves (a character unusual or unknown in the New World Ficus), although the depicted infructescences with achenes and punctations on the fig surface are distinctive of Ficus. Burman (l.c.: 122) synonymized Plumier's drawing with two earlier polynomials: Plukenet's “Ficus Americana, latiore folio venoso ex Curacao” (Phytographia: t. 178, fig. 1. 1692) and Sloane's “Ficus indica, folio oblongo, obtuso, fructu minore, pallide luteo, sphaerico” (Cat. Pl. Jamaica: 189. 1696), but both elements conflict with Linnaeus's protologue. For instance, Plukenet's engraving depicts a sterile stem with five entire, broadly ovate to cordate leaves and more closely resembles F. benghalensis L. (l.c. 1753: 1059) (Hamilton in Trans. Linn. Soc. London 15: 134. 1827). In contrast, Sloane (l.c.) tentatively compared his entity with Plukenet's t. 178, fig. 1 (l.c.) and Kiggelaer's polynomial (Hort. Beaumont.: 17. 1690), the latter recognized as F. religiosa L. (l.c. 1753: 1059) (Hamilton, l.c.: 135). Linnaeus did not include these two additional references in the protologue of F. serrata, likely because of the concise nature of the Systema naturae and probably due to the fact he was working from a copy of a proof plate sent to him by Burman on 19 July 1757, a month before publication of the sixth fascicle of Plantarum Americanarum, as mentioned in a letter preserved at the Linnean Society in London (Linnaean Correspondence, Ref. No. LC/2/195). Linnaeus received plenty of similar proof copies of Plumier's drawings from Burman between 1755 and 1760 and later used them to decorate the walls of the study in his summer home in Hammarby in Sweden, where they can still be seen to this day (Jarvis in Linnean, Special Issue 8: 80. 2008). Four years after Ficus serrata was published, Linnaeus (Sp. Pl., ed. 2: 1515. 1763) abandoned this name and described F. maculata, providing a diagnosis: “FICUS foliis oblongis acuminatis serratis” along with a full citation of Plumier's polynomial “Ficus castaneae folio, fructo globoso maculato Plum. spec. 21” (Plumier, Nov. Pl. Amer. [Cat. Pl. Amer.]: 21. 1703) directly corresponding to his own drawing (now housed at the Library of the National Museum of Natural History in Paris, France) and to Aubriet's copy published by Burman (l.c.) and cited by Linnaeus as “ic. 131. f. 1”. Linnaeus included the provenance “Habitat in America” followed by the symbol “♄” to indicate a woody habit. Linnaeus's choice of the epithet “maculata” likely reflected his principle of selecting a “nomen specificum essentiale” (Stearn in Ray Soc. Publ. 140: 85. 1957), here referring to the distinctively spotted surface of the figs depicted on Plumier's drawing. The absence of additional synonyms or specimens cited in the protologue may represent taxonomic hesitancy or Linnaeus's poor understanding of this species. Unfortunately, Linnaeus's own interleaved copies of both Systema naturae ed. 10 and Species plantarum ed. 2 preserved at the Linnean Society in London contain no annotations in the author's hand which would clarify his intent. Since their publication, both Ficus serrata and F. maculata have been inconsistently applied and generally treated as nomina dubia. For instance, F. maculata has been regarded as nomenclaturally superfluous and illegitimate (e.g., Jarvis, Order out of Chaos: 519. 2007). A strict reading of Art. 52.1–52.3, however, indicates that F. maculata remains legitimate; yet because both names were evidently based on the same Plumier illustration, F. maculata is unambiguously a synonym of F. serrata. Nevertheless, F. maculata continued to appear sporadically in the literature, whereas F. serrata quickly fell into obscurity, likely as a consequence of this early interpretive confusion (e.g., Houttuyn, Nat. Hist. II, 3: 685. 1774; Aublet, Hist. Pl. Guiane 2: 952. 1775; Thunberg, Ficus: 15. 1786; Lamarck, Encycl. 2: 498. 1786; Loureiro, Fl. Cochinch. 2: 666. 1790; Vahl, Enum. Pl. 2: 198. 1805; Willdenow, Sp. Pl. 4: 1152. 1806; Sprengel, Syst. Veg. 3: 784. 1826). Provenance reports were vague, ranging from Santo Domingo (modern Haiti: Lamarck, l.c.; Willdenow, l.c.) to Cochinchina (Vietnam: Loureiro, l.c.), though these records lack any confirmation based on the study of herbarium material. Subsequent interpretations varied. Thunberg (l.c.) first synonymized Ficus maculata with F. hispida L. f. (Suppl. Pl.: 442. 1782), an Old World taxon, although Linnaeus (Syst. Veg. 2: 922. 1784) had recognized them as distinct. Miquel (in London J. Bot. 6: 551. 1847) rejected Thunberg's view, excluding F. maculata from his treatment and later listing it under “Species delendae” (Miquel in Ann. Mus. Bot. Lugduno-Batavi 3: 299. 1867). In contrast, Urban (Symb. Antill. 8: 169. 1920) and Moscoso (Cat. Fl. Doming. 1: 139. 1943) recognized F. serrata as endemic to the Antilles or Haiti, though without herbarium material confirmation. Urban (in Repert. Spec. Nov. Regni Veg. 15: 158. 1918) also made a plausible suggestion that perhaps Plumier mixed the fruits, which are unmistakably figs, with the leaves of another unidentified plant (maybe even from a different family) that he found under the tree depicted on his drawing. Most recent treatments consistently excluded both names from Antillean Ficus (e.g., Warburg in Urban, Symb. Antill. 3: 491. 1903; Howard, Fl. Lesser Antilles 4: 57–64. 1988; Liogier, Fl. Española 8: 452. 1996; Berg & Villavicencio in Ilicifolia 5: 12. 2004). The situation was further complicated by Forsskål's (Fl. Aegypt.-Arab.: 179. 1775) later use of the name Ficus serrata. Forsskål misapplied Linnaeus's American F. serrata to a morphologically similar African-Arabian species, now recognized as F. exasperata Vahl (l.c.: 197), as discussed by Friis (in Kew Bull. 38: 460. 1983), Hepper & Friis (Pl. Pehr Forsskål's Fl. Aegypt.-Arab.: 202. 1994), and Friis & Thulin (in Sci. Danica, Ser. B, Biol. 12: 22. 2025). Consequently, “F. serrata Forssk.” should be regarded as F. serrata sensu Forsskål non L., which lacks nomenclatural status, despite the continued treatment of Forsskål's usage as a later homonym in some global databases (e.g., World Flora Online, Plants of the World Online, and Tropicos; all accessed 14 Oct 2025). During a recent systematic review of Neotropical Ficus, we found no original material unambiguously linked to Linnaeus's concept of F. serrata or F. maculata in any Linnaean or Linnaean-linked herbaria. Stearn (l.c.: 130) suggested that discrepancies between early illustrations, like that of Plumier, and modern specimens might reflect incomplete botanical exploration rather than artistic inaccuracy, but this argument does not apply to the well-studied flora of Haiti. Moreover, no herbarium specimen survives in Paris to corroborate the taxonomic identity of the plant depicted on Plumier's drawing. Since its publication, no Antillean or continental American Ficus with serrate leaves consistent with Linnaeus's diagnosis has been collected or described in the scientific literature, reinforcing the status of both F. serrata and F. maculata as nomina dubia. A single specimen in Geneva's Pre-Linnaean herbarium (i.e., G-PREL barcode G00808713), labeled “Ficus maculata” in Johannes Burman's hand, bears one serrated leaf of uncertain provenance, but this does not qualify as original material (M. Callmander, pers. comm.). Under ICN Art. 7.1, 7.2, 8.1, and 9.3 (Turland & al. in Regnum Veg. 162. 2025), lectotypification would be required to fix the application of both names. The only available elements include (1) Burman's proof plate sent to Linnaeus, which may or may not survive on the wall of Linnaeus's study in Hammarby, Sweden; (2) Burman's published plate; and possibly (3) Claude Aubriet's tracing made for Herman Boerhaave in 1733 (from which Burman's published plate was prepared) to which Linnaeus had access during his stay in Holland in the mid-1730s (Moroni & al. in Taxon 67: 1202–1208. 2018) but he did not make any direct references to it in his own annotated copy of the first edition of the Genera plantarum (Linnaeus, 1737), which includes numerous notes on Plumier's illustrations in Boerhaave's collection (Polhill & Stearn in Taxon 25: 323–325. 1976). Typifying either F. serrata or F. maculata with Plumier's illustration, however, risks attaching these names to an ambiguous element most likely consisting of more than one taxon, possibly even outside the Moraceae family. Additional epitypification using a modern herbarium specimen will also not bring any viable solution, as no known Neotropical fig taxa match Linnaeus's descriptions. Conspicuously spotted figs, however, are common in several Antillean species, such as Ficus americana Aubl. (l.c.: 952), F. citrifolia Mill. (Gard. Dict., ed. 8: Ficus No. 10. 1768), and F. trigonata L. (Pl. Surin.: 17. 1775). Spotted figs also occur in continental American species, including F. pertusa L. f. (l.c.), F. santanderana Dugand (in Caldasia 1(4): 65. 1942), F. yoponensis Desv. (in Ann. Sci. Nat., Bot., sér. 2, 18: 310. 1842), among others, but these taxa do not occur in Haiti, where Plumier most likely made his drawing. Another complication with epitypifying F. serrata or F. maculata is that these names will likely have priority over any other subsequent species names in Ficus because of their relatively early publication dates (i.e., 1759 and 1763). Thus, if the epitype were to represent a taxon described after 1763, which is highly likely as only seven Ficus names were published before that date, it will have to be renamed as either F. serrata or F. maculata. This would be nomenclaturally disruptive if it represents a common species. To avoid future instability, we propose that Ficus serrata L. and its synonym F. maculata L. be formally treated as nomina utique rejicienda under Art. 56. If this proposal is not accepted, a later author might feel compelled to designate an obligate lectotype for these taxa, thereby fixing both names to ambiguous or even conflicting elements. Such a course would perpetuate the current uncertainty surrounding their application and could also risk their misapplication to unrelated taxa in the Neotropics or beyond. This would generate considerable confusion in the taxonomic, floristic, and phylogenetic literature on Ficus and ultimately compromise nomenclatural stability. The authors express their gratitude to Ib Friis (Natural History Museum of Denmark, Copenhagen) for his expert guidance in resolving the nomenclatural status of F. serrata sensu Forsskål. We also wish to thank John Wiersema (Smithsonian Institution, Washington, D.C., U.S.A.) and Melanie Schori (United States Department of Agriculture, Beltsville, Maryland, U.S.A.), whose insightful advice and critical comments have materially strengthened this proposal.