taxonID	type	description	language	source
3B718783A86BFF97FF01FF3342E4FBCD.taxon	materials_examined	TYPE: — PERU. Loreto: Maynas Province, Las Amazonas, Quebrada Sucusari, Explor Napo Camp, 13 April 1991, 03 ° 20 ' S 072 ° 55 ' W, R. Vasquez & N. Jaramillo 15957 (holotype: P- 06863731!; isotypes: B- 101298984!, MO- 2025042!)	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A86BFF97FF01FF3342E4FBCD.taxon	diagnosis	Diagnosis: — F. sytsmae is morphologically similar to F. trachelosyce, sharing a characteristic infundibuliform ostiole. F. sytsmae resembles F. trachelosyce in having a globose receptacle, but it can be readily distinguished from the latter by its leaf blade apex (caudate vs. acuminate), leaf blade base (truncate-cordate vs. acute), longer laminas (10 – 24 cm vs. 8 – 15 cm), longer petioles (3.5 – 13 cm vs. 2 – 3.5 cm), longer stipules (10 – 17 mm vs. 6 – 10 mm), larger fig diameter (12 – 20 mm vs. 10 – 12 mm), and longer ostiole length (5 – 7 mm vs. 2 – 2.5 mm) (Figs. 2, 4).	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A86BFF97FF01FF3342E4FBCD.taxon	description	Description: — Tree 15 – 30 m tall or hemiepiphytic. Leafy twigs 0.5 – 1 cm thick, glabrous; periderm persistent. Terminal stipule 1 – 2 cm long, conical, brown at maturity, glabrous, caducous. Leaf blade (sub) coriaceous, oblongelliptic to elliptic, 10 – 24 x 5 – 13 cm, base truncate to cordate-rounded, apex caudate; both surfaces glabrous; midrib reaching the apex of the lamina; lateral veins brochidodromous, lateral veins 7 – 10 pairs, the first pair of lateral veins diverging from the midrib at an 40 – 45 ° angle, acropetiolar waxy spot at the base of the midrib beneath, not evident when dried; tertiary venation reticulate; petiole 3.5 – 13 cm long, ca. 2 mm thick, glabrous, the epidermis persistent. Figs axillary, solitary or in pairs, green; peduncle 0.8 – 1.3 cm long; epibracts 2, 1 mm long, persistent, the apex rounded, glabrous, brownish in herbarium specimens; receptacle rounded or globose, smooth or wrinkled when dried, 12 – 20 mm in diameter when dry, glabrescent to glabrous, brownish, green at maturity; ostiole infundibuliform, straight, 5 mm long, 5 – 7 mm in diameter. Flowers not seen. Fruit an achene (Fig. 1).	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A86BFF97FF01FF3342E4FBCD.taxon	distribution	Distribution, habitat, and phenology: — Currently, Ficus sytsmae is known to be distributed on the western slopes of the Andes in northeastern Peru (Loreto), northwestern Brazil (Acre), and southeastern Ecuador (Napo), inhabiting lowland primary forests that grow on clay and poorly drained sandy soils at elevations of 140 – 450 m. Populations of Ficus sytsmae have been collected in a fertile state in April, September, and December. Due to the asynchronous flowering of the genus Ficus (Milton 1991; Milton et al. 1982; Windsor et al. 1989), it is possible that other populations may be found in bloom throughout the year. Conservation status: — Ficus sytsmae is assessed as Critically Endangered (EN) based on criterion D, due to the extremely small number of known mature individuals (three) across three widely separated localities in Peru, Brazil, and Ecuador. The Area of Occupancy (AOO) is estimated at <10 km ² and the species is not currently known to occur in any protected areas. Although the Extent of Occurrence (EOO) is large (~ 240,000 km ²), F. sytsmae is known from only three herbarium specimens, and no subpopulations have been documented. Projected threats such as agricultural expansion, logging, and infrastructure development may further reduce habitat quality and population viability. This is a precautionary but justified assignment based on currently available data. Additional fieldwork may result in reclassification under a different category as more data become available. Eponymy: — Dedicated to Kenneth J. Sytsma, North American botanist and professor at the University of Wisconsin-Madison, who contributed to our understanding of Urticalean Rosids and set the stage for diversification studies in the big genus Ficus.	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A86BFF97FF01FF3342E4FBCD.taxon	description	Additional collections (paratypes): — BRAZIL. Acre, Mun. Marechal Thaumaturgo: Rio Juruá, righ bank, “ Mato Grosso ”, ca. 8 ° 58 ’ 16.02 ” S, 72 ° 42 ’ 52.2 ” W, 8 December 2000, D. C. Daly, C. I. Salimon, I. S. Rivero & E. C. Oliveira 10495 (NY- 00677509!). ECUADOR. Napo: Estación Biológica Jatun Sacha, 450 m, 01 ° 04 ’ S 77 ° 37 ’ W, 26 September 1988, Palacios 3028 (NY- 04954975!) (Fig. 3).	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A86BFF97FF01FF3342E4FBCD.taxon	discussion	Discussion: — The new taxon is placed within F. sect. Americanae based on the combination of characters described in the introduction. F. sytsmae is morphologically similar to three species currently placed in the F. pertusa complex (Pelissari et al. 2019). Among them, F. trachelosyce is the most morphologically similar to F. sytsmae, sharing an infundibuliform ostiole but differing in several vegetative characters outlined in the diagnosis. F. sytsmae also resembles F. tubulosa and F. schiedeana in having a protruded ostiole; however, it differs from these taxa by having an infundibuliform ostiole (vs. tubular in F. tubulosa and narrowly prominent in F. schiedeana). The same vegetative characters that distinguish F. sytsmae from F. trachelosyce are also observed in comparisons with the other two species (Table 1). The primary difference in the size and shape of the fig between the new taxon and the other three species suggests that populations of F. sytsmae might be reproductively isolated and, therefore, have a different pollinator. However, this will only be tested through genomic sequencing and field observations of pollination biology. The main author is currently awaiting raw reads from these species and additional taxa from F. sect. Americanae to reconstruct the evolutionary history of this clade. However, characterization of the pollinators in this group has proven elusive and requires further attention, particularly with regard to possible cases of hybridization in this morphologically plastic complex. Vegetatively, the leaves of F. sytsmae are larger and more variable between populations but generally tend to be oblong, compared to those of F. trachelosyce, F. tubulosa, and F. schiedeana which are smaller and usually elliptic. Given the characteristic plasticity of vegetative traits in the F. pertusa complex, we conducted a morphometric analysis of the leaves and figs of the new taxon, along with three of the most similar species. Results are summarized in Figure 4. Leaf shape morphometrics (Figure 4 A – D): — Principal Component Analisis (PCA) reveals that variation in PC 1 (49.4 %) primarily differentiates overall leaf shape along a continuum from obovate to narrowly lanceolate (Fig. 4 B). Meanwhile, PC 2 (34.5 %) accounts for variations in apex and base morphology, ranging from caudate-rounded to long acuminate-truncate (Fig. 4 B). F. sytsmae is distinctly segregated from the other taxa, occupying the upper panel and characterized by broadly oblong leaves with a caudate apex and a cordate-rounded base. A similar but less pronounced pattern is observed in F. schiedeana and F. tubulosa, with their morphospaces predominantly distributed within the upper right and lower left panels, respectively, albeit with some marginal overlap between the two. In contrast, Ficus trachelosyce is not easily distinguishable based solely on leaf shape, displaying substantial overlap with both F. schiedeana and F. tubulosa (Fig. 4 A). While F. sytsmae is largely well segregated, it exhibits marginal overlap with F. tubulosa, with which it shared an overall elliptic leaf shape. However, F. sytsmae can be distinguished by its caudate apex (vs. acuminate in F. tubulosa) and cordate to rounded base (vs. acute in F. tubulosa) (Fig. 4 C). Notable, leaf dimensions (length and width) in F. sytsmae are significantly different from all other species, including F. tubulosa (p-value ≈ 0) (Fig. 4 D, Appendix 1).	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A86BFF97FF01FF3342E4FBCD.taxon	description	Fig shape morphometrics (Figure 4 E – H): — Variation in PC 1 (78.1 %) explains a greater proportion of morphological divergence compared to leaf shape, indicating that fig morphology exhibits stronger interspecific differentiation. Specifically, PC 1 captures variation from tubular ostioles to infundibuliform ones (Fig. 4 F). Meanwhile, PC 2 (10.3 %) describes variation from obovoid figs to globose forms (Fig. 4 F). Ficus sytsmae is well separated from F. tubulosa but remains embedded within the F. trachelosyce morphospace, with partial overlap with F. schiedeana (Fig. 4 E). Ficus trachelosyce and F. tubulosa are overall well differentiated, occupying distinct morphospaces predominantly in the upper right and left quadrants, respectively. In contrast, F. schiedeana is challenging to distinguish based solely on fig shape. While the majority of its morphospace clusters in the lower left quadrant, characterized by globose figs with tubular ostioles, it exhibits substantial variation, resulting in partial overlap with the other three species. The fig of F. sytsmae is most similar to that of F. trachelosyce, sharing an infundibuliform ostiole. However, it can be distinguished by its significantly larger fig dimensions (length and width) than the other three species, including F. trachelosyce (p-value <0.05) (Fig. 4 H, Appendix 1).	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
3B718783A860FF95FF01FB49454FFAED.taxon	materials_examined	TYPE: — MEXICO. Veracruz: near Papantla, Hda. La Laguna, 1828 – 1829, C. J. W. Schiede & F. Deppe 1116 (lectotype, designated by Berg & DeWolf [1975: 258]: U 0004686 [digital image!]; isolectotypes: K 000693798!, B 100347087!) Following Mexico’s independence from the Spanish Crown, Count Albert von Sack, Chamberlain to the King of Prussia, decided to travel to Mexico with the intention of acquiring material from the country to enrich the newly established Berlin University. Ferdinand Deppe, an assistant horticulturist at the royal gardens of Charlottenburg, was chosen as a collector of animals and plants and departed in 1824. After parting ways with the Count in 1825, Deppe requested another travel companion, C. J. W. Schiede from Göttingen, who eventually joined him in 1828. Together, they collected approximately 16,296 plant specimens between 1828 and 1829. The material collected up to May 1829 by Deppe and Schiede was acquired by the museum in Berlin (Stresemann 1954; Ricketson & Pipoly 2003). This batch of botanical collections ultimately came under the supervision of Diederich Franz Leonhard von Schlechtendal (elder), who, in collaboration with L. K. A. von Chamisso, published Plantarum Mexicanarum (Schlechtendal & Chamisso 1831), a work cataloging newly described and revised taxa based on Schiede and Deppe’s collections from their 1828 – 1829 expedition to Mexico. This publication appeared in the botanical journal Linnaea. In this work, Schlechtendal and Chamisso identified specimen no. 1116 as Ficus prinoides Willdenow (1806: 1149). In 1847, the Dutch botanist F. A. W. Miquel described Urostigma schiedeanum Miquel (1847: 539), based on C. J. W. Schiede & F. Deppe 1116. In the protologue for U. schiedeanum, Miquel (1847) explicitly stated that the type material corresponded to that examined by Schlechtendal and Chamisso, referencing it as " Ficus prinoides, Schlechtend. ad pl. Schied. et Deppe in Linnaea VI, p. 357, haud Willd., " with “ haud ” indicating that he was not referring to Ficus prinoides Willd. Many years later, in 1867, Miquel transferred all Urostigma species to Ficus while maintaining U. schiedeanum as a synonym of Ficus ligustrina Kunth & Bouché (1847: 240), a taxon based on Venezuelan collections by Johann Wilhelm Karl Moritz, alongside Ficus prinoides Cham. & Schltdl. Miquel’s attribution of F. prinoides to Chamisso and Schlechtendal likely contributed to subsequent misattributions by various institutions, such as IPNI (The International Plant Names Index; http: // ipni. org /), Missouri Botanical Garden’s Tropicos database (http: // www. tropicos. org /), and World Flora Online (https: // www. worldfloraonline. org /), which have treated the name as an illegitimate homonym. However, Schlechtendal and Chamisso (1831) explicitly cited Willdenow as the original author and thus never intended to publish a new taxon under this name (Heuchert et al. 2017). Following Miquel’s synonymization of U. schiedeanum with F. ligustrina, subsequent taxonomic treatments have not recognized the former as a distinct species. Standley (1917) was the first to include U. schiedeanum as a synonym of Ficus padifolia Kunth (1817: 47). Later, Berg & DeWolf (1975) listed U. schiedeanum Miq. and F. schiediana (Miq.) Miq. as synonyms of F. pertusa. Notably, Miquel (1867) never formally established the combination Ficus schiedeana (see p. 307 under Index generum, specierum et synonymorum). However, according to Articles 41.5 and 41.8 of the International Code of Nomenclature for Algae, Fungi, and Plants (ICN; Turland et al. 2018), Berg & DeWolf (1975) inadvertently made the valid combination Ficus schiedeana (in their work as “ schiediana ”) by explicitly indicating its basionym and providing a complete and direct reference to its author, place of valid publication, page number, and date. Despite this valid publication, F. schiedeana has not been subsequently adopted in the literature. Instead, U. schiedeanum has continued to be treated as a synonym of F. pertusa in Berg & Simonis (2000), Berg & Villavicencio (2003, 2004), and Berg (2012). However, a morphological discrepancy exists between the ostiole descriptions in the protologues of these taxa. Linnaeus filius (1782) described the ostiole of F. pertusa as “ umbilicatis, ” whereas Miquel (1847) characterized the receptacle of U. schiedeanum as “ ore contracto prominulo hiante, ” which translates as “ narrow and prominent ostiole. ” Upon reviewing herbarium specimens and photographic records from iNaturalist, we observed a continuous morphological gradient among certain individuals identified as F. pertusa, some of which exhibit ostiole characteristics aligning with those of F. schiedeana. Thus, we here propose the reinstatement of F. schiedeana (Miq.) Berg & DeWolf as a distinct biological entity. This species is currently known only from Mexico, but its geographic range could extend to Central and South America. Specimens of F. schiedeana examined: — MEXICO. Tenejapa, 1000 m, 1 November 1972, D. E. Breedlove 29314 (CAS 504903 [digital image!]; Chiapas: San Juan de Cancuc, 08 January 1991, E. Santiz Cruz 256 (CAS 504934 [digital image!]; Veracruz: Ignacio de la Llave, 24 December 1966, G. Martinez Calderon 1239 (U 0130164 [digital image!]; Veracruz: Coetzala, 18 47 ’ N 96 54 ’ W, 550 – 700 m, 3 July 1982, G. Diggs, M. Nee & G. Schatz 2733 (USF 173955 [digital image!]; Chiapas: Palenque, 300 m, 29 July 1989, L. R. Landrum & S. S. Landrum 6354 (CAS 504928 [digital image!]; Chiapas: San Fernando, 707 m, 471230.1 1879783.5, 22 May 2009, N. Martinez 2642 (CAS 0018621 [digital image!]); Veracruz: Orizaba, 1000 – 1100 m, 18 51 ’ N 97 03 ’ W, 7 December 1981, M. Nee 23840 (U 0039645 [digital image!]); Chiapas: San Juan de Cancuc, 4700 pies, 9 March 1988, E. Sántiz Cruz 499 (CAS 504932 [digital image!]; San Luis Potosi: Tamazunchale, 900 m, 13 February 1982, P. Tenorio & R. Hernández 046 (USCG 20097 [digital image!]; Chiapas: Yajalón, Chulja, 1000 m, 29 September 1982, A. Méndez 4650 (CAS 504927 [digital image!]).	en	Mitidieri-Rivera, Nicole, Damián-Parizaca, Alexander (2025): Ficus sytsmae (Moraceae), a new species from South America and the reinstatement of Ficus schiedeana. Phytotaxa 708 (2): 167-181, DOI: 10.11646/phytotaxa.708.2.4, URL: https://doi.org/10.11646/phytotaxa.708.2.4
