Dendropanax umbellatus (Ruiz & Pav.) Macbride (1959: 42)

Fernandez-Hilario, Robin, Villanueva-Espinoza, Rosa & Fack, Vinciane, 2020, Redescription and typification of Dendropanax umbellatus (Araliaceae), Phytotaxa 447 (1), pp. 51-60 : 52-58

Dendropanax umbellatus (Ruiz & Pav.) Macbride (1959: 42) ( Fig. 1–4 )

Basionym: Gilibertia umbellata Ruiz & Pavón (1802: 75)

Wangenheimia umbellata (Ruiz & Pav.) Dietrich (1810: 537)

Ginannia umbellata (Ruiz & Pav.) Dietrich (1817: 483)

Type: – PERU. [Huánuco] Muña, Ruiz & Pavón s.n. (lectotype, designated here, BC872843 [digital image!]; isolectotypes BM001008680 [digital image!], MA814084 [digital image!], MA814085 [digital image!], MA814086 [digital image!], MA814087 [digital image!], MA817192 [digital image!], MA817193 [digital image!], MO1612993 [digital image!]) .

Type: – PERU. Amazonas: Prov. Bongará, Dist. Yambrasbamba, bosques aledaños al río Imaza , 1530 m, 05°39’40.22”S ; 77°58’12.75”W, 11–12 January 2019 (fr.), R. Fernandez, R. Villanueva, W. Chuquitucto & A. Wong 1649 (epitype, designated here, MOL!; isoepitypes BRLU!, CPN!, HOXA!, HSP!, USM!) .

Tree up to 15 m tall; outer bark lenticellate and slightly fissured, whitish-brown to white-colored; inner bark slightly aromatic; terminal twigs, leaves, inflorescences and flowers glabrous. Terminal twigs circular in transversal section, 5–7.5 mm diam., striate longitudinally and gray-colored when dry; intrapetiolar stipules reduced up to 1.5 mm long. Leaves alternate, spirally arranged, sometimes subopposite; petioles terete, (1–) 4.8–12.2 (–21.3) cm long; leaf blades plane, coriaceous, with presence of pellucid dots, 9.5–18.1 (–32) × 5.6–14.2 (–22.5) cm, ovate to elliptic, often slightly asymmetric, apex acuminate to acute, base obtuse to rounded (rounded to cordate in young individuals), margin entire to denticulate (distinctly dentate in young individuals), concolorous, lustrous and dark brown when dry, venation brochidodromous and pinnate, sometimes the distal veins eucamptodromous, the first pair branched and suprabasal, originating up to 7 mm from leaf base, secondary veins 4–8, intersecondary veins present, tertiary veins finely reticulated on both surfaces, midrib and secondary veins impressed on the adaxial surface, midrib and secondary veins prominent on the abaxial surface. Inflorescences a simple umbel, sometimes compound, terminal, erect; main axis in simple umbel 23–40 × 2.5–3 mm (3–5 mm wide in fruits), in compound umbel 42–91 × 3–4 mm; with whorled and triangular bracts near the base, 4–10 mm long; (4–) 6–8 (–12) primary branches, (1.7–) 3–4.5 (–7.2) cm long, umbellately or racemosely arranged, each subtended by a triangular bract 5–10 mm long, branches with 1–4 triangular bracts sparse to opposite; umbels (9–) 21–40-flowered. Flowers actinomorphic, light green; the pedicels (12–) 20–30 × 1.5–2 mm; hypanthium 5–6.5 × 4.5–6 mm, campanulate; calyx lobes reduced, denticulate; petals (6–) 8–9 (–10), 3.5 mm long, 2–2.5 mm wide at base, triangular, apex cucullate; stamens (6–) 8–9 (–10), the filaments 3 mm long, the anthers versatile, ca. 1.5 × 1 mm, elliptic; styles 8-9, 0.5–1 mm long. Fruits spheroid, (10–) 15–18 × (10–) 17–22 mm; the stylar column 1.5–3 mm long; fruiting pedicels 1.7–2.5 × 1.5–2 mm; pyrenes 8, 7–12 mm long.

Distribution and habitat. Dendropanax umbellatus is an endemic tree from the montane forests in the Peruvian Andes (Depts. Amazonas, Cajamarca, Huánuco and Pasco) between 1050 to 2133 m in elevation, growing on slight slopes and riparian zones.

Phenology. Flowering takes place from December to February and fruiting from November to January.

Additional specimens examined: PERU. Amazonas: Prov. Bongará, Dist. Jumbilla, along road Jumbilla- Laguna de Pomacochas , 2133 m, 05°47’39”S, 77°55’57”W, 05 November 2012 (fl. bud), H. van der Werff et al. 25085 ( HOXA!) ; Prov. Bongará, Dist. Yambrasbamba, bosques aledaños al río Imaza , 1530 m, 5°39’40.22”S, 77°58’12.75”W, 11-12 January 2019 (fl.), R. Fernandez et al. 1646 ( BRLU!, CPN!, HOXA!, HSP!, MOL!, USM!), (fr.), R. Fernandez et al. 1647 ( BRLU!, HOXA!, HSP!, MOL!), (fr.), R. Fernandez et al. 1648 ( BRLU!, MOL!, HOXA!, HSP!) ; Prov. Bongará, Dist. Yambrasbamba , bosques adyacentes a los caminos, 1900 m, 05°39’31.11”S, 77°57’28.22”W, 12 January 2019 (fl.), R. Fernandez et al. 1684 ( BRLU!, CPUN!, HOXA!, HSP!, MOL!, UFV!, USM!) ; Prov. Bongará, Dist. Yambrasbamba, Centro de Investigación de la ONG Neotropical Primate Conservation y bosque de la comunidad campesina conocido como “El Toro”, 2000 m, 05°39’17.05”S, 77°54’51.30”W, 26-29 November 2018 (sterile), R. Fernandez & W. Chuquitucto 1744 ( MOL!, UFV!) ; Prov. Bongará, Dist. Yambrasbamba , bosques propiedad de Wilder Chiquitucto, 1530 m, 05°39’40.22”S, 77°58’12.75”W, 01 December 2018 (fl. bud), W. Chuquitucto 1 ( HSP!, MOL!, UFV!, USM!) ; Prov. Bongará, Dist. Valera, inmediaciones de catarata Gocta , 2000 m, 06°01’52.95”S, 77°53’01.42”W, 21-30 January 2014 (sterile), R. Fernandez 528 ( HSP!, MOL!) ; Prov. Bongará, Dist. Valera, Cocachimba cerca a la catarata Gocta , 1600 m, 3 September 2013 (fr.), A. Daza 5706 ( MOL!) ; Prov. Luya, Dist. Carropendo, Anexo Tullanya , 1700-2000 m, 26 November 1996 (fr.), J. Campos et al. 3022 ( MOL!, USM!) . Cajamarca: Prov. San Ignacio, Dist. Huarango , El Convenio, 1100-1400 m, 05°13’S, 78°40’W, 02 July 1996 (sterile), J. Campos & E. Rodríguez 2861 ( USM!) ; Prov. San Ignacio, Dist. San José de Lourdes , selva andina, 1650 m, 05°00’43”S, 78°54’09”W, 14 February 2000 (fl., fr.), J. Campos & R. Vásquez 6383 ( MOL!, USM!) . Pasco: Prov. Oxapampa, Dist. Huancabamba, Localidad Grapanazu , sector San Daniel , inmediaciones de los Fundos ganaderos y de cultivos, 2094 m, 10°26’53”S, 75°27’04”W, 14 July 2004 (fr.), J. Perea et al. 1569 ( HOXA!) ; Prov. Oxapampa, Dist. Huancabamba, Parque Nacional Yanachaga-Chemillén , sector Tunqui , zona de amortiguamiento, 1780 m, 24 October 2007 (fr.), A. Monteagudo et al. 15575 ( MOL!, USM!) ; Prov. Oxapampa, Dist. Palcazu, PN Yanachaga-Chemillén , en las nacientes de las quebradas Cashivo y Palmalanbo y naciente del río Lagarto , 1050 m, 10°11’02”S, 75°26’42”W, 20 September 2010 (fr.), R. Rojas & G. Ortiz 7542 ( HOXA!)

Notes. Ruiz and Pavón (1794: 50) established the illegitimate later homonym genus Gilibertia in their “ Florae Peruvianae et Chilensis Prodromus”, without citing any species. They presented only the generic description and an illustration with floral details. Later, they proposed Gilibertia umbellata , which was described as the only representative of Gilibertia in the third volume of the “Flora Peruviana et Chilensis”. The protologue of G. umbellata mentions Muña as the habitat, and an illustration showing a complete plant illustrated by Isidro Galvez. There are nine specimens of Ruiz & Pavón’s collections in the herbaria BC, BM, MA and MO. The specimen BC872843 contains Pavón’s handwriting with the following annotation “ F. Per. G. nov. ”. It also presents incomplete flowers with a fully developed hypanthium and pedicels in a fragment of inflorescence besides a short twig with an attached leaf. The remaining specimens from BM, MA and MO show just sterile twigs or fragments of inflorescences with flower buds. Based on these evidences, we designate as the lectotype the sheet BC872843 following the Art. 8.1 of the ICN ( Turland et al. 2018). This specimen arrived in Barcelona as a result of sales made by Mariano de la Paz Graells to the Real Academia de Ciencias y Artes de Barcelona in 1836 ( Ibáñez et al. 2006). Furthermore, specimens BM001008680 and MO1612993 came to the corresponding herbaria proceeding from sales made by Pavón to Aylmer Bourke Lambert between 1816 and 1824 ( Rodríguez 1994). After Lambert’s death, these materials that were part of his own collection were distributed to different herbaria ( Miller 1970).

In the protologue of Gilibertia umbellata, Ruiz & Pavón (1794: 50) indicated that its flowers are 7-merous, and even 7–9-merous flowers with long pedicels are included in the iconography of the species (see Fig. 1 ). However, Macbride (1959) mentioned that this species also presented 5-merous flowers. Among the collections examined by Macbride (1959: 42) in his revision of Araliaceae for “Flora of Peru ”, negatives of two type specimens were found. The first one regards to a destroyed specimen of Gilibertia pavonii in the herbarium B [F.M. Neg. 3547, http://emuweb. fieldmuseum.org/web/objects/common/webmedia.php?irn=25112] and the second, to a specimen MA814084 with floral buds of Gilibertia umbellata [F.M.Neg.29224,http://emuweb.fieldmuseum.org/web/objects/common/webmedia. php?irn=1786092]. These may be the reasons why Macbride described D. umbellatus as: “ (…) pedicels 5–7 mm long; calyx 7–9-dentate in type, or 5-dentate (…) ” and “ (…) drupe about 5 mm across, globose-ovoid, little narrowed above, shallowly 5-sulcate (…) ”. Although Macbride correctly verified the number of floral parts in MA814084, he could not recognize the dimensions of the fully developed pedicels and fruits of Dendropanax umbellatus . Due to a misreading by Macbride, Gilibertia pavonii was considered a synonym of Dendropanax umbellatus , and since then has remained a cause of confusion in subsequent works (see Brako & Zarucchi 1993; Frodin & Govaerts 2003; Vásquez et al. 2010).

Among original material of Dendropanax umbellatus , four specimens (BM001008680, MA814084, MA814085 and MA814087) present fragments of inflorescences with floral buds. But the lectotype chosen here (BC872843) has a fragment of inflorescence with fully developed hypanthium in incomplete flowers. Furthermore, the remaining specimens consist of twigs or fragments of sterile twigs. Consequently, for the precise application of the name Dendropanax umbellatus , an epitype is designated according to the Art. 9.9 of the ICN ( Turland et al. 2018) which has fully developed pedicels and fruits in complete inflorescences. As a result of the vegetative variability, the most useful characters to distinguish Dendropanax umbellatus from related species are the reproductive characters, such as the length of the main inflorescence axis and pedicels, and mature fruits size.

Dendropanax umbellatus can be variable in leaf blade shape and size, and leaves close to the inflorescences are elliptic and present short petioles, while remaining leaves are ovate and present long petioles. In young or sterile individuals, blades can show atypical dimensions (up to 32 × 22.5 cm in Fernandez 528 and Fernandez & Chuquitucto 1744). Moreover, young individuals have distinctly dentate margins (e.g.: Fernandez & Chuquitucto 1744, Figure 2B ). Dendropanax umbellatus is similar to the Ecuadorian species Dendropanax macrophyllus Cuatrecasas (1951: 308) , but this species differs from the former by having 14–16 flowers per umbel, 6-merous flowers and pedicels 3–6 mm long. In addition, Dendropanax globosus M.J. Cannon & Cannon (1989: 14) , distributed from Costa Rica to Colombia, is morphologically similar to Dendropanax umbellatus . However, Dendropanax globosus is distinguished by the absence of pellucid dots, pedicels 6–10 mm long, and fruits 4–6 × (6–) 7 mm (see Table 1).

*Described as rachis in the protologue

**Measurements taken from JSTOR Global Plants

Gilibertia pavonii Marchal (1878: 248)

Dendropanax pavonii Pav. ex Decaisne & Planchon (1854: 107) nom. nud.

Aralia umbellata Pav. ex Seemann (1868: 138) nom. nud.

Aralia umbellata Pav. nom. herb.

Type : – PERU. Peruvia, Panao and Chinchao, Ruiz & Pavón s.n. (lectotype, designated here, P02441658 [digital image!]; isolectotypes B destroyed, negative at F ( F 0BN003547) [digital image!], BR0000005630776 [digital image!], HAL0117717 [digital image!], MA814080 [digital image!], MA814081 [digital image!], MA814082 [digital image!], MA814083 [digital image!], MA817501 [digital image!], MA817502 [digital image!], MA817503 [digital image!], MA817163 [digital image!], P02441657 [digital image!], P02441659 [digital image!]) .

Notes. When “ Esquisse d´une monographie des Araliacées ” was published by Decaisne & Planchon (1854), they stated that there was a difference among what they considered Dendropanax , a group of plants with 5-merous flowers “ (…) le calice à 5 dents aiguës (…) ” and the genus Gilibertia which was characterized by having 7-merous flowers “ (…) à pétales assez nombreux (généralement 7) (…) ”. In the same publication, Decaisne & Planchon displayed a list of Dendropanax species including Dendropanax pavonii Pavón ex Decaisne & Planchon (1854: 107) , which was alluded to a Pavón collection (“ Aralia umbellata Herb. Pav. ”). This clarify that the concept of Decaisne & Planchon corresponding to Dendropanax pavonii was different from Gilibertia umbellata . Additionally, Decaisne & Planchon just mentioned a collection of Pavón without providing a description, for this reason, the name Dendropanax pavonii is considered a nomen nudum which does not comply with the Art. 38.1 of the ICN ( Turland et al. 2018). Later, Marchal (1878) in his treatment of Gilibertia for “ Flora brasiliensis ” provided a diagnosis, and consequently, the species was validly published under the name Gilibertia pavonii Marchal.

Currently, an unpublished iconography illustrated by José Brunete from the expedition of Ruiz and Pavón, which contains the following annotation “ Aralia umbellata ” (available on JSTOR), along with eight specimens, is housed at MA. However, we designate the specimen P 02441658 as lectotype, conforming to the Art. 8.1 of the ICN ( Turland et al. 2018) because it bears an original label (“ Peruvia . Herb. Pavon. ”). This label refers to the private collections of Pavón acquired by Edmond Boissier in 1841 ( Rodríguez 1994, 1995), which was subsequently donated to the Muséum National d’Histoire Naturelle in 1845 ( O. Poncy pers. comm.). This collection is probably the original material revised by Decaisne & Planchon (1854) for their work “Esquisse d´une monographie des Araliacées”.

Marchal (1878) validly published Gilibertia pavonii with three syntypes: Poeppig 1524 (missing), Ruiz s.n. ( BR 0000005630776, HAL 0117717, P 02441657), and Sello s.n. ( BR 0000005793877). The material cited by Marchal as “ Ruiz s.n. ” matches the collection purchased by Aylmer Bourke Lambert between 1816 and 1824 ( Rodríguez 1994). We conclude that because these three specimens have a label stating “ Ruiz legit ex herbario Lamberti ”. In addition, the specimen BR 0000005630776 bears the label stating “ E. Marchal - Monogr. Araliac. Brasil ” indicating to be part of the material revised for his “ Flora brasiliensis ”.

The type collection of Gilibertia pavonii differs from Dendropanax umbellatus by its elliptic leaves 8.4–13.5 × 4.6–7.9 cm, compound umbel, pedicels 5–7 mm long in fruit, and shallowly 5-sulcate drupes 5 mm across with five pyrenes. These characters are similar to those present in Dendropanax arboreus ( Linnaeus 1759: 967) Decaisne & Planchon (1854: 107) , a widespread species distributed in Mesoamerica and northern South America. Accordingly, Gilbertia pavonii could be treated as synonymous under Dendropanax arboreus , as suggested by other authors (see Macbribe 1959; Frodin’s annotation on Sello s.n.).

Figures

FIGURE 1. Dendropanax umbellatus. Flora Peruviana et Chilensis, Icon. CCCXII (Ruiz & Pavón 1802).

FIGURE 2. Vegetative variability of Dendropanax umbellatus. A. Terminal twig. B–C. Leaf with cordate to rounded base and dentate margin on young individual. D. Leaf with obtuse base and denticulate margin on mature individual. E. Petioles of varying lengths. F. Leaf with an obtuse base on mature individuals. G–H. Leaf with a denticulate margin on mature individuals. Photographs by Robin Fernandez, based on Fernandez & Chuquitucto 1744 (A–C and E), Fernandez et al. 1648 (D and H) and Chuquitucto 1 (F–G).

FIGURE 3. Variability of the inflorescences of Dendropanax umbellatus. A. Simple umbel with floral buds. B. Simple umbel at anthesis. C. Compound umbel with branches racemosely arranged. D. Compound umbel with branches umbellately and racemosely arranged. Photographs by Robin Fernandez, based on Fernandez et al. 1646 (A and B), Fernandez et al. 1684 (C) and Fernandez et al. 1648 (D).

FIGURE 4. Dendropanax umbellatus. A. Bracts on main axis. B. Flowers. C. Immature fruits in compound umbel. D. Mature fruits in simple umbel. Photographs by Robin Fernandez, based on Fernandez et al. 1684 (A), Fernandez et al. 1646 (B), Fernandez et al. 1648 (C) and Fernandez et al. 1647 (D).