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Heisteria austroecuadorica Cornejo, Homeier & C. Ulloa sp. nov. (Fig. 1). Diagnosis: — Species nova affinis Heisteria asplundii, a qua folia angusta, stamina decandra, calycem fructuum basin non occultans differt. Type: — ECUADOR. Zamora Chinchipe: area of the Reserva San Francisco, 3°58’S 79°04’W, 1980 m, 6 September 2005 (fl, fr), J. Homeier & S. Hänel 1609 (holotype: GUAY!; isotypes: GOET!, LOJA!, QCNE-195572!). Description: — Tree, to 20 m tall and 30 cm DBH; milky white latex present in stems and branches; branches ca. 2 mm thick, terete, striate, glabrous. Leaves alternate; petiole 7–16 mm long, rugose, light- to dark-brown, channeled above, glabrous; lamina chartaceous, (4.5–)7–19 × 2–6 cm, elliptic-lanceolate to narrowly elliptic or narrowly lanceolate, apex acuminate to long-caudate, drip-tip 7–12 mm long, base cuneate, margins revolute, dull, smooth, and glabrous on both sides, venation brochidodromous, midrib slightly sulcate above, prominent below, secondary nerves 6 to 9 pairs, prominulous, tertiary venation inconspicuous. Inflorescence ramiflorous, a short axillary fascicle, 3 to 15 flowers per fascicle; bracts cucullate-deltoid, 0.5–1 × 0.5–1 mm, the apex obtuse to broadly obtuse; pedicel 5–7 mm long. Calyx patelliform or cup-shaped, 5-lobed, lobes ca. 0.5 mm long, triangular to broadly triangular, acute at apex, green. Petals 5, ca. 3 × 1 mm, free, densely papillose or shortly pilose on the adaxial side, greenish-white. Stamens 10, filaments ligulate, ca. 2 × 0.3 mm, the 5 inner ones ca. 1.8 mm, all free, anthers ca. 0.5 × 0.8 mm. Nectary absent; ovary oblatesubglobose (in vivo), pyriform (in sicco); style ca. 0.5 mm, stigma slightly 3-lobed. Fruiting pedicel 1.5–2.3 cm long, slightly swollen, 2–3 mm distally. Fruiting calyx slightly 5-lobed, 2.2–2.8 cm in diam., reflexed, stiffly chartaceous, orange to red, bright and shiny above, dull below. Drupe barrel-shape, olive green with maroon at apex (in vivo), 1.2–2 × 1–1.2 cm, rugulose, faintly ribbed longitudinally and tuberculate (from endocarp, in sicco), with a clear, deep, circular depression at the apex. Etymology: —The epithet refers to the geographic pattern of distribution at southeastern Ecuador where the species occurs. Habitat and distribution: — Heisteria austroecuadorica is known from the San Francisco reserve, a forest remnant close to the settlement of El Líbano north of the Podocarpus National Park, and the Jamboe river basin in southeastern Ecuador, between 1000 to 2050 masl; all three localities are on the Loja-Zamora road. The area is characterized by a rugged terrain and small-scale structural and floristic vegetation changes, which result mainly from the decreasing soil fertility along ravine-slope-ridge gradients (Werner & Homeier 2015). The newly described species is restricted to the ravines and lower slope positions with more fertile soils within the species-rich evergreen lower montane forests below 2050 masl, that reach up to 30 m height (Homeier et al. 2008, 2010). In the ravines and on the lower slopes of this specific forest type, Heisteria austroecuadorica is accompanied by several tree species that have been described from this zone in the last years, stressing the biological importance of this area. Some examples are: Byrsonima homeieri Anderson (2007: 97), Critoniopsis zamorensis Haro-Carrión & Robinson (2008: 17), Meliosma longepedicellata Cornejo (2009: 93), Magnolia zamorana Arroyo et al. (2013: 507), Meriania franciscana Ulloa Ulloa & Homeier (2008: 383), Naucleopsis francisci Berg & Homeier (2010: 197), Ocotea homeieri van der Werff (2020: 210) and Quiina zamorensis Schneider & Zizka (2012: 265). All of these aforementioned species have a narrow pattern of distribution that is restricted to the particularly diverse Amotape-Huancabamba zone (Weigend 2002, 2004), south of the Girón-Paute dry valley in southern Ecuador (Jørgensen et al. 1999). With its relatively high-density wood (0.72 g /cm 3), large fruits and slow diameter growth rates, Heisteria austroecuadorica combines characteristics typical of a late successional tree species. Phenology: —Flowering was observed in June, September, October and December; fruiting material was collected in April, September and October. There are no existing observations on flower pollination or fruit dispersal. Conservation status: —This species is uncommon in the Reserva San Francisco: Homeier has only found a total of six trees (out of almost 4000) within three recently established one-hectare plots. At a second location, outside the Reserve, in El Líbano, only one tree has been seen. A third locality is represented by one collection only in the Jamboe river basin just further down at slightly lower elevation. Based on the very small area of occurrence (12 km 2), an extent of occurrence of 10.3 km 2, the existence of only three localities, only one of which is protected within the private Reserva San Francisco, we assign a provisional category of Endangered (EN B2a(iv)), following the IUCN (2022) criteria. Paratypes: — Ecuador: Zamora Chinchipe: Reserva San Francisco, road Loja-Zamora, ca. 30 km from Loja, 3°58’S 79°04’W, 1800–2050 m, June 2005 (fl.), O. Cabrera & N. León 869 (HUTPL, fragm. in GUAY); same location, 1840 m, 20 October 2001 (fr.), J. Homeier 1025 (LOJA, QCNE, MO, GOET); same location, 1910 m, 11 September 2003 (sterile), J. Homeier & W. Quizhpe 1261 (GOET, MO, QCNE); same location, 1910 m, 20 October 2013 (fl.), J. Homeier 5098 (HUTPL, QCA, GOET); same location, 2040 m, 5 December 2016 (fl.), J. Homeier & J. Peña 5820 (HUTPL, QCA, GOET). Road Loja-Zamora, El Líbano, 4°5’S 78°59’W, 1720 m, April 2011 (fr.), J. Homeier, F.A. Werner & D. Cárate 4713 (LOJA, QCA, QCNE, GOET). Cuenca del río Jamboe, 4°08’S 78°56’W, 1000 m, September 1999 (fl, fr), W. Palacios 14881 (MO). Discussion: —The fruits of Heisteria austroecuadorica are characterized by the apex presenting a circular depression, a key character in Sleumer’s treatment (1984) that is only found in three other species in tropical America: H. amphoricarpa Sleumer (1984: 49), H. asplundii, and H. skutchii Sleumer (1984: 50). Heisteria amphoricarpa, a Brazilian species, has a unique urn-shaped calyx enveloping most of the drupe and flowers with five stamens, while H. skutchii, a Costa Rican species with 10 stamens, has much wider, patent fruiting calyx (over 30 mm diameter). In Ecuador, the fruits of Heisteria austroecuadorica are relatively large with a depressed apex like those of H. asplundii, a tree endemic to northwestern Ecuador. However, H. austroecuadorica immediately differs from H. asplundii by its narrower leaves, with blades elliptic-lanceolate to narrowly elliptic or narrowly lanceolate, 2–6 cm wide (versus leaves always with broadly elliptic blades, 10–15 cm wide in H. asplundii), the flowers with 10 stamens (vs. 5 stamens), the smaller fruiting calyx, up to 2.8 cm and covering just the base of the drupe (vs. up to 4.8 cm and covering the basal 1/3 of the drupe); additionally, these two species are widely separated on opposite sides of the Andes mountains.

Cornejo, Xavier, Homeier, Jürgen, Ulloa, Carmen Ulloa (2023): Heisteria austroecuadorica (Erythropalaceae): A new species from southeastern Ecuador. Phytotaxa 599 (3): 201-206, DOI: 10.11646/phytotaxa.599.3.7, URL: http://dx.doi.org/10.11646/phytotaxa.599.3.7

Following recent mimosoid phylogenetic and phylogenomic studies demonstrating the non-monophyly of the genus Albizia, we present a new molecular phylogeny focused on the neotropical species in the genus, with much denser taxon sampling than previous studies. Our aims were to test the monophyly of the neotropical section Arthrosamanea, resolve species relationships, and gain insights into the evolution of fruit morphology. We perform a Bayesian phylogenetic analysis of sequences of nuclear internal and external transcribed spacer regions and trace the evolution of fruit dehiscence and lomentiform pods. Our results find further support for the non-monophyly of the genus Albizia, and confirm the previously proposed segregation of Hesperalbizia, Hydrochorea, Balizia and Pseudosamanea. All species that were sampled from section Arthrosamanea form a clade that is sister to a clade composed of Jupunba, Punjuba, Balizia and Hydrochorea. We find that lomentiform fruits are independently derived from indehiscent septate fruits in both Hydrochorea and section Arthrosamanea. Our results show that morphological adaptations to hydrochory, associated with shifts into seasonally flooded habitats, have occurred several times independently in different geographic areas and different lineages within the ingoid clade. This suggests that environmental conditions have likely played a key role in the evolution of fruit types in Albizia and related genera. We resurrect the name Pseudalbizzia to accommodate the species of section Arthrosamanea, except for two species that were not sampled here but have been shown in other studies to be more closely related to other ingoid genera and we restrict the name Albizia s.s. to the species from Africa, Madagascar, Asia, Australia, and the Pacific. Twenty-one new nomenclatural combinations in Pseudalbizzia are proposed, including 16 species and 5 infraspecific varietal names. In addition to the type species Pseudalbizzia berteroana, the genus has 17 species distributed across tropical regions of the Americas, including the Caribbean. Finally, a new infrageneric classification into five sections is proposed and a distribution map of the species of Pseudalbizzia is presented.

Eschweilera brevipetiolata S.A. Mori & Cornejo, sp. nov. (Fig. 1) This new species of Eschweilera is similar to E. integricalyx S.A. Mori but differs by the cauliflorous (vs. ramiflorous) inflorescences, longer pedicels, 1–1.3 cm (vs. ca. 0.5 cm long), calyces with triangular and well-defined sepals (vs. calyces nearly entire), leaves sessile to subsessile, the petioles absent to ca. 1 cm long (vs. petioles 1.5–2.5 cm long), and shorter height of the trees, 6 to 16 m tall (vs. canopy trees). Type: — COLOMBIA. Valle del Cauca: Bajo Calima; Consesión Pulpapel / Buenaventura, carretera al Dindo, 3°55’ N 77°00’ W, ca. 100 m, 25 Sep 1986 (fl), M. Monsalve 1169 (holotype: CUVC-25917!; isotypes: MO-3701257!, MO-3701347!, NY-00853327!). Tree to 16 m tall, the trunk and bark unknown. Stems glabrous, sparsely lenticellate, cracked or shallowly fissured when dry. Leaves: sessile to very shortly petiolate, the petioles shallowly channeled adaxially, rounded abaxially, hemispherical in cross section, up to ca. 10 mm long, 7–8 mm diam., with many unicellular, papillose, light brown trichomes; blades oblanceolate to oblanceolate-elliptic, 50–75 × 18–30 cm, chartaceous, with abundant reddishbrown punctations abaxially, the base obtuse to rounded or truncate, the margins entire, the apex acuminate; venation eucamptodromous towards base, brochidodromous towards apex, without conspicuous marginal veins, the midrib prominent to flat toward base adaxially, markedly prominent abaxially, densely papillose at least on the lower half of its length, the secondary veins in 20–33 pairs, slightly prominent adaxially, salient abaxially, the marginal vein conspicuous, the tertiary veins reticulate. Inflorescences cauliflorous unbranched to 1-branched, the rachis 1.5–7 cm long, rather thin, ± straight to suberect, glabrous or glabrescent, lenticellate with age; pedicels 10–14 × ca. 1.5–2 mm, truncate to articulation, glabrous or glabrescent, drying blackish. Flowers ca. 3.5 cm diam.; hypanthium glabrous, sparsely lenticellate; calyx with 6 lobes, the lobes triangular to deltoid, 2–3 × 3–4 mm, ascendant at anthesis, smooth (dry) abaxially, the bases valvate, arising from fused calyx rim; petals ± oblong, 2–2.5 × ca. 1 cm, pink; androecial hood with four coils, ca. 1.5 cm wide, pink; vestigial stamens on outside of coils; staminal ring with ca. 300 stamens, the filaments ca. 1.5 mm long, clavate, the anthers 0.5–0.7 mm long; ovary 2-locular, with ca. 10 ovules per locule, the summit widely obconical, the style not distinguished from ovary summit. Fruits depressed globose, ca. 5 × 4 cm, the calyx lobes persistent in young fruits, woody, the calycine ring not conspicuous, the supra-calycine zone erect, the infra-calycine zone rounded to pedicel, the operculum convex, somewhat prolonged to umbonate at apex, the pericarp 2–3 mm thick, brown when dry. Seeds usually wedge-shaped in cross section, the outer surface hemispherical, the two sides flat, ca. 4 × 3 cm, the veins visible, plain, the seed coat smooth, the arils unknown. Discussion: — Eschweilera brevipetiolata resembles E. integricalyx S.A. Mori, another species endemic to the Choco in Colombia, but it differs from the latter by the cauliflorous (vs. ramiflorous) arrangement of inflorescences, pedicels 1–1.3 cm long (vs. ca. 0.5 cm long), calyces with triangular and well-defined sepals (vs. calyces nearly entire), leaves sessile to subsessile, the petioles absent to ca. 1 cm long (vs. petioles 1.5–2.5 cm long), and shorter height of the trees, 6 to 16 m tall (vs. canopy trees). Due to the large blades, Eschweilera brevipetiolata may resemble E. sclerophylla Cuatrec. (1951: 92), another species endemic to western Colombia, but the latter species differs from it by the petiolate (1–1.3 cm long vs. sessile or nearly sessile) leaves, axillary or terminal (vs. cauliflorous) inflorescences, large and yellow (vs. smaller and pink) flowers, and truncate (vs. tapered) hypanthia. Etymology: —The epithet refers to the very short petiole that vegetatively characterizes this taxonomic novelty. Common names: —Guasco (Monsalve 1349). Habitat and distribution: —An understory to medium-sized tree of mature, wet to pluvial forest in non-flooded areas characterized by palms, such as Oenocarpus bataua Mart. (1823: 23) and Socratea exorrhiza (Mart.) H. Wendl. (1860: 103). It is known only from western Colombia in the Bajo Calima area, department of Valle del Cauca, from 50–100 m elevation. Phenology: — Flowers have been collected in late September, and may be present until early October, and fruits have been collected in Mar, Apr, Jul, Ago, and Nov. Conservation status: — Eschweilera brevipetiolata occurs in the Bajo Calima area and is represented by a small population with several mature individuals. It is suggested that this new species be assigned the IUCN conservation status of near threatened (NT) (IUCN, 2022). Paratypes: — COLOMBIA. Valle del Cauca: Bajo Calima Concession, ca. 16 km NW Buenaventura, at end of Gasolina Rd. Juanchaco área, BV-82, wet tropical forest on 20–45° slopes, yellow clay over alluvial, 77°10’W 3°50’N, 50 m, 28 May 1987 (st), D. Faber-Langendoen et al. 712 (CUVC!), 727 (NY!); 77°02’W 3°55’N, 50 m, 11 Jul 1984 (st), A . Gentry et al. 47914 (CUVC!, NY!); carretera Hans, 100 m, 2 Oct 1987 (fl), M . Monsalve 1910 (CUVC!); carretera La Gasolina, 100 m, 26 Nov 1986 (fr), M . Monsalve 1349 (CUVC!). Dindo area, 76°58’W 3°59’N, 50 m, 20 Jul 1984 (fr), A . Gentry & M. Monsalve 48417 (CUVC!, NY!); 77°02’W 3°59’N, 50 m, 26 Mar 1986 (fr), A . Gentry et al. 53640 (CUVC!, NY!).

Eschweilera silverstonei Cornejo, sp. nov. (Fig. 2) This new species of Eschweilera is similar to E. integrifolia but differs by the more developed, paniculate-pyramidal, multiflorous, terminal inflorescences that are ca. 10 cm long (vs. ramiflorous, mostly unbranched, shortly racemose, pauciflorous, and up to ca. 4.5 cm long), and greenish-yellow flowers (vs. purple to creamish-white). Type: — COLOMBIA. Valle del Cauca: Bajo Calima; Consesión Pulpapel / Buenaventura, 3°55’ N 77°00’ W, ca. 100 m, 9 Aug 1984 (fl), M. Monsalve 214 (holotype: CUVC-19929!; isotype; NY-02152729!). Tree ca. 5 m tall, the trunk and bark unknown. Stems glabrous, sparsely lenticellate. Leaves: shortly petiolate, the petioles shallowly channeled adaxially, rounded abaxially, hemispherical in cross section, 8–13 mm long, ca. 3 mm diam., glabrous or glabrescent; blades oblong-elliptic, 20–25 × 5–7 cm, chartaceous, with abundant brown punctations abaxially, the base obtuse to rounded, the margins entire, the apex acuminate; venation eucamptodromous towards base, brochidodromous towards apex, without conspicuous marginal veins, the midrib flat adaxially, prominent abaxially, glabrous or nearly so, the secondary veins in 13–16 pairs, slightly prominent on both sides, the tertiary veins reticulate. Inflorescences terminal, pyramidal-paniculate, the rachis ca. 4 cm long, ± straight, glabrescent, lenticellate with age; pedicels 5–6 x 2–3 mm, tapered to articulation, lenticellate, puberulous, drying blackish. Flowers ca. 3 cm diam.; hypanthium glabrous, truncate at summit; calyx with 6 lobes, the lobes triangular to deltoid, 3–4 x 2.5–3.5 mm, ascendant at anthesis, smooth (dry) abaxially, the bases barely imbricate, arising from fused calyx rim; petals suborbicular to ± oblong 1.5–2 x 1–1.5 cm, greenish white; androecial hood with three coils, ca. 1.5 cm wide, greenish white; vestigial stamens on outside of coils; staminal ring with ca. 300 stamens, the filaments 0.3–0.5 mm long (dry), clavate, the anthers ca. 0.5 mm long (dry); the ovary summit slightly raised over hypanthium, the style ca. 1 mm long, distinguished from ovary summit, stylar collar present. Fruits, seeds, and arils not known. Discussion: — Eschweilera silverstonei resembles E. integrifolia but differs from the latter species by the more developed, paniculate-pyramidal, terminal, multiflorous inflorescences (bearing over 30 flowers each), that are ca. 10 cm long (vs. ramiflorous, mostly unbranched, shortly racemose, pauciflorous inflorescences (bearing up to 4 flowers each), that are up to 4.5 cm long, and greenish-yellow flowers (vs. purple to creamish-white). This statement is based on the numerous field observations of E. integrifolia in the type locality and several additional places where this species occurs in western Ecuador. Etymology: —The epithet honors Phillip Silverstone-Sopkin (1939–2018), who studied the taxonomy of Amaryllidaceae and was an enthusiastic researcher of the Colombian flora. He was the former Director of the CUVC herbarium at the Universidad del Valle in the city of Cali, Colombia. Common names: —Guascanato (Monsalve 214). Habitat and distribution: — An understory tree of mature, wet to pluvial forest in non-flooded areas. It is known only from western Colombia in the Bajo Calima area, in the department of Valle del Cauca, at ca. 100 m elevation. Phenology: — Flowers have been collected in August. Conservation status: — Eschweilera silverstonei occurs in the Bajo Calima area. Although the status of its populations is not currently known, due to the absence of new collections for over three decades, we suggest that this new species be assigned the IUCN conservation status of near threatened (NT) (IUCN, 2022). Paratypes: —Known only from the type collection., Published as part of Cornejo, Xavier, 2023, Eschweilera brevipetiolata and E. silverstonei: Two new species of Lecythidaceae from western Colombia, pp. 287-292 in Phytotaxa 585 (4) on pages 290-292, DOI: 10.11646/phytotaxa.585.4.5, http://zenodo.org/record/7703867, {"references":["IUCN Standards and Petitions Committee (2022) Guidelines for Using the IUCN Red List Categories and Criteria. Version 15. Prepared by the Standards and Petitions Committee. Available from: http: // www. iucnredlist. org / documents / RedListGuidelines. pdf (accessed 1 July 2022)"]}