Cremastosperma pedunculatum (Diels) R.E.Fr.

Trees or shrubs (0.5–)1.5–20 m tall; young twigs and petioles glabrous to densely covered with appressed or erect, simple, whitish to golden, up to 1 mm long hairs. Leaves distichous, simple, entire, petiolate, exstipulate; lamina elliptic to obovate or narrowly so, index 1.6–5, chartaceous to coriaceous, glabrous (rarely sparsely covered with appressed or erect, simple, up to 1 mm long hairs) above, glabrous to densely hairy (particularly at the base and on veins) below, base acute, obtuse or rounded, rarely subcordate to cordate, apex acuminate, sometimes caudate, rarely obtuse to acute, extreme tip rounded, venation brochidodromous, primary vein raised over entire leaf length above with an often conspicuous longitudinal groove particularly in the basal half, secondary veins 5–20(–30) on either side of the primary vein, often with 1–6 intersecondary veins, running parallel to primary vein for a short distance, thereafter angles with primary vein either increasing or decreasing towards the apex (or consistent), sometimes branching, often forming distinct loops, smallest distance between loops and margin 1–7 mm, tertiary veins percurrent (or reticulate). Inflorescence of single flowers or occasionally up to 8 in a rhipidium, pendant, clustered in groups of up to 7, terminal on short axillary shoots (i.e. peduncles) on leafy or leafless twigs, older branches or on the main trunk (then often on brachyblasts); 1-several lower bracts, deltate to depressed ovate, rarely narrowly elliptic, leafy, rounded to acute, soon falling off or persistent; single upper bract attached to pedicel, ovate to deltate, acute to obtuse; flower buds open or closed in development, when closed (ovoid to triangular) broadly to depressed ovoid; peduncles, pedicels, outer sides of bracts, sepals and petals glabrous to densely covered with appressed or erect, simple, up to 1 mm long hairs, bracts, sepals and petals ciliate. Flowers actinomorphic, bisexual, with one whorl of free or slightly connate, imbricate, sepals and two whorls of free, imbricate, petals, green, creamy or yellow in vivo, often black in sicco; sepals and petals thin at margins, occasionally with prominent venation; sepals three, much smaller than petals; petals six, the outer ones ovate, elliptic or broadly so, the inner ones elliptic to obovate or narrowly so, stamens numerous (ca. 100), spirally arranged, extrorse, inserted on and below a ventral ridge encircling a central depression in the receptacle in which the carpels are inserted, 1–2 mm long, connective appendage transversely rhombic-hexagonal; carpels 20–40, spirally arranged, free, ovary 1-locular, glabrous or hairy, with 1 basal, lateral or apical ovule (reported by Van Heusden 1992), stigma sessile. Fruit apocarpous, monocarps 5–40, stipitate, mostly asymmetrical, sometimes strongly so, sometimes with an (often excentric) apicule, green maturing mostly through red to brown or black in vivo, light brown to black in sicco. Seeds 1, lateral or apical (reported by Van Setten & Koek-Noorman 1992), ellipsoid to globose, yellow to reddish-brown, surface deeply to shallowly pitted, lacking an aril, with a raised or sunken raphe encircling seed longitudinally (diagonally), regularly (or more sinuously), ruminations spiniform.

34 species in the Neotropics: from southern Costa Rica in the north to Bolivia in the south. Most species are distributed in regions surrounding the Andean mountain range, two in coastal Venezuela (Cremastosperma macrocarpum Maas and C. venezuelanum Pirie), one in French Guiana (C. brevipes (DC.) R.E.Fr.) and one widespread across Brazil, south of the Amazon River (C. monospermum (Rusby) R.E.Fr.).

Lowland to premontane tropical wet forest, inundated areas and terra firme. At elevations of 0–2000 m. Flowers and fruit of species of Cremastosperma appear similar in overall morphology and ontology to those of other Annonaceae demonstrated or presumed to be beetle-pollinated and/or bird-dispersed. We have observed in various species that the inner petals form a loose pollination chamber when mature, similar to that observed in, for example, Guatteria (Gottsberger 1970); and flowers of Malmeoideae genera in general are visited at least predominantly by small beetles (Saunders 2012). Similarly, we have observed in various species that the fruits become fleshy at maturity and often present a colour contrast between, for example, black monocarps and bright red stipes, representing a classic bird/monkey dispersal syndrome (Gautier-Hion et al. 1985). However, we are unaware of detailed studies of either pollination biology or seed dispersal in species of Cremastosperma.

Figure 4. 

Distribution map of Cremastosperma showing the four disjunct areas of the distribution: 1 the Chocó/Darién/western Ecuador region north into Central America 2 the tropical Andes (including a rough demarcation between areas occupied by northern/lowland and southern/montane clades) 3 coastal Venezuela 4 French Guiana.

Figure 5. 

Summary phylogeny of Malmeeae, adapted from Chatrou et al. (2018). A putative synapomorphy for the the Malmea, Pseudoxandra and Cremastosperma clade is indicated; one node that is not subject to significant support is indicated with an asterisk.

Figure 6. 

Phylogenetic hypotheses for Cremastosperma based on rbcL, matK, trnT-F, psbA-trnH, ndhF and pseudtrnL-F (adapted from Pirie et al. (2018)). A Excluding taxa for which ndhF was unavailable B including all taxa. Topologies and branch lengths are of the best scoring ML trees with scale in substitutions per site. Branch lengths subtending the ingroup are not to scale. Clade support is indicated: ML and parsimony bootstrap percentages (above; left and right, respectively) and Bayesian posterior probabilities (below). Geographically restricted clades/lineages are highlighted with different colours and labelled on the right.

Most similar to C. pedunculatum, from which it differs by the branching inflorescence, shorter pedicels and shorter stipes and the absence of indument on the pedicels. Differs from the only two other Cremastosperma species with branching inflorescences by the absence of indument on flowers and fruits and shape of the monocarps (compared to generally hairy C. cauliflorum with globose to transversely broadly ellipsoid monocarps) and stipes shorter than monocarps (much longer than monocarps in C. napoense).

ECUADOR. Zamora-Chinchipe: Palanda, Región de la Cordillera del Cóndor, sector sur, Parroquia San Francisco de Vergel, Cuenca alta del Río Vergel, Pica Sangola, 16 Mar 2005, Quizhpe, W. et al. 1088 (holotype: MO! [MO-2985962]; isotypes: MO! [MO-2985963], LOJA, U! [U 0249915]).

Tree 6–20 m tall; young twigs and petioles glabrous to sparsely covered with appressed golden brown hairs 0.1–0.2 mm long. Leaves: petioles 5–9 by 1.5–2 mm; lamina elliptic to obovate, 17–27 by 6–7.5 cm (leaf index 2.7–2.9), chartaceous, patchy blackish-brown with darker primary vein, glabrous, base acute to obtuse, apex acuminate (acumen ca. 6 mm long), primary vein raised over entire of leaf length, ca. 2 mm wide at widest point, secondary veins 9–10, intersecondary veins 3–4, distance between from 5–10 mm at the base to 15–25 mm closer to the apex, angles with primary vein from 30–40° at the base to 40–50° closer to the apex, not branching, forming indistinct loops, smallest distance between loops and margin 2–3 mm, tertiary veins somewhat reticulate. Inflorescence of 1–2 flowers, branching, clustered in groups of up to 2, on leafy and leafless twigs; transition between short axillary shoot and pedicel unclear, combined structure 26–33 by ca. 1.5 mm at the base, ca. 4 mm at the apex (in fruit; only young buds seen), short axillary shoot and pedicels glabrous; 1 lower bract, broadly elliptic, ca. 2 by 1.5 mm, rounded, only present in early bud, glabrous, ciliate; upper bract directly subtending flower in bud, midway along pedicel in fruit, shallowly triamgular, ca. 2 by 3 mm, rounded, glabrous, ciliate; closed flower buds depressed ovoid, blackish-brown in sicco, sepals and petals glabrous and ciliate; mature flowers not seen, flowers reported as greenish-yellow in vivo. Monocarps 6–17, ellipsoid, symmetrical, 19–30 by 15–25 mm, no obvious apicule, black in vivo, blackish-brown in sicco; stipes 10–14 by 1.5 mm at the base increasing to ca. 4 mm at the apex; fruiting receptacle ellipsoid to transverse elipsoid, 6–12 mm diam.; monocarps, stipes and receptacle glabrous. Seeds ellipsoid, reddish-brown, pitted to grooved, 15–16 by 10–11 mm, raphe sunken, encircling seed longitudinally, ruminations spiniform.

Ecuador (Zamora-Chinchipe), Peru (Cajamarca)

In dense and secondary forest. At elevations of 1700 and 2200 m. Flowering: March; fruiting: March and June.

Branching inflorescences are unusual in the genus, previously described in only two species: C. cauliflorum and C. napoense.

The epithet “alticola” refers to the high elevations at which the species has been found, which is unusual for the genus (and also the family in general).

Neither of the only two known collections of C. alticola was collected in protected areas and one was reported to be found in close proximity to human habitation. Given this low area of occupancy and the likely ongoing decline in area, extent and/or quality of the habitat, we classify the species as Endangered [EN] (Table 1).

Peru. Cajamarca: San Ignacio, Distr. San José de Lourdes, Perea & Flores 2540 (AMAZ, HUT, L, MO, MOL, USM, WAG),

Map 2. 

Distribution of Cremastosperma alticola Pirie & Chatrou, C. antioquense Pirie, C. chococola Pirie, C. dolichopodum Pirie & Maas, C. longipes Pirie, C. novogranatense R.E.Fr., C. osicola Pirie & Chatrou, C. panamense Maas and C. stenophyllum Pirie

Figure 8. 

Cremastosperma alticola Pirie & Chatrou. Fruiting specimen (Quizhpe et al. 1088).

COLOMBIA, Antioquia: Mun. Anorí, Corregimiento Providencia, Buenos Aires, 4 km from Providencia, 500–700 m a.s.l., 10 Dec 1972, Soejarto, D.D. 3586 (holotype: COL! [COL000334459]; isotypes: F! [V0054279F], GH! [GH00257162], HUA, MEDEL! [MEDEL000017], MO! [MO-047625]).

Tree ca. 5 m tall; young twigs and petioles sparsely covered with appressed brown hairs up to 0.2 mm long or glabrous. Leaves: petioles 7–10 by 2–3 mm; lamina elliptic, 16–27 by 6–9.5 cm (index 2.3–2.8), chartaceous, drying to a mosaic of brown and lighter green on both sides, glabrous on both sides, base obtuse, apex acuminate (acumen 10–15 mm long), primary vein grooved in the basal half, 1–1.5 mm wide at widest point, secondary veins 8–11, intersecondary veins occasional, distance between from 10 mm at the base to 50 mm closer to the apex, angles with primary vein from 50° at the base to 70° closer to the apex, forming distinct loops, smallest distance between loops and margin 3–4 mm, tertiary veins more or less percurrent. Inflorescence of single flowers, axillary on leafy twigs or from main trunk, then solitary or clustered in groups of at least two on brachyblasts; peduncles ca. 2 by 1.5 mm (in flower), 2–3 by 1.5–2 mm (in fruit); pedicels 20–28 by ca. 1 mm at the base, 1.5–2 mm at the apex (in flower), 20–40 by ca. 2 mm at the base, ca. 3 mm at the apex (in fruit), peduncles and pedicels sparsely covered with appressed whitish-golden hairs to 0.2 mm long; 2 lower bracts, deltate, ca. 1 mm long, obtuse, soon falling off; upper bract attached around halfway along pedicel, deltate, ca. 1 mm long, obtuse, outer side of upper and lower bracts rather densely to densely covered with appressed whitish-golden hairs to 0.2 mm long; closed flower buds not seen; flowers light green, stamens and carpels yellowish or pinkish in vivo, petals dark brown, contrasting to lighter colour of sepals and pedicels in sicco; sepals fused at base, deltate, appressed, 2–2.5 by 2–2.5 mm, acute, soon falling off, sparsely to rather densely covered with appressed whitish-golden hairs to 0.2 mm long; outer petals elliptic, ca. 12 by 8 mm, inner petals elliptic, 10–12 by 5–6 mm, outer side of outer and inner petals rather densely covered with appressed whitish-golden hairs to 0.2 mm long; receptacle depressed ovoid; androecium 5–7 mm diam., stamens ca. 1 mm long, connective appendage 0.5–0.7 mm wide, glabrous; gynoecium 1–1.5 mm diam., carpels ca. 1.5 mm long, glabrous. Monocarps ca. 10, ellipsoid to broadly so, strongly asymmetrical, 13–14 by 11 mm, orange to deep red, maturing to black in vivo, dark reddish-brown in sicco, with an excentric apicule; stipes orange to deep red in vivo, ca. 20 by 1.5 mm; fruiting receptacle depressed ovoid, ca. 6 mm diam; monocarps, stipes and receptacle glabrous. Seeds ellipsoid, reddish-brown with dark pits each surrounded by a raised rim, ca. 12 by 9 mm, raphe sunken, regular.

Colombia (Antioquia).

Primary forest. At elevations of 500–700 m. Flowering: February, fruiting: December.

The distinctive strongly asymmetrical monocarps of Cremastosperma antioquense could only be confused with those of C. chococola,and collections of both species display cauliflory (though not exclusively so in C. antioquense) with inflorescences inserted on similar brachyblasts. However, C. chococola can easily be distinguished from C. antioquense by its somewhat smaller, narrowly elliptic leaves with typical pinkish-brown colour on the underside (no such colour contrast in the brown/green-drying leaves of C. antioquense) and by the absence of hairs on the pedicels. The flowers of C. antioquense are superficially similar to those of C. awaense, particularly in the dimensions of the sepals and petals and lengths of the pedicels. However, petals of C. antioquense are uniformly covered in indument in contrast to the distinctive indument patterning on those of C. awaense and the fruits of the two species are more distinct: in contrast to C. awaense, the monocarps of C. antioquense are smaller, shorter than the stipes, strongly asymmetrical and entirely glabrous. In addition, none of the collections of C. awaense display cauliflory, a condition found in both of the two collections of C. antioquense.

None of the only three known collections of C. antioquense was found in protected areas. Given the low area of occupancy and a likely ongoing decline in area, extent and/or quality of the habitat, we propose to classify the species as Endangered [EN] (Table 1).

COLOMBIA. Antioquia: Providencia, 7°30'N, 74°40'W, 500–700 m a.s.l., 12 Feb 1971, Soejarto 2798 (COL, GH, HUA); Anorí, Vereda “La Esperanza”, 7°11'03"N, 75°01'53"W, 8 Nov 1999, Tuberquia et al. 1416 (COL).

Figure 9. 

Cremastosperma antioquense Pirie. a leaf and flower b fruit (aSoejarto 2798bSoejarto 3586).

ECUADOR. Carchi: Maldonado, parish of Tobar Donoso, Ethnic Reserve Awá, Sabalera, 900 m a.s.l., 22 Nov 1992, Aulestia, C. 842 (holotype: QCNE! [QCNE87829]; isotypes: MO! [MO-1609731], U! [U0012222]), US! [US00901537].

Tree 4–15(-20) m tall, 8–25 cm diam.; young twigs and petioles sparsely to rather densely covered with appressed golden hairs to 0.3 mm long. Leaves: petioles 4–11(–15) by 1.5–3 mm; lamina elliptic to slightly obovate or narrowly so, 17–33 by 5.5–13 cm (index 2.2–3.6), chartaceous, brown/grey green above, darker below, veins on underside dark brown, glabrous above, veins sparsely to rather densely covered with appressed golden hairs to 0.3 mm long below, base obtuse to acute, apex acuminate (acumen 10–20 mm long), primary vein 1–3 mm wide at widest point, secondary veins 7–11, intersecondary veins occasional, distance between from 10 mm at the base to up to 60 mm closer to the apex, angles with primary vein from 45–50° at the base to 55–60° closer to the apex, forming loops in the apical half, smallest distance between loops and margin 1–3 mm, tertiary veins slightly reticulate. Inflorescence of single, solitary flowers, axillary on leafy or leafless twigs; peduncles ca. 1.5 by 1 mm (in flower), 1.5–3 by 1–2 mm (in fruit); pedicels 27–28 by ca. 1 mm (in flower), 35–60 by 1–2 mm (in fruit), peduncles and pedicels rather densely to densely covered with appressed golden hairs to 0.3 mm long; single lower bract, broadly elliptic, 1–2 by 1–1.5 mm, obtuse, soon falling off, outer side densely covered with appressed golden hairs to 0.3 mm long; upper bract attached around midway along the pedicel, broadly elliptic, 1–2.5 by 1–2 mm, obtuse, rather densely to densely covered with appressed golden hairs to 0.3 mm long; closed flower buds depressed ovoid; flowers green or cream in vivo, blackish in sicco; sepals free, deltate, reflexed (appressed in bud), 2–2.5 by 2–2.5 mm, obtuse, soon falling off, outer side rather densely to densely covered with appressed golden hairs to 0.3 mm long; outer petals elliptic to broadly elliptic, 10–15 by 8–9 mm, inner petals elliptic, 10–15 by 5–6 mm, sparsely to rather densely covered with appressed golden hairs to 0.2 mm long on the outer side, denser at the base and in a band leading from the base to the apex of the petals; stamens 1–1.5 mm long, connective appendage ca. 1 mm wide; gynoecium ca. 2 mm diam., carpels 30–40, 1–2 mm long, sparsely covered with golden, <0.1 mm long hairs. Monocarps 10–12(–20), ellipsoid, slightly asymmetrical, 22–28 by 12–17 mm, brown in sicco, with an excentric apicule or rarely a nipple-like protuberance; stipes 11–24 by 1–1.5 mm; fruiting receptacle depressed ovoid, 3.5–8 mm diam. monocarps, stipes and receptacle very sparsely to sparsely covered with appressed white hairs <0.1 mm long. Seeds ellipsoid, asymmetrical, yellow-orange with shallow pits, ca. 19 by 11 mm, raphe sunken, regular.

Pacific coast of Colombia (Chocó, Nariño) and Ecuador (Carchi, Esmeraldas).

Primary humid to premontane tropical forest. At elevations of 0–2000 m. Flowering: January, September and November; fruiting: January, February and June to September.

Colombia: Guasca negra (Romero-Castañeda 3369). Ecuador: Cargadera negra (Tipaz et al. 1718), Castaña negro (Aulestia, C. et al. 1842), Huasca negra (Quelal et al. 191), Teuug teiug (Tipaz et al. 1428).

Cremastosperma awaense can be distinguished by the unique pattern of indument on the outer sides of the petals; denser at base and in a line leading to the petal apex. The sparse indument of very short (<0.1mm) hairs on the monocarps and stipes are not visible without magnification and the fruits appear glabrous. This character is also exhibited by some specimens of C. westrae. C. awaense can easily be distinguished from both C. westrae and the geographically closer C. stenophyllum Pirie on the basis of the length of the pedicel. The pedicels of C. westrae are shorter (not exceeding 17 mm) and those of C. stenophyllum longer (ca. 45 mm in comparison to 27–28 mm in flower).

Although the species is distributed across a moderately wide area (EOO >20,000), C. awaense is rather rarely collected (AOO <500 km²) and half of the specimens known to us are from within just one ethnic reserve in Ecuador which reportedly does not confer protected status to the biota. Decline in non-protected areas may lead to a considerable reduction in EOO and hence we assign the category Near Threatened [NT] (Table 1).

COLOMBIA. Chocó: Termales, between Jobi and Arusi, 5°37'N, 77°25'W, 5–50 m a.s.l., 31 Jan 1995, Betancur et al. 6043 (COL, HUA, U, US). ECUADOR. Carchi: Reserva Etnica Awá-Camumbí, 0°53'N, 78°16'W, 1700–1900 m a.s.l., 20 Jul 1991, Quelal et al. 191 (U); Tulcán, 0°53'N, 78°20'W, 1600 m a.s.l., 20 Sep 1991, Rubio et al. 2181 (MO); Reserva Etnica Awá, sector Sabalera, 1°00'N, 78°24'W, 650–1000 m a.s.l., 19 Jun 1992, Tipaz et al. 1428 (U); Reserva Etnica Awá, 0°53'N, 78°25'W, 1800 m a.s.l., 17 Aug 1992, Tipaz et al. 1718 (U); Lita-Alto Tambo Road, ca. 20 km past Lita, 0°55'N, 78°30'W, 550 m a.s.l., 26 Jun 1991, Van der Werff et al. 12045 (QCNE, U). Esmeraldas: Reserva Etnica Awá, 1°08'N, 78°33'W, 200 m a.s.l., 21 Sep 1992, C. Aulestia et al. 637 (U); Bravito, 0°35'N, 79°02'W, 600 m a.s.l., 9 Sep 1998, X. Cornejo & Bonifaz 6451 (U); Lita-San Lorenzo road, 1°05'N, 78°40'W, 300–500 m a.s.l., 12 May 1990, Gentry et al. 70042 (U); Eloy Elfaro, 0°49'N, 78°45'W, 250 m a.s.l., 23 Oct 1993, Tirado et al. 591 (MO, US); Río Santiago, 0°49'S, 78°54'W, 200 m a.s.l., 17 Jul 1994, Tirado et al. 1083 (U).

Map 3. 

Distribution of Cremastosperma awaense Pirie, C. dolichocarpum Pirie, C. magdalenae Pirie, C. napoense Pirie, C. pacificum R.E.Fr. and C. westrae Pirie

Figure 10. 

Cremastosperma awaense Pirie. a flower b leaf and fruit (aAulestia 842bVan der Werff 12045).

Most similar to C. pendulum, C. confusum and C. monospermum (all species also found in central and southern Peru, with glabrous flowers and fruit and smallish leaves), from which it differs in the unusually short stipes; further differs from C. monospermum in the shape of the flower bud (depressed ovoid, compared to broadly ovoid-triangular) and from C. monospermum and C. pendulum in the shorter pedicels.

PERU, San Martín: Huinguillo, 3 Mar 1962, Woytkowski, F. 7128 (holotype: MO! [MO-047731]; isotypes: GH, UC).

Tree ca. 8 m tall; young twigs and petioles glabrous. Leaves: petioles 4–7 by 1–2 mm; lamina elliptic to obovate, 11–17 by 4.5–7 cm (leaf index 2.3–2.4), chartaceous, olive green, more greyish above, glabrous, base acute to obtuse, apex acuminate (acumen 8–10 mm long), primary vein raised over entire leaf length, ca. 1 mm wide at widest point, glabrous, secondary veins 5–6, intersecondary veins ca. 1–2, distance between varying along length (15 mm to ca. 25 mm), angles with primary vein from ca.70° at the base to ca.60° closer to the apex, not branching, forming distinct loops, smallest distance between loops and margin 3–4 mm, tertiary veins reticulate. Inflorescence of single flowers, axillary; short axillary shoot, ca. 1 by 0.5 mm (in flower), ca. 2 by 1 mm (in fruit); pedicels 21–28 by 1 mm at the base, 2.5 mm at the apex (in flower), ca. 44 by 1–3 mm at the base, ca. 3 mm at the apex (in fruit), short axillary shoot and pedicels glabrous; 2 lower bracts, soon falling off; upper bract attached at between one third and half of the pedicel length, deltate, ca. 1 mm long by ca. 1 mm diam., apex rounded, ciliate; closed flower buds depressed ovoid; flowers creamy white in vivo, dark brown in sicco; sepals connate for ca. 1 mm, deltate, somewhat recurved, 2–2.5 by 2.5–3 mm, rounded, persistent, glabrous; outer petals ovate, 11–13 by 8–9 mm, inner petals elliptic to ovate, 8–12 by 4–5 mm, glabrous, ciliate; receptacle ovoid, apex concave; androecium ca. 3 mm diam., stamens ca. 100, 1.5–2 mm long, connective appendage ca. 1 mm wide; gynoecium ca. 2 mm diam., carpels ca. 15, glabrous. Monocarps ca. 13, ellipsoid, slightly asymmetrical, 7–9 by 6–7 mm, without apicule, blackish-brown in sicco; stipes ca. 2 by ca. 2 mm; fruiting receptacle elipsoid, 6 mm diam.; monocarps, stipes and receptacle glabrous. Seeds not seen.

Peru (San Martín).

Forest. At an elevation of ca. 500 m. Flowering and fruiting: March.

Cremastosperma brachypodum, known only from the type, is distinctive within Cremastosperma due to the unusual shortness of the stipes, but otherwise typical of species of the southern/montane clade in the absence of indument on all parts.

The specific epithet “brachypodum” is derived from the Greek brachy (short) and podum (-stalked), referring to the short stipes.

This new species is only known from a single collection dating back to 1962 in a non-protected, rural area. Critically endangered [CR] (Table 1).

Figure 11. 

Cremastosperma brachypodum Pirie & Chatrou. Flowering and fruiting specimen (Woytkowski 7128).

FRENCH GUIANA, without location Martin s.n. (lectotype: G! [designated in Pirie, Kankainen & Maas, 2005], isotypes: BM, K! (uncertain whether this specimens represents an isotype) [K000485530]).

Cremastosperma poiteaui (Diels) R.E.Fr., Acta Horti Bergiani 10: 328. 1931.

Guatteria poiteaui Diels, Notizbl. Bot. Gart. Berlin-Dahlem 11: 74. 1931.

FRENCH GUIANA, without location, 1819–1821, Poiteau, A. s.n. (holotype: G! [G00237255 G00237256]; isotypes: B! [B 10 0242372], F! [F932330], S! [S-R-7016]).

Tree 4–20 m tall, 3–20 cm diam.; young twigs and petioles glabrous to sparsely covered with appressed white/yellowish hairs to 0.2 mm long. Leaves: petioles 4–9(–12) by 1.5–4 mm; lamina elliptic to obovate or narrowly so, 18–39 by 7–15 cm (index 2–3.5), chartaceous, brown/green with a reddish tinge on both sides (particularly on the veins on the underside), darker above, glabrous on both sides, base obtuse to rounded, rarely acute, apex acuminate (acumen 5–20 mm long), primary vein 1.5–3 mm wide at widest point, secondary veins 9–15, intersecondary veins 0–2, distance between from 5–10 mm at the base to 35 mm closer to the apex, angles with primary vein from 45–60° at the base to 55–70° closer to the apex, not branching, forming mostly distinct loops, smallest distance between loops and margin 1–3 mm, tertiary veins percurrent. Inflorescence of single flowers, solitary or clustered in groups of 2, on leafless twigs; peduncles 1–2 by 1–2 mm (in flower), 2–5 by ca. 2 mm (in fruit), sparsely to rather densely covered with appressed white/yellowish ca. 0.1 mm long hairs; pedicels 15–20 by 1–2 mm at the base (in flower), 18–23 by ca. 2 mm (in fruit), sparsely covered with appressed white/yellowish hairs ca. 0.1 mm long or glabrous; 2 lower bracts, deltate, basal to 0.2 mm long, apical 0.3–0.5 mm long, obtuse, mostly persistent, rather densely covered with appressed white/yellowish hairs ca. 0.1 mm long; upper bract mostly attached midway along pedicel, deltate to broadly ovate, 1–2 by ca. 1 mm, obtuse or emarginate, sparsely covered with appressed white/yellowish hairs ca. 0.1 mm long; closed flower buds depressed ovoid, opening in development; flowers green, sometimes tinged with red around margins of petals or creamy yellow in vivo, reddish or dark brown in sicco, sepals and petals glabrous; sepals connate for 0.5–1 mm, broadly ovate, recurved, rarely appressed, 3–4[-5] by 3.5–5 mm, obtuse, soon falling off; outer petals elliptic, 12–22 by 7–12 mm, rounded, inner petals narrowly obovate to narrowly elliptic, 10–24 by 4–7 mm, obtuse; androecium ca. 6 mm diam.; stamens 1.4–1.6 mm long, connective appendage 0.5–0.7 mm wide; gynoecium ca. 1 mm diam.; carpels ca. 25, ca. 2.5 mm long, glabrous. Monocarps 7–17, ellipsoid, slightly asymmetrical, 11–17 by 9–11 mm, green maturing to red, reddish-brown, dark purple or black in vivo, blackish or reddish-brown in sicco, with an excentric apicule; stipes green maturing to red in vivo, 7–14 by 1–1.5(3) mm; fruiting receptacle depressed ovoid, 4–8 mm diam., monocarps, stipes and receptacle glabrous. Seeds broadly ellipsoid to globose, yellowish or orange-brown, slightly pitted, ca. 8 by 7–8 mm, raphe raised within a sunken groove, somewhat irregular.

French Guiana, region of Saül and Nouragues.

Primary moist forest. At elevations of 200–800 m. Flowering: February, March, May and October; fruiting: May – October, December and January.

French Guiana: Apélému (Wayapi; Grenand 1509, Jacqemin 2349), Maman yawé (Créole; Moretti 12).

Cremastosperma brevipes is the only species of the genus found in the Guianas. The leaves, when dried, have a characteristic reddish tinge (particularly on the underside). This species is similar to C. venezuelanum, but differing in particular by the acuminate as opposed to acute or obtuse leaf apex of C. venezuelanum, smaller sepals and monocarps and shorter stipes. Its use as fish bait has been reported. As of many Annonaceae, the bark of C. brevipes has been described as aromatic: on collection of Riera 668, a peppery smell is reported.

C. brevipes has been collected regularly, occurring over a fairly wide area, including the protected area of Les Nouragues. Least concern [LC] (Table 1).

. FRENCH GUIANA Rivière Arataye, Saut Pararé, 4°02'N, 52°42'W, 3 Dec 1985, Barrier 5166 (CAY, L, P, U); Saül, Mont Galbao, 3°36'N, 53°16'W, 525–700 m a.s.l., 11 Sep 1994, Boom 10812 (CAY, NY, U); Mont Inéri, 4°22'N, 52°10'W, 10 m a.s.l., 9 Sep 1997, Cremers 15331 (U); Trésor Reserve, Creek 4, middle part, 4°35'N, 52°16'W, Ek et al. 1615 (U); Régina, Montagne Tortue, 4°18'N, 52°22'W, 200–450 m a.s.l., 17 Jun 1988, Feuillet et al. 10239 (U); Piste de Saint-Elie, 5°20'N, 53°00'W, 29 Nov 1987, De Granville 10176 (CAY, P, U, US); Approuague, 4°28'N, 52°02'W, 28 Jul 1997, Hequet 432 (P, U); Pedra Alice, River Oiapoque, 3°40'N, 52°01'W, 17 Sep 1960, Irwin et al. 47562 (NY, U); Route de Bélizon, Les Eaux Claires-Saül, 3°37'N, 53°12'W, 200 m a.s.l., 9 Feb 1993, Maas et al. 8064 (MO, U); Saül, Piste de Galbao, 4°36'89"N, 54°16'25"W, 28 Aug 2003, Munzinger & Heuret 1818 (P); Crique Cacao, 2°20'N, 53°12'W, 2 May 1987, Prévost & Sabatier 2284 (CAY, U); Montagne de Kaw, 4°33'N, 52°10'W, 21 Feb 1998, Prévost 3446 (NY, U); Les Nouragues, 4°03'N, 52°42'W, 8 Oct 1992, Riera 1564 (P); Les Nouragues, basin of River Approuague, 4°04'57"N, 52°40'39"W, 89–140 m a.s.l., Oct 2001, Scharf 76 (U).

Map 4. 

Distribution of Cremastosperma brevipes (DC.) R.E.Fr.; C. gracilipes R.E.Fr.; C. monospermum (Rusby) R.E.Fr.

Figure 12. 

Cremastosperma brevipes (DC.) R.E.Fr. a fruiting specimen b flower buds (aBoom 10812bPrévost 3446).

PERU, Amazonas: Bagua, Distr. Imaza, community Yamayakat, trail to Putuim, 360 m a.s.l., 22 Nov 2003, Pirie, M.D. et al. 71 (holotype: U! [two sheets U0121238, U0121239]; isotypes: AAU!, AMAZ!, CUZ!, E! [E00268265], F! [V0047939F], HAO!, HUT!, K! [K000580475], MO! [MO-1459050], MOL!, NY! [NY00689082], US! [US00901687], USM! [USM000035], WU! [WU0038419]).

Tree 2–10 m tall; young twigs and petioles densely covered with mainly erect golden hairs up to 1 mm long. Leaves: petioles 3–7 mm by 2.5–3 mm; lamina elliptic or narrowly so to slightly obovate, 17–28 by 6–11 cm (index 2.4–3.5), chartaceous, mid-brown, occasionally slightly grey above (immature leaves drying black), sparsely covered with mainly erect golden hairs up to 1 mm long or glabrous above, densely so on edge of lamina and on all veins below, base rounded to subcordate, apex acuminate (acumen 5–20 mm long), primary, secondary and tertiary veins sunken in depressions in leaf surface, primary vein 1.5–2 mm wide at widest point, densely covered with mainly erect golden hairs up to 1 mm long above and below, secondary veins 15–20 (intersecondary veins rare), distance between from 6 mm at the base to 16 mm closer to the apex, angles with primary vein consistently around 60–70°, occasionally branching, forming distinct loops, smallest distance between loops and margin 1–1.5 mm; tertiary veins mostly percurrent. Inflorescences of single, successively produced, flowers, axillary on leafy branches, on leafless branches and produced from the main trunk (then on brachyblasts); peduncles and pedicels sparsely to rather densely covered with mainly erect golden hairs up to 1 mm long, peduncles 17–20 by 1–1.5 mm (in flower), 18–25 by 1.5 mm (in fruit); pedicels up to 120 by 1 mm at the base (in flower), 110–150 by 1.5 mm (in fruit); bracts densely covered with mainly erect golden hairs up to 1 mm long, single lower bract, elliptic to ovate, ca. 2.5 by 1 mm, acute, persistent or falling off; upper bract within central third of pedicel length, elliptic to ovate, 2–3 by 1–2 mm, acute; flower buds depressed ovoid, developing to ovoid before opening; flowers green, maturing to yellow with a basal orange patch on the outside of the outer petals in vivo, golden brown in sicco, outer sides and apical portion of the inner sides of petals and outer sides of sepals densely covered in appressed golden hairs up to 1 mm long, inner sides otherwise glabrous; sepals basally connate, deltate, 5–7 by 6 mm, acute, soon falling off, outer petals broadly ovate, ca. 18 by 15 mm, inner petals ovate, concave, ca. 25 by 12 mm; androecium ca. 5 mm diam., stamens 1–1.5 mm long, connective appendage ca. 0.5 mm wide; gynoecium [ca. 2.5] mm diam., glabrous. Monocarps 8–10, dark brown in sicco, ellipsoid, slightly asymmetrical, ca. 15 by 11 mm, often with an excentric apicule; stipes 14–16 by 1.5 mm; fruiting receptacle 5–6 mm diam., monocarps, stipes and receptacle sparsely to moderately densely covered with erect golden hairs up to 0.2 mm long. Seeds ellipsoid, orange-brown, shallowly pitted, ca.13 by 10 mm, raphe raised, regular.

Ecuador (Morona-Santiago) and Peru (Amazonas).

Primary forest, on red clay. At elevations of 300–500 m. Flowering: February and November; fruiting: November and June.

Cremastosperma bullatum can easily be distinguished from all other species of Cremastosperma by any one of the number of unique and striking characteristics it displays. The leaf blade has a corrugated (bullate) appearance, both in the field and when pressed, which is due to the deeply sunken nature of the primary, secondary and tertiary venation. The indument present on many of its parts is far longer than in any other species in the genus and, also uniquely in the genus, densely inserted in a halo-like formation around the leaf margin. Other notable characteristics are the unusually long pedicel, the orange colouring of the base of the outer petals of mature flowers, the inner petals considerably longer than the outer petals and the rounded to subcordate shape of the leaf base.

Cremastosperma bullatum is known from a small number of locations, within a limited area of northern Peru and adjacent Ecuador outside of protected areas. Endangered [EN] (Table 1).

ECUADOR. Morona-Santiago: Región de la Cordillera del Cóndor, 3°05'13"S, 78°04'23"W, 380 m a.s.l., 1 Jun 2006, Wisum & Kajekai 446 (US). PERU. Amazonas: Bagua, Yamayakat, 4°55'S, 78°19'W, 320 m a.s.l., 20 Jan 1996, Jaramillo et al. 942 (MO, U); Bagua, Yamayakat, trail to Putuim, 5°03'09"S, 78°20'58"W, 343 m a.s.l., 22 Nov 2003, Pirie et al. 66 (HAO, U, USM); Bagua, Putuim, 5°01'44"S, 78°22'43"W, 339–359 m a.s.l., 25 Nov 2003, Pirie et al. 94 (AMAZ, CUZ, HAO, HUT, MO, U, USM).

Map 5. 

Distribution of Cremastosperma bullatum Pirie; C. cauliflorum R.E.Fr.; C. cenepense Pirie & Zapata; and C. longicuspe R.E.Fr.

Figure 13. 

a–b Cremastosperma brevipes (DC.) R.E.Fr. a flowers (Maas et al. 8064; photo PJMM) b fruit (Mori et al 22721; photo Scott Mori) c–e C. bullatum Pirie c, e Flowering specimen (Pirie et al. 94; photos c MDP e Robin van Velzen) d leaf base showing bullate corrugations of the lamina and long golden indument (Pirie et al. 71; photo: MDP) f, g C. cauliflorum R.E.Fr. Flowering specimens (fMaas et al. 9029, photo PJMM gChatrou et al. 224, photo LWC) h–j C. dolichocarpum Pirie. Flowering and fruiting specimen (Pedraza et al. 2146; photos: María F. González).

PERU, Loreto: Mishuyacu, near Iquitos, 100 m a.s.l., Feb-Mar 1930, Klug, G. 902 (holotype: B! [B 10 0242371]; isotypes: F! [F0054580F], NY! [NY00025860], S! [S-R-6959], US! [US00104263]).

Tree 2–20 m tall, 4–25 cm diam.; young twigs and petioles glabrous to rather densely covered with appressed or erect golden hairs to 0.5 mm long. Leaves: petioles 4–12(–16) by 2–4(–6) mm; lamina elliptic to obovate or narrowly so, (14–)20–61 by 5–14(–22) cm (index 2.3–3.7), chartaceous, olive/brown green above, darker below, glabrous above except for base of primary vein sparsely covered with appressed or erect hairs to 0.3 mm long, base, primary and secondary veins sparsely to rather densely covered with appressed or erect golden hairs to 0.5 mm long below, base acute to obtuse, apex acuminate (acumen 5–45 mm long), primary vein verrucose (particularly at the base), deeply grooved for most of length, 1.5–3.5(–5) mm wide at widest point, secondary veins (6–)10–17, occasionally 1–2 intersecondary veins, distance between from 4 mm at the base to up to 40 mm closer to the apex, angles with primary vein from 45–70° at the base to 45–60° closer to the apex, not branching, forming mostly distinct loops, smallest distance between loops and margin 1–5 mm, tertiary veins percurrent. Inflorescence of 1–5 flowers, branching, solitary or clustered in groups of up to 7, on thick leafless twigs or on main trunk (then often on brachyblasts); peduncles 3–12(–15) by 1–1.5(–3) mm (in flower), 3–15 by 1–3 mm (in fruit); pedicels 10–45 by 1–3 mm at the base (in flower), 15–45 by 1–3 mm (in fruit), peduncles and pedicels rather densely to densely covered with mainly erect golden hairs ca. 0.3 mm long, often with hairs more densely covering the articulation point between shoot and pedicel; single lower bract (from the axil of which short shoots develop bearing new flowers), deltate, 1.5–2 mm long, acute, soon falling off, densely covered with mostly appressed golden hairs to 0.3 mm long; upper bract attached around midway along pedicel, broadly to very broadly ovate or deltate, 2–4 mm long, obtuse or acute, outer side densely covered with appressed or erect golden hairs to 0.3 mm long; closed flower buds depressed ovoid, opening in development; flowers (pale) green, creamy white, greenish-yellow or yellow in vivo, brownish-yellow or brown with orange, dark brown or black base in sicco, outer side of sepals and petals densely covered with erect or appressed golden hairs to 0.4 mm long, inner side of sepals and petals sparsely to rather densely covered with erect hairs to 0.4 mm long or glabrous, base glabrous; sepals free, broadly to very broadly ovate-deltate, mostly recurved, 3–5 by 4–6 mm, obtuse, soon falling off; outer petals elliptic to broadly elliptic, 10–25(–32) by 9–17 mm, inner petals elliptic, 11–21(–32) by 6–11 mm; androecium 7–10 mm diam., stamens 1.5–2 mm long, connective appendage 0.7–1 mm wide; gynoecium 2–3 mm diam., carpels ca. 40, 2–2.3 mm long, sparsely to rather densely covered with mostly appressed golden hairs to 0.2 mm long. Monocarps 9–41, globose to transversely broadly ellipsoid, slightly asymmetrical, 8–13 by 10–14 mm, green maturing to orange, red, brown and black in vivo, blackish-brown or brown in sicco, sometimes with an apicule at or near the apex; stipes 7–23(–32) by 1–2 mm; fruiting receptacle depressed ovoid, 4–11 mm diam; monocarps, stipes and receptacle rather densely covered with erect golden hairs to 0.2 mm long. Seeds broadly ellipsoid to globose, orange, pitted, 9–10 by 9–10 mm, raphe sunken, regular.

Amazonian Colombia (Amazonas, Putamayo), Ecuador (Morona-Santiago, Napo, Pastaza, Sucumbíos), Peru (Loreto) and Brazil (Acre, Amazonas).

Moist primary forest, mostly non-inundated areas, on clayey or lateritic soil or white sand. At elevations of 100–500 m. Flowering: June, August, October-February; fruiting: May-February.

Colombia: Espintana blanca (Raffauf 102); Jiobo ñaatraje duceju (J. Murillo 565). Ecuador: Mantach (Shuar; Warush Juwa RBAE 119), Moncapatahue (Huaorani; M. Aulestia 3238), Nanguehue (M. Aulestia 3395), Piton (Quech; Cazalet et al. 7528), Uñetahue (Huaorani; Rubio 812), Uñitague (Huaorani; Espinosa 399). Peru: Bara (Rimachi 480), Bara caspi (McDaniel 20761, Rimachi 2397), Espintana (Cheta 6/173, Rimachi 2397), Mantaach (Achual Jivaro; W.H. Lewis 12049).

Cremastosperma cauliflorum is one of only three species of the genus displaying a branched inflorescence. It can be discerned from C. napoense and C. alticola by the presence of indument on the monocarps and stipes and by the greater length and density of hairs on the inflorescences. The wood is aromatic, flowers reported as vanilla scented.

Cremastosperma cauliflorum is one of the more widespread and abundant species of the genus, found in protected areas in Colombia and Ecuador, as well as in Northern Peru. Least concern [LC] (Table 1).

BRAZIL. Acre: Mun. Mancio Lima, Rio Moa, 7°25'S, 73°38'W, 12 Oct 1989, Cid Ferreira et al. 10019 (NY); Cruzeiro do Sul-Boa Fé road, km 12, 7°34'S, 72°44'W, 2001, Maas et al. 9029 (U); Mun. Cruzeiro do Sul, BR-307, 7°34'S, 72°45'W, 2001, Maas et al. 9251 (U). COLOMBIA. Amazonas: Tarapaca, 3°02'S, 70°00'W, 100 m a.s.l., 30 Jun 1992, Rudas et al. 4514 (MO); Parque Nacional Amacayacu, 3°45'S, 70°15'W, 100 m a.s.l., 8 Aug 1989, Vásquez et al. 12655 (MO). Putamayo: Puerto Leguízamo, Puerto Leguízamo-La Tagua road, 0°92'00"S, 74°46'40"W, 19 Jan 2000, Suárez 1331 (COL). ECUADOR. Morona-Santiago: Taisha, 2°23'S, 77°30'W, 500 m a.s.l., 14 Jun 1980, Brandbyge et al. 31854 (AAU, U); El Centro Shuar Pampants, 2°47'S, 77°36'W, 300 m a.s.l., 11 Sep 1985, Warush Juwa RBAE 119 (U); Cordillera de Cutucú, 2°06'48"S, 77°44'39"W, 610 m a.s.l., 16 Jul 2007, Wisum & Kajekai 1165 (WAG). Napo: Reserva Etnica Huaorani, 0°59'S, 76°12'W, 235 m a.s.l., 18 Jan 1995, Aulestia & Omehuat 3238 (QCNE); Río Wai si ayá, 0°15'S, 76°21'W, 300 m a.s.l., 13 Aug 1981, Brandbyge et al. 33503 (AAU, MO, NY, U); Río Aguarico, E of mouth of Río Cuyabeno, 0°16'S, 75°54'W, 200 m a.s.l., 20 Feb 1980, Holm-Nielsen et al. 21501 (AAU, K, MO, U); Cuyabeno Wildlife Reserve, 0°29'S, 75°32'W, 230 m a.s.l., 25 Sep 1991, Palacios 7602 (U). Pastaza: Pozo petrolero ‘Danta 2’ de UNOCAL, 1°47'S, 76°48'W, 365 m a.s.l., 1 Oct 1990, Espinoza & Coba 399 (MO, U); Pozo petrolero ‘Ramirez’, 1°32'S, 76°51'W, 300 m a.s.l., 21 Feb 1990, Zak 5272 (U). PERU. Loreto: Yanamono Explorama Reserve, 3°27'S, 72°51'W, 100–150 m a.s.l., 31 Dec 1998, Chatrou et al. 224 (MOL, U); Allpahuayo-IIAP, 3°50'S, 73°25'W, 150 m a.s.l., 2 Jan 1999, Chatrou et al. 233 (L, MOL); Prov. Requena (locality unknown), 4°50'S, 73°45'W, 170 m a.s.l., 8 Aug 1985, Cheta 6/ 173 (K); Prov. Maynas, NW of Zona Protectado Pucacuro, 2°08'13"S, 75°08'58"W, 160–270 m a.s.l., 26 Aug 2006, Dávila et al. 2784 (L); Nauta-Iquitos road, 4°30'S, 73°32'W, 140 m a.s.l., 27 Jun 1979, C. Díaz 1229 (F, MO, U); Caserío Gamitana, Reserva del Río Mazán, 3°30'S, 73°10'W, 116 m a.s.l., 21 Jun 1990, Grández et al. 1591 (LPB, MO, U); Prov. Maynas, NW of Area de Conservación Ampiyacu, 2°52'40"S, 73°00'46"W, 150–160 m a.s.l., 16 Oct 2009, Huamantupa et al. 12928 (L); Las Amazonas, ExplorNapo Camp, 3°20'S, 72°55'W, 100–140 m a.s.l., 15 Feb 1991, Pipoly et al. 12982 (MO, U); Prov. Requena, Quebrada Yanayacu-Río Tapiche, 6°15'49"S, 73°54'31"W, 140–180 m a.s.l., 16 Oct 2014, Ríos et al. 4491 (F, L); Andoas, 2°55'S, 76°25'W, 210 m a.s.l., 5 Jun 1981, Vásquez & N. Jaramillo 1938 (F, MO, U); Maniti, Recreo, 3°42'S, 72°50'W, 115 m a.s.l., 14 May 1988, Vásquez 10637 (F, MOL, NY, U); Iquitos-Nauta road, 4°10'S, 73°20'W, 150 m a.s.l., 14 Dec 1988, Vásquez & N. Jaramillo 11423 (MO, U, USM); Puerto Almendras, 3°48'S, 73°25'W, 122 m a.s.l., 20 Jun 1989, Vásquez & T. Soto 12358 (MO, U, USM).

Figure 14. 

Cremastosperma bullatum Pirie a leaf b leaf base c flower d fruit (a–cVásquez et al. 24891dJaramillo, N. et al. 972).

Figure 15. 

Cremastosperma cauliflorum R.E.Fr. a fruiting specimen b inflorescence (aPrance et al. 24094bVasquez & N. Jaramillo 11423).

PERU, Amazonas: prov. Condorcanqui, Río Cenepa region, community Mamayaque, 11 Aug 1997, Rojas, R. et al. 269 (holotype: U! (barcode U0123477]; isotypes: AMAZ, HUT, MO [MO-1664376], USM).

Tree ca. 10 m tall; young twigs and petioles sparsely (axillary buds densely) covered with appressed golden hairs ca. 0.1 mm long. Leaves: petioles 4–7 by 1–2 mm; lamina elliptic to narrowly so, 12–22 by 4–8 cm (index 2.7–3), chartaceous, grey-yellow green above, light brown or yellowish-green below, glabrous on both sides, base cordate to subcordate, apex acuminate (acumen 8–10 mm long), primary vein 1–1.5 mm wide at widest point, secondary veins 7–12, intersecondary veins occasional, distance between from 2–5 mm at the base to 15–25(35) mm closer to the apex, angles with primary vein from 80–90° at the base to 50–60° closer to the apex, forming distinct loops, smallest distance between loops and margin 2–5 mm, tertiary veins more or less percurrent. Inflorescence of single, solitary flowers, axillary on leafy twigs; peduncles ca. 2 by 2 mm (in fruit); pedicels ca. 8 by 2 mm at the base (in fruit), peduncles and pedicels sparsely covered with appressed golden hairs ca. 0.1 mm long; 2 lower bracts, soon falling off; upper bract attached midway along pedicel, soon falling off; closed flower buds and flowers not observed. Monocarps 8–10, blackish-brown in sicco, ellipsoid, slightly asymmetric, 14–15 by 9–11 mm, with an excentric apicule; stipes 7–8 mm by ca. 1.5 mm; fruiting receptacle 4–7 mm diam.; monocarps, stipes and receptacle rather densely covered with appressed golden hairs ca. 0.1 mm long. Seeds ellipsoid, golden brown shallowly wrinkled (immature), ca. 12 by 7 mm, raphe sunken, regular.

Peru (Amazonas, in the area of the Cenepa River, a tributary of the Marañon River).

Primary forest. At elevations of 250–400 m. Flowering: not recorded; fruiting: July and August.

Peru: Yais (Amarun; R. Rojas et al. 0255, 0269)

Cremastosperma cenepense is most likely to be confused with C. yamayakatense and C. gracilipes, which are more commonly collected in northern Peru and also characterised by relatively small leaves and fruits. It differs in the shape of the leaf base (cordate or subcordate as opposed to acute in C. yamayakatense and C. gracilipes), the indument on the fruits (rather dense as opposed to almost always absent in C. yamayakatense) and lengths of the pedicel (shorter than that of C. gracilipes) and stipes (shorter than those of C. yamayakatense).

Three of the only four known collections of C. cenepense are from more or less the same locality and none was found in protected areas. Given the low area of occupancy and a likely ongoing decline in area, extent and/or quality of the habitat, we propose to classify the species as Endangered [EN] (Table 1).

PERU. Amazonas: Río Cenepa region, Quebrada Nahem, 780 ft a.s.l., 15 July 1974, Kayap 1078 (MO, U); Río Cenepa region, community Mamayaque, 4°34'S, 78°14'W, 400 m a.s.l., 9 Aug 1997, R. Rojas et al. 0 255 (U); Río Cenepa region, community Mamayaque, 4°37'08"S, 78°13'46"W, 400 m a.s.l., 18 Aug 1997, Rojas et al. 0351 (MO).

Figure 16. 

Cremastosperma cenepense Pirie & Zapata. a fruiting twig b leaf base c fruit (a, cRojas 269; bKayap 1078).

COLOMBIA, Chocó: Alto de Buey, 500–1200 m a.s.l., 8 Jan 1973, Gentry, A.H. & Forero, E. 7286 (holotype: MO! [MO-047629]; isotype: COL! [COL000214771]).

Tree ca. 5 m tall; young twigs and petioles glabrous. Leaves: petioles 5–8 by 1.5–2 mm; lamina narrowly elliptic, 11–20 by 4–5.5 cm (index 3.7–4), chartaceous, dark/olive brown, shiny above, lighter pinkish-brown, matt below, glabrous above and below, base acute to cuneate, apex acuminate (acumen 7–10 mm long), primary vein ca. 1 mm wide at widest point, verrucose below, secondary veins 8–10, no intersecondary veins, distance between from 5 mm at the base to 30 mm closer to the apex, angles with primary vein from ca. 60° at the base to 60–70° closer to the apex, forming distinct loops, smallest distance between loops and margin 2.5–3.5 mm, tertiary veins reticulate. Inflorescence of single flowers, solitary or clustered in groups of at least two, on brachyblasts on the main trunk; peduncles, 2–3 by 1–1.5 mm (in fruit); pedicels 38–42 by 1 mm at the base, 1 mm at the apex (in fruit), peduncles and pedicels glabrous; lower bract(s) not observed; upper bract attached within basal half of pedicel, ovate, ca. 1 by 0.7 mm, obtuse, glabrous; flowers not observed. Monocarps 10–13, ellipsoid, strongly asymmetrical (stipes inserted within basal half of longest axis), 13–14 by 10–11 mm, with an excentric, to 0.2 mm long, apicule, green maturing through red to dark blue in vivo, dark brown in sicco; stipes 15–18 by ca. 1 mm increasing to 1.5 diam. when mature; fruiting receptacle depressed ovoid, 4–5 mm diam.; monocarps, stipes and receptacle glabrous. Seeds ellipsoid, orange-brown, pitted, 9–11 by 6–8 mm, raphe sunken, regular.

Pacific coast of Colombia (Chocó).

Tropical wet forest. At elevations of 0–1200 m. Flowering: not recorded; fruiting: January and June.

The strongly asymmetric monocarps of Cremastosperma chococola resemble those of C. antioquense: see above for distinctions.

Only three collections of C. chococola are known to us, from different localities but within a restricted region and only one of which is in a protected area. Endangered [EN] (Table 1).

COLOMBIA. Chocó: Punta Lanas, 1 September 1989, Espina et al. 3188 (HUA, MO); Parque Nacional Utría, 6°20'N, 77°20'W, 0–100 m a.s.l., 5 Jun 1990, F. García & Agualimpia 390 (MO).

Figure 17. 

Cremastosperma chococola Pirie. a leaf and fruit (Gentry & Forero 7286).

Cremastosperma confusum is most similar to C. monospermum, which produces effectively indistinguishable small fruits on slender pedicels. C. confusum differs from C. monospermum in the flower buds, which open in development, unlike in C. monospermum in which flower buds remain closed with a characteristic roughly triangular shape. The flowers of C. confusum are more similar to those of C. leiophyllum, from which it differs in the shape and colour of the monocarps, which, unlike those of C. leiophyllum, do not dry black and are not asymmetrical.

PERU, Madre de Dios: Tambopata, Explorer’s Inn, near the confluence of Río Tambopata and Río La Torre, 39 km SW of Puerto Maldonado, along the Big Tree Trail, 21 Jan 1989, Smith, S.F. et al. 1578 (holotype: USM; isotypes: U! [U0012270], NY).

Tree or shrub 3–8 m tall, 3–20 cm diam.; young twigs and petioles sparsely covered with appressed whitish hairs to 0.3 mm long. Leaves: petioles 7–10 by 1.5–3 mm; lamina elliptic to obovate or narrowly so, 10–28(–34) by 7–10(–12) cm (index 1.9–3.5), chartaceous to coriaceous, green (or greenish-brown), darker above, lighter or more brown with darker or reddish veins below, glabrous above, sparsely covered with appressed whitish hairs to 0.3 mm long particularly on veins below, base acute to obtuse (rarely rounded), apex acuminate (acumen 6–20 mm long), primary vein 1.5–3 mm wide at widest point, secondary veins 7–10, intersecondary veins 0–1, distance between from 11–20 mm at the base to 12–32 mm closer to the apex, angles with primary vein from 70–80° at the base to 40–50° closer to the apex, rarely branching, mostly forming distinct loops, smallest distance between loops and margin 2–5 mm, tertiary veins mostly percurrent. Inflorescence of single (very rarely branching) flowers, solitary, axillary on leafy or leafless twigs or thicker branches; peduncles (1-)3–5 by 1–2 mm (in flower), 3–9 by ca. 2 mm (in fruit), rather densely covered with appressed to erect whitish hairs to 0.2 mm long; pedicels 20–45(–70) by ca. 1 mm at the base (in flower), 22–80 by 1–2.5 mm (in fruit), pink, purple or reddish in vivo, glabrous; 2 lower bracts of unequal dimensions, basal lower bract depressed ovate, ca. 0.5 by 1 mm, obtuse, sometimes persistent, densely covered with appressed to erect whitish hairs to 0.2 mm long, apical lower bract elliptic, ca. 1.5 by 1 mm, obtuse, sometimes persistent, sparsely to rather densely covered with appressed to erect whitish hairs to 0.2 mm long; upper bract attached around midway along pedicel, (broadly) ovate, 1.5–2 by ca. 1–1.5 mm, obtuse or rounded, persistent, sparsely covered with appressed to erect whitish hairs to 0.2 mm long; closed flower buds depressed ovoid, opening in development; flowers green maturing to yellow or white and yellowish at base in vivo, dark to yellowish light brown sometimes tinged with red, darker at the base in sicco, sepals and petals glabrous; sepals free, deltate to triangular, appressed to recurved, ca. 3 by 2–3 mm, acute or obtuse, soon falling off or sometimes briefly persistent; outer petals elliptic, 14–17 by 6–8 mm, inner petals elliptic to narrowly so, 16–17 by 5–7 mm; androecium ca. 6 mm diam., connective appendage to 0.8 mm wide; gynoecium ca. 1.5 mm diam., carpels glabrous. Monocarps 10–25(–32), ellipsoid (broadly so when immature), slightly asymmetrical, 8–12 by 6–8 mm, green maturing to greenish-purple and brown in vivo, light to dark brown or blackish in sicco, with an excentric apicule; stipes 8–13 by 2 mm; fruiting receptacle 4–10 mm diam.; monocarps, stipes and receptacle glabrous. Seeds ellipsoid, light brown, shallowly pitted, ca. 11 by 6 mm, raphe sunken, regular.

Bolivia (La Paz), Peru (Cuzco, Madre de Dios).

Primary and secondary moist and wet forest, occasionally on floodplains. At elevations of 210–670 m. Flowering: August – December; fruiting: January – May, August and October.

A number of the specimens of Cremastosperma confusum have in the past been identified as C. leiophyllum or C. monospermum, two well known species found relatively nearby in Bolivia and widespread across Bolivia, Brazil and Peru, respectively. The distributions of C. confusum and C. leiophyllum do not overlap, but in the area of Madre de Dios and adjacent La Paz, Bolivia, C. monospermum and C. confusum may both occur and, in this area in particular, non-flowering specimens of the two are often not discernible. These species are closely related as well as morphologically similar and further data to test their boundaries and the potential for gene-flow between species/populations are warranted. Variation in the size, shape and texture of leaves and length of pedicel of C. confusum is relatively wide, with specimens collected in Cuzco in particular exhibiting larger leaves.

The species is named C. confusum because of the past confusion caused by its similarities to different other nearby species and the lack of unambiguous diagnostic characters for the fruiting material.

Cremastosperma confusum, found in southern Peru and the Las Paz region in northern Bolivia, has a fairly wide EOO and is not uncommon. It is also found within protected areas. Least concern [LC] (Table 1).

BOLIVIA. La Paz: Ixiamas, 13°08'34"S, 68°03'59"W, 18 October 2009, Couvreur 159 (L, NY); Ixiamas, 13°00'39"S, 68°04'17"W, 29 October 2009, Couvreur 262 (L, NY); Ixiamas, 13°59'16"S, 67°48'24"W, 29 October 2009, Couvreur 267 (L, NY). PERU. Cuzco: Camisea, Campamento San Martín-C, 11°47'08"S, 72°41'57"W, 467 m a.s.l., 12 January 1997, Acevedo-Rodríguez et al. 8635 (MO, P, U, USM); Prov. Cuzco (locality unknown), 11°47'00"S, 72°41'57"W, 467 m a.s.l., 26 Jan 1997, Acevedo-Rodríguez et al. 9132 (USM); La Convención, Manguyari, 12°47'S, 72°40'W, 670 m a.s.l., 3 Feb 1989, Núñez Vargas et al. 10146 (MO, U); Camanti, 13°13'S, 70°45'W, 643 m a.s.l., 18 Feb 1991, Núñez Vargas 12951 (U, USM); La Convención, Distr. Echarati, 11°41'S, 73°00'W, 350 m a.s.l., 18 Apr 1998, Núñez Vargas, et al. 21737 (USM); Camanti, Maniri, 13°17'S, 70°48'W, 720 m a.s.l., 15 Oct 1990, Timaná 1010 (MO); Camanti, 13°17'19"S, 70°46'27"W, 26 Feb 2007, Valenzuela Gamarra, L et al. 8643 (CUZ, HUT, MO, USM). Madre de Dios: Parque Nacional Manu, Cocha Cashu, 11°52'S, 71°22'W, 29 Sep 1976, Foster & Terborgh 5083 (US); Parque Nacional Manu, Tayakome, 11°41'S, 71°36'W, 350–400 m a.s.l., 28 Sep 1986, Foster & Achille 11495 (U, USM); Tambopata Reserve, Río Tambopata, 12°50'S, 69°17'W, 250 m a.s.l., 5 Mar 1981, Gentry et al. 32009 (MO); Shintuya-Salvación road, 12°40'S, 71°15'W, 500 m a.s.l., 14 May 1984, Knapp & Mallet 6450 (F, NY, US); Las Piedras, 12°36'23"S, 69°04'54"W, 17 Aug 2004, Suclli & Huamantupa 1943 (MO); Tambopata Reserve, Explorer’s Inn, 12°47'S, 69°41'W, 270 m a.s.l., 21 Sep 1998, Vásquez Chávez et al. 25605 (L, MO, MOL); Tambopata Reserve, 12°15'S, 69°17'W, 260 m a.s.l., 21 Nov 1984, Young & Stratton 219 (MO, U). Puno: Ridge between Río Candamo and Río Guacamayo, 13°30'S, 69°50'W, 400–600 m a.s.l., 22 May 1992, Gentry et al. 76947 (MO).

Map 6. 

Distribution of Cremastosperma confusum Pirie, C. leiophyllum R.E.Fr.; and C. oblongum R.E.Fr.

Figure 18. 

Cremastosperma confusum Pirie. a fruiting and flowering specimen b flower (aSmith, S. et al. 1578bSmith, S. et al. 794).

COLOMBIA, Antioquia: Frontino, Nutibara, upper watershed of Río Cuevas, 15 Jul 1986. Sánchez, D. et al. 415 (holotype: U [U0012253]; isotypes: COL! [COL000221555], MEDEL! [MEDEL000018, MEDEL000019]).

Tree 6–18 m tall, 15–22 cm diam.; young twigs and petioles sparsely covered with white-yellow appressed hairs 0.3–0.5 mm long. Leaves: petioles 3–6 by 1.5–2 mm, often with warts extending up primary vein; axillary buds densely covered with white-yellow appressed hairs 0.3–0.5 mm long; lamina narrowly elliptic to elliptic, 14–24.5 by 6–10 cm (index 1.6–3.1), chartaceous to subcoriaceous, mid-dark brown above, lighter below, glabrous above, sparsely covered with white-yellow appressed hairs 0.3–0.5 mm long (particularly on veins) below, base obtuse-acute (narrowly cuneate), apex acuminate (acumen 5–15 mm long), primary vein not conspicuously grooved, 1–2 mm wide at widest point, secondary veins (5–)7–9(–11), intersecondary veins occasional, distance between from 5 mm at the base to 30 mm closer to the apex, angles with primary vein 40–50° at the base to 50–70° closer to the apex, not branching, forming distinct loops in the apical half-two thirds, smallest distance between loops and margin 1.5–4 mm, tertiary veins percurrent with significant reticulation. Inflorescences of single flowers solitary or clustered in groups of 2 (or more), produced from leafy twigs or leafless branches; peduncles of two internodes, the second 1.2–4 by 1 mm (in flower), approx. 2 by 2 mm (in fruit); pedicels 28–47 by 1 mm at the base, 1–1.5 mm at the apex (in flower), 40–55 by 1.5–2 mm at the base, 2–2.5 mm at the apex (in fruit); peduncles and pedicels rather densely covered with white-yellow appressed hairs 0.3–0.5 mm long; two lower bracts (one on each internode), the apical one persisting later into flowering, 1–1.5 by 0.7–1 mm, broadly ovate, obtuse, soon falling off, rather densely covered with white-yellow appressed hairs 0.3–0.5 mm long; upper bract broadly to narrowly ovate, 1–3.5 by 1–2 mm, obtuse, persistent, densely covered with white-yellow appressed hairs 0.3–0.5 mm long; closed flower buds depressed ovoid; flowers green maturing to yellow in vivo, brown outside and black inside in sicco; sepals free, ovate, appressed, 3–4 mm long, obtuse, occasionally persistent on less mature fruit, densely covered with white-yellow appressed hairs 0.3–0.5 mm long; outer petals ovate to broadly so, 10–15 by 9–11 mm, inner petals ovate, 10–16 by 7–8 mm densely covered with white-yellow appressed hairs 0.3–0.5 mm long; receptacle ovoid to depressed ovoid; androecium 3–5 mm diam., stamens 1–1.2 mm long, connective appendage ca. 1 mm wide; gynoecium 1.5–2 mm diam., carpels 0.5–0.6 mm long, glabrous. Monocarps 10–20 (fully ripe fruit not observed), black in sicco, ellipsoid or narrowly so, 27–28 by 11–12 mm, with an excentric apicule (obvious only in immature fruit); stipes 17–19 by 1.5–2 mm; fruiting receptacle broadly ovoid, 4–5 mm diam.; monocarps and stipes sparsely covered with golden appressed hairs <0.1mm long or glabrous, receptacle often sparsely covered with white-yellow appressed hairs 0.3–0.5 mm long. Seeds ellipsoid to narrowly so, ca. 17 by 7 mm, dark brown and wrinkled, raphe sunken, encircling seed diagonally.

Colombia (Antioquia, northern and western foothills of the Cordillera Occidental; one specimen potentially representing the species from Riseralda [Betancur & al. 33011]).

Forest, at elevations of 1200–1500 m. Flowering: May and December; fruiting: May, July and December.

Cremastosperma dolichocarpum can be distinguished from other species of Cremastosperma by the unique long-ellipsoid monocarps after which the species is named and identified even when sterile by the conspicuous axillary buds with dense indument.

Most of the collections of Cremastosperma dolichocarpum are from two national parks (Parque Nacional Las Orquídeas and Parque Nacional Paramillo), but overall they represent a small number of populations within a restricted area. Endangered [EN] (Table 1).

COLOMBIA. Antioquia: Parque Nacional Las Orquídeas, 6°32'N, 76°14'W, 1450 m a.s.l., 13 Aug 1993, Cogollo et al. 6345 (MO); Parque Nacional Paramillo, Río San Jorge, 7°15'N, 75°55'W, 1560 m a.s.l., 3 Mar 1993, Gentry et al. 79039 (U); Urrao, 6°29'N, 76°14'W, 1300 m a.s.l., 11 Dec 1992, Pipoly et al. 16951 (MO).

Figure 19. 

Cremastosperma dolichocarpum Pirie. a flower b leaf and fruit (aCallejas 3110bSánchez 415).

Differs from most species of the genus in the greater length of the pedicels (ca. 170 mm in flower; 250 mm in fruit). Compared to C. longipes (), differs by the habit (shrub ca. 3m tall, compared with a tree >4.5 m tall), smaller leaves (up to 35 cm long, compared with 35–60 cm long) and lack of indument on all parts.

PERU, Pasco: Oxapampa, Distr. Palcazú, Communidad Nativa Alto Lagarto – Villa Progreso. Reserva Comunal Yanesha, 500 m a.s.l., 9 Feb 2009, Rojas, R. 6486 (holotype: USM; isotypes: HOXA, L! [L4318001], MO! [MO-2967424]).

Shrub ca. 3 m tall; young twigs and petioles sparsely covered with appressed hairs 0.1–0.2 mm long when very young, soon completely glabrous. Leaves: petioles 5–12 by 2–3 mm; lamina narrowly oblong-elliptic to oblong-elliptic, 22–38 by 11–16 cm (index 2–3), coriaceous, greyish above, brownish below, upper side glabrous, lower side glabrous except for some scattered hairs along primary nerve, base obtuse, apex acuminate (acumen ca. 10 mm long), extreme tip obtuse, primary vein raised over all of its length, 2–3 mm wide at widest point, glabrous, secondary veins 10–14, intersecondary veins absent, distance between from 8–10 mm at the base to 40 mm in the centre to 10 mm closer to the apex, angles with primary vein from 50–55° at the base to 45–50° closer to the apex, not branching, forming distinct loops, smallest distance between loops and margin 1–4 mm, tertiary veins percurrent. Inflorescence of single flowers, on older branches or potentially cauliflorous; combined short axillary shoot and pedicel 170–260 by 0.5–1 mm at the base, 2 mm at the apex (in flower), 250–260 mm by 0.5–1 mm at the base, ca. 2 mm at the apex (in fruit), short axillary shoot and pedicels glabrous; 1 lower bract, broadly triangular, ca. 1.5 by 2 mm, obtuse, persistent, glabrous but margins densely covered with brown hairs ca. 1 mm long; upper bract attached at ca. 1/3 from base, broadly triangular, ca. 1.5 by 2 mm, obtuse, glabrous, ciliate; closed flower buds not seen; flowers yellow tinged red in vivo, dark brown in sicco, glabrous; sepals free, depressed ovate, patent to reflexed, ca. 2 by 3.5 mm, obtuse, persistent, glabrous but margins covered with hairs ca. 0.1 mm long; petals grooved at base, outer petals narrowly elliptic to narrowly obovate, 25–30 by 7–10 mm, inner petals narrowly elliptic to narrowly obovate, 7–10 by 3–4 mm, glabrous; receptacle broadly oblongoid; intact androecium not observed, stamens 2–2.5 mm long, connective appendage ca. 1× 0.2 mm wide; gynoecium ca. 4 mm diam., carpels ca. 25, ca. 1 mm long, glabrous. Monocarps ca. 3, ellipsoid, obtuse, symmetrical, 20–22 by 15 mm, no obvious apicule, distally green to proximally reddish in vivo, blackish in sicco; stipes 19–21 by 1 to 2 mm subtending monocarp; fruiting receptacle oblongoid, ca. 5 mm diam., glabrous; monocarps, stipes and receptacle glabrous. Seeds not seen.

Peru (Pasco).

Primary forest remnants. At an elevation of ca. 500 m. Flowering and fruiting: February.

Cremastosperma dolichopodum is only known from two specimens collected at the same locality, but the length of the pedicel – a generally reliable character within the genus – greatly exceeds that of any similar known species.

The epithet “dolichopodum”, from the Greek dolicho (long) and podum (-stalked) refers to the unusual length of the pedicel.

Of the three known collections of Cremastosperma dolichopodum, two are from the same locality, within a protected area, the third from a roadside remnant of primary forest within the same region. Given the extent of collections of other species in the region (with regular documentation of the much earlier described and more widely distributed C. oblongum and C. pendulum) C. dolichopodum must be naturally rare as well of restricted distribution. Endangered [EN] (Table 1).

PERU. Pasco: Pozuzo, Alto Lagarto to Pozuzo Alto Victoria road, 10°07'09"S, 75°29'25"W, 1500 m a.s.l., 29 June 2008, Rojas & Ortíz 5758 (HOXA, MO, USM); Palcazú, Communidad Nativa Alto Lagarto, 10°08'00"S, 75°22'06"W, 500 m a.s.l., 9 Sep 2009, Rojas & Ortíz 6485 (HOXA, L, MO, USM).

Figure 20. 

Cremastosperma dolichopodum Pirie & Maas. Fruiting specimen (Rojas & Ortíz 6486).

PERU, Ost-Peru, Regenwald von Ost-Peru: Stromgebiet des Marañon von Iquitos aufwärtsbis zur Santiago Mündung am Pongo de Manseriche, oberer Marañon; unterhalb des Pongo de Manseriche, flutfreier Hochwald, 155 m a.s.l., 13 Dec 1924, Tessmann, G. 4748 (holotype: B! [B 10 0242370]; isotype: S! [S-R-6960]).

Tree or shrub 0.5–10 m tall; young twigs and petioles glabrous to sparsely covered with appressed brown hairs to 0.4 mm long. Leaves: petioles 2–8 by 1–2.5 mm; lamina elliptic to obovate or narrowly so, 11–28 by 3–10 cm (index 2–4(–4.7)), chartaceous, (pale) greyish- or brownish-green on both sides, often more greyish above, glabrous on both sides, base acute to obtuse or rounded, apex caudate (cusp 10–35 mm long), primary vein lightly grooved for basal third, 1–1.5 mm wide at widest point, more or less verrucose below, secondary veins 8–17, often 1–3 intersecondary veins, distance between from ca. 5 mm at the base to up to 25(-30) mm closer to the apex, angles with primary vein rather variable, from 45–80° at the base to 60–80° closer to the apex, forming distinct loops, smallest distance between loops and margin 2–6.5 mm, tertiary veins percurrent. Inflorescence of single, solitary flowers, on leafy twigs; peduncles 1–4 by ca. 1 mm (in flower), 2–5 by 1.5–2 mm (in fruit); pedicels (12–)15–25 by 1 mm at the base (in flower), 14–30 by 1–1.5 mm (in fruit), peduncles and pedicels rather densely covered with more or less erect brown hairs 0.2 mm long; 2 lower bracts, deltate, ca. 1 mm long, soon falling off, rather densely covered with more or less erect brown hairs 0.2 mm long; upper bract attached around midway along the pedicel, ovate or broadly so, 1–3 by ca. 1 mm, obtuse or acute, outer side sparsely to rather densely covered with appressed or erect whitish hairs to 0.2 mm long; closed flower buds depressed ovoid, opening loosely in development; flowers green to greenish-yellow, pale yellow or cream in vivo, dark brown with a lighter brown calyx in sicco, outer sides of sepals and petals sparsely to rather densely covered with erect or appressed whitish hairs to 0.2 mm long, inner sides glabrous to sparsely covered with appressed whitish hairs to 0.2 mm long (or inner petals papillate); sepals free, broadly ovate to deltate, recurved, 3–4 by 2.5–4 mm, obtuse, soon falling off; outer petals (broadly) elliptic to ovate, 9–15 by 7–12 mm, inner petals elliptic, obovate or narrowly so, 8–16 by 4–7 mm; androecium ca. 5 mm diam., stamens 1.2–1.5 mm long, connective appendage 0.7–0.8 mm wide; gynoecium ca. 2 mm diam., carpels ca. 25, ca. 2.2 mm long, sparsely covered with erect golden hairs 0.1 mm long. Monocarps 3–23, ellipsoid, slightly asymmetrical, 10–15 by 7–9 mm, with an excentric apicule, green maturing to pink or yellow through to red, purple and black in vivo, reddish or dark brown in sicco; stipes green maturing to pink or yellow to red in vivo, 7–17 by 1–1.5 mm, increasing to 3 mm diam. when mature; fruiting receptacle 3–8 mm diam.; monocarps, stipes and receptacle sparsely to rather densely covered with erect whitish hairs 0.1 mm long. Seeds ellipsoid, orange-brown, shallowly pitted, 5–8 by 3.5–6 mm, raphe sunken, regular.

Amazonian Colombia (Amazonas, Caquetá, Putamayo), Ecuador (Napo, Pastaza) and Peru (Loreto).

Primary forest, but also secondary, inundated and non-inundated forest. At elevations of 100–500 m. Flowering: January and April – August; fruiting: throughout the year.

Ecuador: Ambi cara caspi (Hurtado 3019); Ansuelo caspi muyo (Lawesson et al. 39560), Ayacara (Whitmore 871), Daycabome (Huaorani; M. Aulestia et al. 1726), Muncapatamo (Huaorani; Espinoza et al. 578).

Cremastosperma gracilipes most closely resembles C. microcarpum. The hairs on the flowers are shorter and less dense, which results in their drying a darker brown. The leaves are further generally distinctive in the shape of the apex (markedly caudate with an often long drip-tip) and in the green colour they consistently retain on drying. However, none of these characteristics is easy to define objectively or usefully and, although the geographic distributions of the two species are somewhat different (with C. microcarpum extending further into lowland Amazonia), there is apparent overlap. The species are closely related and further data to test their boundaries and the potential for gene-flow between species/populations is warranted. The leaves of C. longicuspe are similar to those of C. gracilipes, but in contrast to C. gracilipes, both flowers and fruit are entirely glabrous.

Cremastosperma gracilipes is one of the more widespread and abundant species of the genus, including occurrences in protected areas in Colombia and Ecuador, as well as in Peru. Least concern [LC] (Table 1).

COLOMBIA. Amazonas: Parque Nacional Amacayacu, 3°45'S, 70°15'W, 100 m a.s.l., 9 Aug 1989, Vásquez et al. 12675 (U). ECUADOR. Napo: Yasuní Forest Reserve, 0°40'S, 76°28'W, 240–310 m a.s.l., 27 Jun 1995, Acevedo-Rodríguez 7543 (US); Añangu, Río Napo, 0°30'S, 76°25'W, 300 m a.s.l., 9 Apr 1982, Balslev 2418 (QCA); Comuna San José de Payamino, 0°30'S, 77°18'W, 300 m a.s.l., 1 Dec 1983, Balslev et al. 4634 (AAU, NY); Punto Aguarico, 0°05'N, 76°59'W, 300 m a.s.l., 6 Apr 1980, Brandbyge et al. 30472 (AAU, MO, NY, U, WU); San Pablo de Los Secoyas, 0°15'S, 76°21'W, 300 m a.s.l., 7 Aug 1980, Brandbyge et al. 32589 (AAU, U); Reserva Biológica Jatun Sacha, 1°04'S, 77°36'W, 450 m a.s.l., 17 Jan 1987, Cerón 702 (MO, U); El Chuncho Floristic Reserve, 0°30'S, 77°01'W, 250 m a.s.l., 1 Oct 1987, Cerón et al. 2321 (MO); El Coca-Los Sachas Road, 0°25'S, 76°55'W, 250 m a.s.l., 8 Oct 1987, Cerón & Neill 2428 (MO, U); Parque Nacional Sumaco, 0°24'S, 77°23'W, 540 m a.s.l., 6 Nov 1996, Dik 1732 (MO); La Joya de los Sachas, 0°25'S, 76°37'W, 250 m a.s.l., 14 Sep 1992, Gudiño & Grefa 1787 (MO); San José de Payamino, 40 km W of Coca, 0°30'S, 77°20'W, 300–600 m a.s.l., 23 Apr 1984, Irvine 817 (F); Parque Nacional Yasuní, Añangu, 0°31'S, 76°23'W, 270 m a.s.l., 18 Feb 1983, Luna et al. 9057 (NY, U); Reserva Etnica Huaorani, 0°39'45"S, 76°40'00"W, 300 m a.s.l., Naranjo & B. Freire 493 (MO); Via Loreto, 4 km W of Río Payamino, 1°28'S, 77°02'W, 250 m a.s.l., 3 Aug 1986, Neill et al. 7195 (MO); Río Tiputini, 0°43'S, 76°57'W, 300 m a.s.l., 21 Jul 1991, Ollgaard et al. 99056 (AAU); Parque Nacional Yasuní, 0°26'S, 76°35'W, 250 m a.s.l., 7 Jun 1994, Pitman 195 (U); Comunidad Kichwa El Eden, 0°31'35"S, 76°05'40"W, 200 m a.s.l., D. Reyes & Carillo 691 (MO); Estación Científica Yasuní, 0°38'S, 76°30'W, 200–300 m a.s.l., 27 Aug 1995, Romoleroux & Grefa 1834 (U); Sector Huashito, 0°20'S, 77°05'W, 250 m a.s.l., 3 Nov 1989, Rubio 320 (MO); Río Pacuno, Bimbino, 0°40'S, 77°20'W, 300 m a.s.l., 19 Oct 1960, Whitmore 737 (U); INIAP-Payamino Experimental Station, 0°26'S, 77°01'W, 250 m a.s.l., 3 Sep 1986, Zaruma 581 (AAU, MO, NY, U). Pastaza: Pandanuque, S of oil well Villano 2 de ARCO, 1°28'S, 77°27'W, 550 m a.s.l., 30 Aug 1997, Alvarez et al. 2405 (MO); Pozo petrolero ‘Namoyacu’ de UNOCAL, 1°40'S, 76°57'W, 290 m a.s.l., 13 Nov 1990, Espinoza et al. 578 (MO); Pozo petrolero ‘Masaramu’ de UNOCAL, 0°44'S, 76°52'W, 400 m a.s.l., 1 May 1990, Gudiño 310 (MO); Pozo petrolero Villano 2 de ARCO, 1°25'S, 77°20'W, 400 m a.s.l., 1 Dec 1991, F. Hurtado 3019 (U); Lorocachi, 1°38'S, 75°58'W, 200 m a.s.l., 24 May 1980, J. Jaramillo et al. 30773 (AAU, U); Río Acaro or Challuayacu, 1°23'S, 77°25'W, 360 m a.s.l., 19 Jan 1998, Neill et al. 11095 (MO); Río Pastaza, 2°20'S, 76°55'W, 285 m a.s.l., 24 Jul 1980, Ollgaard et al. 35223 (AAU); Río Curaray, 1°30'S, 76°32'W, 230 m a.s.l., 3 Sep 1985, Palacios & Neill 783 (MO, U). PERU. Loreto: Oleoducto Secundario road betw. Bartra 1 & 4, 2°30'S, 75°45'W, 200 m a.s.l., 15 Sep 1979, C. Díaz & N. Jaramillo 1411 (U); Prov. Maynas, NE of Comun Serafin Filomeno, 4°08'S, 72°55'W, 120–130 m a.s.l., 7 May 1991, Grández et al. 2488 (MO); Distr. Nauta, “20 Enero”, 4°39'12"S, 73°49'18"W, 150 m a.s.l., 22 Jun 2006, Huamantupa & N. Smith 7746 (MO, WAG); Río Ampiyacu, Pebas and vicinity, 3°10'S, 71°49'W, 4 May 1977, Plowman et al. 7248 (GH); Prov. Requena, Distr. Tapiche, Quebrada Yanayacu, 6°15'49"S, 73°54'31"W, 15 Oct 2014, Ríos et al. 4422 (F); Andoas, 2°55'S, 76°25'W, 180 m a.s.l., 9 Sep 1983, Vásquez 4423 (MO); Yanamono Explorama Reserve, 3°30'S, 73°05'W, 108 m a.s.l., 25 Oct 1989, Vásquez & N. Jaramillo 13007 (MO, U); Allpahuayo-IIAP, 4°10'S, 73°30'W, 150–180 m a.s.l., 11 Jul 1991, Vásquez et al. 17376 (MO).

Figure 21. 

Cremastosperma gracilipes R.E.Fr. a fruiting specimen b flower (aHurtado 3019bPalacios 1651).

BOLIVIA, La Paz: Mapiri, San Carlos, 850 m a.s.l., 2 Dec 1926, Buchtien, O. 705 (holotype: B! [B 10 0242369]; isotypes: HBG, MO! [MO-0477531], S! [S-R-7017], US).

Annona nitida Ruiz & Pav., Anales Instit. Bot. Cavanilles 17: 429, t. 488. 1959. Non Martius (1841), nom. nud.

Guatteria rusbyi J.F.Macbr., Publ. Field Columbian Mus., Bot Ser. 4: 171. 1929.

Guatteria lucida Rusby, Mem. New York Bot. Gard. 7: 245. 1927. Non C. Presl.

BOLIVIA, Beni: Covendo 630 m a.s.l., 26 Aug 1921. White, O.E. 913 (holotype: NY! [NY00026024]).

Shrub or tree 3–20 m tall, 3–18 cm diam.; young twigs and petioles glabrous. Leaves: petioles 4–12 by 1–3(–4) mm; lamina obovate to elliptic or narrowly so, 12–28 by 4–9(–12) cm (index 2–3.9), chartaceous, often green or greenish-brown above and below, more greyish above with darker or reddish veins, glabrous on both sides, base acute to obtuse, apex acuminate (acumen 5–15 mm long), primary vein 1.5–3 mm wide at widest point, verrucose, secondary veins 7–13, intersecondary veins 1–6, distance between from 14–24 mm at the base to 10–25 mm closer to the apex, angles with primary vein from 60–80° at the base to 40–50° closer to the apex, not branching forming distinct loops, smallest distance between loops and margin 2–4(–6) mm, tertiary veins percurrent. Inflorescence of single flowers, solitary or clustered in groups of up to 4, on older, leafless twigs; peduncles 1–2 by 1–2 mm (in flower), 2–4 by 1.5–3 mm (in fruit), sparsely covered with appressed golden <0.1 mm long hairs or glabrous; pedicels 18–34 by 1–1.5 mm at the base (in flower), 18–34(–43) by 1–3 mm (in fruit), glabrous; 1–3 lower bracts, depressed ovate, ca. 0.5 by 1 mm, obtuse, soon falling off, glabrous; upper bract attached around midway along pedicel, ovate to broadly so, ca. 1.5 by 1 mm, obtuse, glabrous; closed flower buds depressed ovoid, opening early in development; flowers green maturing to yellow or creamy yellow in vivo, dark yellow, reddish-brown or dark brown in sicco; sepals free, very broadly ovate-triangular, recurved, 2–3 by 2–3 mm, obtuse, soon falling off, sepals and petals glabrous; outer petals elliptic, 12–15 by 8–11 mm, rounded, inner petals elliptic, 13–15 by 6–8 mm; androecium ca. 7 mm diam., pinkish in vivo, stamens 1.4–1.8 mm long, connective appendage ca. 0.8 mm wide; gynoecium ca. 2 mm diam., carpels 2–2.2 mm long, glabrous. Monocarps, stipes and receptacle glabrous, monocarps 6–30, ellipsoid, asymmetrical, 14–17 by 8–9 mm, green maturing to yellow, orange-red and red in vivo, black (reddish-brown when immature) in sicco, with an excentric apicule; stipes 16–26 by 1–1.5 mm; fruiting receptacle 4–9 mm diam. Seeds ellipsoid, light brown, pitted ca. 12 by 7 mm, raphe sunken, regular.

Bolivia (Beni, Cochabamba, La Paz, Santa Cruz).

Mostly in primary wet or moist forest, also in mildly disturbed areas, often on slopes or terraces, on sandstone soils. At elevations of 200–1000 m. Flowering: February, May, July through September, November and December; fruiting: February through August, November and December.

Chocolatillo (Meneces & Terceros 395), Chocolatillo Negro (D.N. Smith et al. 14006, 14058), Eye (Youras; Naessany 106), Quií-quií (Hinojosa & Seidel 1313).

Cremastosperma leiophyllum is the most southerly distributed species of the genus and. of the two found in Bolivia, the only endemic. Bud development in C. leiophyllum is open (as opposed to that of C. monospermum). It most closely resembles C. spec B (which is not found in Bolivia), from which it can best be distinguished by the characteristic shape (asymmetrical, the stipes thickening somewhat where they meet the monocarps) and colour (blackish) of the mature fruits when dried.

Although a regional endemic, Cremastosperma leiophyllum has been collected regularly over a faily broad area of Bolivia including two national parks. Least concern [LC] (Table 1).

BOLIVIA. Beni: Serrania del Pilón Lajas, Yucumo, 15°15'S, 67°00'W, 850–900 m a.s.l., 3 May 1991, Killeen et al. 3080 (USZ); Serranía del Pilón Lajas, Yucumo, 15°13'S, 67°03'W, 760–870 m a.s.l., 19 May 1989, D.N. Smith et al. 13253 (LPB, MO, U, USZ). Cochabamba: Puerto Aurora, Región del Chapare, 16°50'S, 65°10'W, 28 Jun 1989, Naessany 106 (LPB, U); Río Blanco, 17°12'S, 64°25'W, 240 m a.s.l., 31 May 2000, Seidel 3543 (LPB). La Paz: San Carlos, Sarampiuni, 13°00'S, 65°00'W, 500 m a.s.l., 12 Mar 1927, Buchtien 706 (B, S); Río Quiquibey, 15°29'S, 67°04'W, 1000 m a.s.l., 11 Nov 1990, M.A. Lewis 37979 (MO, U); Alto Beni, Santa Ana, 15°37'S, 67°25'W, 500 m a.s.l., 28 Jun 1990, Seidel et al. 2878 (LPB, U); Consata, 15°20'S, 68°31'W, 1300 m a.s.l., 15 Dec 1981, Solomon et al. 6606 (LPB, MO, U). Pando: Campamento Gomero Pingo de Oro, 11°31'36"S, 69°06'11"W, 21 Oct 1999, Paniagua 2119 (F). Santa Cruz: Velasco, N of Puerto Frey, 14°39'50"S, 61°09'33"W, 210 m a.s.l., 22 Sep 1995, P.F. Foster 230 (WAG); Ayacucho Forest Reserve, 17°00'S, 63°00'W, 240 m a.s.l., 14 Apr 1976, Meneces & Terceros 395 (MO, NY); Choré Forest Reserve, Río Ibabo, 16°35'S, 64°31'W, 180 m a.s.l., 16 Aug 1990, Neill & Quevedo 9334 (LPB, MO, U, USZ); Parque Nacional Amboró, 17°33'S, 63°44'W, 360 m a.s.l., 15 Nov 1991, I.G. Vargas et al. 1113 (F, LPB, NY, USZ).

Figure 22. 

Cremastosperma leiophyllum R.E.Fr. a fruiting specimen b flower (a, bSeidel & Schulte 2265)

Figure 23. 

a, b Cremastosperma leiophyllum R.E.Fr. a flower b fruit (Pirie et al. 2; photo LWC) c, d Cremastosperma megalophyllum R.E.Fr. c habit (Maas et al. 8577, photo: PJMM), fruit (Maas et al. 8595, photo PJMM) e–g C. microcarpum R.E.Fr. e fruit (Maas et al. 6281; photo PJMM) f, g flower (Maas et al 8222; photo PJMM) h–j C. monospermum (Rusby) R.E.Fr. h habit i leaves j flower bud (Pirie et al. 5; photos LWC) k, l C. oblongum R.E.Fr. k flower l fruit (Maas et al. 9148; photos PJMM) m–o C. osicola. m fruit n flower o leaf and flower with scale bar (photos Reinaldo Aguilar).

PERU, Loreto: Maynas, anno 1831. Poeppig, E.F. s.n. (lectotype (designated in Pirie, Kankainen & Maas, 2005): LE; isotype: S! [S-R-6962]).

Cremastosperma killipii R.E.Fr., Kongl. Svenska Vetenskapsakad. Handl. 24: 3, pl. 1 a-b. 1948.

PERU, Loreto: Yurimaguas, lower Río Huallaga, 135 m a.s.l., Aug-Sep 1929, Killip, E.P. & Smith, A.C 29020 (holotype: US! [US00104264], isotype: S! [S-R-6961]).

Tree or shrub 1.5–20 m tall; young twigs and petioles sparsely covered with appressed whitish or golden hairs to 0.2 mm long. Leaves: petioles 4–14 by 1–3 mm; lamina elliptic, obovate or narrowly so, 10–27 by 3–11 cm (index 1.5–5), chartaceous, green or greyish-green above, green or brownish-green below, glabrous above, very sparsely covered with appressed whitish hairs to 0.2 mm long particularly on veins below, base acute, rarely obtuse, apex caudate (cusp 20–35 mm long), primary vein 1–2 mm wide at widest point, secondary veins 7–15, intersecondary veins 0–1(–4), distance between from 7–15 mm at the base to 9–18 mm closer to the apex, angles with primary vein from 60–80° at the base to 40–50° closer to the apex, rarely branching, forming distinct loops, smallest distance between loops and margin 2–4 mm, tertiary veins more or less percurrent. Inflorescence of single flowers solitary (or clustered in groups of 2), on leafy twigs; peduncles 2–5 by ca. 1 mm (in flower), 4–10 by 1–2 mm (in fruit), sparsely to rather densely covered with appressed golden hairs to 0.2 mm long; pedicels 10–14 by 1–1.5 mm at the base (in flower), 11–20 by 1–2 mm (in fruit), sparsely covered with appressed golden hairs to 0.2 mm long or glabrous; 2 lower bracts of unequal dimensions, basal lower bract deltate, ca. 0.5 by 0.5 mm, acute, soon falling off, apical lower bract narrowly elliptic, ca. 1.5 by 0.5 mm, rounded, soon falling off, lower bracts sparsely covered with appressed golden hairs to 0.1 mm long or glabrous; upper bract attached near base or midway along pedicel, ovate, ca. 2 by ca. 1 mm, acute, sparsely covered with appressed golden hairs to 0.1 mm long or glabrous; closed flower buds not seen, buds opening loosely in development; flowers yellowish in vivo, brown in sicco; sepals free, deltate, appressed or recurved, 3–4 by 3–4 mm, acute, soon falling off, sepals and petals glabrous; outer petals broadly elliptic, 10–12 by 9–12 mm, inner petals broadly elliptic, ca. 11 by 10 mm; androecium not seen; gynoecium not seen. Monocarps 6–13(–36), ellipsoid, slightly asymmetrical, 12–13 by 8–10 mm, white, red, deep red or deep purple in vivo, reddish-brown to dark brown or black in sicco, with an excentric apicule when unripe; stipes 10–19 by 2 mm; fruiting receptacle 4–5(–9) mm diam.; monocarps, stipes and receptacle glabrous. Seeds ellipsoid, orange or reddish-brown, 8–9 by 5–6 mm, raphe sunken, regular.

Ecuador (Napo); Peru (Loreto, San Martín), most collections found in the basin of the Río Huallaga.

Primary tropical wet forest, on sandy or white sand soil. At elevations of 140–200 m. Flowering: August-October; fruiting: February-June, August, September and November.

Ecuador: Moncapatahue (Huaorani; M. Aulestia & Gonti 2072). Peru: Anonilla (J. Ruíz et al. 1748).

Cremastosperma longicuspe most closely resembles C. gracilipes, particularly in the shape of the leaf. However, in contrast to C. gracilipes, the fruits and flowers are glabrous. From the limited material available, the flower buds appear to be loosely, rather than widely open. In describing C. killipii, Fries (1948) noted its similarity to C. longicuspe. The leaves of the type specimen of C. killipii are unusually broad, but leaf and other characters otherwise fall within the variation found in C. longicuspe, including the notable caudate apex.

Cremastosperma longicuspe is known from a scattering of collections across a reasonably wide area; none within protected areas and with no new collections of the species since September 2000. Given the low AOO and ongoing decline in habitats in the region, it is assigned: Vulnerable [VU] (Table 1).

ECUADOR. Napo: Reserva Etnica Huaorani, 0°55'S, 76°09'W, 250 m a.s.l., 20 Mar 1994, Aulestia & Gonti 2072 (QCNE, U). PERU. Loreto: Río Huasaga, 3°20'S, 76°20'W, 185 m a.s.l., Lewis et al. 11166 (USM); Puranchim, Río Sinchiyacu, 2°50'S, 76°55'W, 200 m a.s.l., 21 Nov 1986, Lewis 12186 (MO, USM); Andoas, 2°55'S, 76°25'W, 180 m a.s.l., 3 Nov 1983, Vásquez & N. Jaramillo 4559 (MO, U). San Martín: Santa Rosa de Davidcillo, Tarapoto-Yurimaguas road, 6°16'S, 76°17'W, 200 m a.s.l., 22 Apr 1986, Knapp & Mallet 7178 (F, U, USM); Shapaja-Chazuta road, Río Huallaga, 6°36'S, 76°10'W, 250–300 m a.s.l., 4 Aug 1986, Knapp 7864 (F, MO, NY, U, USM); Tarapoto-Yurimaguas Road, km 46, 6°24'S, 76°18'W, 350 m a.s.l., 30 Aug 1986, Knapp & Mallet 8203 (USM).

Figure 24. 

Cremastosperma longicuspe R.E.Fr. a fruiting specimen b flower (avan der Werff 10161bPoeppig s.n.).

COLOMBIA, Chocó: San José del Palmar, mouth of Río Torito (tributary of Río Hábita), west slope, 3 Mar 1980, Forero, E. et al. 6576 (holotype: COL! [COL000334460]; isotype: MO! [MO-047638]).

Tree 4.5–15 m tall; young twigs and petioles black, verrucose, sparsely to rather densely covered with white-golden appressed hairs ca. 0.4 mm long. Leaves: petioles 10–15 mm long, 2.5–7 mm diam.; lamina narrowly elliptic to elliptic, 35–60 by 10–25 cm (leaf index 2.3–3), chartacous to subcoriacous, olive/dark brown above, lighter below, glabrous above, sparsely covered with white-golden appressed hairs ca. 0.4 mm long on veins below (densely so developing leaves), base acute, apex acuminate (acumen 10–15 mm long), primary vein deeply grooved in basal half, 2–6 mm wide at widest point; secondary veins 10–16, intersecondary veins rare, distance between from 10 mm at the base to 80 mm closer to the apex, angles with primary vein 45° at the base to 60–70° closer to the apex, not branching, forming distinct loops in the apical half to third of the leaf, smallest distance between loops and margin 3–4 mm; tertiary veins mainly percurrent. Inflorescences of single, pendulous flowers, produced from leafless branches; peduncles 5–8 by ca. 1 mm (in flower), ca. 4 by 2 mm (in fruit); pedicels 90 (less mature) – 210 by 1 mm at the base, 1.5 at the apex (in flower), ca. 240 by 2 mm at the base, 3 mm at the apex (in fruit); peduncles and pedicels sparsely to rather densely covered with white-golden appressed hairs ca. 0.4 mm long; single lower bract, broadly elliptic, 1–2 by ca. 1 mm, acute, soon falling off, densely covered with white-golden appressed hairs ca. 0.4 mm long; upper bract attached on lower half of pedicel, elliptic, 1.5–3 by ca. 1 mm, acute, densely covered with white-golden appressed hairs ca. 0.4 mm long; closed flower buds not seen; flowers green (immature) in vivo, medium brown in sicco; sepals free, triangular to broadly trullate, appressed, 3–4.5 mm long, acute, soon falling off, rather densely to densely covered with white-golden appressed hairs ca. 0.4 mm long; outer petals elliptic, ca. 22 by 12 mm, inner petals narrowly elliptic, ca. 22 by 6 mm, sparsely to rather densely covered with white-golden appressed hairs ca. 0.4 mm long; stamens ca. 1.2 mm long, connective appendage ca. 1 mm wide. Monocarps ca. 20, black in sicco, ellipsoid, slightly asymmetrical, ca. 20 by 12 mm, without an apicule; stipes ca. 25 by 2 mm; fruiting receptacle ovoid, 8 mm diam.; monocarps and stipes glabrous, receptacle sparsely covered with white-golden appressed hairs ca. 0.4 mm long. Seeds ellipsoid, 18–20 by 8–9 mm, orange/brown, with many shallow pits, raphe slightly raised, encircling seed longitudinally.

Pacific coast of Colombia (Chocó, Riseralda) and Ecuador (Esmeraldas).

Humid lowland to premontane forest. At elevations of 280–1400 m. Flowering: January and March; fruiting: September.

Cremastosperma longipes can easily be distinguished from most other species of the genus by the exceptional length of the pedicel, after which the species is named. The flowers and fruits of most species of Cremastosperma are borne on pedicels less than 50 mm long, with rare exceptions such as C. pedunculatum and C. bullatum never exceeding 150 mm in length, significantly shorter than those of C. longipes. The only species with pedicels of a comparable length is the newly described C. dolichopodum, which differs from C. longipes in the lack of indument on the flowers and receptacle. In addition, leaves of C. longipes are unusually large, equalling the maximum dimensions observed in C. megalophyllum, a more densely collected species from Amazonian Colombia, Ecuador and Peru.

Cremastosperma longipes is only known from three collections from widely spaced localities outside of protected areas. Vulnerable [VU] (Table 1).

COLOMBIA. Riseralda: Geguadas-Puerto de Oro road, 800–1400 m a.s.l., 13 Sep 1991, J.L. Fernández et al. 8872 (COL). ECUADOR. Esmeraldas: Fila de Bilsa, 0°37'N, 79°51'W, 280 m a.s.l., 30 Jan 1991, Gentry et al. 72995 (F, MO).

Figure 25. 

Cremastosperma longipes Pirie. a monocarp (with stipe) and seeds b leaf, flower and flower bud (aFernández 8872bForero et al. 6576).

VENEZUELA, Falcón: Sierra de San Luis, above Santa María, 1300 m, 26 Jul 1979, van der Werff, H.H. & Vera, B., Flora Falcón 937 (holotype: U! [U0000246]).

Tree 5–10 m tall; young twigs and petioles glabrous. Leaves: petioles 3–10 by 1–3 mm; lamina narrowly elliptic to elliptic, 10–30 by 4–9(12) cm (index 2.3–3.7), chartaceous, green, brownish-green or brown on both sides, darker above, glabrous on both sides, base obtuse to rounded, apex acuminate (acumen 5–10 mm long), primary vein grooved in basal half, 1–2 mm wide at widest point, verrucose, secondary veins 6–12, intersecondary veins 1–2, distance between from 7–15 mm at the base to 7–23 mm closer to the apex, angles with primary vein from 60–80° at the base to 45–60° closer to the apex, not branching, forming mostly distinct loops, smallest distance between loops and margin 1–5 mm, tertiary veins mostly percurrent. Inflorescence of single flowers, on leafy twigs; peduncles ca. 1 by 0.5–1 mm (in flower), 1–2 by 1–2 mm (in fruit), sparsely covered in appressed golden hairs to 0.1 mm long or glabrous; pedicels 35–45 by 0.5–1 mm at the base up to 2.5 mm at the apex (in flower), 40–65 by 1–1.5 mm at the base up to 5 mm at the apex (in fruit), glabrous; single lower bract, depressed ovate, ca. 0.5 by 1 mm, rounded, soon falling off, rather densely covered with appressed golden hairs to 0.1 mm long; upper bract at or near base of pedicel, ovate, ca. 1.5 by 0.8 mm, obtuse, glabrous; closed flower buds not seen; flowers pale greenish-yellow with green base or cream-coloured in vivo, black in sicco, sepals and petals glabrous; sepals free, broadly to depressed ovate, recurved, 1.5–2 by 1.5–2 mm, obtuse, soon falling off; outer petals ovate, 11–14 by 8–9 mm, obtuse, inner petals obovate, 15–16 by ca. 7 mm, obtuse; androecium ca. 6.5 mm diam., gynoecium not seen. Monocarps 7–18, ellipsoid, slightly asymmetrical, 18–24 by 12–14 mm, very small strongly excentric apicule, green maturing to yellowish-brown, brown or purple-black in vivo, reddish-brown or dark brown in sicco; stipes 7–14 by ca. 2 mm; fruiting receptacle 5–8 mm diam.; monocarps, stipes and receptacle glabrous. Seeds ellipsoid, orange-brown, shallowly pitted, ca. 20 by 8 mm, raphe sunken, regular.

Venezuela (Carabobo, Falcón, Yaracuy).

Primary or secondary evergreen cloud forest. At elevations of 700–1500 m. Flowering: July; fruiting: March, May-July, October and December.

One of only two species of Cremastosperma found in Venezuela, C. macrocarpum can most easily be distinguished from C. venezuelanum by its smaller leaves (10–30 cm as opposed to 30–53 cm long) and longer pedicels (40–65 mm as opposed to 16–22 mm in fruit).

Cremastosperma macrocarpum is known from seven collections representing five localities outside of protected areas, the most recent having been collected in 1991. Endangered [EN] (Table 1).

VENEZUELA. Carabobo: Río Morón valley, 10°17'N, 68°10'W, 700–1100 m a.s.l., 3 May 1991, W. Diaz & Niño 274 (NY, U). Falcón: Cerro Galicia, 1500 m a.s.l., 10 Jun 1978, T. Ruíz Z. et al. 3499 (U). Yaracuy: Río Carabobo, 10°26'N, 68°49'W, 800–1200 m a.s.l., 31 Mar 1980, Liesner & A. González 9763 (MO, U); San Felipe, 10°15'N, 68°29'W, 1200 m a.s.l., 7 Dec 1980, Steyermark 123804 (U).

Figure 26. 

Cremastosperma macrocarpum Maas. a fruiting specimen b flower (a, bvan der Werff & Vera, Flora Falcón 937).

COLOMBIA, Antioquia: San Luis, Medellín-Bogotá highway, 8.1 km E of bridge over Río Caldera, 13 Mar 1983, Escobar, L.A. de & Folsom, J.P. 3309 (holotype: NY!, 2 sheets [NY00759135, NY00759134]; isotypes: HUA! [HUA0000058], U! [U0012249]).

Tree 3–7 m tall; young twigs glabrous. Leaves: petioles 6–14 mm long, 2–3 mm diam.; lamina narrowly elliptic to elliptic, 20–28 by 7–9 cm (index 2.5–3.1), chartaceous to subcoriaceous, olive to more lime green or brown above, darker below, glabrous on both sides, base obtuse to acute, apex acuminate (acumen 5–10 mm long), primary vein deeply grooved in basal 1/2 – 3/4, occasionally verrucose below, ca. 2 mm wide at widest point, glabrous, secondary veins 9–14, often 2 or 3 intersecondary veins, distance between from 4–5 mm at the base to 20–35 mm closer to the apex, angles with primary vein from 40–50° at the base to 70–80° closer to the apex, occasionally branching, occasionally forming more or less indistinct loops in the apical half, smallest distance between loops and margin 2–4 mm, tertiary veins rather reticulate. Inflorescences of single flowers, solitary or clustered in groups of two, axillary on leafy twigs; peduncles 2–3 by ca. 2 mm (in fruit); pedicels ca. 7 by 1 mm at the base, 1.5 mm at the apex (in flower), 16–20 by 1.5–2 mm at the base, ca. 3 mm at the apex (in fruit), peduncles and pedicels glabrous, two lower bracts, the apical one depressed triangular, ca. 1.5 by 2 mm, acute, persistent, glabrous, upper bract attached around midway along pedicel, broadly ovate to deltate, 0.5–2.5 by 1–2.5 mm, actute to obtuse, glabrous; closed flower buds not seen; flowers in vivo immature light green, black in sicco, sepals and petals glabrous; sepals fused for basal 1 mm, broadly to very broadly ovate, appressed, (2–) 5–7 by (2–)–5 mm, acute, mostly persistent, outer petals elliptic, ca. 12 by 7 mm, inner petals narrowly elliptic, ca. 12 by 5 m; androecium 2.5–2.7 mm diam., stamens ca. 0.7 mm long, connective appendage 0.3–0.4 mm wide. Monocarps 20–30, globose, symmetrical, 12–13 by 12–13 mm, green maturing to red in vivo, black in sicco, with a slightly excentric, 0.25 mm long apicule; stipes (immature) 9–10 by 1–1.5 mm; fruiting receptacle depressed ovoid, 7–9 mm diam. (only immature fruits seen); monocarps, stipes and receptacle glabrous. Seeds globose, shallowly pitted with a papery outer layer, ca. 13 by 11 mm, orange-brown, raphe neither raised nor sunken, regular.

Colombia (Antioquia, west side of the Magdalena valley).

Disturbed primary or secondary forest. At elevations of 670–1200 m. Flowering and fruiting: March.

Cremastosperma magdalenae can be distinguished from other species of the genus by the combination of globose monocarps and large sepals which mostly persist into fruiting (one slightly differing collection, Cárdenas 2899, displays immature fuits with smaller sepals only persistent on one of the two duplicates studied). Also noteworthy are the relatively short pedicels and the absence of indument on all parts. The absence of hairs on fruits and flowers reveals the blackish colour typical of specimens of Cremastosperma upon drying. Both C. panamense and C. pacificum (a species found on the Pacific coast of Colombia) also lack indument, but, amongst other differences, the sepals of both species are much smaller and do not persist into fruiting.

Cremastosperma magdalenae is known from just four collections representing similar non-protected localities, three close to the highway between Medellín and Bogotá. Endangered [EN] (Table 1).

. COLOMBIA. Antioquia: Al Prodigio, 6°06'N, 74°48'W, 350 m a.s.l., 26 Jun 1990, Cárdenas et al. 2899 (COL, MO); Mun. San Luis, Medellín-Bogotá road, Río Samaná, 6°00'N, 74°50'W, 670 m a.s.l., 19 Mar 1982, Hernandez 251 (COL, HUA); Mun. San Luis, Medellín-Bogotá road, Río Caldera, 6°00'N, 74°50'W, 1000 m a.s.l., 13 Mar 1983, Juncosa et al. 736 (MO).

Figure 27. 

Cremastosperma magdalenae Pirie. a flower b fruiting twig and cross section of seed (aHernandez 251bEscobar & Folsom 3309).

PERU, Loreto: Boca de Pebas, 23 Oct 1927, Ducke, A. RB19620 (holotype: S! [S-R-6963]; isotypes: B! [B 10 0242368], RB! [RB00534080, RB00534176]).

Tree or shrub 3–15 m tall, 2–15(–35) cm diam.; young twigs and petioles glabrous. Leaves: petioles 5–22 by 1.5–9 mm; lamina obovate to elliptic or narrowly so, 13–57 by 3–26 cm (index 2–5.1), chartaceous to coriaceous, shiny on both sides, secondary veins often impressed above (giving slightly bullate appearance), (dark) greyish-green or brown above, more brown or green below, glabrous on both sides, base obtuse to rounded, rarely acute or decurrent, apex acuminate (acumen (5-)10–30 mm long), primary vein conspicuously grooved in basal half, 1–5 mm wide at widest point, secondary veins (5–)8–21, intersecondary veins often 1–2(–3), distance between from 5–25 mm at the base to 9–35(–43) mm closer to the apex, angles with primary vein from (30–)45–70(–80)° at the base to (30–)40–75° closer to the apex, rarely branching, forming distinct loops, smallest distance between loops and margin 1–5 mm, tertiary veins percurrent. Inflorescence of single flowers, solitary or clustered in groups of up to 3, on leafy or leafless twigs or on the main trunk; short axillary shoot, 4–7(–8) by 0.7–1 mm (in flower), 4–10 by 1.5–4 mm (in fruit), glabrous or sparsely covered with whitish appressed hairs to 0.1 mm long; pedicels 10–20(–32) by ca. 1 mm at the base, up to 2(–2.5) mm at the apex (in flower), 15–30(–40) by 1.5–3 mm at the base, up to 4 mm at the apex (in fruit), glabrous; 3 lower bracts, deltate, ca. 1 by 1 mm, acute, soon falling off, sparsely covered with whitish appressed hairs to 0.1 mm long or glabrous; upper bract attached in apical half of pedicel, broadly ovate, 1–3 by 1–2 mm, obtuse, glabrous; closed flower buds depressed ovoid, opening loosely in development; flowers green maturing to yellow in vivo, black in sicco, sepals and petals glabrous; sepals free or connate for 1 mm, broadly ovate to triangular, appressed, open and conspicuous whilst petals still closed in young buds, 4–6 by 4–6 mm, obtuse to acute, sometimes briefly or partly persistent; outer petals broadly elliptic, 11–18 by 9–15 mm, obtuse, inner petals obovate, 10–16 by 5–7 mm, obtuse; androecium diam. unknown, stamens 1.3–1.8 mm long, connective appendage 0.7–1 mm wide; gynoecium diam. unknown, carpels 1.5–2 mm long. Monocarps 6–32, ellipsoid to broadly ellipsoid, (slightly) asymmetrical, 12–20 by 9–14 mm, green maturing to yellow, orange, purple and black in vivo, reddish or dark brown or black in sicco, often with an excentric apicule; stipes 10–30 by 1–2 mm; fruiting receptacle 3–12 mm diam.; monocarps, stipes and receptacle glabrous. Seeds broadly ellipsoid, reddish-orange, pitted, ca. 12 by 9–10 mm, raphe raised (more so when seeds immature), regular.

Amazonian Colombia (Caquetá, Putamayo), Ecuador (Morona-Santiago, Napo, Pastaza, Sucumbíos, Zamora-Chinchipe) and Peru (Amazonas, Loreto).

Primary and secondary premontane or lowland rainforest, sometimes inundated, on red (oxisols/lateritic) or sometimes volcanic soils. At elevations of 100–1200 m. Flowering: April-June, September-January; fruiting: throughout the year.

Ecuador: Ayacara (Quichua; A. Alvarado 298), Cucha casa caspi (Hurtado 3005), Caramoyu (Whitmore 854), Ichilla cara caspi (small bark tree; D. Irvine 374), Lynshtimoia (Quechna; Whitmore 717), Mandachi (Shuar; W. Palacios 10278), Oñetahue (the plant itself), Oñetahuemo (the fruit) (Huaorani; J.S. Miller et al. 794), Tara caspi (Quichua; Grijalva & Grefa 305) or T’zinytala (Quichua; W. Palacios 10278). Peru: Bara (Rimachi Y. 1027).

Despite its name, the size of leaves of Cremastosperma megalophyllum varies from large to relatively small with respect to those of other species of the genus. This variation is also apparent in the size of the fruits, which are similar to those of C. napoense, but which in contrast are never borne on a branching inflorescence. C. megalophyllum is best distinguished by the shape of the glabrous, black-drying flowers: the large sepals open earlier and to a greater extent than the petals (in contrast to those of C. napoense, bud development of which is open from an early stage and the lighter colour (particularly of the pedicel) of which indicates the presence of indument).

Cremastosperma megalophyllum is one of the more widespread and abundant species of the genus, including occurrences in protected areas in Ecuador. Least concern [LC] (Table 1).

COLOMBIA. Caquetá: Mocoa, 2°00'N, 76°00'W, 4 May 1899, Sprague 351 (K). ECUADOR. Morona-Santiago: Misión Salesiana-Shuar, 3°25'S, 78°35'W, 800 m a.s.l., 8 Jun 1986, Zaruma & Arguello 518 (MO, U). Napo: Yasuní Forest Reserve, 0°40'S, 76°22'W, 200 m a.s.l., 25 Jun 1995, Acevedo-Rodríguez et al. 7523 (US); Hollín-Loreto road, foothills of Volcán Sumaco, 0°44'S, 77°28'W, 600 m a.s.l., 26 Jun 1989, Alvarado 298 (AAU, GB, MO, NY, U); San Pablo de Los Secoyas, 0°15'S, 76°21'W, 300 m a.s.l., 6 Aug 1980, Brandbyge et al. 32566 (AAU, U); Dureno, Comunidad Indigena Cofan, 0°02'S, 76°42'W, 350 m a.s.l., 22 Sep 1986, Cerón 404 (AAU, MO, U); Reserva Biológica Jatun Sacha, 1°04'S, 77°36'W, 450 m a.s.l., 17 Jan 1987, Cerón 612 (MO, QAME, U); Galeras Cordillera, E of Río Pusuno, 0°52'55"S, 77°34'15"W, 900–1200 m a.s.l., 19 Jun 2009, Couvreur et al. 130 (L); La Joya de los Sachas, 0°25'S, 76°37'W, 220 m a.s.l., 23 Nov 1992, A. Grijalva & Grefa 305 (U); Río Suno, W of Río Napo, 0°42'S, 77°10'W, 400 m a.s.l., 23 Jun 1968, Holm-Nielsen et al. 903 (AAU); San José de Payamino, 40 km W of Coca, 0°30'S, 77°20'W, 300–600 m a.s.l., 3 Nov 1983, Irvine 374 (F); Reserva Etnica Huaorani, 0°39'45"S, 76°40'00"W, 300 m a.s.l., Naranjo & B. Freire 399 (MO); Cañón de los Monos, 15 km N of Coca, 0°20'S, 77°01'W, 250 m a.s.l., 5 Apr 1985, Neill et al. 6359 (MO, QAME, U); Coca-Loreto road. 10 km W of Río Payamino, 0°35'S, 77°10'W, 275 m a.s.l., 10 Oct 1987, Neill et al. 7883 (MO, U); INIAP-Payamino Experimental Station, 0°25'S, 77°10'W, 250 m a.s.l., 18 Nov 1991, Neill et al. 10005 (MO); Coca-Loreto road, 8 km after Loreto, 0°35'S, 77°20'W, 800 m a.s.l., 8 Jun 1987, Palacios 1620 (MO, U); Río Payamino, road to Loreto, 0°26'S, 77°06'W, 300 m a.s.l., 15 Dec 1987, Palacios 2311 (MO, U); Puerto de Misahuallí, 1°04'S, 77°37'W, 450 m a.s.l., 7 Sep 1988, Palacios 2937 (U); El Chuncho Floristic Reserve, 0°25'S, 77°01'W, 250 m a.s.l., 23 May 1993, Palacios 10663 (U); Via Los Zorros, Sol Naciente, 0°40'S, 77°07'W, 250 m a.s.l., Aug 1993, Palacios 11016 (U); Río Napo, 20 km W of Coca, 0°35'S, 77°03'W, 300 m a.s.l., 22 Apr 1985, Stein et al. 2577 (AAU, MO, NY, U); Río Tutapishco, 0°36'S, 77°22'W, 480 m a.s.l., 27 Jan 1996, H. Vargas & Cerda 671 (U); Parque Nacional Sumaco, 0°08'S, 77°08'W, 400 m a.s.l., 30 Sep 1996, H. Vargas & Alvarado 1025 (MO); Estación Científica Yasuní, 0°38'S, 76°30'W, 200–300 m a.s.l., 24 Apr 2002, Villa et al. 1499 (F, U); Ridge W of Hac. Cotapino, parallel to Río Cotapino, 0°40'S, 77°20'W, 360 m a.s.l., 18 Oct 1960, Whitmore 717 (K); Payamino-Loreto road, 4–6 km from R Areas, 0°26'S, 77°02'W, 250 m a.s.l., 13 Sep 1986, Zaruma 657 (MO); Parque Nacional Yasuní, 0°31'S, 76°32'W, 240 m a.s.l., 6 Mar 1993, Zuleta 175 (U). Pastaza: Pozo petrolero ‘Masaramu’ de UNOCAL, 0°44'S, 76°52'W, 390 m a.s.l., 1 May 1990, Espinoza 129 (MO, NY, U); Río Landayacu, 1°34'S, 77°25'W, 580 m a.s.l., 25 Nov 1990, Gudiño 1123 (U); Curaray, NE of Destacamento, 1°21'S, 76°56'W, 250 m a.s.l., 19 Mar 1980, Holm-Nielsen 22082 (AAU, U); Pozo petrolero Villano 2 de ARCO, 1°25'S, 77°20'W, 400 m a.s.l., 1 Dec 1991, F. Hurtado 3005 (U); Río Tigüeno, 1°16'S, 77°11'W, 200 m a.s.l., 4 Jun 1995, J.S. Miller et al. 794 (U); Pozo Petrolero Villano 2, 1°29'S, 77°27'W, 500 m a.s.l., 24 Jul 1992, Palacios 10278 (U); Vía Auca, 115 km S of Coca, 6 km S of Río Tigüino, 1°15'S, 76°55'W, 320 m a.s.l., 31 Mar 1989, Zak & Rubio 4202 (MO, U). Sucumbíos: Rio Aguarico, Zabalo village, 0°21'24"S, 75°39'56"W, 22 Nov 1998, Aguinda et al. 458 (F); Campo Bermejo 6 Norte, 0°14'S, 77°13'W, 1050 m a.s.l., 23 Mar 1990, Cerón et al. 9324 (U); Reserva Cuyabeno, 0°00'S, 76°12'W, 265 m a.s.l., 1 Apr 1988, R. Valencia et al. 67352 (AAU, U); Dureno, 0°02'07"S, 76°45'17"W, 250 m a.s.l., 31 May 2007, Vriesendorp 428 (L). Zamora-Chinchipe: Jamboe Bajo, 4°05'S, 78°55'W, 1100 m a.s.l., 3 Nov 1996, Clark et al. 3158 (U); Cordillera del Condór, 4°26'14"S, 78°37'12"W, 900–1400 m a.s.l., 19 Jun 2005, Quizhpe et al. 1505 (MO, U). PERU. Amazonas: Atalaia do Norte. Río Javari, 4°33'S, 71°40'W, 2 Jan 1989, Cid Ferreira et al. 9959 (NY). Loreto: Río Marañón, San Rafael, 3°46'S, 73°03'W, 11 Sep 1989, Daly et al. 6186 (MO, NY, U); Alto Amazonas, 2°55'S, 76°25'W, 210 m a.s.l., 6 Jun 1981, Vásquez & N. Jaramillo 1969 (MO); Miraflores, Quebreda Tamshiyacu, 4°15'S, 72°40'W, 200 m a.s.l., 24 Feb 1986, Vásquez & N. Jaramillo 7237 (MO, U); Yanamono Explorama Reserve, 3°30'S, 72°50'W, 106 m a.s.l., 28 Sep 1988, Vásquez 11076 (F, U); Iquitos-Nauta road, 4°30'S, 73°30'W, 130 m a.s.l., 8 Nov 1988, Vásquez et al. 11192 (MO, U); Las Amazonas, ExplorNapo Camp, 3°15'S, 72°54'W, 140 m a.s.l., 27 Jun 1991, Vásquez et al. 16885 (MO, U).

Map 7. 

Distribution of Cremastosperma megalophyllum R.E.Fr.; C. microcarpum R.E.Fr.; C. pedunculatum (Diels) R.E.Fr.; and C. peruvianum R.E.Fr.

Figure 28. 

Cremastosperma megalophyllum R.E.Fr. a fruiting specimen b flower buds (aZuleta 175bPalacios 3270).

BRAZIL, Amazonas: Mun. Humaitá, Rio Madeira, near Tres Casas, 18 Sep 1934, Krukoff, B.A. 6151 (holotype: S! [S-R-6964]; isotypes: A! [00039263], F! [V0054581F], G! [G00237254], GB! [GB-0047034], K! [K000485529], MICH! [MICH1286075], MO! [MO-047716], NY! [NY00025862], U! [U0000248], US! [US00811272, US00901600]).

Tree or shrub 2–20 m tall, 2.5–18.5 cm diam.; young twigs and petioles glabrous to rather densely covered with appressed and/or erect white or golden hairs 0.3–0.4 mm long. Leaves: petioles 2–12 by 1.5–3 mm; lamina narrowly elliptic to elliptic or narrowly obovate, 13–31 by 4–12 cm (index 1.8–3.8(–4.8)), chartaceous, green, greyish- or brownish-green or brown on both sides, shiny above, venation below often yellowish, glabrous above, glabrous to rather densely covered with appressed or erect white hairs to 0.2 mm long at the base and on primary and secondary veins below, base acute to obtuse, rarely rounded or narrowly cuneate, apex acuminate (acumen 10–40 mm long), primary vein 1–2 mm wide at widest point, more or less verrucose on both sides, lightly grooved for around half of length, secondary veins 7–15, often 1–4 intersecondary veins, distance between from ca. 5 mm at the base to ca. 20 mm closer to the apex, angles with primary vein mostly from 45–60° at the base to 60–80° closer to the apex, not branching, forming distinct loops, smallest distance between loops and margin 2–7 mm, tertiary veins percurrent. Inflorescence of single flowers, solitary or clustered in groups of up to 3, on leafy or leafless twigs; peduncles (3–)5–15 by ca. 1 mm (in flower), 4–15 by 1–2 mm (in fruit); pedicels (5–)12–24 by ca. 1 mm at the base (in flower), 10–25 by 1–2 mm (in fruit), peduncles and pedicels sparsely to rather densely covered with appressed or erect whitish hairs to 0.3 mm long; 1 to several lower bract(s), the basal-most small and scale-like, those more apical mostly (long) elliptic, occasionally leaf-like, 2–6(–60) by ca. 1 mm, acute, soon falling off, rather densely covered with appressed white hairs to 0.3 mm long; upper bract attached mostly on the basal half of the pedicel, ovate to deltate, 1.5–2.5 by 1–1.5 mm, obtuse or acute, sparsely to rather densely covered with appressed or erect golden hairs to 0.3 mm long; closed flower buds very broadly to depressed ovoid, opening loosely in development; flowers green, maturing to brown, (pale) yellow, cream or white outside, cream or yellow inside, sepals green or dark brown outside, green with a pink base inside in vivo, pale (orange-) brown or brown with dark or reddish-brown base in sicco, sepals and petals rather densely to densely covered with appressed or erect golden hairs (whitish close to the edges) to 0.3 mm long; sepals free or connate for 0.5 mm, broadly ovate to deltate, not reflexed, 3–4[-6] by 2.5–4[-6] mm, obtuse, soon falling off; outer petals ovate to very broadly ovate, rounded, 11–18[-19] by 10–17 mm, inner petals elliptic to (narrowly) obovate, obtuse, 10–16[-22] by 5–8[-10] mm; androecium ca. 7 mm diam., stamens 1.3–1.5 mm long, connective appendage 0.6–0.8 mm wide; gynoecium ca. 1 mm diam., carpels 2–2.5[-2.9] long, sparsely covered with erect whitish hairs to 0.1 mm long. Monocarps (8-)17–33, ellipsoid to broadly ellipsoid, asymmetrical, 8–11 by 6–8 mm, often with an oblique longitudinal groove corresponding to the seed raphe, green maturing to pink or orange through purple or brownish-red, brown and black in vivo, dark or reddish-brown in sicco, with an excentric apicule, monocarps, stipes and receptacle sparsely to rather densely covered with erect whitish hairs ca. 0.1 mm long; stipes 8–16 by 1 mm; fruiting receptacle 4–8 mm diam. Seeds broadly ellipsoid, orange brown, pitted, 6–8 by 5–6 mm, raphe sunken, somewhat irregular.

Amazonian Colombia (Amazonas), Ecuador (Morona-Santiago), Peru (Amazonas, Loreto) and Brazil (Amazonas).

Forest inundated by white (várzea) or black (tahuampa) water, on yellowish, lateritic soil. At elevations of 80–200 m. Flowering: March, July and September; fruiting: throughout the year.

Peru: Bara (McDaniel et al. 17020), Barra caspi (McDaniel et al. 20677), churú yais (F. Dominguez 52, 59), Hicojilla (Schunke V. 6412), Icoja (M.E. Mathias et al. 5510), Yais (Huambisa; Huashikat 321, 493), Zorro Caspi Blanco (Freitas 8).

Cremastosperma microcarpum resembles most closely C. gracilipes, from which it differs in the denser, longer hairs on the flowers and the generally acuminate as opposed to caudate leaf apex (but see discussion under that species). The hairy flower resembles somewhat those of C. cauliflorum, but which cannot be confused as C. microcarpum never exhibits a branching inflorescence. In addition, the monocarps of C. cauliflorum are larger than those of C. microcarpum and characteristically globose to transversely broadly ellipsoid as opposed to ellipsoid.

Cremastosperma microcarpum is one of the more widespread and abundant species of the genus, including occurrences in protected areas in Colombia. Least concern [LC] (Table 1).

BRAZIL. Amazonas: Rio Embira, 7°30'S, 70°15'W, 10 Jun 1933, Krukoff 4748 (F, G, K, MICH, MO, NY, S, U). COLOMBIA. Amazonas: Parque Nacional Amacayacu, 3°01'S, 70°02'W, 100 m a.s.l., 21 Jun 1991, Rudas et al. 2260 (MO). ECUADOR. Morona-Santiago: Santiago-Río Morona road, E of Santiago, 2°58'24"S, 77°49'36"W, 322 m a.s.l., 10 Jul 2004, Croat 90749 (MO, U). PERU. Amazonas: Río Santiago valley, Quebrada Caterpiza, 4°00'S, 77°00'W, 200 m a.s.l., 3 Sep 1979, Huashikat 321 (MO). Loreto: Yanamono Explorama Reserve, 3°27'S, 72°51'W, 100–150 m a.s.l., 30 Dec 1998, Chatrou et al. 208 (MOL, U); Nauta-Parimari, 5°00'S, 74°15'W, 90 m a.s.l., 1993, Del Carpio 2213 (MO); Jenaro Herrera, 4°55'S, 73°40'W, 125 m a.s.l., 26 Oct 1988, Freitas 8 (U); Mishana, Río Nanay, 3°50'S, 73°30'W, 130 m a.s.l., 26 Feb 1979, Gentry & Aronson 25111 (U, USM); Quistacocha, near Iquitos, 3°45'S, 73°20'W, 140 m a.s.l., 13 Mar 1981, Gentry et al. 32153 (MO, NY, U, USM); Río Samiria, Flor de Yarina, 5°02'S, 74°30'W, 140–160 m a.s.l., 4 Aug 1982, Gentry et al. 38085 (MO, U); Reserva Nacional Pacaya-Samiria, 3°18'S, 74°50'W, 130 m a.s.l., 22 Oct 1990, Grández & N. Jaramillo 2032 (MO); Braga, near Caño Supay, Río Ucayali, 4°55'S, 73°44'W, 110 m a.s.l., 27 Oct 1994, Maas et al. 8222 (L, U, USM, WU); Mishana, Río Nanay, Quebreda San Pedro, 3°55'S, 73°35'W, 130 m a.s.l., 26 Sep 1986, Vásquez & N. Jaramillo 8010 (MO, U); Mariscal Castilla, Caballo cocha, 3°55'S, 70°30'W, 106 m a.s.l., 14 Jul 1987, Vásquez & N. Jaramillo 9350 (F, MO, U, USM); Sapuena, Jenaro Herrera, 4°50'S, 73°45'W, 170 m a.s.l., 17 Nov 1987, Vásquez & N. Jaramillo 10088 (MO, U, USM).

Figure 29. 

Cremastosperma microcarpum R.E.Fr. a leaves b flowers and flower buds c fruit (a, bMaas et al. 6281cVásquez & Jaramillo 9350).

BOLIVIA, La Paz: San Buenaventura, 470 m a.s.l., 12 Nov 1901, Williams, R.S. 670 (holotype: NY [NY00025876]; isotype: K! [K000485528]).

Cremastosperma juruense R.E.Fr., Acta Horti Bergiani 12: 282. 1937.

BRAZIL, Amazonas: Basin of Rio Juruá; near mouth of Rio Embira (tributary of Rio Tarauacá), 6 Jun 1933, Krukoff, B.A. 4697 (holotype: NY! [NY00025861]; isotype: S! [S-R-7933]).

Cremastosperma monospermum (Rusby) R.E.Fr. var. brachypodum R.E.Fr., Acta Horti Bergiani 12: 559. 1939.

BRAZIL, Pará: Boa Vista on the Tapajós River, 5 May 1929; Dahlgren & Sella 162 (holotype: S! [S-R-7011]; isotypes: B! [B 10 0242367], F! [V0054582F]).

Tree or shrub, 1–12 m tall, 4–10 cm diam.; young twigs and petioles glabrous. Leaves: petioles 5–10 by 1–3 mm; lamina obovate, elliptic or narrowly so, (8-)10–35 by 4–12 cm (index 2–3.5), chartaceous, green to brown, darker above, veins often reddish below, glabrous on both sides or rarely sparsely covered with appressed whitish to 0.4 mm long hairs on primary vein below, base acute to obtuse, rarely narrowly cuneate, mostly decurrent, apex acuminate to caudate (acumen/cusp 5–30 mm long), primary vein 1–2 mm wide at widest point, secondary veins 6–10, intersecondary veins 0–3, distance between from 10–18 mm at the base to 12–24 mm closer to the apex, angles with primary vein from 70–80° at the base to 40–50° closer to the apex, rarely branching, forming mostly distinct loops, smallest distance between loops and margin 2–6 mm, tertiary veins mostly reticulate. Inflorescence of single flowers solitary (or clustered in groups of 2), on leafy or leafless twigs; peduncles 2–10 by 0.5–1 mm (in flower), 2–10 by 1.5–2 mm (in fruit), sparsely covered with appressed to erect golden to 0.1 mm long hairs or glabrous; pedicels 40–50(-70) by 0.5–1 mm at the base (in flower), (12-)22–73 by 1–1.5 mm (in fruit), green or reddish in vivo, glabrous; 2 lower bracts, deltate, ca. 0.8 by 0.8 mm (occasionally large and leafy), obtuse, soon falling off, sparsely covered with appressed golden 0.1 mm long hairs or glabrous; upper bract mostly attached around half way along pedicel, narrowly ovate or deltate, 1–2.5 by ca. 1 mm, acute, obtuse, rounded or truncate, outer side sparsely covered with appressed golden 0.1 mm long hairs or glabrous; closed flower buds broadly ovoid-triangular, remaining closed (or nearly so) throughout development; flowers green, maturing to creamy yellow, yellow or orange in vivo, dark or reddish-brown or black in sicco; sepals and petals glabrous, sepals free or connate for ca. 1 mm, broadly ovate or deltate, appressed, patent or recurved, 2–4 by 2–3 mm, acute or obtuse, mostly soon falling off; outer petals ovate, 9–14 by 6–8 mm, inner petals elliptic to ovate or narrowly so, 10–13 by 4–5 mm; androecium not seen; gynoecium not seen. Monocarps 10–29, ellipsoid to broadly so, slightly asymmetrical, 9–11 by 7–8 mm, green maturing to pink, maroon, red or (blue-) black in vivo, brown, dark or reddish-brown or black in sicco, with an excentric apicule; stipes (6-)8–15 by 1 mm; fruiting receptacle 3–8 mm diam., monocarps, stipes and receptacle glabrous. Seeds broadly ovoid, reddish-brown, pitted, pits appear black with raised rim, 8–10 by 6–7 mm, raphe sunken, regular.

Bolivia (Beni, La Paz, Pando), Brazil (Acre, Pará, Rondônia) and widespread across Peru.

Primary and secondary lowland forest, occasionally on poorly drained soils or brown latosols. At elevations of 200–500 m. Flowering: April, July – December; fruiting: throughout the year.

Bolivia: Yohisi (Chacobo; Boom 5039), huabu midha (Bourdy 1828). Peru: Ayacbara (Schunke V. 2554).

Cremastosperma monospermum is the most widespread species of the genus – the only one found both along the eastern foothills of the Andes as far south as Bolivia and across Brazil south of the Amazon. It is best distinguished by the shape of the flower bud: roughly triangular with an obtuse apex, apparently remaining closed throughout development, with the petals not opening fully even at maturity. In most other species of the genus, including the otherwise very similar C. confusum (see notes under that species), the flower bud opens during development. Cremastosperma pendulum and C. yamayakatense also exhibit glabrous, closed flower buds, but the shape of both is depressed ovoid. The latter also has a short, sturdy pedicel, very different to C. monospermum, the flower of which is borne on a slender and often long (though rather variable) pedicel.

The authors do not consider it useful to recognise sub-specific taxa within C. monospermum. The variation in stipe length and thickness represented by the type of var. brachypodum, described by Fries (1939), falls within that of the species as a whole and is therefore synonymised here.

Cremastosperma monospermum is the most widespread and abundant species of the genus, including occurrences in protected areas in Brazil and Peru. Least concern [LC] (Table 1).

BOLIVIA. Beni: Chácobo, 11°45'S, 66°02'W, 200 m a.s.l., 15 Apr 1984, Boom 5039 (K, LPB, MO, NY, U); Río Beni above Río Qui, 14°48'S, 67°23'W, 320 m a.s.l., 22 May 1990, Daly et al. 6573 (LPB, MO, NY, U); Río Beni, Rurrenabaque, 14°28'S, 67°30'W, 320 m a.s.l., 25 May 1990, Daly et al. 6630 (NY, U); San Borja, 15°10'S, 66°28'W, 250 m a.s.l., 23 Nov 1988, R.B. Foster et al. 12549 (F, LPB, U); Bosque de Chimanes, 15°30'S, 66°15'W, 250 m a.s.l., 27 Oct 1989, R.B. Foster & Terceros 13422 (U); Prov. José Ballivián, Carinavi-San Borja road, 15°17'S, 67°04'W, 850–900 m a.s.l., 1 Nov 1989, D.N. Smith & V. Garcia 13834 (LPB); Bosque de Producción Permanente Chimanes, 15°10'S, 66°37'W, 260 m a.s.l., 25 Aug 1990, D.N. Smith et al. 14216 (LPB, MO). La Paz: Luisita, 13°50'S, 67°15'W, 180 m a.s.l., 27 Feb 1984, Beck & Haase 10092 (LPB, MO, U); Santa Fé, 13°40'S, 68°12'W, 250 m a.s.l., 5 Aug 1995, DeWalt et al. 662 (LPB, U, WU); Puerto Muscoso, 13°01'S, 68°50'W, 190 m a.s.l., 25 Jun 1995, Helme & Kruger 742 (LPB, MO, U); Alto Beni, Sapecho, 15°30'S, 67°20'W, 550 m a.s.l., 3 Apr 1989, Seidel 2674 (LPB, U). Pando: Manuripi, 11°40'S, 68°00'W, 18 Aug 1990, G. Gonzales 53 (LPB); Puerto América, 11°35'S, 68°03'W, 180 m a.s.l., 9 Sep 1995, Jardim 2408 (U); Agua Clara, 11°44'20"S, 67°02'40"W, 13 Nov 2001, Pirie et al. 4 (U); Puerto Rico, road to Cobija, 11°03'25"S, 67°47'46"W, 175 m a.s.l., 15 Nov 2001, Pirie et al. 5 (U). BRAZIL. Acre: Fazenda Bom Sossego, 7°40'S, 73°09'W, 27 Sep 1985, Campbell et al. 9026 (U); Seringal Porongaba, 10°51'S, 68°48'W, 23 May 1991, Daly et al. 6686 (MO, NY, U); Mun. Rio Branco, BR 317, 10°30'S, 67°45'W, 8 Jun 1991, Daly et al. 6909 (MO, NY, U); Reserva Extrativista do Alto Jurua, 8°55'S, 72°31'W, 11 Mar 1992, Daly et al. 7347 (MO, NY, U); Mun. Porto Acre, 9°45'S, 67°38'W, 3 Nov 1993, Daly et al. 8043 (NY, U); Mun. Cruzeiro do Sul, 8°13'41"S, 73°02'22"W, 11 May 2003, Daly et al. 11719 (NY); Mun. Tarauacá, Seringal Universo, 8°25'S, 71°19'W, 15 Jun 1995, Figueiredo et al. 853 (U); Rio Muru, Seringal Vitoria Velha, 8°26'S, 70°49'W, 20 Jun 1995, Figueiredo et al. 906 (U); Mun. Sena Madureira, Rio Macaua, 9°24'S, 68°54'W, 3 Apr 1994, L. Lima et al. 582 (NY); Rio Branco-Porto Acre road, 10°00'S, 67°50'W, 11 Oct 1980, Nelson 680 (U); Riozinho do Andirá, colocação Curitiba, 9°43'45"S, 68°08'53"W, 6 Jun 1998, A.R.S. Oliveira et al. 488 (WAG); Porto Walter, Rio Juruá Mirim, 8°07'S, 72°49'W, 31 May 1994, Silveira et al. 767 (NY, U); Tarauacá, 9°25'45"S, 68°40'00"W, 18 Jun 2006, Silveira et al. 3796 (L); Mun. Tarauacá, Rio Liberdade, 7°18'49"S, 66°01'46"W, 18 Jun 2006, Silveira et al. 3811 (L); Floresta Estadual do Antimary, 9°25'45"S, 68°04'00"W, 8 Jul 2006, Silveira et al. 4171 (L). Mato Grosso: Mun. Alta Floresta, 10°07'S, 57°30'W, 29 Sep 1985, Cid Ferreira et al. 6301 (F, GH, K, MICH, NY, U, US). Pará: Itaituba, 7°40'S, 55°15'W, 14 May 1983, M.N. Silva 322 (MO, NY, U, US); Serra dos Carajás, 5°49'S, 50°32'W, 225–250 m a.s.l., 15 Jun 1982, Sperling et al. 6198 (GH, K, MO, NY, U). Rondônia: Mun. Costa Marques, Chapada dos Parecís, 11°12'S, 62°63'W, 14 Jun 1984, Cid Ferreira et al. 4507 (MO, NY, U); Espigão de Oeste, 11°12'S, 60°61'W, 20 Jun 1984, Cid Ferreira et al. 4673 (F, GH, MO, NY, U, US); Ariquemes, 10°00'S, 62°59'W, 200 m a.s.l., 15 Mar 1987, Nee 34402 (U); Campo Novo, 10°38'S, 63°37'W, 300 m a.s.l., 23 Apr 1987, Nee 34992 (MO); Porto Velho, Campo Novo, road to Ariquemes, 10°35'S, 63°30'W, 300 m a.s.l., 25 Apr 1987, Nee 35033 (U); Ariquemes, Mineração Mibrasa, 10°35'S, 63°35'W, 15 May 1982, Teixeira 480 (MO, U). PERU. Huánuco: Pucallpa, western Sira mountain, 9°28'S, 74°47'W, 680 m a.s.l., 7 Apr 1988, Wallnöfer 112 7488 (U). Loreto: Quebrada Shesha, Río Abujao, 8°20'S, 73°45'W, 14 Dec 1978, C. Díaz et al. 825 (NY, U); Shucushayacu, Río Huallaga, 6°01'S, 75°55'W, 180 m a.s.l., 11 Oct 1985, Gentry et al. 52233 (MO, U); Pongo de Manseriche, 4°26'S, 77°34'W, 650 m a.s.l., 25 Nov 1997, R. Rojas et al. 666 (F, MO); Canchahuayo, Río Ucayali, 7°05'S, 75°10'W, 500 m a.s.l., 25 Nov 1985, Vásquez et al. 6932 (F, U); Yanamono Explorama Reserve, 3°28'S, 72°50'W, 106 m a.s.l., 3 Oct 1989, Vásquez & N. Jaramillo 12829 (MO, MOL, U). Madre de Dios: Tambopata, Río Heath, 12°39'S, 68°44'W, 210 m a.s.l., 22 May 1996, M. Aguilar & Castro 764 (U, USM); Cuzco Amazónico, 15 km ENE of Puerto Maldonado, 12°35'S, 69°05'W, 200 m a.s.l., 14 Dec 1989, Gentry et al. 68663 (MO, USM); Inkaterra Ecological Reserve, 12°35'S, 69°09'W, 190 m a.s.l., 30 Aug 2006, Monteagudo et al. 12749 (WAG); Tambopata, 12°33'S, 69°03'W, 200 m a.s.l., 27 May 1989, Núñez et al. 10589 (MO); Cuzco Amazónico Lodge, 12°35'S, 69°03'W, 200 m a.s.l., 13 May 1990, Núñez & Timaná 12152 (U); Collpa de Blanquillo, Río Madre de Dios, 12°27'S, 70°41'W, 200 m a.s.l., 11 May 1995, Núñez 16303 (USM); Cuzco Amazónico, 12°05'S, 69°03'W, 200 m a.s.l., 21 Jun 1989, Phillips et al. 545 (MO, U); Las Piedras, Cusco Amazónico, 12°29'S, 69°03'W, 200 m a.s.l., 25 Feb 1991, Timaná 1537 (MO); Puerto Maldonado, Cusco Amazónico, 12°32'S, 69°03'W, 220 m a.s.l., 20 May 2003, L. Valencia & Suclli 2205 (U). Ucayali: Peru-Brazil border, Quebrada Sapallal, 8°02'S, 73°55'W, 260 m a.s.l., 19 Jun 1987, Gentry & C. Diaz 58434 (USM); Prov. Purus, Distr. Purus, Río Caranja, 10°04'S, 71°06'W, 325 m a.s.l., 17 Jul 1998, Graham 605 (F); Prov. Coronel Portillo, Iparia, el Sira, 9°25'57"S, 74°32'47"W, 350–400 m a.s.l., 21 Sep 2007, Graham 4706 (L); Prov. Coronel Portillo, Calleria, 8°09'13"S, 74°15'48"W, 150–175 m a.s.l., 26 Feb 2003, Schunke V. 15223 (F).

Figure 30. 

Cremastosperma monospermum (Rusby) R.E.Fr. a fruiting specimen b flower buds (aNuñez & Timaná 12152bCid Ferreira et al. 6301).