NEW SPECIES OF CREMASTOSPERMA (ANNONACEAE) FROM COLOMBIA, ECUADOR, AND PANAMA

The genus Cremastosperma R.E. Fr. can most easily be distinguished from other Neotropical Annonaceae with apocarpous, stipitate fruits by its midrib which is raised on the upper side with a mostly conspicuous longitudinal groove. The individual distributions of its species are restricted to four (disjunct) areas: 1) the Chocó/Darién/western Ecuador region (the narrow tropical zone to the west of the Andean mountain chain on the Pacific Ocean side of north-western South America) plus Central America; 2) the tropical Andes (including forests on the eastern side of the Andes extending from Colombia through eastern Ecuador and Peru as far south as Bolivia); 3) coastal Venezuela; and 4) French Guiana. Since the description of the genus by Fries (1930), further work by Fries (1931, 1934, 1937, 1939, 1948, 1950), Maas et al. (1986) and Pirie & Zapata (2004) brought the total number of species of Cremastosperma to 21. Of these, 16 represent species distributed only in the tropical Andes region, with only one each found in Venezuela and French Guiana, respectively, and three in the Chocó/Darién/western Ecuador region and Central America combined. Although Chatrou & Pirie (2005) increased the number of species of Cremasto­ sperma known from Venezuela to two, recent systematic research has revealed a far greater underestimation of species diversity represented in particular by collections from the Chocó/Darién/western Ecuador region. The general increase in numbers of collections of Cremastosperma made since the time of Fries has been restricted in areas of north-western South America. Far fewer were available to the author than for comparable areas on the Amazonian side of the Andean mountain chain. Nevertheless, it is the opinion of the author that the degree of morphological differentiation clear


INTRODUCTION
The genus Cremastosperma R.E.Fr. can most easily be distinguished from other Neotropical Annonaceae with apocarpous, stipitate fruits by its midrib which is raised on the upper side with a mostly conspicuous longitudinal groove.The individual distributions of its species are restricted to four (disjunct) areas: 1) the Chocó/Darién/western Ecuador region (the narrow tropical zone to the west of the Andean mountain chain on the Pacific Ocean side of north-western South America) plus Central America; 2) the tropical Andes (including forests on the eastern side of the Andes extending from Colombia through eastern Ecuador and Peru as far south as Bolivia); 3) coastal Venezuela; and 4) French Guiana.
Since the description of the genus by Fries (1930), further work by Fries (1931Fries ( , 1934Fries ( , 1937Fries ( , 1939Fries ( , 1948Fries ( , 1950)), Maas et al. (1986) and Pirie & Zapata (2004) brought the total number of species of Cremastosperma to 21.Of these, 16 represent species distributed only in the tropical Andes region, with only one each found in Venezuela and French Guiana, respectively, and three in the Chocó/Darién/western Ecuador region and Central America combined.
Although Chatrou & Pirie (2005) increased the number of species of Cremasto sperma known from Venezuela to two, recent systematic research has revealed a far greater underestimation of species diversity represented in particular by collections from the Chocó/Darién/western Ecuador region.The general increase in numbers of collections of Cremastosperma made since the time of Fries has been restricted in areas of north-western South America.Far fewer were available to the author than for comparable areas on the Amazonian side of the Andean mountain chain.Nevertheless, it is the opinion of the author that the degree of morphological differentiation clear 13a.Pedicels ≤ 28 mm long (in flower).Petals appear glabrous, but with dense hairs at base and in a line leading to the petal apex . . . . . . . . . . . . . . . 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 c. 2 by 1.5 mm (in flower), 2-3 by 1.5-2 mm (in fruit); pedicels 20-28 by c. 1 mm diam.at the base, 1.5-2 mm diam.at the apex (in flower), 20-40 by c. 2 mm diam.at the base, c. 3 mm diam.at the apex (in fruit), peduncles and pedicels sparsely covered with appressed whitish golden to 0.2 mm long hairs; 2 lower bracts, deltate, c. 1 mm long, obtuse, caducous; upper bract attached around halfway along pedicel, deltate, 1 mm long, obtuse, outer side of upper and lower bracts rather densely to densely covered with appressed whitish golden to 0.2 mm long hairs; 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, caducous, sparsely to rather densely covered with appressed whitish golden to 0.2 mm long hairs; outer petals elliptic, c. 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 to 0.2 mm long hairs; receptacle depressed ovoid; androecium 5-7 mm diam., stamens c. 1 mm long, connective appendage roughly rhombic, 0.5-0.7 mm wide, glabrous; gynoecium 1-1.5 mm diam., carpels 30-40, c. 1.5 mm long, glabrous.Monocarps c. 10, ellipsoid, strongly asymmetrical, 13-14 by c. 11 mm, with an excentral, to 0.2 mm long, apicule, orange to deep red, maturing to black in vivo, dark reddish brown in sicco; stipes orange to deep red in vivo, c. 20 by 1.5 mm, monocarps and stipes glabrous; fruiting receptacle depressed ovoid, c. 6 mm diam., glabrous.Seeds ellipsoid, reddish brown with dark pits each surrounded by a raised rim, c. 12 by 9 mm, raphe impressed, encircling seed longitudinally.
Note -Cremastosperma antioquense is similar to C. awaense, particularly in the appearance of the flowers.However, 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.
Distribution -Pacific coast of Ecuador (Esmeraldas and Carchi) and Colombia (Nariño and Chocó).Habitat & Ecology -Sea level to almost 2000 m elevation in primary humid to premontane tropical forest.Flowering: January, September, November; fruiting: January, February, June to September.
Etymology -Named after the ethnic reserve 'Awá', the type locality in Carchi, Ecuador, in which many of the collections of this species were made.
Note -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 (< 0.1 mm) 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.Cremastosperma awaense can easily be distinguished from both C. westrae and the geographically closer C. stenophyllum on the basis of the length of the pedicel.That of C. westrae is shorter (not exceeding 17 mm) and that of C. stenophyllum longer (c.45 mm in comparison to 27-28 mm in flower).
Etymology -Cremastosperma chococola is so named in reference to its being found only in the Colombian department of Chocó.
Note -The strongly asymmetric monocarps of C. chococola resemble those of C. antioquense, 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 by its small, narrowly elliptic leaves with typical pinkish brown colour on the underside, and by the absence of hairs on the pedicels.
Habitat & Ecology -At elevations of 1200-1500 m.Flowering: December, May; fruiting: December, May, July.Note -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.
Notes -Cremastosperma longipes can easily be distinguished from 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 (Diels) R.E.Fr. and C. bullatum Pirie never exceeding 150 mm in length, significantly shorter than those of C. longipes.In addition, leaves of C. longipes are unusually large, equalling the maximum dimensions observed in C. megalophyllum R.E.Fr., a more densely collected species from Amazonian Colombia, Ecuador, and Peru.
Etymology -Cremastosperma magdalenae is named after the river Magdalena which flows north-east between the central and oriental cordilleras of the northern Andes mountain range in Colombia.It is the only species of Cremastosperma known from this valley.
Note -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 fruits with smaller sepals only persistent on one of the two duplicates studied).Also noteworthy are the relatively short pedicels, the short shoot subtending the pedicel comprising of two internodes (indicated by the presence of two lower bracts), and the absence of indument on all parts.The absence of hairs on fruits and flowers reveal 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.Pirie,; Map 2 Inflorescentia composita atque monocarpiis glabris distinctum.C. caulifloro simile sed petalis indumento minus denso pilis brevioribus differt.-Typus: Alvarado 267 (holo U; iso AAU, MO, QCNE), Ecuador, Napo, Archidonia Cantón, foothills south of Volcano Sumaco, km 50 on Hollín -Loreto road, community Huahua Sumaco, 3 May 1989 (fr).
Habitat & Ecology -Primary pluvial pre-montane forest, often on volcanic soils but also reported growing on limestone.At elevations of 600-1300 m.Flowering: September, November, December, February; fruiting: August to December, March to May.
Note -The characteristic pattern of indument on the inner petals of C. napoense appears to be unique for the genus.The species can be further distinguished by the combination of a branching inflorescence and glabrous fruits.The only other species in the genus with such an inflorescence is C. cauliflorum R.E.Fr., which differs both in the presence of brown indument on the (characteristic depressed ovoid) monocarps and in the dense hairs on the flowers.Tree c. 10 m tall, c. 20 cm diam.; young twigs and petioles rather densely covered with appressed, golden, to 0.2 mm long hairs.Leaves: petioles 5-9 by 2-2.5 mm; lamina narrowly elliptic, 25-30 by 6.5-8 cm (index 3.8-4.3),chartaceous, minutely verrucose, greyish green above, green below, very sparsely covered with appressed yellowish white, to 0.2 mm long hairs below and on veins above, base acute, apex acuminate (acumen 20-25 mm long), primary vein 1-1.5 mm wide at widest point, verrucose, secondary veins 8-10, intersecondary veins occasional, distance between from 5 mm at the base to 50 mm closer to the apex, angles with primary vein from 45-55° at the base to 70-80° closer to the apex, not branching, not forming loops, tertiary veins with some reticulation.Inflorescence of single flowers (1 flower observed) on brachyblasts on thicker twigs or branches; peduncle c. 1.5 by c. 1 mm (in flower); pedicels c. 45 by c. 1 mm (in flower), peduncles and pedicels and outer side of bracts (densely), sepals (densely), and petals (sparsely to rather densely) covered with appressed, yellowish white, to 0.2 mm long hairs; 2 lower bracts, deltate, c. 1 mm long, obtuse; upper bract attached on basal half of pedicel, ovate, c. 1.5 by 0.8 mm, acute; flowers green, maturing to yellow in vivo, light brown with dark brown patches at the base of the petals in sicco; sepals deltate, 2 mm long, obtuse; outer petals elliptic, c. 18 by 8 mm, inner petals elliptic, c. 18 mm long (diam.unknown).Fruit not seen.
Habitat & Ecology -Secondary vegetation with primary elements.Flowering: January.Note -Only two collections of C. stenophyllum, one of which sterile, have been observed by the author.However, these are consistently distinct from all other species of the genus.Cremastosperma stenophyllum can be distinguished even when sterile by the conspicuously green-drying, narrowly elliptic long acuminate leaves.The flower somewhat resembles those of C. awaense, but both the pedicel and leaf acumen are longer and C. stenophyllum also lacks the distinctive pattern of indument on the petals of C. awaense: the hairs are instead evenly distributed on the outer surfaces.
Etymology -Cremastosperma westrae is named after the Dutch botanist Lubbert Y.Th.Westra, whose lifelong dedication to plant systematics includes, in the last two decades, a great contribution to the taxonomy of Neotropical Annonaceae.His encyclopaedic knowledge remains an invaluable and much appreciated resource to colleagues at the Utrecht branch of the National Herbarium of the Netherlands and further afield.
Notes -Cremastosperma westrae is most similar to C. novogranatense: it differs in having longer stipes, smaller sepals, and less dense, shorter hairs on the petals.The shape of the fruits of C. pacificum bear a resemblance to those of C. westrae.