IncreasIng dIversIty In the specIes-rIch genus guatterIa ( annonaceae )

A large amount of unidentified material of the Neotropical genus Guatteria (Annonaceae) is lying on herbarium shelves around the world. Here twelve species are described in an attempt to reduce this amount of unidentified material. Next to this, previously unidentified material is now assigned to G. asplundiana, G. microcarpa, G. pastazae, and G. scalarinervia, four previously described but poorly known species, thus providing a clearer circumscription of these species. Furthermore, G. discolor and G. megalophylla appear to have a wider distribution than previously thought. Lastly, G. macrantha, a curious species only known from the type, is excluded from the genus and referred to Uvaria. With the description of twelve new species and the synonymizing of one, the total number of species in Guatteria adds up to over 300, thus rivalling Inga (Fabaceae) and Ocotea (Lauraceae) for the status of largest genus of Neotropical trees.

species mentioned.they named the genus after Don Juan Bautista Guátteri, who "died a few months ago, in the bloom of his life".they state that because of his death "the Royal Botanical Gardens of Parma (Italy) now lacked a very distinguished Professor and a very honourable person".In 1798, four species were included (Guatteria glauca, G. hirsuta, G. ovalis (now Ruizodendron ovale), and G. pendula (now Cremastosperma pendulum; Ruiz López & Pavón 1798)), without typification of the generic name (the typification method did not yet exist in those days).Hutchinson (1923) designated G. glauca as lectotype (for a further discussion about conservation and lectotypification of Guatteria see Van setten & Maas (1990).
Almost twenty years after Ruiz López & Pavón's initial description, Dunal (1817) wrote a monograph of the family of Annonaceae in which he mentioned 20 Guatteria species.At present his Asian species are no longer considered to belong to the genus (as said, Guatteria is exclusively Neotropical).Dunal described them as such based on the superficially similar monocarps.A further six of Dunal's species have been synonymized into other American Annonaceae genera, leaving only G. guianensis (his illegitimate G. aberemoa), G. eriopoda, G. glauca, G. hirsuta, and G. ouregou (incl. G. podocarpa DC.) as true members of the genus.other early contributions to the taxonomy of Guatteria were made by De saint-Hilaire (1824 -1825; 5 new species), Von schlechtendahl (1834; 5 new species), Von Martius (1841; 20 new species), triana & Planchon (1862; 11 new species), Diels (1905Diels ( , 1906Diels ( , 1924aDiels ( , b, 1927Diels ( , 1931;;20 species), and Fries (1938;12 new species).Because of these efforts, at the start of the revision work by Robert Fries (published in 1939 and the only revision to date), already 151 species of Guatteria had been described.Fries synonymized six of these and transferred 36 species to a different genus.As an addition he placed two species in Guatteria that had been described earlier in other genera of Annonaceae (G.axilliflora (DC.)R.E.Fr. was described as Annona axilliflora DC., and G. guianensis (Aubl.)R.E.Fr. as Aberemoa guianensis Aubl.).He himself then described 106 new species in his revision (Fries 1939).these additions made Guatteria, with 217 species, already one of the largest Annonaceae genera at that time.In order to create some order in the taxonomy of Guatteria Fries erected 30 sections (table 1; Fries 1939).Later he recognised two subgenera: Anomalantha (containing only Guatteria anomala) and Guatteria (containing all other species in the genus; Fries 1955).several years after that (Fries 1959b) he reduced the number of sections to 22, merging several not well circumscribed ones (table 1).
since Fries' 1939 treatment, the number of species has gradually increased, mainly because of work by Fries himself (Fries 1947(Fries , 1948a(Fries , b, 1950(Fries , 1952(Fries , 1956(Fries , 1957a(Fries , b, 1959a;;Maguire et al. 1957Maguire et al. , 1960;;totalling 36 new species).Apart from several more described species, larger recent contributions were made by Maas et al. (1988;4 new species), scharf et al. (2005;2006a, b; 12 new species), and Erkens et al. (2006;5 new species).the total number of species currently recognised is 286, and with the merging of three small, related genera with Guatteria (Erkens & Maas 2008) reaches 294. the genus already rivals Inga (Fabaceae; c. 300 spp.) and Ocotea (Lauraceae; c. 300 spp.) for the status of largest genus of Neotropical trees.yet many more new species are probably awaiting description because a large amount of unidentified material of Guatteria is lying on herbarium shelves around the world, c. 1500 specimens at the Utrecht herbarium alone.The twelve species described here are a first step in an effort to reduce the amount of unidentified material.Furthermore, material so far undetermined is now assigned to four previously described but poorly known species, thus providing a clearer circumscription of these species.Also, two species have a wider distributional range than previously thought.Lastly, one species only known from the type is assigned to another genus.the total number of accepted names in Guatteria, including the new species described elsewhere in this issue (scharf et al. 2008), therefore now adds up to 307.

MAtERIAL AND MEtHoDs
Herbarium material was investigated from the following herbaria: CUVC, GoEt, k, MAD, Mo, Ny, PR, U, and WIs.Measurements as a rule were made of dried material.Measurements of material in spirit are given between accolades { }.Colour indica-  Fries (1939) and revised later (Fries 1955;1959).Monotypic sections Acrantha (G.terminalis) and Pycnantha (G.elongata) were described in 1939 but were not mentioned in subsequent publications.therefore, whether Fries submerged or abolished these sections is not clear.Shrub or tree 2-30 m tall, 5-40 cm diam., trunk fissured, slash orange (in material from Acre); young twigs densely to sparsely covered with appressed hairs, soon glabrous.
Notes -Guatteria arenicola is a small-leaved species from the white sand campina or campinarana vegetations in Brazil (Acre), and forests on white sand in Peru (san Martín).It is quite noteworthy by its tiny, prominently veined, coriaceous leaves, and its very small flowers, which have strongly recurved pedicels.Another characteristic of this species is the presence of bark with an orange slash, a feature not seen in any other representative of the genus Guatteria.
Distribution -Pacific Coast of Colombia (El Valle).
Notes -Guatteria argentea is very easily recognizable by its silvery indument on many parts of the plant and by its obovate, densely verruculose leaves with an attenuate base.
Distribution -Colombia (Antioquia) and Amazonian Ecuador and Peru.
Habitat & Ecology -In premontane or montane forest.At elevations of 300 -2300 m.Flowering and fruiting: throughout the year.
Notes -Guatteria asplundiana has a fruit which resembles that of some species of Unonopsis.It is particularly characterized by its relatively large monocarps, which are, moreover, often wrinkled after drying, probably due to the fact that the seeds are much smaller than the monocarps.some material collected in the Ecuadorian state of Napo (named 'Guatteria sp. 10, large monocarps' by Chatrou et al. (1997)) is aberrant in having larger than average monocarps (17-35 by 17-30 mm) and seeds (10-14 by 8-14 mm), and a thicker monocarp wall (0.5-1 mm).As all leaf and flower characters fall within the range of variability of G. asplundiana we have refrained from giving this material a status of its own.It is cited under a separate heading and in the identification list this material is printed in bold type.tree c. 20 m tall, 30 cm diam., slash orange-brown; young twigs densely covered with erect and appressed, pale brown, somewhat curly hairs.Leaves: petiole 2-4 mm long, 2-3 mm diam.; lamina narrowly oblong-elliptic to narrowly elliptic, 17-27 by 7.5-10 cm (leaf index 2.6-2.9),chartaceous, not verruculose, shiny, green above, green below, sparsely covered with erect and appressed, pale brown, somewhat curly hairs above, rather densely so below, base obtuse, oblique, apex acute to shortly and very gradually, bluntly acuminate (acumen 5-10 mm long), primary vein impressed above, secondary veins distinct, 10-15 on either side of primary vein, prominent above, smallest distance between loops and margin 2-3 mm.Flowers in axils of leaves or on older branchlets, solitary; pedicels 40-55 mm long, 1-2 mm diam., articulated at 1/10-1/6 from the base, with 4-6 minute bracts below articulation, densely to rather densely covered with erect and appressed, pale brown, somewhat curly hairs; flower buds not seen; sepals free, ovate, c. 6 by 4 mm, patent to reflexed, outer side densely covered with appressed hairs; petals pale green inside, greenish brown outside in vivo, unequal, outer ones oblong-elliptic, 13-16 by 5-7 mm, inner ones elliptic to elliptic-oblong, 13-20 by 7-11 mm, outer side densely covered with (mostly) appressed hairs; stamens c. 2 mm long, connective shield finely papillate-hairy to subglabrous.Monocarps and seeds not seen.
Habitat & Ecology -In mixed forest on hills, on heavy clay.At elevations of 100-250 m.Flowering: April; fruiting: unknown.
Note -In the 'Florula de las Reservas Biológicas de Iquitos, Peru' (Vásquez 1997) mention is made of an Ephedranthus sp.A, accompanied by a short description.Although we were unable to examine the herbarium material we feel certain to judge by the description that Ephedranthus sp.A is identical with our G. confusa.this is the reason why the epithet 'confusa' was chosen.Guatteria discolor can be recognized by slightly pointed flower buds and relatively long (20-25 mm), shortly stipitate monocarps (stipes 2-6 mm long).In the herbarium it can be recognized by thick, 2-coloured leaves with a reddish indument on the lower side.
Habitat & Ecology -In non-inundated forest (from lowland rain forest to premontane forest), sometimes in periodically inundated forest.At elevations of 150-1800 m.Flowering: throughout the year; fruiting: from May to November.
Vernacular names -Bolivia: Piraquina, oyshobo (yuracare).Peru: Atzmiriqui.Notes -Guatteria duodecima can at first glance be distinguished by relatively narrow and verruculose leaves, usually narrowed both toward the base and toward the apex, and densely to rather densely covered with appressed, almost silvery hairs on the lower side.other distinctive features of this species are found in the shortly pedicellate flowers and monocarps which are always longer that the stipes.
It is named duodecima or 'twelfth' in reference to Chatrou et al. (1997) who lists this as unidentified species and given number 'sp.12'.
Note -It is quite amazing that G. flabellata has never been described before as it is well collected and as it has very distinct differentiating characters.It is quite noteworthy by its large, thick, brown drying leaves, with the veins impressed above and with a cordate to obtuse base; the petals are very well marked by the quite distinct impressed venation at their outer side; furthermore the petals are distinctly narrowed toward the base (almost clawed), whereas the apex is often somewhat emarginate.The specific epithet refers to the fan-shaped (fan = flabellum) petals.Note -Guatteria flagelliflora is unique by its flagellate inflorescence, a feature not reported so far in Guatteria.Liana or creeping shrub; young twigs with a zigzag pattern, sparsely covered with appressed hairs, soon glabrous.Leaves: petiole 3-5 mm long, 1-2 mm diam.; lamina narrowly elliptic, 6-10 by 1.5-2.5 cm (leaf index 3.5-4.5),coriaceous, not verruculose, dull, greyish green above, pale brown below, glabrous above, except for some hairs along primary vein, sparsely covered with appressed hairs below, base acute, apex long-acute, primary vein impressed above, secondary veins indistinct, 13-16 on either side of primary vein, flat above, smallest distance between loops and margin 1-2 mm.Flowers (only one available) in axil of leaf near (aborted) top of branchlet, single; pedicels 15-25 mm long, 1 mm diam., articulated at 1/4-1/2 from the base, bracts soon falling off, densely covered with appressed, brown hairs; flower buds subglobose; sepals free, broadly ovate, c. 6 by 5.5 mm, outer side densely covered with appressed hairs; petals pale yellow in vivo, narrowly elliptic to narrowly elliptic-obovate, outer ones c. 13 by 5 mm, inner ones c. 15 by 6 mm, outer side densely covered with appressed hairs; stamens c. 1.5 mm long, connective shield glabrous.Monocarps and seeds not seen.
Habitat & Ecology -In low, primary forest, with many epiphytes, and with a thick layer of moss, roots, and leaves, on sandstone soil.At elevations of 320-1300 m.Flowering: February and August; fruiting: unknown.Note -Guatteria fractiflexa is one of the two liana species in the genus, the other being G. scandens, a species quite common in the Guianas.Guatteria fractiflexa is quite remarkable by the zigzag branches (hence the specific epithet).
Habitat & Ecology -In non-inundated, primary, sometimes premontane, wet forest, on clayey, red soil.At elevations of 300-1300 m.Flowering: March to May and september to November; fruiting: throughout the year.
Vernacular name -Ecuador: Cara huasca (Quichua).Notes -Guatteria gentryi has been named after our friend and dear colleague, the late Alwyn ('Al') Gentry, one of the greatest Neotropical taxonomists of the last century.
Habitat & Ecology -In non-inundated forest, on stony soil covered with lichens and mosses.At elevations of c. 100 m.Flowering: unknown; fruiting: November.
Note -Guatteria japurensis is most distinctive by the long and slender fruiting pedicels, which are very unusual in Guatteria.Another noteworthy feature is found in its leaves which are indistinctly veined and which are rugulose on the upper surface.Guatteria macrantha has been described by Presl (1835) as having stellate hairs and inflorescences opposite the leaves.these characteristics are not found in any species of Guatteria.Although the monocarps are unknown and the locality is uncertain ('habitat in Mexico?'), the specimen can be identified as Uvaria (pers.comm.P. keßler).Uvaria does not occur in Mexico, but this collection was probably made somewhere else.It is known that many of Haenke's labels containing the locality information have become misplaced, and that the localities given by Presl can, therefore, not always be trusted (stearn 1938).For instance, species now known to be exclusively American are recorded by Presl from the Philippines, and vice versa.Map 4 Guatteria megalophylla Diels (1905) 127.
Habitat & Ecology -In premontane and montane forest, on soils derived from sandstone substrate or on red clay.At elevations of 300-1800(-2200) m.Flowering: from November to May; fruiting: from July to March.
Vernacular names -Peru: Palo yais, Wampu yais, yais.Notes -Guatteria pastazae was previously known from only 2 collections near Mera.It is well recognizable by its verruculose leaves with obscure venation and an attenuate leaf base, combined with shortly pedicellate flowers and shortly stipitate monocarps.
Guatteria pastazae shows some features of G. puncticulata R.E.Fr., a species occurring in the lowlands of Peru (Loreto) and Brazil (Amazonas), like the verruculose leaves and obscure leaf venation.It differs, however, by its much shorter stipes (stipes shorter than to almost equalling monocarp length) versus stipes longer than monocarp.
A specimen from Peru (Rodríguez R. & Cruz A. 2053) might be placed near G. pastazae, but it differs by fruiting pedicels 20-25 mm long and an articulation at 1/5-1/4 from the base, and by distinctly smaller leaves.
Habitat & Ecology -In premontane wet forest.At elevations of 250 -1000 m.Flowering: May, June, september to November; fruiting: February to June.
Vernacular name -Ecuador: Cargadera negra, Degteiug, tilalde (Awapit).Note -Guatteria venosa belongs to Fries's section Mecocarpus by its distinct marginal vein, the presence of verruculi on the leaves and by the short stipes.It is distinguished from other species of section Mecocarpus by an unusually high number of secondary veins (hence the specific name) and by the few verruculi on the leaves compared to other species in the section.