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Sintaxones en los bosques semideciduos de la Sierra aestra, Cuba Oriental.

Semi-deciduous forests syntaxa from Sierra Maestra, Eastern Cuba.

INTRODUCTION

Semi-deciduous forests (CAPOATE & BERAZAIN, 1984; Reyes, 2006, 2011-2012), also known as seasonal evergreen forests (DEL RISCO, 1995), were the most extensive in the flat and hilly areas of the Cuban archipelago (BORHIDI, 1996) where they formed mature communities in relative equilibrium.

Degrading human intervention occurred due to deforestation and fire; the latter played a major role in the driest areas. When exploitation in altered oecotopes corresponding to this type of forest is excluded, a syngenetic evolution by enrichment and competition among species is also developed, up to reach a stage of secondary tree shrubwood, in which few species dominate the coverage. Then, they constitute an Early secondary community or Homeostasis I (sensu BUDOWSKI, 1985; CAPOATE & al., 1988; REYES & ACOSTA, 2004a, b), and establish at the same time, a stage of relative equilibrium (Reyes & al., 2004b). Later, other species come to gradually replace the existing ones, creating new Late secondary communities (Fiera II and Homeostasis II), which are different regarding degradation and ecological conditions. The number of tree species, the degree of cenotic complexity and the dominance of species of advanced stages change with the successional stage.

Low areas from Sierra Maestra have been heavily exploited (crop and / or pasture), thus offering variations on existing plant communities. LEON (1946) and Samek (1974) established an altitudinal zonation and named those low areas as "Yayales" and "Planar floor" respectively. Likewise, Reyes (2006) said semi-deciduous forests were developed there, so they will be treated as "floor of semi-deciduous forest and scrubs". The aim of this paper is to describe the main currently existing forest syntaxa on the vegetation floor before mentioned.

MATERIALS AND METHODS

DESCRIPTION OF THE STUDY AREA

The "floor of semi-deciduous forest and scrubs" from Sierra Maestra--where this kind of forest evolves--presents typically tropical ecological conditions; the weather is warm and the mean temperature is about 24[grados]C. The relative humidity varies from 80 to 85%, whereas the evaporation averaging from 1500 to 1900 mm. The annual rainfall for this area is from 800 to 1200 mm per year. It has two different seasons: one with a sharp little rainy season (November to April), ranging from 300 to 400 mm and the other, the rainy season (May to October), in which precipitation goes from 800 to 1 000 mm (VILAMAJO & al., 1989; MONTENEGRO, 1991a, b, c, d).

Geologically the study area is generally located in El Cobre Group (MENDEZ & al, 1994.) and in siallitic brown soils or reddish brown fersiallitic soils (sensu RENDA, 1989; HERNANDEZ & al, 1994). The areas corresponding to the Coastal Terraces in Southern Sierra Maestra (NUNEZ & VINA BAYES, 1989) are mainly composed by highly percolating Pliocene marl from La Cruz Formation (Cuban - Hungarian Commission, 1976), and by shallow to very shallow soils.

METHODOLOGY

In a minimum area of 400 [m.sup.2], vegetation inventories (stands, samples) were made by means of the BRAUN-BLANQUET (1921, 1951, 1964) method; using the SAMEK (1973) experiences, they were numbered and marked on the map 1:50 000 (Figure 1). There were also performed some observations of the sample's surroundings and ecological conditions: altitude, exposure, slope and soil (type, depth, texture, structure, stoniness, humidity, etc). organization of vegetation inventories and syntaxa separation were performed by means of phytocoenological methods (SCAMONI & PASSARGE, 1959, 1963). Homogeneity according to the law of RAUNKIAER (1934) was then confirmed. Species with grades of presence IV and V were considered for characteristic combination, and for subassociations and variants the differential combinations (SAMEK, 1973; Mueller -DOMBOIS & ELLEMBERG, 1974).

[FIGURA 1 OMITIR]

Generally, in the description of layers ([E.sub.1]--herbaceous, [E.sub.2]--shrub and [E.sub.3]--canopy layers) and synusiae (Li--lianas, Ep--epiphytes) the following categories of species presence were established: constants, present in 80 % or more of the inventories; frequently, present in 60 to 79 %; less frequent, present in 30 to 59 %; casual, present from 15 to 29 %. Species with values lower than 15 % were not named in the description. Reyes & Fornaris (2011) agreement that vegetable earth layers are: L withered leaves, F incompleted decomposition (leaching) and H humus.

Categorization and naming of syntaxa was performed according to the International Code of Phytosociological Nomenclature (WEBER & al, 2000). Flora de Cuba vols. 1-5 (LEON, 1946; LEON & ALAIN, 1951, 1953, 1957; ALAIN, 1964) was used to name species and corrections were made following modern publications (GUTIERREZ, 2000; HAGEN, 2007; ACEVEDO-RODRIGUEZ & STRONG, 2012). Collected specimen are in Herbarium BSC.

RESULTS

A large ecosystem diversity can be found, even at the classes, orders and alliances levels due to the great range of ecological conditions on the "floor of semi-deciduous forests and scrub" from the Sierra Maestra, where these syntaxa developed. Hence it is considered a complex phytosociological arrangement, which is observed at the end of work.

EUGENIO MALEOLENTIS-DIPHOLIDETALIA SALICIFOLIAE (Reyes 2005) Reyes stat. nov. (Holotypus: Eugenio-Dipholidion salicifoliae Reyes 2005).

Composition & characteristic species. Sideroxylon salicifolium, Gymnanthes lucida, Eugenia maleolens, E. axillaris, Bursera simaruba, Exostema caribaeum, Colubrina elliptica, Erythroxylum havanense, E. rotundifolium, Maytenus buxifolia, Amyris elemifera, Chiococca alba, Stigmaphyllon sagreanum, etc.

Physiognomy. Microphyll semi-deciduous forest, low, the arboreal layer (canopy layer) reaches between 7 to 12 m in high and covers from 30 to 100 %.

Ecology and distribution. It occupies siallitic brown soils from calcareous marl in the Coastal Terraces from Sierra Maestra and brown soils on the limestones from the north side of such mountain group. The rainfall range between 700 and 1 200 mm with a distinct dry season. It takes place on the "floor of semi-deciduous forest and scrubs" from Sierra Maestra.

Syntaxonomy. REYES & MARTINEZ (2005) placed the alliance Eugenio-Dipholidion salicifoliae Reyes 2005 in the class Gerascantho-Burseretea simarubae Borhidi 1996 and in the order Gerascantho-Burseretalia simarubae Borhidi 1996. Due to a better correspondence such alliance was transferred to the class Thrinaceto-Plumerietea obtusae Borhidi 1996, which raises its rank (Article 27 a) to the order Eugenio maleolentis-Dipholidetalia salicifoliae (Reyes, 2005) Reyes stat. nov. (Recommendation 46H) and equally maintains the alliance Eugenio-Dipholidion salicifoliae Reyes 2005 with their associations, original diagnosis and unaltered types.

Eugenio-Dipholidion salicifoliae Reyes 2005.

Canello winteranae-Sideroxyletum salicifolii

Reyes & Acosta ass. nova hoc loco (Table 1, holotypus rel. 2)

Physiognomy & floristic composition. This syntaxon constitutes a phase of syngenetic development, which corresponds to a Late successional community, i.e., the stage known as Fiera II, when arboreal species surpass the layer seen in Homeostasis I and begin interacting with one another, whereas the pioneers diminish their vitality. The canopy layer generally covers from 70 to 90 % and ranges between 9 and 12 m tall. Floristically, it is quite heterogeneous as it has averaged about 11 species by inventory (occasionally 15). It was found a total of 41 species that reach this layer. The most common species are: Sideroxylon salicifolium, Canella winterana, Acacia macracantha and Senna atomaria. With less coverage is also found Bur sera simaruba. The most abundant species in the shrub layer are Eugenia axillaris, Coccothrinax fragans and Randia aculeata are also observed. While Scleria lithosperma, Chiococca alba, Zapoteca gracilis, Paspalum breve, Lasiacis divaricata, Lantana montevidensis, Zanthoxylum fagara and Croton lucidus were the most abundant species in the herbaceous layer. The characteristic combination is shown in Tables 1-3 and consists of 21 species.

Ecology and distribution. The studied areas are hills between 20 and J00 m asl. Most are located at the north side of the Coastal Terraces of the South of Sierra Maestra (x- 6J0.708, y- 151.079), so their secondary watershed and dell preferably have north-northwest direction and Northwest's to a lesser degree. Geologically these hills are composed by Pliocene marl from La Cruz Formation. Soils are brown (carbonate), from shallow to very shallow and even react with saliva. They are usually clay loam, sandy clay or sandy, stony in greater or lesser level, with small and medium stones. The most frequent inclination is between 20 and 40 degrees, the micro-relief on the slope is generally flat. Regarding the litter, the L layer reaches about 2 cm; the F layer is generally absent and the humus is missing. The earthworm activity (which is higher in more developed successional stages) is very important in the incorporation of humus to the soil. Where such activity is greater, a mixed layer of soil (1 to 1.5 cm) and half-decomposed litter is formed, thus affecting the ecosystem nutrient recycling. The annual mean temperature is around 26[grados] C and the average maximum is in the order of 32[grados] C. The annual precipitation is about 800 mm and the yearly evaporation is between 1 700 and 1 900 mm.

It has three sub-associations: typicum (= simaroubetosum laevis), trichilietosum hirtae and diospyretosum grisebachii. Differences are caused by the degree of successional development and the ecological conditions, mainly the exposure and position on the slope.

typicum

It is the most advanced community in Fiera II. The main tree pioneers generally show diminished vitality and poor or very poor regeneration. However, the thickness of Sideroxylon salicifolium and Eugenia axillaris is very high in the arboreal layer. The proportion of trees seedlings in shrub and herbaceous layers is high. It covers 50 to 80 % and 40 to 70 % respectively, which also confirm the advanced successional stage. It is generally observed in exposures to the West and in the mid to upper hillside, so the relative height is greater than 20 m. The soil is very shallow, sometimes gravelly and with some stones on the surface. Earthworm, great activity is also considerable. Due to its inclination, position on the slope and exposure, it can be considered as a dry oecotope.

Two variants are found: typicum and Caesalpinia vesicaria (Table 1).

trichilietosum hirtae Reyes subass. nova hoc loco (Table 2, holotypus rel. 1)

This subassociation is an early stage of Late successional community; where Acacia macracantha and Senna atomaria still play an important role in the canopy layer; and the thickness of Sideroxylon salicifolium and Eugenia axillaris is still very low. The microphanerophytes and nanophanerophytes number is important in the shrub and herbaceous layers. The first covers between 25 and 70 % and the second from 10 to 90 %. Preferentially they occupy the slope bottom and are directed towards the North (NE-NW). Therefore, the water balance is more favourable here than in other sub-associations. Three variants with weak floristic differences are presented: typicum, with Tournefortia volubilis and with Trema micrantha (Table 2).

diospyretosum grisebachii Reyes subass. nova hoc loco

(Table 3, holotypus rel. 3)

This syntaxon has a more advanced successional stage than the previous. In its arboreal layer, pioneer species reach less thickness; whereas Sideroxylon salicifolium and Eugenia axillaris are important. In shrub and herbaceous layers, which typically cover from 50 to 80 % and from 40 to 70 % respectively, the number of tree seedlings increases compared to the previous syntaxon. It can be found in the lower slopes of the whole association, since only occasionally exceeds 20 degrees of inclination. The relative height is usually greater than 20 m. Two variants are presented: a typicum with a mean of 35.7 species and another variant of Zapoteca gracilis richer with 45.2 average, which preferentially occupies the lower parts of the slopes (Table 3).

Sideroxyletum salicifolio-foetidissimi Reyes & Acosta ass. nova hoc loco (Table 4, holotypus rel. J)

Physiognomy & floristic composition. This microphyll semi-deciduous forest was altered by intense selective logging, which has changed its structure, yet maintains its floristic composition. The canopy layer is 7 to J0 m high, so it is considered a low forest. Its cover varies from 30 to 50 % and it has also emerging trees (emergents) of about J5 m. Sideroxylon foetidissimum subsp. foetidissimum distinguishes for being constant and abundant. Bursera simaruba is also constant and locally abundant. Ocotea coriacea (sometimes abundant), Chrysophyllum oliviforme, Sideroxylon salicifolium and Exostema caribaeum (locally abundant) are frequent. Shrub layer is dense and has from 80 to 90 % coverage. They are constant and abundant: Eugenia sp. and Comocladia dentata, Exostema caribaeum, Adelia ricinella, Casearia hirsuta and Erythroxylum havanense (locally abundant) are equally constant. Ocotea coriacea (abundant), Alvaradoa amorphoides, Erythroxylum rotundifolium, Zanthoxylum fagara, Bursera simaruba and Chrysophyllum oliviforme are frequent. The herbaceous layer is sparse and has coverage of 20 to 30 %. Sideroxylon foetidissimum subsp. foetidissimum, S. salicifolium, Picramnia pentandra, Eugenia sp., Varronia globosa, Zapoteca gracilis, Desmodium incanum, Scleria lithosperma and Commelina elegans are constant. Concerning lianas, Stigmaphyllon sagreanum, Pisonia aculeata, Gouania lupuloides, Smilax havanensis, Serjania diversifolia and Chiococca alba (locally abundant) are constant. Selenicereus grandiflorus and Tragia hexandra are frequent. Regarding the epiphytes, only Tillandsia fasciculata is frequent. Litter has an L layer of 2 to 2.5 cm, and an F of 0.5 to 1 cm. The H layer is thin and forms a root mat of about a centimetre in some places. The strata and synusiae composition is shown in Table 4.

Ecology and distribution. It shows in "Charco Redondo" geological formation. The soil is brown, very shallow and sometimes rocky up to 80 %. The slope of the surface is small (5 %) and exposure is varied. Altitude varies from 230 to 240 m a.s.l.

Climatically, it is in the area where the influence of Sierra Maestra ends. The mean annual temperature ranges from 24 to 26[grados]C, with an average maximum of 32 to 34[grados]C and an average minimum of 18 to 20[grados]C. Rains in the area are between 1 000 and 1 200 mm, distributed in a rainy season from May to October and a less rainy season from November to April. It rains 40 to 60 days a year. The middling relative humidity is 75 to 80 % and the evaporation varies from 2 000 to 2 200 mm. The study was conducted at the place known as "El Granizo" (x: 565.474, y: J75.766), between Jiguani and Baire, in the hills of Northern edge of Sierra Maestra.

Oxandro lanceolatae-Burseretum simarubae Reyes & Acosta ass. nova hoc loco (Table 5, holotypus rel. 4)

Physiognomy & floristic composition. Although this vegetation type (mesophyll semideciduous forest) has been exploited for livestock development, the analyzed area has a high degree of naturalness. The canopy layer has a coverage ranging from 50 to 90 % and a height between 7 and 15 m. Bursera simaruba, Oxandra lanceolada, Cordia gerascanthus and Piscidia piscipula are constant. As frequent: Sideroxylon foetidissimum subsp. foetidissimum, Cupania americana, Guazuma ulmifolia, Spondias mombin, Phyllostylon brasiliensis and Lysiloma sabicu. As less frequent: Drypetes alba, Gymnanthes lucida, Cupania glabra, Zuelania guidonia, Poeppigia procera, Roystonea regia, Trophis racemosa, Sideroxylon salicifolium and Carpodiptera cubensis. The shrub layer has a coverage ranging from 40 to 90 %. Constant species are: Erythroxylum havanense and Picramnia pentandra. Are frequent species Coccothrinax gundlachii and Trichilia hirta. As less frequent Capparis flexuosa, Casearia sylvestris var. myricoides and Zapoteca gracilis. The herbaceous layer is relatively poor. It covers between 30 and 50 % of the surface, and only has Olyra latifolia as constant. There are in the category of less frequent: Oeceoclades maculata, Pharus glaber, Urochloa humidicola and Rivina humilis. Lianas in turn occur as follows: Serjania diversifolia, Gouania lupuloides, Pisonia aculeata and Stigmaphyllon sagreanum are constant. Epiphytics are underrepresented, showing Tillandsia recurvata as constant and T. usneoides as occasional. The composition of life forms in this syntaxon is as follows: 40.5 % of the species are trees, 24.3 % are shrubs, 13.5 % make up the herbaceous layer, 2.7 % are ferns, 16.2 % are lianas and 4.1 % are epiphytes. Litter is unevenly distributed. The L layer ranges between 1.5 and 3 cm; occasionally more, but in isolated nano-depressions it can reach 15 cm. F and H layers are almost imperceptible, and only the last is observed mixed with the horizon A.

Two variants are presented: typicum and Poeppigia procera.

typicum variant. It tends to be in greater soil depth. It does not have a well defined differential combination, but there are missing many species observed in the other variant.

Poeppigia procera variant. This syntaxon is preferably found in very shallow soils. The differential combination is very well defined (see Table 5).

Ecology and distribution. The micro-relief is generally flat. It locally presents rocky outcrops on the surface. The slope is also very variable, ranging between 20 and 40 degrees. The exposure has two main trends: one towards east and the other to the south. The most important soil is the reddish brown fersiallitic, whose depth varies from very shallow to moderately deep; well drained, both internal and external. The annual rainfall in this area is 1000 to 1200 mm per year, having two distinct seasons: the less rainy season (November to April) with an average of 300 to 400 mm and the rainy (May to October) with ranges from 800 to 1000 mm. Altitude varies between 120 and 200 m asl. This syntaxon occurs on the southern slope of Sierra Maestra mountain range. At present, the most extensive relicts are located near the mouth of Turquino River (x: 508.527, y:144.674).

Zanthoxylo fagarae - Guazumetum ulmifoliae Reyes & Acosta ass. nova hoc loco (Table 6, holotypus rel. 1)

Physiognomy & floristic composition. This secondary syntaxon is a late successional community known as 'Fiera II'. The canopy layer increases in size along with the development of the forest, from 6-8 m to 10-12 m and 100 % of cover. A real stratification between arboreal and shrub layers is missing in the lower parts. The most abundant species are Guazuma ulmifolia, Zanthoxylum fagara and Senna atomaria; sometimes, Casearia sylvestris var. sylvestris and Cupania americana.

The herbaceous layer covers between 30 and 60 %. The most abundant species vary locally and they are: Erythroxylon havanense, Samanea saman and Spermacoce laevis. They are also accompanied by a large number of species with low constancy. This community has a rich characteristic combination, which can be seen in Table 6. Litter is often presented in a dispersed way. Layers L and F are mixed in the structurally less evolved parts, and they are no more than a centimetre. On the contrary, in the more developed parts, the L layer is about 1.5 cm and F, 0.5 cm. H layer is missing.

Ecology and distribution. It lies on rocks from the undifferentiated El Cobre Group. The soils are yellowish brown, very shallow, with sharp sheet erosion and about 5 % of stones on the surface. Such surface is somewhat irregular and preferably occupies the middle and lower basins. It was studied in the environs of the town El Cobre (Santiago de Cuba, x: 592.474, y: 156.371).

Oxandro lanceolatae-Spondiadetum mombin Reyes & Acosta ass. nova hoc loco (Table 7, holotypus rel. 1)

Physiognomy & floristic composition. The canopy layer has two sub-strata: the upper is about 20 m, with emergents of 25 m (sometimes this layer reaches the latter height); and the lower, which contrary is between 8 and 10 m, occasionally 12 to 15 m. Both cover 100% of the surface.

This layer is very prolific, its arrangement is shown in Table 7. Coverage of shrub layer range from 20 to 30% and is composed almost entirely of tree species. The herbaceous layer is poor and extends over 10% of the surface. Nearly half of its species are also trees. The humus layers are not well developed. The L layer is between 3.5 and 5 cm thick, covering about 80 % of the surface. F layer is almost imperceptible, scattered among the rocks and gravel, while H layer is missing.

Ecology and distribution. This association is relatively scarce and was studied in the middle of Turquino River (x: 506.532, y: 148.853). It comes near the river, at the bottom of the slope and therefore receives the contribution of its runoff. Besides, being in that landscape position, it is influenced by the night gravitational winds; therefore the water balance is favourable. The weather is warm. The mean annual temperature is around 24[grados]C. The relative humidity varies between 80 and 85 %. The annual rainfall is between 1400 and 1600 mm, while evaporation ranges from 1500 to 1700 mm. The study area corresponds to rocks from El Cobre Group. The soil is siallitic brown, deep, with gravel in the profile (20 %). It also has 40 to 100 % of rocks on the surface.

DISCUSSION

The Sierra Maestra is the mountain range with the highest peaks of the Cuban archipelago (Pico Turquino with 1972 m a.s.l). That's why it shows a vegetation altitudinal zonation from the cloud forests in the upper parts up to the semi-deciduous forests and shrubs in the lowest (Reyes, 2006, 2011-2012). Interestingly, a zonation with similar altitudes (up to 2000 m a.s.l) was described in the Venezuelan Guayana (HUBER, 1995). The difference in altitudinal floors (zones) is produced by the diversity of ecological conditions caused by altitude, mainly due to moisture dissimilarities and its interrelation with temperature. The semi-deciduous forests that develop in Sierra Maestra below 500 m asl, i.e., in flat, hilly and premontane areas ("floor of semi-deciduous forest and scrub") change in mesophyllous and microphyllous, according to the ecological conditions. Mesophyllou's develop in the most favourable oecotopes (mainly due to moisture), while microphyllous, in drier and oligotrophic areas. They likewise form a syntaxa complex in correspondence with the oecotopes properties (Reyes & al, 2004a, b). Since the mesophyll semi-deciduous forests--the most anthropised vegetation type of Sierra Maestra--the relicts where it maintains an acceptable structural and floristic composition are scattered. In turn, middle zones are occupied by economic areas and different successional stages. A microphyll semi-deciduous forest with many sclerophyllous species locally presents in stressful environmental conditions, caused by a poor soil development and / or a marked water deficit, often with accentuated wind affection (BORHIDI, 1996).

CONCLUSIONS

According to the wide range of ecological conditions, both geological and / or soil and climatic; variation occurs in mesophyll and microphyll semi-deciduous forests (the last under the most stressful conditions). These forests are different classes. This fact and the syngenetic evolution stages produce great syntaxonomical diversity, since two classes, four orders, five alliances, 11 associations, 12 subassociations and 11 variants are observed.

SYNTAXONOMICAL SCHEME

THRINACETO-PLUMERIETEA OBTUSAE Borhidi 1996

Eugenio-Dipholidetalia salicifoliae (Reyes 2005) Reyes stat. nov.

Eugenio-Dipholidion salicifoliae Reyes 2005

Sideroxyletum salicifolio-foetidissimi Reyes & Acosta ass. nov.

Canello winteranae-Sideroxyletum salicifolii Reyes & Acosta ass. nov.

trichilietosum hirtae Reyes subass. nov.

var. typicum

var. Tournefortia volubilis

var. Trema micrantha

diospyretosum grisebachii Reyes subass. nov.

var. typicum

var. Zapoteca gracilis

GUAZUMO-CEIBETEA PENTANDRAE Borhidi 1996

Lonchocarpo sericeo-Ceibetalia pentandrae Borhidi & Muniz 1996

Guazumo-Cupanion glabrae Borhidi & Del Risco in Borhidi 1991

Oxandro lanceolatae-Burseretum simarubae Reyes & Acosta ass. nov.

var. typicum

var. Poeppigia procera

Oxandro-Burseretalia Borhidi & Muniz in Borhidi 1991

Trichilio-Guazumion ulmifoliae Reyes 2004

Zanthoxylo fagarae-Guazumetum ulmifoliae Reyes & Acosta ass. nov.

Picramnio-Ocoteion coriaceae Reyes 2004

Oxandro lanceolatae-Spondiadetum mombin Reyes & Acosta ass. nov.

Clarification. Reyes & Acosta (2012) exposed in Lazaroa 33 (pp. 114) the alliance Bactri cubensis-Pinion cubensis Reyes 2012, whose spelling must be corrected (Article 41b) to Bactrio cubensis-Pinion cubensis Reyes 2012, due to the incorrect vocal union.

doi: 10.5209/rev_LAZA.2014.v35.43800

Received: 26 December 2013

Accepted: 17 October 2014

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Reyes, O.J.--2006--Clasificacion de la vegetacion de la Sierra Maestra--Biodiversidad de Cuba Oriental, vol. VIII. pp. 23-41. Ed. Academia.

Reyes, O.J.--2011-2012--Clasificacion de la vegetacion de la Region Oriental de Cuba--Rev. Jard. Bot. Nal. Univ. Habana 32-33: 59-71.

Reyes, O.J. & Martinez Quesada, E.--2005--Fitocenosis presentes en las areas costeras del sur de la Sierra Maestra. IV. Asociaciones Colubrino ellipticae-Gymnanthetum lucidae y Eugenio-Dipholidetum salicifoliae--For. Veracruzana 7(1): 47-52.

Reyes, O.J., Martinez Quesada, E. & Acosta Cantillo, F.--2004a--Comunidades secundarias de la Region Oriental de Cuba. I. Clase Bothriochloetea pertusae Reyes --Biodiversidad de Cuba Oriental, vol. VII, pp. 92-102. Ed. Academia.

Reyes, O.J., Acosta Cantillo, F. & Martinez Quesada, E.--2004b--Comunidades secundarias de la Region Oriental de Cuba. II. Clase Senno - Acacietea macracanthae Reyes--Editorial Academia. Biodiversidad de Cuba Oriental, Vol. VII. pp. 102-115.

Reyes, O.J. & Acosta Cantillo, F.--2004a--Comunidades secundarias de la Region Oriental de Cuba. III. Asociaciones Trichilio hirtae - Guazumetum ulmifoliae, Cupanio --Guazumetum ulmifoliae y Helictereti semitrilobae--Colubrinetum ellipticae--Editorial Academia. Biodiversidad de Cuba Oriental, Vol. VII. pp. 115-126.

Reyes, O.J. & Acosta Cantillo, F.--2004b--Comunidades secundarias de la Region Oriental de Cuba. IV. Asociaciones Cupanio glabrae - Ocoteetum coriaceae, Ocoteo coriaceae - Alvaradoetum arborescentis y Guareo guidoniae --Dendropanacetum arborei--Editorial Academia. Biodiversidad de Cuba Oriental, Vol. VII. pp. 126-139.

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Orlando J. Reyes & Felix Acosta Cantillo (*)

* Eastern Center of Ecosystems and Biodiversity (BIOECO). Ministry of Science, Technology and Environment (CITMA). Jose A. Saco Nr. 601, esq. Barnada. Santiago de Cuba, Cuba. E mail: joel@bioeco.ciges.inf.cu
Table 1

Canello winteranae-Sideroxyletum salicifolii ass. nova
var. typicum (1-3), var. Caesalpinia vesicaria (4-6)
(Eugenio-Dipholidion salicifoliae, Eugenio maleolentis
-Dipholidetalia salicifoliae, Thrinaceto-Plumerietea
obtusae)

Inclination ([grados])              40    20    20   30    20    25
Exposition                          WSW   SSW   N    NNW   SSW   NNE
Canopy layer-[E.sub.3] (%)          70    60    70   70    80    70
Shrub layer-[E.sub.2] (%)           70    80    80   80    50    60
Herbaceous layer-[E.sub.1] (%)      60    70    50   50    40    30
N. species                          30    27    22   37    37    30
Releve N.                            1     2    3     4     5     6

Characteristics

[E.sub.3]-Sideroxylon                1     3    3     2     2     3
  salicifolium
Canella winterana                          +    +     +     r     +
Acacia macracantha                   4     r    +     +     r     r
Senna atomaria                       2     r    +     r     1
Bursera simaruba                     r     +    r     +     +     +
[E.sub.2]-Eugenia axillaris          r     2    2     2     2     2
Randia aculeata                      1     +    1     +     +     2
Lantana montevidensis                1          +     +     r     r
Comocladia dentata                                    +     r     +
Bourreria virgata                    +                            r
[E.sub.1]-Coccothrinax fragans       1     +    +     +           1
Croton lucidus                       3     3    1     1     1     1
Zapoteca gracilis                    +     r    r     r     1
Zanthoxylum fagara                         +    +     r     r     r
Scleria lithosperma                  r     2    1     2           2
Paspalum breve                             2          1     1     1
Lasiacis divaricata                  r     +          +     +
Li-Tragia hexandra                         +    +     r     r     +
L-Stigmaphyllon sagreanum            +     r          r     +     +
Chiococca alba                       r     1          1     1     +
Smilax havanensis                    r     +          +     r     +

Differentials of typicum

[E.sub.3]-Simarouba laevis                 1    1     +     r     +
[E.sub.2]-Amyris elemifera           r     2    1           +     1
Li-Tragia volubilis                        +          r     r     +

Differentials of trichilietosum hirtae and diospyretosum grisebachii

Li-Salacia nipensis                  1                      r     r

Differentials of trichilietosum hirtae

[E.sub.1]-Erythroxylum                                r     +
  havanense
Commelina erecta                                      r     +

Differentials of diospyretosum grisebachii

[E.sub.3]-Exostema caribaeum         r     1          1     +     1
Colubrina elliptica                  1                2     2     r
Thouinia patentinervis                                      r     r
[E.sub.1]-Lantana camara             +                            r
Spermacoce laevis                                     r     r
Li-Passiflora suberosa               +                r     r     r

Releve N.                            1     2    3     4     5     6
Differentials of variants
[E.sub.2]- Caesalpinia vesicaria                      3     3     +
Capparis flexuosa                                     r     r
Companions
[E.sub.1]- Oplonia tetrasticha             1                r     1
Varronia globosa subsp. humilis            r          r
Melochia nodiflora                   r                +
Li- Passiflora foetida               r                      r
Distictis rhynchocarpa                     +          1     +
Ep- Tillandsia fasciculata           r     r

Other species: Euphorbia heterophylla, Tillandsia recurvata and
Casearia hirsuta r, Callicarpa resinosa +, Both-riochloapertusa
4 in 1; Erithalis fruticosa + in 2; Diospyrus grisebachii,
Sideroxylon foetidissimum subsp. foetidissimum, Ehretia tinifolia
and Piper richardianum r, Ocotea coriacea and Cupania glabra 1
in 3; Triopteris rigida +, Urochloa humidicola 1, Turnera ulmifolia
and Citharexylum ternatum r in 4; Erytrhroxylum rotundifolium +,
Desmodium incanum and Pisonia aculeata r in 5; Rochefortia
acanthophora + in 6.

Localities: all releves were collected in the hills at the north
side of the Coastal Terraces of the South of Sierra Maestra,
between 20 and 100 m asl. (x: 610.708, y: 151.079); holotypus
rel. 2.

Table 2

Canello winteranae-Sideroxyletum salicifolii trichilietosum
hirtae subass. nova var. typicum (1), var. Tournefortia
volubilis (2, 3), var. Trema micrantha (4-6), (Eugenio
-Dipholidion salicifoliae, Eugenio maleolentis-Dipholidetalia
salicifoliae, Thrinaceto-Plumerietea obtusae)

Inclination ([grados])              30   10    30   35   30   45
Exposure                            NW   ENE   NE   NW   NE   NNW
Canopy layer-[E.sub.3] (%)          90   90    90   80   70   80
Shrub layer-[E.sub.2] (%)           30   25    50   60   70   60
Herbaceous layer-[E.sub.1] (%)      70   90    60   40   15   10
N. species                          31   40    47   42   36   35
Releve N.                           1     2    3    4    5     6

Characteristics

[E.sub.3]-Sideroxylon               +     +    +    2    1
  salicifolium
Canella winterana                   r     r    r    +    1     r
Acacia macracantha                  4     3    4    1    +     2
Senna atomaria                      +     2    3    1    1     +
Bursera simaruba                    +     r         r    +     1
[E.sub.2]-Eugenia axillaris         +          1    3    3     2
Comocladia dentata                  r     2    r    1    r
Bourreria virgata                   +     r    +    +    r     r
Lantana montevidensis                          +    r
Randia aculeata                           r    r    r          +
[E.sub.1]-Coccothrinax fragans            r    r    r    r     2
Scleria lithosperma                 1     2    1    1    2     +
Lasiacis divaricata                 1     1    2    r          +
Paspalum breve                      +     3    r    1    +
Zapoteca gracilis                   +     1              1     +
Croton lucidus                                      3          1
Zanthoxylum fagara                        r         r
L-Stigmaphyllon sagreanum           +     r    +    r          1
Chiococca alba                      2     1    +    1    1     1
Smilax havanensis                   +     r    +               +
Tragia hexandra                                     +    +

Differentials of Trichilietosum hirtae

[E.sub.3]-Trichilia hirta           +     3    r    3    1     r
Sideroxylon foetidissimum           +     r    1    +    +     +
  subsp. Foetidissimum
Cordia alba                         +     r    2    2          +
[E.sub.2]-Ehretia tinifolia         +     2    +    1    +     +
Turnera ulmifolia                   1     r    1    r          +
[E.sub.1]-Erythroxylum havanense    +     r    +    1    1     +
Chrysophyllum oliviforme            r     r    r    r    1
Casearia hirsuta                          r    +    r    1     r
Urochloa humidicola                 1     2    2    r    +     1
Heliotropium indicum                      r    +    r          +
Commelina erecta                    +     r    +               r
Desmanthus virgatus                       1    r    r    1
Li-Gouania lupuloides               2     1    r    1    1     +
Serjania diversifolia               +          +    r    r     +
Salacia nipensis                    +          +    r    r
Ep-Tillandsia recurvata                   r    r    r    1

Differentials of Diospyretosum grisebachii and Simaroubetosum laevis

[E.sub.3]-Colubrina elliptica                            1     2
Li-Passiflora suberosa                         r               r

Differentials of Diospyretosum grisebachii

[E.sub.1]-Lantana camara                  r    +               r
Euphorbia heterophylla              r          r

Differentials of variants

[E.sub.1]-Bourreria                       +    1
  setoso-hispida
Tournefortia volubilis                    r    r
[E.sub.1]-Trema micrantha                           r    1     r
Cupania glabra                                      +    2
Li-Pisonia aculeata                       r    r

Companions

[E.sub.3]-Lonchocarpus                         +    1          +
  domingensis
[E.sub.2]-Adelia ricinella                r         +
[E.sub.1]-Desmodium incanum                    +         r
Varronia globosa subsp. humilis                r    1
Li-Cissampelos pareira                         r    1    1
Centrosema virginianum              r          2
Cissus verticillata                       r         r

Other species: Callicarpa resinosa r in 1 Passiflorafoetida,
Spermacoce laevis and Tillandsiafasciculata r, Zuelania guidonia + in
2; Bothriochloa pertusa and Rochefortia acanthophora +, Melochia
nodiflora and Varronia lineata r in 3; Ocotea coriacea r in 4;
Diospyrus grisebachii and Lonchocarpus longipes +, Caesalpinia
vesicaria 1, Erytrhroxylum rotundifolium and Capparis flexuosa r
in 5; Oplonia tetrasticha +, Citharexylum ternatum and Triopteris
rigida r in 6. Localities: all releves were collected in the hills
at the South of Sierra Maestra, between 20 and 100 m asl; holotypus
rel. J.

Table 3

Canello winteranae--Sideroxyletum salicifolii diospyretosum
grisebachii subass. nova var. typicum (1-4), var. Zapoteca
gracilis (5-8) (Eugenio-Dipholidion salicifoliae, Eugenio
maleolentis-Dipholidetalia salicifoliae, Thrinaceto
-Plumerietea obtusae)

Inclination ([grados])                    20     5    35
Exposition                          NE    NNW   NE    NNE
Canopy layer-[E.sub.3] (%)          50    80    80    80
Shrub layer-[E.sub.2] (%)           80    50    50    60
Herbaceous layer-[E.sub.1] (%)      30    40    70    40
N. species                          38    41    31    33
Releve N.                            1     2     3     4

Characteristics

[E.sub.3]-Sideroxylon                3     2     3     4
  salicifolium
Canella winterana                    r     r           1
Acacia macracantha                   1     +     +     +
Senna atomaria                       r     +     +
Bursera simaruba                     +     +     r     +
[E.sub.2]-Eugenia axillaris          4     4     3     1
Comocladia dentata                   1     1           1
Randia aculeata                      +     +     +     +
Lantana montevidensis                +     r     r     +
Zanthoxylum fagara                   +     r     r
Bourreria virgata                    r           +
[E.sub.1]-Coccothrinax fragans       +     +           +
Croton lucidus                       3     2     3     2
Lasiacis divaricata                  1     1           +
Paspalum breve                             1     1     r
Scleria lithosperma                  2     2     2     1
Li-Tragia hexandra                   +     +     +     +
Chiococca alba                       1     1     1     1
Smilax havanensis                    r     r     r     r
Stigmaphyllon sagreanum              r     r     r

Differentials of trichilietosum hirtae and diospyretosum
grisebachii

[E.sub.3]-Sideroxylon                            r
  foetidissimum subsp.
  Foetidissimum
Li-Salacia nipensis                        r           +
Gouania lupuloides                         1

Differentials of diospyretosum grisebachii and
simaroubetosum laevis

[E.sub.3]-Exostema caribaeum         +     r     +     2
Colubrina elliptica                  r     1     2     2
Li-Passiflora suberosa               r     r           r

Differentials of diospyretosum grisebachii

[E.sub.3]-Diospyrus grisebachii      r     +     +     r
Thouinia patentinervis                     r           r
[E.sub.2]-Trema micrantha                  r     +
Erithalis fruticosa                  1     1     r     +
Lantana camara                       r
[E.sub.1]-Spermacoce laevis          r     r
Bourreria setoso-hispida                   r     r
Varronia lineata                     +     r
Euphorbia heterophylla               r     r
Li-Cissus verticillata               1     +     1

Differentials of trichilietosum hirtae

[E.sub.2]-Erythroxylum havanense
Chrysophyllum oliviforme
[E.sub.1]-Urochloa humidicola              1
Casearia hirsuta                                       +
Li-Serjania diversifolia                         r

Differential of simaroubetosum laevis

[E.sub.3]-Amyris elemifera                       +     1

Differentials of variants

Ej-Zapoteca gracilis
Varronia globosa subsp.
  humilis
Heliotropium indicum
Desmodium incanum
Desmanthus virgatus
Melochia nodiflora
Commelina erecta
Li-Distictis rhynchocarpa
Triopteris rigida

Companions

[E.sub.3]-Zuelania guidonia                            r
Lonchocarpus longipes                                  r
Cupania glabra                             r
Ocotea coriacea                            r
Citharexylum ternatum                +
[E.sub.2]-Rochefortia                r
  acanthophora
Caesalpinia vesicaria                r
Adelia ricinella                           r
Ej-Morinda royoc                           r     r     r
Erytrhroxylum rotundifolium          r           +     +
Tournefortia volubilis                                 r
Wissadula amplissima                             r
Piper richardianum                                     r
Callicarpa resinosa                                    r
Oplonia tetrasticha                  1           +
Li-Centrosema virginianum
Pisonia aculeata                     +
Passiflora foetida                   r
Ep-Tillandsia fasciculata            r
Tillandsia recurvata                 r     r

Inclination ([grados])                    30    20     5
Exposition                                NNE   NW    NNE
Canopy layer-[E.sub.3] (%)          80    70    80    70
Shrub layer-[E.sub.2] (%)           70    80    80    60
Herbaceous layer-[E.sub.1] (%)      90    50    50    40
N. species                          51    38    51    41
Releve N.                            5     6     7     8

Characteristics

[E.sub.3]-Sideroxylon                1     4     2     4
  salicifolium
Canella winterana                    r     r     r     +
Acacia macracantha                   3     +     4
Senna atomaria                       2     +     3     r
Bursera simaruba                     r     +     +     r
[E.sub.2]-Eugenia axillaris          3     4     +     1
Comocladia dentata                   3     1     r     +
Randia aculeata                      +     r     +     +
Lantana montevidensis                2     1
Zanthoxylum fagara                   r     +     r
Bourreria virgata                    +           +     1
[E.sub.1]-Coccothrinax fragans       1     r     r     +
Croton lucidus                             2     4     1
Lasiacis divaricata                  +           r     +
Paspalum breve                       3           2     1
Scleria lithosperma                  4
Li-Tragia hexandra                   +     r           +
Chiococca alba                       1     1     +     1
Smilax havanensis                    1           r     +
Stigmaphyllon sagreanum              +     r     r

Differentials of trichilietosum hirtae and diospyretosum
grisebachii

[E.sub.3]-Sideroxylon                r     r     r     +
  foetidissimum subsp.
  Foetidissimum
Li-Salacia nipensis                  +     r           r
Gouania lupuloides                         1     +     1

Differentials of diospyretosum grisebachii and simaroubetosum
laevis

[E.sub.3]-Exostema caribaeum         +           r     r
Colubrina elliptica                  +                 2
Li-Passiflora suberosa                     r     r

Differentials of diospyretosum grisebachii

[E.sub.3]-Diospyrus grisebachii      r           +     2
Thouinia patentinervis               r           r     r
[E.sub.2]-Trema micrantha                  r     r     r
Erithalis fruticosa                  +                 2
Lantana camara                       r     r     r     r
[E.sub.1]-Spermacoce laevis          +           r     +
Bourreria setoso-hispida             r     r           r
Varronia lineata                     3     r     +     r
Euphorbia heterophylla               r     r     r     r
Li-Cissus verticillata               +     1

Differentials of trichilietosum hirtae

[E.sub.2]-Erythroxylum havanense     r     +     1
Chrysophyllum oliviforme             r     1     +
[E.sub.1]-Urochloa humidicola        1           +
Casearia hirsuta                                 +
Li-Serjania diversifolia                   r     r

Differential of simaroubetosum laevis

[E.sub.3]-Amyris elemifera                       r

Differentials of variants

Ej-Zapoteca gracilis                 +     1     3     2
Varronia globosa subsp.              +     r     r
  humilis
Heliotropium indicum                 r           r
Desmodium incanum                    r                 r
Desmanthus virgatus                  r           r
Melochia nodiflora                         r     r
Commelina erecta                           r     r
Li-Distictis rhynchocarpa            +     r           1
Triopteris rigida                    r     r     +     r

Companions

[E.sub.3]-Zuelania guidonia                            +
Lonchocarpus longipes                +                 +
Cupania glabra                                   1
Ocotea coriacea                            1
Citharexylum ternatum                      r     r
[E.sub.2]-Rochefortia                                  r
  acanthophora
Caesalpinia vesicaria                            1     r
Adelia ricinella                                 r
Ej-Morinda royoc                     r
Erytrhroxylum rotundifolium                            r
Tournefortia volubilis               r     r
Wissadula amplissima                       r     r
Piper richardianum                               +
Callicarpa resinosa                  +
Oplonia tetrasticha                              r
Li-Centrosema virginianum            +                 r
Pisonia aculeata                                 +
Passiflora foetida                                     r
Ep-Tillandsia fasciculata            +                 r
Tillandsia recurvata                             r

Other species: Cissampelos pareira and Turnera ulmifolia r,
Bothriochloa pertusa 2 in 1 Ehretia tinifolia + in 3;
Lonchocarpus domingensis r in 6.

Localities: all releves were collected in the hills at the South
of Sierra Maestra, between 20 and !00 m asl; holotypus rel. 3.

Table 4

Sideroxyletum salicifolio-foetidissimi ass. nova
(Eugenio-Dipholidion salicifoliae, Eugenio
maleolentis-Dipholidetalia salicifoliae,
Thrinaceto-Plumerietea obtusae)

Altitude (m a.s.l.)                  230      240      230
Exposure                              E       NE        W
Canopy layer-[E.sub.3] (%)           30       40       50
Shrub layer-[E.sub.2] (%)            80       90       80
Herbaceous layer-[E.sub.1] (%)       20       30       20
N. species                           54       45       39
Releve N.                             1        2        3

Characteristics

[E.sub.3,1]-Sideroxylon               2        2        3
  foetidissimum
Bursera simaruba                      1        1        +
[E.sub.3,2]-Sideroxylon               1        1        +
  salicifolium
Ocotea coriacea                       1        2        1
Exostema caribaeum                    1        +        1
Exothea paniculata                             2        1
[E.sub.3,2,1]-Comocladia              2        2        1
  dentata
Cupania glabra                        r        +        +
Adelia ricinella                      1        +        1
Alvaradoa amorphoides                 r        r        +
[E.sub.2]-Eugenia sp.                 3        2        4
Guettarda sp.                         1        +
Senna atomaria                        r                 +
Zanthoxylum fagara                    r                 r
[E.sub.2,1]-Picramnia                 r        +        +
  pentandra
Casearia hirsuta                      +        +        +
Erythtroxylum havanense               2        1        1
E. rotundifolium                      +        +        +
Varronia globosa                      r        r        r
Capparis flexuosa                     r        r
Coccoloba diversifolia                r        +
Erithalis fruticosa                   r        r
Chrysophylum oliviforme               1                 +
[E.sub.1]-Zapoteca gracilis           r        +        +
Desmodium incanum                     1        +        +
Scleria lithosperma                   +        r        1
Commelina elegans                     +        r        r
Rivina humilis                                 r        r
Oeceoclades maculata                           1        +
Lasiacis divaricata                   r        r
Selenicereus grandiflorus             +        r
Agave underwoodi                      r                 r
Urochloa humidicola               [r.sub.0]         [+.sub.0]
Li-Pisonia aculeata                   1        +        +
Smilax havanensis                     1        1        +
Stigmaphyllon sagreanum               1        +        +
Gouania lupuloides                    1        r        r
Serjania diversifolia                 +        +        r
Chiococca alba                        1        1        +
Turbina corymbosa                     1        r
Tragia hexandra                       +        r
Lantana camara                        r                 r
Ep-Tillandsia fasciculata             r                 r

Other species: Cedrela odorata, Clusia rosea,
Lonchocarpus domingensis, Colubrina elliptica, Citharexylum
caudatum, Waltheria indica, Eupatorium sp., Tabebuia
sp., Melicoccus bijugatus and Distictis rhynchocarpa r,
Colubrina ferruginosa, Hyperbaena longiuscula and
Morinda royoc +, Bothriochloa pertusa 2 in 1; Swietenia
mahagoni 2, Gymnanthes lucida, Drypetes alba and
Hyperbaena longiuscula +, Palicourea sp., Canella alba,
Vitis tiliaefolia and Abrus precatorius r in 2; Thouinia
sp. 1, Rhamnaceae, Zamia sp. and Rutaceae r in 3.
Localities: All releves collected at the hills of northern
Sierra Maestra ('El Granizo'; x: 565.474, y: 175.766),
between Jiguani and Baire; holotypus ass. rel. 1.

Table 5

Oxandro lanceolatae-Burseretum simarubae ass. nova
var. typicum (1-4), var. Poeppigiaprocera (5-9)
(Guazumo-Cupanion glabrae, Lonchocarpo sericeo
-Ceibetalia pentandrae, Guazumo-Ceibetea pentandrae)

Altitude (m a.s.l.)              160   155   150   145   120
Inclination ([grados])           35    40    30    35    25
Exposition                       NE    SW     S    NE    SW
Canopy layer--[E.sub.3] (%)      90    70    50    80    70
Shrub layer-[E.sub.2] (%)        50    50    80    40    80
N. species                       27    27    29    25    38
Releve N.                         1     2     3     4     5

Characteristics

[E.sub.3]-Bursera simaruba        3     3     2     1     3
Oxandra lanceolata                3     2     3     +     3
Cordia gerascanthus               2     1     r           r
Piscidia piscipula                r     r     +     +     1
Sideroxylon foetidissimum         1     r                 1
Cupania americana                 r                 r     r
Guazuma ulmifolia                             +     +     r
Spondias mombin                   r     1     r     r
Phyllostylon brasiliensis         +           2     +     2
Lysyloma sabicu                   r     r                 r
[E.sub.2]-Erythroxylum            r     r     +     r     1
  havanense
Picramnia pentandra               2     r     r           1
Coccothrinax gundlachii           r     2     r     3     1
Trichilia hirta                   r           r           r
Ej-Olyra latifolia                2     1     1     1     r
Oeceoclades maculata              +           r           r
Li-Serjania diversifolia          r     r     +     r     r
Gouania lupuloides var            r     +     +     +     +
  lupuloides
Pisonia aculeata                  r     r     +     +     +
Stigmaphyllon sagreanum           r     r     r     r     r
Ep-Tillandsia recurvata                 r     r     r     r

Differentials

[E.sub.3]-Drypetes alba           1
Gymnanthes lucida
Cupania glabra var. glabra
Zuelania guidonia
Poeppigia procera
Roystonea regia
Trophis racemosa
Sideroxylon salicifolium                                  r
Carpodiptera cubensis
Exothea paniculata
Geoffroea inermis
[E.sub.1]-Pharus glaber
Li-Trichostigma octandrum
[E.sub.3]-Calycophyllum                       r     1     +
  candidissimun
Collubrina reclinata                          r     1     r
[E.sub.2]-Capparis flexuosa             r     r     r     r
[E.sub.1]-Urochloa humidicola           1     2     r
Chlorophora tinctoria                   r     +           1
Eugenia sp.                                   +     2     +
Commelina elegans                             +     r     r

Accompaniers

[E.sub.3]-Prunus myrtifolia
Ficus membranacea                                         r
[E.sub.2]-Casearia sylvestris     r           r
  var. myricoides
Senna atomaria                                r           r
[E.sub.1]-Rivina humilis          r                       r
Zapoteca gracilis                       2     +
Hyperbaena paucinervis            r
Malpighia suberosa                                  r
Croton lucidus                          1           2
Desmodium incanum                                         r
Varronia leptoclada                                 +
Petiveria alliacea                      r                 r
Li-Triopteris rigida                                      r
Passiflora suberosa                                       r
  var. pallida
Cissus verticillata                                       r
Chiococca alba                          +
Pithecoctenium echinatum                                  r
Ep-Tillandsia usneoides           r

Altitude (m a.s.l.)              150   195   185   200
Inclination ([grados])           30    40    20    32
Exposition                       NE    ENE   ESE   SE
Canopy layer-[E.sub.3] (%)       75    60    70    80
Shrub layer-[E.sub.2] (%)        90    70    80    60
N. species                       43    32    35    35
Releve N.                         6     7     8     9

Characteristics

[E.sub.3]-Bursera simaruba        2     2     2     2
Oxandra lanceolata                3     3     2     2
Cordia gerascanthus               1     +     +     r
Piscidia piscipula                1     r           r
Sideroxylon foetidissimum         1     1     1     r
Cupania americana                 1     +     1     1
Guazuma ulmifolia                 r     1     1     1
Spondias mombin                   r           1
Phyllostylon brasiliensis         1     1
Lysyloma sabicu                   r           +     r
[E.sub.2]-Erythroxylum            1     r     +     r
  havanense
Picramnia pentandra               1     1     r     r
Coccothrinax gundlachii           +     r
Trichilia hirta                   r           r     r
Ej-Olyra latifolia                2     +     1     +
Oeceoclades maculata              r           r     r
Li-Serjania diversifolia          r     r     r     r
Gouania lupuloides var            r           1     +
  lupuloides
Pisonia aculeata                  r     +     +
Stigmaphyllon sagreanum           r     r     r     r
Ep-Tillandsia recurvata           +     r     r     r

Differentials

[E.sub.3]-Drypetes alba           +     2     1     3
Gymnanthes lucida                 2     1     +     1
Cupania glabra var. glabra        1     r     +     r
Zuelania guidonia                 +     +     +     1
Poeppigia procera                 r     r     r     r
Roystonea regia                   r     r     r     r
Trophis racemosa                  r     r     r     r
Sideroxylon salicifolium          r     r     r     r
Carpodiptera cubensis             +     r     r
Exothea paniculata                1     1           +
Geoffroea inermis                 r           1     +
[E.sub.1]-Pharus glaber           2     2     2     r
Li-Trichostigma octandrum         r           r     r
[E.sub.3]-Calycophyllum
  candidissimun
Collubrina reclinata
[E.sub.2]-Capparis flexuosa                         r
[E.sub.1]-Urochloa humidicola
Chlorophora tinctoria
Eugenia sp.
Commelina elegans

Accompaniers

[E.sub.3]-Prunus myrtifolia                   r     r
Ficus membranacea                                   r
[E.sub.2]-Casearia sylvestris     r     r           r
  var. myricoides
Senna atomaria
[E.sub.1]-Rivina humilis          r
Zapoteca gracilis                 +     +
Hyperbaena paucinervis                  r
Malpighia suberosa                      r
Croton lucidus
Desmodium incanum
Varronia leptoclada
Petiveria alliacea
Li-Triopteris rigida              r
Passiflora suberosa               r
  var. pallida
Cissus verticillata               r
Chiococca alba
Pithecoctenium echinatum          r
Ep-Tillandsia usneoides           r

Other species: Zanthoxylum martinicensis r in 1; Polypodium
polipodiodes and Castela spinosa r and Belairia spinosa
+ in 2; Selenicereus grandifloras r in 3; Commelina diffusa
+ and Ceiba pentandra r in 4; Oplismenus setarius 1 in 5;
Hyperbaena sp. r in 6; Chrysophyllum oliviforme r in 7;
Palicourea sp. and Tabebuia angustata r in 8.

Localities: All releves were collected on the southern
slope of Sierra Maestra mountain range near the mouth of
Turquino River (x: 508.527, y:144.674) in altitude between
120 and 200 m asl; holotypus ass. rel. 4.

Table 6

Zanthoxylo fagarae-Guazumetum ulmifoliae ass. nova
(Trichilio-Guazumion ulmifoliae, Oxandro-
Burseretalia, Guazumo-Ceibetea pentandrae)

Altitude (m a.s.l.)               225   220   240
Exposition                        SE    NE     N
Inclination ([grados])            20    30    15
[E.sub.3]+ [E.sub.2] (%)          100   100
[E.sub.3]-Canopy layer (%)                    100
[E.sub.2]-Shrub layer (%)                     20
[E.sub.1]-Herbaceous layer (%)    30    50    60
N. species                        38    53    48
Releve N.                          1     2     3

Characteristics

[E.sub.3,2]-Guazuma ulmifolia      2     3     4
Zanthoxylum fagara                 2     1     +
Senna atomaria                     3     3     3
Acacia macracantha                 1     r     +
Samanea saman                      3     +     1
Casearia hirsuta                   +     r     2
Chrysophyllum oliviforme           r     1     r
Cupania americana                  r     2     +
Cupania glabra                     r     +     +
Chromolaena odorata                r     r     r
Phyllostylon brasiliensis          1     1
Leucaena leucocephala              r     r
[E.sub.2]-Eugenia floribunda             +     +
Picramnia pentandra                      1     1
Lonchocarpus blainii                     +     r
Casearia sylvestris                      3     1
  var. sylvestris
Psidium guajava                          r     +
[E.sub.1]-Erythroxylum             +     2     3
  havanense
Rivina humilis                     +     r     +
Commelina elegans                  r     r     +
Petiveria alliacea                 +     r     r
Abutilon sp.                       r     r     r
Pavonia typhalea                   r     r     +
Trichilia hirta                    r           1
Spermacoce laevis                  +           3
Desmanthus virgatus                r           +
Phyla nodiflora                    r           +
Turnera ulmifolia                  +           r
Rauvolfia tetraphylla              r           +
Desmodium incanum                        r     +
Adiantum trapeciforme                    2     +
Varronia globosa ssp. humilis      r     r
Oeceoclades maculata               r     r
Li-Gouania lupuloides              1     2     +
Pisonia aculeata                   1     1     2
Serjania diversifolia              +     2     3
Turbina corymbosa                  r     +     1
Trichostigma octandrum             +           2
Smilax havanense                   r           r
Lasiasis divaricata                      r     +
Cissampelos pareira                      1     +
Chiococca alba                           +     r
Tournefortia hyrsutissima                1     +
Centrosema sp.                           r     r
Stigmaphyllum lineare                    r     r
Stigmaphyllum sagreanum            r     +

Other species: Sterculia apetala 3, Harrisia eriophora
+, Commelina sp. Malpighia sp. and Passiflora sp. r
in 1; Guarea guidonia 2, Zanthoxylum martinicense and
Faramea occidentalis 1, Cecropiapeltata, Citharexylum
sp., Spondias mombin, Ocotea coriacea, Desmodium
sp., Roystonea regia, Zyzygium jambos, Urena lobata,
Lantana camara, Genipa americana, Abrus precatorius,
Hippocratea volubilis r in 2; Cedrela odorata 1,
Poeppigia procera, Spigelia anthelmia, Gastrococos crispa,
Melochia nodiflora, Sideroxylon foetidissimum subsp.
foetidissimum and Tragia volubilis r in 3.
Localities: All releves were collected in Santiago de
Cuba: El Cobre (x: 592.474, y: 156.371); holotypus ass.
rel. 1.

Table 7

Oxandro lanceolatae-Spondiadetum mombin ass. nova
(Picramnio-Ocoteion coriaceae, Oxandro-Burseretalia,
Guazumo-Ceibetea pentandrae)

Altitude (m a.s.l.)          200   190
Inclination ([grados])        8    35
Exposition                    E     N
Canopy layer-[E.sub.3] (%)   100   100
Shrub layer-[E.sub.2] (%)    30    20
Herbaceous layer-[E.sub.1]   10    10
  (%)
N. species                   62    48
Releve N.                     1     2

Characteristics

E3- Spondias mombin           3     1
Oxandra lanceolata            3     2
Cecropia peltata              2     1
Ocotea coriacea               2     +
Guazuma ulmifolia             2     1
Guarea guidonia               2     1
Cupania americana             +     +
Cupania glabra                r     1
Roystonia regia               r     r
Sideroxylon foetidissimum     r     r
  subsp. foetidissimum
Ficus membranacea             1     +
Trichilia hirta               +     r
Trophis racemosa              1     r
Zanthoxylum martinicense      2     +
Pseudolmedia spuria           +     3
Dendropanax arboreus          +     3
Zuelania guidonia             +     r
Bursera simaruba              +     r
Geoffroea inermis             r     r
Casearia hirsuta              r     r
[E.sub.2]-Calophyllum         1     r
  antillanum
Cordia gerascanthus           +     r
Picramnia pentandra           +     r
[E.sub.1]-Erythroxylum        +     r
  havanense
Chrysophyllum oliviforme      r     r
Psychotria sp.                +     r
Oplismenus setarius           r     r
Lithacne pauciflora           +     +
Pharus glaber                 +     r
Oeceoclades maculata          r     r
Adiantum trapeziforme         r     +
Li-Abrus precatorius          r     r
Serjania diversifolia         r     +
Pisonia aculeata              r     r
Cissus verticillata           r     r
Trichostigma octandrum        r     r
Gouania lupuloides            r     r
Hyperbaena sp.                r     +
Ep- Tillandsia fasciculata    r     r
Tillandsia usneoides          r     2

Other species: Cojoba arborea, Calycophyllum
candidissimum, Thouinia sp., Piscidiapiscipula,
Ficus suffucans, Eugenia sp, Casearia sylvestris
var. sylvestris, Erythroxylum areolatum,
Poeppigia procera, Eugenia floribunda,
Pavonia spinifex, Petiveria alliacea, Capparis flexuosa,
Philodendron lacerum, Pithecoctenium echinatum,
Guzmania monostachya, Trichocentrum undulatum, Olyra
latifolia and Tillandsia recurvata r, Clusia rosea, Licaria
jamaicensis and Anthurium cubense + in 1; Drypetes alba
3, Buchenavia capitata (tetraphylla) 2, Prunus
occidentalis, P. myrtifolia, Urera baccifera and
Vitis tiliaefolia r, Sapium jamaicense and Eugenia
axillaris + in 2. Localities: All releves were
collected in the middle of Turquino River (x: 506.532,
y: 148.853); holotypus ass. rel. 1.
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Author:Reyes, Orlando J.; Acosta Cantillo, Felix
Publication:Lazaroa
Date:Jan 1, 2014
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