Original article

Phenology of flora components in the use of natural and traditional medicine in the community of Verraco, Santiago de Cuba, Cuba

0000-0002-1792-6323Miguel Ángel Castell Puchades^{1  *}  , 0000-0003-3107-1242Yenisei Revilla Gongora¹  , 0000-0001-7120-0024Gustavo Polanco Durán¹  , 0000-0002-6623-8404Yamila Baró Bou¹ 

¹Centro Oriental de Ecosistemas y Biodiversidad, Santiago de Cuba. Cuba.

Abstract

Based on ethnobotanical studies carried out in the community of Verraco, belonging to the Baconao Biosphere Reserve, province of Santiago de Cuba, the components of the flora used in natural and traditional medicine were identified in situ, and the reproductive phenology of each of the identified species was determined. The objective of the work was to present the flowering and fruiting patterns of flora species that are components of natural and traditional medicine in the coastal community of Verraco, which belongs to the Baconao Biosphere Reserve. A total of 82 species of flowering plants used by the community for natural and traditional medicine were recorded, which are grouped into 75 genera and 49 botanical families. Of the total species studied, 46 % (38) were recorded with flowers and fruits, while only 12 % (10) and 39 % (32) were determined in the flowering and fruiting stages, respectively. Of the total species, only four are endemic and 24 % are naturalized, while 19 % correspond to non-indigenous but possibly naturalized species. Forty species flowers, coinciding with the rainy season, while 36 bear fruit during the same period. In the dry or low rainfall stage, a total of 57 species were recorded to flower in this period. A total of 27 species present a flowering stage of more than four months, while 21 species bear fruit in a similar stage of time. Only one species was recorded that flowers and bears fruit all year round: Heliotropium angiospermum. A total of 21 species present synchrony, as both phenological periods coincide in the same reproductive stage.

Key words: Ethnobotanical perception studies; Reproductive phenology; Medicinal plants.

Introduction

The phenology of plant species is one of the main studies to be carried out to understand the flora of an ecosystem ^{(Ortiz, 1990)}. Phenological studies in general, and especially the flowering and fruiting phases, are essential to determine the optimal time to collect seeds and thus guarantee the management of medicinal, forest, food, melliferous and other species, contributing to possible conservation programs ^{(Albert-Puentes et al., 1995)}.

Several ethnobotanical studies have been carried out in various coastal communities in the country, with emphasis on the use of traditional medicine in Cuba, including those of Rosette et al. (2019) and ^{Polanco et al., (2011)}, for the Guanahacabibes and Baconao Biosphere Reserves, respectively; among others ^{Vilamajó (1984)} , ^{Vilamajó (1985)}, ^{Vilamajó and Menéndez (1988)}, ^{Figueredo et al., (2010)}, ^{Figueredo et al., (2013)}. The aim of the work was to publicize the flowering and fruiting patterns of flora species that are components of natural and traditional medicine in the coastal community of Verraco, part of the Baconao Biosphere Reserve.

The population's knowledge and monitoring of the phenology of flora components in the use of natural and traditional medicine enables them to make appropriate use of plants for the treatment and cure of different conditions, and allows them to draw up strategies for the appropriate sowing and cultivation of these species.

Materials and methods

Based on the ethnobotanical studies carried out in the community of Verraco by ^{Polanco et al., (2011)}, and on the floristic list of ^{Figueredo et al., (2013)}, the components of the flora that are used in natural and traditional medicine were selected and their phenology was determined in situ, especially the flowering and fruiting phases of each of the identified species, according to the methodology of ^{Albert-Puentes et al., (1993ª)}.

Monthly monitoring was carried out between 2015 and 2016 in the community of Verraco, located in the geographical coordinates X: 639 080, Y: 139 020, belonging to the Baconao Biosphere Reserve, province of Santiago de Cuba, and the phenological field observations were complemented with reviews of materials belonging to the Spermatophyte Section of the Herbarium BSC "Dr. Jorge Sierra Calzado" of the Centro Oriental de Ecosistemas y Biodiversidad of Santiago de Cuba, noting the phenological data.

As for the duration of flowering and fruiting, the species were classified according to ^{Castillo and Carabias (1982)} as:

• Short = species whose flowering or fruiting period is < 4 months.

• Long = species whose flowering or fruiting period is ≤ 4 months.

According to the flowering period, the criteria of ^{Sarmiento and Monasterio (1983)} were followed, where:

• Species with continuous flowering, are those that bloom all year round.

• Early flowering species, those that bloom at the beginning of the rainy season (spring).

• Delayed-flowering species are those that begin in the second half of the rainy season, and extend through the end of that season.

• Late-blooming species are those that flower in the dry season.

• Opportunistic species: those that are able to flower in any period, as long as the environmental conditions are favorable to them.

For the analysis of the distribution of plant species and the updating of taxonomic nomenclature, the criteria of ^{Greuter and Rankin were followed (2017)}.

Results and discussion

According to the ethnobotanical perception studies carried out in the community of Verraco, a total of 82 species of flowering plants (angiosperms) were recorded, which are used by the community for the use of natural and traditional medicine, which are grouped into 75 genera and 49 botanical families (Table 1).

Table 1 - Phenology of the species with medicinal use by the Verraco community  

Family	Scientific name	Common name	Distribution	Phenology
				Flowering	Fructification
Acanthaceae	Dicliptera sexangularis (L.) Juss.	árnica	Cu - Esp, Ja, PRc	Jan-Sep	Jan-Sep
	Justicia pectoralis Jacq.	carpintero	PCu - PEsp, PJa	Ap
Aizoaceae	Tetragonia tetragonioides (Pallas) Kuntze	espinaca	NCu	May
Amaranthaceae	Chenopodium ambrosioides L.	apasote	PCu - PEsp PJa	Ap-June
Anacardiaceae	Mangifera indica L.	mango	NCu - PEsp PJa	Jan-Ap	Ap-Aug
Annonaceae	Annona muricata L.	guanábana	PCu - PEsp PJa		Nov
	Annona squamosa L.	anón	CCu - CEsp CJa	Jun	Aug-Sep
Apiaceae	Petroselinum crispum (Mill.) Nyman ex A.W. Hill	perejil	NCu	Jun	Aug-Sep
	Pimpinella anisum L.	anís	NCu	Jun	Aug-Sept
Arecaceae	Acrocomia crispa (Kunth) C.F. Baker ex Becc	corojo	˚Cu	Jun
	Cocos nucifera L.	coco	PCu - PEsp PPRc	Nov-Mar	Nov-Mar
Asteraceae	Bidens pilosus L.	romerillo	Cu - Esp Ja PRc	Mar	Oct
	Isocarpha oppositifolia (L.) Cass.	manzanilla	Cu - Ja AmN AmC AmS	Feb
	Pluchea carolinensis (Jacq.) G. Don	salvia	Cu - Esp Ja PRc	Jan-Jun	Oct-Nov
	Vernonanthura havanensis (DC.) H. Rob.	rompezaragüey	˚Cu	Jan, Mar/Nov
	Xanthium strumarium L.	guisazo de Baracoa	PCu - AmN NVM		Mar/May/Jul
Boraginaceae	Heliotropium angiospermum Murray	alacrancillo	Cu - Esp Ja PRc	Whole a year	Whole a year
Brassicaceae	Nasturtium officinale W. T Aiton	berro	NCu(CuW(Art May	May-Sep	Feb
Bromeliaceae	Bromelia pinguin L.	piña ratón	NCu - Esp Ja PRc	May-Oct	Nov-Feb
Burseraceae	Bursera graveolens (Kunth) Triana& Planch	sasafrás	PCu - AmN AmC	Dec-March	Dec-Mar
	Bursera simaruba (L.) Sarg.	almácigo	Cu - Esp Ja PRc	Feb-Sep	October-Mar
	Protium cubense (Rose) Urb.	copal	˚Cu		Apr
Cactaceae	Nopalea cochenillifera (L.) Salm-Dyck	tuna	NCu - PEsp PJa	Dec-Mar	Oct
Canellaceae	Canella winterana (L.) Gaertn.	cúrbana	Cu - Esp Ja PRc	Jun-Sep
Capparaceae	Capparis frondosa Jacq	raíz de berraco	Cu (CuE(Ho Gu)) -	Jun	Aug
Caricaceae	Carica papaya L.	fruta bomba, papaya	CCu - CEsp CJa	Feb-Mar
Commelinaceae	Tradescantia spathacea Sw.	barquito, cordobán	NCu - PEsp PPRc	Ap-Jun
Costaceae	Cheilocostus speciosus (J. König) C. Specht	caña mexicana	NCu - PEsp PJa	May
Crassulaceae	Kalanchoe pinnata (Lam.) Pers.	hoja de aire	NCu - PEsp PJa	Jan-Mar	Ap-Jun
Cucurbitaceae	Momordica charantia L.	cundeamor	NCu - PEsp PJa	Mar-Jun	Mar/Sep-Oct
Cyperaceae	Cyperus rotundus L.	caramaná	PCu - PEsp PJa	May-Oct	May-Oct
Erythroxylaceae	Erythroxylum havanense Jacq. var. havanense	jibá	Cu - Esp Men AmS	Jan-Ap	Mar-May
Euphorbiaceae	Euphorbia serpens Kunth	coronilla	Cu - Esp Ja PRc	Ap
	Jatropha gossypiifolia L.	túa túa	Cu - Esp Ja PRc	May-Sep	Jun-Oct
	Jatropha sp.	chaya	NCu	Nov
Fabaceae	Caesalpinia bahamensis Lam.subsp. bahamensis	palo de brasil	Cu (CuW(Mat IJ)	Apr-Jun/Sep-Dec
	Cassia grandis L.	cañándonga	Cu(CuW(Art) AmS	Feb-May	May-Aug
	Desmodium incanum DC.	amor seco	Cu - Esp Ja PRc Men Cay AmN AmC AmS	Jul	Jul
	Senna occidentalis (L.) Link	platanillo	DCu(CuW(PR* Art Hab* May Mat IJ)	Feb-Ap	Feb-Ap
	Tamarindus indica L.	tamarindo	PCu(CCuW CCuC	Dec-Mar	Mar-May
Lamiaceae	Mentha spicata L.	hierba buena	CCu(CuW(Hab*)) -	Mar
	Ocimum basilicum L.	albahaca morada	NCu(CuW(PR* Art Hab* Mat)	Ap
	Origanum majorana L.	mejorana	PCu(CCuW(Hab*)	Feb-Ap
	Plectranthus amboinicus (Lour.) Spreng.	orégano	PCu(CCuW(PR* Art Hab* May Mat IJ)	Ap
	Plectranthus tomentosus Benth.	meprobamato		Mar	Jun
	Rosmarinus officinalis L.	romero	PCu(CCuW CCuC	Mar
	Vitex agnus-castus L.	vencedor	C Cu(CuW(Hab*)	Nov
Lauraceae	Persea americana Mill.	aguacate	PCu(CuW(Art Mat) CuC(SS) CuE(Ho	Mar-May	Jun-Sept
Lythraceae	Lawsonia inermis L.	resedá	PCu - pHab* CEsp		Ap and Sep
	Bastardia viscosa (L.) Kunth var. viscosa	malva bruja	Cu(CuW(PR* Art Hab* May Mat IJ)	Nov
Malvaceae	Gossypium arboreum L.	algodón	-Cu - VM		Feb/Ap/Jun/ Nov/Dec
	Guazuma ulmifolia Lam.	guásima	Cu(CuW(PR* Art Hab* May IJ)	Oct-Dec	Oct-Feb
	Hibiscus rosa-sinensis L.	marpacífico rojo	NCu	Mar-Jun
	Talipariti elatum (Sw.) Fryxell	majagua	NCu	Ap-Jun
Meliaceae	Azadirachta indica A. Juss.	árbol del nim	PCu(CuW(Hab*)) -		Sep
	Trichilia hirta L.	jubabán	Cu(CuW(PR* Art Hab* May Mat	Jan-June	Jan/Ab/Jun/May/Dec
Moringaceae	Moringa oleifera Lam.	moringa	NCu(PCuW(Art cHab*) CCuC(Ci SS)	Mar	May
Myrtaceae	Eucalyptus sp.	eucalipto		Jan/May	Jan-Feb/Oct
	Psidium guajava L.	guayaba	DCu - Esp Ja PRc	May/Jun	Ap-Jul
Oleaceae	Jasminum fluminense Vell.	jazmín de 5 pétalos	NCu(CuW(Mat) CuC(VC LT)	Jan/Sept Aug
Papaveraceae	Argemone mexicana L.	cardo santo	DCu(CuW(PR* Hab* Mat) CuC(VC Ci)	Feb-Ap
Pasifloraceae	Turnera ulmifolia L.	marilope o tapón	PPRc Men Bah Cay	Mar/Ap	Mar/Nov
Phyllanthaceae	Phyllanthus tenuicaulis Müll. Arg. subsp. tenuicaulis	fruta o huevo escondido	˚Cu	Feb-Ap	Feb-June
Phytolaccaceae	Petiveria alliacea Lin.	anamú	Cu(CuW(PR* Hab* May IJ) CuC(VC Ci SS	Ab/Jun/Ago/ Nov/Dec
Piperaceae	Piper auritum Kunth	anizón	DCu(CuW(Hab*) CuC(Ci SS) CuE(Gr	Ap-June	Jun-Aug
Plantaginaceae	Plantago major L.	llantén	NCu(CuW(PR* Art Hab* May Mat)	Feb-Jun	Feb-Jun
Poaceae	Cymbopogon citratus (D. C.) Stapf.	caña santa, hierba de calentura	CCu - CEsp CJa CPRc CMen CBah	Oct	Dec
Polygonaceae	Coccoloba uvifera (L.) L.	uva caleta	Cu - Esp Ja PRc	Feb-Ap
Punicaceae	Punica granatum L.	granada	C Cu - C Esp C Ja C PRc C Men		Oct-Dec
Rhamnaceae	Colubrina elliptica (Sw.) Brizicki & Stern	carbonero	Cu - Esp Ja PRc Men Bah Cay AmN	Ap-Jun
Rutaceae	Citrus x aurantium L.	naranja agria	NCu(CuW(Hab* Mat) CuC(Ci Cam)		Jan-May
	Citrus x limon (L.) Osbeck	limón	CCu - CEsp CJa		Jan/Jun/Oct/ Nov
	Citrus reticulata Blanco	mandarina	CCu - CEsp CJa CPRc CAmN		Dec-Jan
	Ruta chalepensis L.	ruda	(C)Cu - CEsp CPRc CMen VM	Mar	Jun
Sambucaceae	Sambucus canadensis L.	sauco blanco	NCu	Feb-May
Solanaceae	Solanum americanum Mill.	yerba mora	Cu(CuW(PR* Art Hab* May Mat IJ)	Mar-May/Oct	Mar-May/Jul/Oct-Nov
	Solanum torvum Sw.	prendejera	Cu (CuW(PR* Art Hab* May Mat IJ) CuC(VC Ci SS	May/Nov	Jan/May Oct/Nov
Urticaceae	Pilea depressa (Sw.) Blume	lloviznita	CPRc CMen CAmC CAmS Cu - Esp Ja	Ap
Verbenaceae	Lippia alba (Miller) N. E. Br.	menta americana	Cu (CuW(PR* Art Hab* May Mat)	Ap/Jun-Jul
	Stachytarpheta jamaicensis (L.) Vahl.	verbena	Cu (CuW(PR* Art Hab* May Mat IJ)	Feb-May	Jan/Jul/Oct-Nov
Vitaceae	Cissus verticillata (L.) Nicolson & Jarvis subsp. verticillata	bejuco ubí	Cu - Esp Ja	Ap
Xanthorrhoeaceae	Aloe vera (L.) N. L. Burm.	sábila	PCu - PEsp PJa	Feb-May	Ap-Jun

Legend: AmC: Central América, AmN: North América (including México), AmS: South América (including Trinidad, Tobago, Curazao, Margarita, etc.), Art: Artemisa province, Bah: Bahamas, CA: Ciego de Ávila province, Cam: Camagüey province, Cay: Islas Caymán, Ci: Cienfuegos province, CuC: central Cuba, CuE: Easter Cuba, CuW: Western Cuba, Esp: Hispaniola, Gr: Granma province, Gu: Guantánamo province, Hab*: La Habana province, Ho: Holguín province, IJ: Pine Island, Ja: Jamaica, Mat: Matanzas province, May: Mayabeque province, Men: minor Antillas (including Virgens Island), PR*: Pinar del Río province, PRc: Puerto Rico, SS: Sancti Spíritus province, VC: Villa Clara province, VM: Old World (including Australia e Pacific Island), Endemic in Cuba, pointing by error, D: Present but doubtfully indigenous, N: naturalized, P: no indigenous but likely naturalized, C: widely cultivated, (C): Occasionally Cultivated or currently not cultivated, Jan: January, Feb: February, Mar: March, Ap: April, May: May, Jun: June, Aug: August, Sep: September, Oct: October, Nov: November, Dec: December.

These results are congruent with ^{Polanco et al., (2011)}, ^{Rosete et al., (2019)}, and Nina (2017), who in different ethnobotanical studies on the use of biodiversity for medicinal purposes report that community members use mainly curative and aromatic plant species to address certain health conditions, using not only those available in the natural environment, but also those grown in home gardens and farms, largely using arvenous species.

From the total number of species studied, 46 % (38) were recorded with flowers and fruits, while only 12 % (10) and 39 % (32) were determined in the flowering and fruiting stages, respectively (Table 1). According to ^{Polanco et al., (2011)}, the fruits and flowers of medicinal plants represent, after the leaves, the organs most used by the inhabitants of this community to combat the various conditions and diseases.

The best represented botanical families are: Lamiaceae with seven species, followed by Asteraceae and Fabaceae with five, while the genera with the largest number of infra-generic taxa were Citrus (Rutaceae) with three species, followed by the genera Annona (Annonaceae), Bursera (Burseraceae), Jatropha (Euphorbiaceae), Plectranthus (Lamiaceae), and Solanum (Solanaceae), with two in each case. In ethnobotanical studies conducted in the community of Gran Piedra, ^{Hernández (2001)} reported the Fabaceae and Asteraceae families as the most represented for their medicinal uses.

From the total of species, only four are endemic: Acrocomia crispa C.F. Baker ex Becc, Vernonanthura havanensis (DC.) H. Rob., Protium cubense Urb. and Phyllanthus tenuicaulis Müll. Arg. subsp. tenuicaulis. 24 % of the identified species are naturalized (many of them are fruit trees), while 19 % correspond to non-indigenous but possibly naturalized species, according to ^{Greuter and Rankin (2017)}. A total of eight species are cultivated.

There is a report on the floristic diversity of Verraco ^{(Figueredo et al., 2013)}, where 57 % of the total plant species grow in the surroundings of the community and only 44 % are cultivated by the inhabitants.

^{Pelicié and Hernández (1985)} obtained 91 % of endemism at the ethnobotanical level in that coastal sector (Verraco-Cazonal); later, in 1988, they updated the list of medicinal plants for that locality, identifying 59 species with 27 % of endemism.

According to ^{Albert-Puentes et al., (1993)}, 40 species flowers during the rainy season from May to October, including Bromelia pinguin L. and Cyperus rotundus L., while 36 species bear fruit during the same period, such as Solanun torvum Sw. and Solanum americanum Mill., both of the family Solanaceae.

In the dry or low rainfall stage, a total of 57 species were recorded that bloom during this period, although many of them overlap this phenophase during the months of April and May, which mark the end of the dry stage and the beginning of the rainy stage ^{(Bermúdez and Durán, 1991)}. The possible dependence of the phenological patterns on environmental factors, as well as their relationship with other plants and animals was pointed out by ^{Ramírez and Brito (1987)}.

Figure 1 shows the categories of species according to the period in which they bloom, highlighting in this aspect the Late Blooming (FTA) with 31 species, followed by the opportunistic species (CP) with 24, those that bloom at any stage of the year.

For ^{Opler et al., (1976)}, the phenological phases are closely related to rainfall, given that the seasonality in the tropics is fundamentally given by this climatic factor. ^{Bawa et al., (1990)} reported spatial and temporal variations in phenological patterns revealed in individual and community studies over two decades; however, the factors determining these patterns remain unknown ^{(Bawa and NG., (1990)}.

Fig. 1. - Species categories according to flowering period, FC: continuous flowering, FTE: early flowering, FR: delayed flowering, FTA: late flowering, CP: opportunistic species  

A plant may not develop all its phenological stages if it grows in different climatic conditions than its region of origin. ^{Abu-Asab et al., (2001)} report extensions of the flowering period in 89 species in the Washington DC area, analyzing 30-year records; correlating these observations directly with local increases in minimum temperatures.

According to ^{Castillo and Carabias (1982)}, and as shown in Figure 2, a total of 45 species (55 % of the total recorded) were identified that bloom in a period of less than four months, as is the case of Argemone mexicana L.(Figure 3); that is, they could be defined as short-lived species. In the same category there are 26 species (32 % of the total recorded), whose fruiting stage does not exceed the same number of months.

Fig. 2 - Species categories according to flowering and fruiting time. FLC: short period flowering, FLL: long period flowering, FRC: short period fruiting, FRL: long period fruiting  

A total of 27 species have a long flowering period and 21 species bear fruit over a period of more than 4 months, as they are: Nasturtium officinale W. T Aiton, Cassia grandis L., Mangifera indica L., among others; many of them are edible and cultivated. Only one species has been recorded that flowers and bears fruit all year round: Heliotropium angiospermum Murria.

Some species are recorded that overlap both phenological periods: Dicliptera sexangularis (L.) Juss, from January to September and C. rotundus, from May to October; both are herbaceous species. Other species, such as Cocos nucifera Lin., and Trichilia hirta L.(Figure 4), present their flowering and fruiting phases in the same period, although the latter presents several fruiting peaks (Table 1).

Fig. 3. - Argemone mexicana L.  

Fig. 4. - Trichillia hirta L.  

Other authors, such as ^{Albert-Puentes et al. (1993)}, carried out phenological studies where they obtained similar results of overlapping with T. hirta(Figure 4) and other tree species such as: Prunus occidentalis Sw. and Matayba apetala f. oppositifolia (A. Rich.) Radlk. Similar fruiting patterns were found by ^{Sánchez et al. (2009)} for species analogous to those reported in these localities.

A total of 21 species are synchronized, that is, they flower and bear fruit at the same time, at the same time, as they are: Bursera graveolens Triana & Planch , and Plantago major L.; ^{Augspurger (1983)} defined synchronism as an escape of plants from flower predators and then from seeds, so it is an adaptive strategy for successful pollination and seed dispersal ^{(Steven et al., 1987)}.

For ^{Castillo and Carabias (1982)}, one aspect to take into account is the variability of phenological patterns (not only between species and years, but between individuals of the same species, so that not all individuals belonging to the same species flower and bear fruit simultaneously, and sometimes not even in the same year).

Bursera simaruba (L.) Sarg. (seedling), extends its flowering for nine months, while fructification is recorded for six months. These results show a very different behavior to the ones obtained by ^{Hechavarría et al. (2000)}, in Itabo, Matanzas, where fruits are harvested only in July.

It is known that many species, such as Carica papaya L., are able to revert their reproductive habits in extreme conditions, which probably is a mechanism for the perpetuation of the species; for that reason it is important to complement the phenological observations in the field with the herbarium records, because according to Croat (1969) individuals can bloom in a year out of season, producing a lengthening of the period of the observed phenophase.

Conclusions

In the community of Verraco, Baconao Biosphere Reserve, 87 % and 61 % of the components of the flora for medicinal and traditional use are recorded with flowers and fruits, which enables the inhabitants, based on their knowledge of these plant phenophases, to make appropriate use of them for the treatment and cure of various conditions, and to be able to trace strategies for the planting and cultivation of these medicinal species

Acknowledgements

Thanks are due to the national project "Components of biological diversity used by Cuban families in natural and traditional medicine", which is part of the Natural and Traditional Medicine Program of the Ministry of Public Health (MINSAP), for its logistical support for the development of this research.

Referencias bibliográficas

ABU ASAB, M.S., PETERSON, P.M., SHETLER, S.G. y ORLI, S.S., 2001. Earlier plant flowering in spring as a response to global warming in the Washington, DC, area. Biodiversity and Conservation [en línea], vol. 10, pp. 597-612. [Consulta:22/05/2020]. ISSN 0960-3115. Disponible en: Disponible en: http://repository.si.edu/xmlui/handle/10088/3371 .

ALBERT PUENTES, D., LÓPEZ, A., RODRÍGUEZ, M. y DUARTE, M., 1995. Recursos fitogenéticos forestales I. Familia Meliaceae. Fontqueria, vol. 42, pp. 329-351.

ALBERT PUENTES, D., LÓPEZ, A. y ROUDNÁ, M., 1993. Observaciones fenológicas en árboles tropicales. Consideraciones metodológicas. Fontqueria, vol. 36, pp. 257-263.

AUGSPURGER, C.K., 1983. Phenology, Flowering Synchrony, and Fruit Set of Six Neotropical Shrubs. Biotropica [en línea], vol. 15, no. 4, pp. 257-267. [Consulta: 22/05/2020]. ISSN 0006-3606. DOI 10.2307/2387650. Disponible en: Disponible en: https://www.jstor.org/stable/2387650 . JSTOR

BAWA, K.S. y HADLEY, M., 1990. Reproductive Ecology of Tropical Forest Plants [en línea]. S.l.: Unesco. ISBN 978-0-929858-22-7. Disponible en: https://books.google.com.cu/books/about/Reproductive_Ecology_of_Tropical_Forest.html?id=AuHwAAAAMAAJ&redir_esc=y.

BERMÚDEZ, G. y DURÁN, T., 1991. Mapas de precipitación. Atlas de Santiago de Cuba. Cuba: Academia de Ciencias de Cuba.

CASTILLO, S. y CARABIAS, J., 1982. Ecología de la vegetación de las dunas costeras: fenología. Biótica [en línea], vol. 7, no. 4, pp. 551-568. [Consulta: 22/05/2020]. Disponible en: Disponible en: https://biblat.unam.mx/es/revista/biotica-mexico-d-f/6 .

DE STEVEN, D., WINDSOR, D.M., PUTZ, F.E. y DE LEON, B., 1987. Vegetative and reproductive phenologies of a palm assemblage in Panama. Biotropica [en línea], vol. 19, no. 4, pp. 342-356. [Consulta: 28/05/2020]. ISSN 0006-3606. DOI 10.2307/2388632. Disponible en: Disponible en: https://www.jstor.org/stable/2388632 . JSTOR

FIGUEREDO CARDONA, M., POLANCO DURÁN, G. y REVILLA GÓNGORA, Y., 2010. Estudio para la conservación de las plantas útiles de las Terrazas Costeras de la Reserva de la Biosfera Baconao, Santiago de Cuba, Cuba. Sistemas Biocognitivos Tradicionales: Paradigmas en la Conservación Biológica y el Fortalecimiento Cultural [en línea]. México: Asociación Etnobiológica Méxicana, A. C., pp. 36-40. ISBN 978-607-482-095-9. Disponible en: https://www.researchgate.net/publication/280309669_Sistemas_Biocognitivos_Tradicionales_Paradigmas_en_la_Conservacion_Biologica_y_el_Fortalecimiento_Cultural.

FIGUEREDO, l.M., ACOSTA, F., CASTELL M.A., y POLANCO G. 2013. Objetos de conservación de la flora y la vegetación de los cerros calizos costeros de la Reserva de la Biosfera Baconao, Santiago de Cuba. Foresta Veracruzana vol 15 no.1 pp. 9-24. Disponible en: https://www.redalyc.org/jatsRepo/1813/181359681003/html/index.html

GREUTER, W. y RANKIN RODRÍGUEZ, R., 2017. Vascular plants of Cuba. A preliminary checklist. Second, updated Edition of The Spermatophyta of Cuba, with Pteridophyta added [en línea]. Berlin: Botanischer Garten und Botanisches Museum Berlin. ISBN 978-3-946292-18-0. Disponible en: https://www.researchgate.net/publication/321886706_Vascular_plants_of_Cuba_A_preliminary_checklist_Second_updated_Edition_of_The_Spermatophyta_of_Cuba_with_Pteridophyta_added /link/5a381f76458515919e71ed8a/download.

HECHAVARRÍA, O., RODRÍGUEZ, E., MORALES, N., VERA, N., ESPÍN, G., CORRALES, B., FUENTES, V., y PÉREZ, A. 2000. Calendario fenológico de 51 especies forestales de Cuba. Mejoramiento Genético y Semillas Forestales. Revista Forestal Centroamericana 30, Turrialba, Costa Rica.p.5-8.

HERNÁNDEZ, C.J. 2001. Uso de las plantas. En: Diversidad Biológica de los Macizos Montañosos Orientales. Tomo II. Programa Científico-Técnico Nacional. Desarrollo Sostenible de la Montaña. 549 p.

OPLER, P.A., FRANKIE, G.W. y BAKER, H.G., 1976. Rainfall as a Factor in the Release, Timing, and Synchronization of Anthesis by Tropical Trees and Shrubs. Journal of Biogeography [en línea], vol. 3, no. 3, pp. 231-236. [Consulta: 26/05/2020]. ISSN 0305-0270. DOI 10.2307/3038013. Disponible en: Disponible en: https://www.jstor.org/stable/3038013 . JSTOR

ORTIZ, R., 1990. Fenología de árboles en un bosque semideciduo. Acta Botánica Venezuélica [en línea], vol. 16, no. 1, pp. 93-116. [Consulta: 26/05/2020]. ISSN 0084-5906. Disponible en: Disponible en: https://www.jstor.org/stable/41740491 . JSTOR

PELICIÉ, O. y HERNÁNDEZ, J., 1985. Estudio etnobotánico del matorral xeromorfo costero comprendido entre las playas Verraco y Cazonal de Santiago de Cuba. En: Meeting Name: Simposio de Botánica, Memorias: Primer Simposio de Botánica, Palacio de las Convenciones, Editora de la Academia de Ciencias de Cuba, Ciudad de La Habana, Cuba [en línea] Disponible en: https://koha.inpa.gov.br/cgi-bin/koha/opac-detail.pl?biblionumber=13571.580.63 S612

POLANCO, D. G., SIMÓN, R. F. R., MORELl B. A., y ECHEZARRETA C. C. M. 2011. Uso, consumo y demanda de las plantas medicinales en la ciudad de Santiago de Cuba. Ediciones UO. Santiago de Cuba, Cuba. ISBN: 978-959-207-416-3.

RAMÍREZ, N. y BRITO, Y., 1987. Patrones de floración y fructificación en una comunidad pantanosa tipo morichal (Calabozo-Guàrico, Venezuela). Acta Científica Venezolana [en línea], vol. 38, pp. 276-281. Disponible en: https://www.researchgate.net/publication/281627160_Ramirez_N_Y_Brito_1987_Patrones_de_floracion_y_fructificacion_en_una_comunidad_pantanosa_tipo_morichal_Calabozo -Guarico_Venezuela_Acta_Cientifica_Venezolana_38_276-281.

ROSETE, S., RICARDO, N. y NÁPOLES, R., 2019. Productos forestales no maderables en la Reserva de la Biosfera Península de Guanahacabibes. Biodiversidad, usos tradicionales y conservación de los productos forestales no maderables en Cuba [en línea]. S.l.: s.n., pp. 34-67. ISBN 978-84-9717-376-6. Disponible en: https://www.researchgate.net/publication/335971002_Biodiversidad_usos_tradicionales_y_conservacion_de_los_productos_forestales_no_maderables_en_Cuba.

SÁNCHEZ, J.A., MUÑOZ, B. y MONTEJO, L., 2009. Rasgos de semillas de árboles en un bosque siempreverde tropical de la Sierra del Rosario, Cuba. Pastos y Forrajes [en línea], vol. 32, no. 2, pp. 1-1. [Consulta: 28/05/2020]. ISSN 0864-0394. Disponible en: Disponible en: http://scielo.sld.cu/scielo.php?script=sci_abstract&pid=S0864-03942009000200007&lng=es&nrm=iso&tlng=es .

SARMIENTO, G. y MONASTERIO, M., 1983. Life Form and Phenology. Tropical Savannas. Amsterdam: Elsevier, pp. 79-108.

VILAMAJÓ, D., 1984. Comportamiento fenológico de especies del estrato arbóreo de un bosque siempreverde. Ciencias, vol. 11, pp. 79-92.

VILAMAJÓ, D., 1985. Comportamiento fenológico de especies de vegetación secundaria. Ciencias, vol. 13, pp. 51-56.

VILAMAJÓ, D. y MENÉNDEZ, L., 1988. Fenología de algunas especies importantes en Sierra del Rosario. En: Google-Books-ID: np1JGwAACAAJ, Ecología de los bosques siempreverdes de la Sierra del Rosario, Cuba: Proyecto MAB no. 1, 1974-1987 [en línea]. La Habana: Oficina Regional de Ciencia y Tecnología de la Unesco para América Latina y el Caribe, pp. 243-260. Disponible en: https://books.google.com.cu/books/about/Ecolog%C3%ADa_de_los_bosques_siempreverdes_d.html?id=np1JGwAACAAJ&redir_esc=y.

Received: May 18, 2020; Accepted: July 03, 2020

^*Autor para la correspondencia: [e-mail]

Los autores declaran no tener conflictos de intereses.

Los autores han participado en la redacción del trabajo y análisis de los documentos.