Ecology, Phenology and Conservation Issues of Six Threatened or Remarkable Rubiaceae in the Lesser Antilles ()
1. Introduction
The Lesser Antilles archipelago located in the Caribbean is composed of a chain of main islands and a myriad of islets, extending from the Virgin Islands in the north to Trinidad and Tobago in the south. These islands are the result of the subduction process between the North American and Caribbean plates, which led to the formation of an external arc composed of ancient limestone islands with low relief, an internal arc still active, of volcanic origin with more marked relief, separated by an intermediate arc resulting from underwater volcanic activities since the Miocene [1]-[3]. All of these islands, although of modest areas, benefit from a tropical climate and are home to exceptional biodiversity, contributing to the richness of the Caribbean global biodiversity hotspot [4]-[10].
The mountainous reliefs of the volcanic islands of the archipelago condition a staggered pattern of precipitation, and climate variations with altitude induce an altitudinal bioclimatic gradient, going from the dry coastline to the hyper-humid summits [7] [11]-[13]. This gradient, combined with a great diversity of soils, favors the formation of multiple habitats and the development of specialized flora, giving rise to a great variety of plant levels [7] [9] [11]-[15]. However, since colonization, the anthropization of these islands has continued to intensify, endangering the richness of their ecosystems [16]-[20]. Urbanization, agriculture, deforestation, pollution, invasive species and current climate change generate major ecological risks, threatening the stability, diversity and functioning of natural environments [7] [17] [19] [20]-[22].
The sustainable management of natural resources in the Lesser Antilles archipelago has long been hampered by a lack of institutional coordination and persistent socio-economic inequalities, limiting the effectiveness of regional and local conservation strategies [19] [23] [24]. Despite their ecological richness, these island ecosystems remain particularly vulnerable, due to their isolation, small size, and increasing anthropogenic pressure [7] [9] [16] [25]. Martinique, a volcanic island located in the heart of the archipelago, is home to rich but vulnerable ecosystems, subject to strong anthropogenic pressures and various ecological risks that can lead to the erosion of this biodiversity [7] [16] [17] [20] [23] [25]-[27]. In this context, the Ethnobotanical Park of the town of Marin located on the southern Caribbean coast of Martinique, at the place called “Rivière habitation”, plays a strategic role in ex situ conservation, that is to say the preservation of threatened or remarkable species outside their natural environment.
There are six Rubiaceae preserved in this park: Erithalis fruticosa, Genipa americana, Guettarda scabra, Ixora ferrea, Randia aculeata and Randia nitida. As part of a doctoral project carried out on the Rubiaceae family, between 2015 and 2020 at the University of the Antilles, a study was specifically devoted to these six Rubiaceae. By combining phenological observations, floristic inventories and bibliographic analysis, the objective was to better understand their ecological requirements and their development rates. This approach aims to strengthen their in situ conservation, by promoting their maintenance in their natural habitat, but also to consider ecological restoration actions, including their reintroduction into their natural habitats for species threatened with extinction.
2. Materials and Methods
2.1. Bibliographic Data
The Rubiaceae constitute one of the largest families of flowering plants in the world and are among the five most represented families in the Lesser Antilles archipelago [4] [6] [26]-[28]. Their morphological and physiological diversity allows them to occupy a wide range of habitats, from dry, rocky or sandy coastlines to humid forests located at high altitudes, demonstrating a remarkable capacity for adaptation [4] [6] [8] [26]-[27] [29]. The volcanic islands of the archipelago such as Martinique present a heterogeneity of habitats and forest ecosystems ranging from the coast to the summits which can accommodate this diversity of species of the Rubiaceae family (Figure 1 and Figure 2).
Figure 1. Location of the lesser Antilles archipelago and Martinique (Caribbean).
In the ecosystems of the Lesser Antilles, although the majority of these species are introduced or naturalized, they perform important ecological functions. Some actively participate in the dynamics of natural regeneration of degraded plant formations, thus contributing to the resilience of tropical island forests [7] [9] [26] [27]. This functionality makes them valuable allies to be taken into account in ecological restoration programs. According to the IUCN (International Union for Conservation of Nature), 11 categories are used to assess the extinction risk of a species and define its conservation status [30]. They apply at different scales (global, regional, etc.), meaning that a species can have a global status and another regional status. These categories are: Globally Extinct (EX), Extinct in the Wild (EW), Regionally Extinct (RE), Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT), Least Concern (LC), Data Deficient (DD), Not Applicable (NA) and Not Evaluated (NE) [30]. Among the species of the Rubiaceae family present in the French Antilles, some Rubiaceae thus have a conservation status of concern. This is the case of Randia nitida (or Randia armata), classified as critically endangered for decades and preserved in the ethnobotanical park of the town of Marin in the south of Martinique (Figure 3) [16] [20] [30].
The six species studied here illustrate the ecological and functional diversity of this family within the natural ecosystems of the Lesser Antilles archipelago (Table 1) [6] [20] [26] [27] [30]. Indeed, Erithalis fruticosa and Randia aculeata, two relatively common species, thrive on dry coastlines and in xerophilous forests [6]
Figure 2. Schematic representation of the bioclimatic and plant stages in the Lesser Antilles (Amerindian period and present day) [13].
[26] [27]. Both are said to be valued for their traditional uses: Randia aculeata for its medicinal properties, particularly for treating snake bites, and Erithalis fruticosa for its Amerindian cultural uses, the resin was once used to make torches [7] [31] [32]. In contrast, Genipa americana and Ixora ferrea are rarer and occupy mesophilous to hygrophilous habitats [6] [26] [27]. The former is well known for its multiple uses: its fruit is edible, yielding a blackish dye used by Amerindians for their rituals, and has antibacterial properties [33] [34]. The latter may be primarily ornamental, like several other species of the same genus [35].
Guettarda scabra, although fairly common locally, is classified as critically endangered globally [30]. It grows in dry, often degraded areas, in xerophilous and xero-mesophilous forests [26] [27]. The genus provides a heavy wood prized for construction, while folk medicine uses it to treat various inflammatory or digestive disorders [8] [36].
Finally, Randia nitida appears to be the most threatened [6] [26] [27] [30]. This rare species, adapted to xero-mesophilous to mesophilous forests, is believed to have nutritional (edible fruit) and medicinal (healing, inflammation, digestive disorders) benefits, reinforcing the urgent need for conservation measures [8] [37]-[40].
Figure 3. Location of the Ethnobotanical Park in the town of Marin, Martinique.
According to the most recent flora of the region, these six species also present varied phenological rhythms (flowering and fruiting staggered according to the species), which would allow for the consideration of differentiated management strategies (Figure 4) [6]. All of these elements together can contribute to the development of targeted conservation plans or to the development of restoration programs adapted to the ecological and cultural specificities of each taxon.
Table 1. Bibliographic data [6] [20] [26] [30].
Scientific name |
Threat status worldwide (INPN) |
Highest threat status in the french antilles (INPN) |
Abundance according to Fournet J., 2002 |
Remarkable uses |
Ecological profile |
Flowering period |
Fruiting period |
Erithalis fruticosa (Shrub) |
LC—Least Concern |
LC—Least Concern |
Common |
Cultural, religious, and utilitarian: Very hard and durable wood. The resin is flammable and was once used by Amerindians as torches. Medicinal: Measles and wounds, treating a newborn’s bleeding navel, etc.. |
Dry coastline. Altitude: 0 to 80 meters. |
All year round. |
- |
Genipa americana (Tree) |
Not Assessed |
Not Assessed |
Fairly rare |
Cultural and religious: Plant formerly used by Amerindians for ceremonies and blackish dye. Medicinal: Antibacterial. Food: Edible fruit. |
Mesophilous vegetation layer, absent from primary mesophilous forest. Often planted. Altitude: 20 to 200 meters. |
September-October. And often June-July. |
|
Guettarda scabra (Tree) |
CR—Critically Endangered |
LC—Least Concern |
Fairly common |
Utilitarian uses: Hard, heavy wood, poles for construction. Medicinal: Stomach problems, general inflammation. |
Lower hills and dry, rocky coastline. Most common in xerophilous forest and xero-mesophilous forest, in degraded areas. Altitude: 0 to 100, even 500 meters. |
Almost all year round, intermittently. |
|
Ixora ferrea (Shrub) |
- |
LC—Least Concern |
Fairly rare |
Ornamental plant. Utilitarian uses: hard and heavy wood, stakes, firewood, etc. |
Mesophilous and hygrophilous forest. Altitude: 0 to 150, even 700 meters. |
March-April |
September |
Randia aculeata (Shrub) |
LC—Least Concern |
LC—Least Concern |
Common |
Utilitarian uses: Brown wood, hard, heavy, strong, durable. Manufacture of fishing rods, Christmas trees, blue dye (fruit pulp)... Medicinal: Against snake bites, chronic pain, cancer, bone fever, wound care and infections... |
Degraded or secondary xerophilous and
xero-mesophilous forest. Lowland dry region, especially on the coast. Altitude: 0 to 150 meters. |
Especially during long days. |
- |
Randia nitida or armata (Tree) |
- |
CR—Critically Endangered |
Fairly rare |
Food: edible fruit. Medicinal: helps with healing, to treat inflammatory diseases, diarrhea, etc. |
Xero-mesophilous and mesophilous forest. Altitude: 0 to 200 meters. |
- |
- |
![]()
Figure 4. Illustrations of the flowers, fruits and stems of the six Rubiaceae studied.
2.2. Methods
This study is based on a methodological approach combining field observations, floristic inventories and bibliographic research, with the aim of enriching our knowledge on the phenological rhythms and ecological requirements of six remarkable or threatened species of the Rubiaceae family, present in Martinique and preserved in the Ethnobotanical Park of the town of Marin. Thus, between June 2016 and February 2020, monthly observations were carried out in this park to monitor the phenological cycles of these six species. Each month, a single visit made it possible to record, for all individuals, the main flowering and fruiting phases, as well as to qualify stem growth and the evolution of tree height using a descriptive gradient (slow, moderate, rapid, sustained, continuous or regular growth). However, the quantities of flowers and fruits produced were not estimated. Although this protocol of a single random monthly visit may underdetect very short-term blooms, repeating these observations over several years reduces this bias because they were cross-referenced with bibliographic data, thereby limiting potential underdetections and strengthening the reliability of the phenological trends described.
These data were enriched by the reuse of a database of 120 floristic inventories carried out randomly in different bioclimates and associated forests, across Martinique between 2015 and 2020, initially mobilized as part of doctoral work on the ecology of Rubiaceae, within the UMR ESPACE DEV-BIORECA laboratory of the University of the Antilles [26]. The areas of the transects (or minimum areas) varied according to the plant communities studied; on average from 240 square meters to more than 1000 square meters, generally from the dry to humid bioclimate, due to greater floristic richness in the forest formations of more humid bioclimates. These inventories were revisited in order to precisely identify the occurrences of the six targeted species, and to extract information such as their ecological profile and ecosystem affinity (belonging to types of plant formations and bioclimates), their ecological importance (population dynamics), as well as their ecological temperament (i.e. their behavior with respect to environmental gradients).
At the same time, a bibliographic review was conducted using scientific literature or online scientific databases (Google Scholar, ResearchGate, and the National Inventory of Natural Heritage—INPN) in order to gather existing knowledge on these species and thus describe the material part of our study. All of these elements made it possible to constitute a fairly comprehensive database for the analysis of the ecological dynamics specific to each species.
3. Results
3.1. Field Observations (June 2016-February 2020)
The town of “Marin”, where the ethnobotanical park is located, is characterized by a tropical climate with an average annual temperature of 26.5˚C, oscillating between 24˚C and 29˚C, average annual rainfall of approximately 1800 mm, annual evapotranspiration of approximately 1675 mm, and annual insolation of approximately 2800 hours (Table 2) [41].
It is in this climatic context that the six species of the Rubiaceae family, a total of 33 individuals, were monitored from June 2016 to February 2020 at the Ethnobotanical Park. The 33 individuals correspond to all representatives of the six target species present in the park at the time of the study. This comprehensive park-wide sampling integrates different physiognomies and developmental stages, including both bushes, shrubs and young growing trees, making it representative of the ex situ populations maintained there. Although limited in size, the dataset provides original ecological and phenological trends, valuable for conservation purposes. However, any inferences at a broader regional scale must be made with caution. Genipa americana, initially in shrub form for the majority of individuals, has evolved into a tree with rather rapid and sustained growth (Table 3). Conversely, Ixora ferrea and Erithalis fruticosa showed fairly slow growth to reach the shrub stage, while Guettarda scabra, Randia aculeata and Randia nitida, showed moderate growth to become trees or shrubs (Table 3).
Table 2. Main climatic characteristics of the commune of Marin (Source: Météo-France and IGN).
Average annual minimum temperature |
Average annual maximum temperature |
Average annual temperature |
Average annual rainfall |
Annual evapotranspiration |
Annual insolation |
24˚C |
29˚C |
26.5˚C |
≃1800 mm |
≃1675 mm |
≃2800 hours |
Table 3. Growth characteristics of the 6 Rubiaceae preserved and observed at the Marin Ethnobotanical Park.
Scientific name |
Number of individuals |
Initial physiognomy (2016) |
Terminal physiognomy (2020) |
Annual growth type of stems |
Annual growth type of individual heights |
Erithalis fruticosa |
3 |
1 shrub |
Shrub |
Slow and continuous |
Slow and continuous |
2 bushes |
Genipa americana |
9 |
8 shrubs |
Tree |
Fast and sustained |
Fast and sustained |
1 bush |
Guettarda scabra |
4 |
2 trees |
Tree |
Moderate |
Moderate |
2 bushes |
Ixora ferrea |
2 |
1 bush |
Shrub |
Slow and continuous |
Slow and continuous |
Randia aculeata |
2 |
2 bushes |
Shrub |
Moderate |
Moderate |
Randia nitida or armata |
13 |
13 bushes |
Tree |
Moderate |
Moderate |
Figure 5 shows the spatial distribution of observed individuals, mostly grouped by species within the park. While this distribution does not appear to directly impact the growth rates or phenological cycles of the species, it nevertheless reflects a diversity of microenvironments, linked to the hilly topography of the site, located in a low-lying area and in close proximity to a watercourse. Thus, Erithalis fruticosa presents a flowering that appears almost continuous throughout the year, accompanied by intermittent fruiting equally distributed throughout the year (Table 4). Genipa americana flowers mainly during the rainy season, while its fruiting appears asynchronous and can extend over a large part of the year (Table 4).
Guettarda scabra flowers more during the shoulder season and the rainy period, resulting in fruiting that extends beyond the flowering peak, almost until the end of the year (Table 4). Ixora ferrea has a relatively brief flowering period early in the year during the dry season, with some late fruits observed thereafter, while Randia aculeata appears not to follow a strict cycle: flowers and fruits can be observed at different times of the year (Table 4). Finally, Randia nitida briefly loses its leaves before flowering, suggesting a floral peak at the end of the dry season, extending into the rainy season. Fruits, on the other hand, are visible throughout the year and mature at variable times (Table 4).
Figure 5. Location of the sites of the 6 Rubiaceae preserved at the Marin Ethnobotanical Park (several individuals of the same species are located close to each other).
3.2. Reuse of Floristic Inventory Data (2015-2020)
Between 2015 and 2020, 120 floristic inventories were carried out in Martinique and made it possible to note the presence of these six Rubiaceae in natural environments [26] [27].
The minimum areas of the transects carried out varied from 240 to more than 1000 m2 and covered a diversity of plant formations presented in Table 5 (Appendix 1 and Appendix 2).
Table 4. Phenological phases of flower and fruit production of the 6 Rubiaceae. Table produced according to the model of Lobo Segura, J. A., et al., in 2007 [42].
Species |
Phases |
|
Dry season |
Shoulder season |
Rainy season |
Shoulder season |
January |
February |
March |
April |
May |
June |
Juily |
August |
September |
October |
November |
December |
Erithalis fruticosa |
Flowering |
|
|
|
|
|
|
|
|
|
|
|
|
Fruiting |
|
|
|
|
|
|
|
|
|
|
|
|
Genipa americana |
Flowering |
|
|
|
|
|
|
|
|
|
|
|
|
Fruiting |
|
|
|
|
|
|
|
|
|
|
|
|
Guettarda scabra |
Flowering |
|
|
|
|
|
|
|
|
|
|
|
|
Fruiting |
|
|
|
|
|
|
|
|
|
|
|
|
Ixora ferrea |
Flowering |
|
|
|
|
|
|
|
|
|
|
|
|
Fruiting |
|
|
|
|
|
|
|
|
|
|
|
|
Randia aculeata |
Flowering |
|
|
|
|
|
|
|
|
|
|
|
|
Fruiting |
|
|
|
|
|
|
|
|
|
|
|
|
Randia nitida or armata |
Flowering |
|
|
|
|
|
|
|
|
|
|
|
|
Fruiting |
|
|
|
|
|
|
|
|
|
|
|
|
Table 5. Summary table of the characteristics of the 120 floristic inventories carried out in Martinique from 2015 to 2020 (Appendix 1 and Appendix 2) [26] [27].
|
Lower plant level |
Medium plant level |
|
Mangrove |
Back- mangrove |
Backshore |
F.S.S.T.F.X |
F.T.X.M |
F.S.S.T.T |
F.S.S.T.T |
F.O.S.S.T |
F.O.S.M.T |
Number of inventories |
5 |
9 |
11 |
55 |
10 |
17 |
1 |
5 |
7 |
Total area (m2) |
3150 |
10790 |
6065 |
31110 |
6610 |
12560 |
500 |
3950 |
6250 |
Minimum inventory area (m2) |
400 to 950 |
420 to 2200 |
240 to 800 |
350 to 1000 |
500 to 1000 |
450 to 1000 |
500 |
600 to 1000 |
700 to 1050 |
Number of inventories per commune in Martinique |
(3) Trinité, (1) Robert, (1)
Sainte-Anne |
(9) Marin |
(2) Trinité, (2) Vauclin, (3) Marin, (4) Sainte-Anne |
(2) Case Pilote, (3) Schoelcher, (12) Vauclin, (10) Diamant, (14) Marin, (14)
Sainte-Anne |
(2) Schoelcher, (3) Diamant, (5) Marin |
(9) Schoelcher, (3) Marin, (5) Vauclin |
(1) Diamant |
(1) Sainte-Marie, (1) Case Pilote, (2)
Sainte-Luce, (1) Marin |
(3) Sainte-Marie, (1) Case Pilote, (3) Fort-de-France |
Inventory team |
Members of the UMR SPACE DEV-BIORECA laboratory |
F.S.S.T.F.X: Tropical seasonal evergreen and xeric facies plant formations (xerophilous), F.T.X.M: Transitional xero-mesophilous plant formations (tropical seasonal evergreen), F.S.S.T.T: Typical tropical seasonal evergreen plant formations (mesophilous), F.O.S.S.T: Tropical seasonal ombro-evergreen plant formations (mesohygrophilous), F.O.S.M.T: Tropical sub-montane rainforest plant formations (hygrophilous).
Table 6 shows that Guettarda scabra was the most common species with 1271 individuals, followed by Erithalis fruticosa with 972 and Randia aculeata with 893. Genipa americana, Ixora ferrea and Randia nitida were rarer, with 2, 25 and 28 individuals respectively. The presence of seedlings for most of these species in several stations indicates a certain capacity for regeneration.
Table 7 describes their ecological temperament: Erithalis fruticosa and Guettarda scabra (and even Genipa americana) are strongly heliophilous, Randia aculeata shows a more mixed behavior, while Ixora ferrea and Randia nitida show a preference for more shaded conditions. The majority of these species belong to the lower or even middle strata of our forests with the exception of Guettarda scabra which can contribute to the canopy species (Table 7).
Table 6. Number of inventory stations and number of individuals of the 6 Rubiaceae (Appendix 1 and Appendix 2) [26].
Species |
Type |
Number of adult individuals (excluding regenerations
and dead trees) |
Ecological significance |
Stations where the species is present (excluding regenerations and
seedlings) |
Stations where the species is present at the regeneration/ seedling stage |
Erithalis fruticosa |
Shrub |
972 |
very low |
S27, S31, S33, S39, S47, S50, S52, S55, S56, S57, S59, S60, S61, S62, S64, S65, S66, S67, S83, S84, S85, S86, S95, S96, S97 |
S31, S33, S34, S35, S47, S50, S56, S61, S67, S83, S85, S86, S95, S96 |
Genipa americana |
Tree |
2 |
very low |
S110 |
- |
Guettarda scabra |
Tree |
1271 |
very low |
S46, S47, S55, S56, S57, S60, S61, S62, S64, S65, S66, S67, S82, S83, S86, S108, S106, S116, S118 |
S46, S47, S56, S61, S63, S67, S83, S85, S103 |
Ixora ferrea |
Shrub |
25 |
very low |
S88, S89, S90, S98, S103, S104, S107, S108 |
S89, S98, S103, S107 |
Randia aculeata |
Shrub |
893 |
very low |
S6, S8, S9, S13, S20, S21, S22, S23, S32, S38, S44, S45, S46, S47, S49, S56, S57, S58, S59, S60, S61, S65, S66, S67, S83, S85, S86, S97, S100, S101, S103, S110, S111, S112, S114, S116, S118 |
S6, S25, S32, S44, S45, S47, S56, S58, S61, S63, S67, S83, S85, S86, S97, S100, S103, S115, S117 |
Randia nitida or armata |
Tree |
28 |
very low |
S28, S29, S30 |
S71 |
Ecological importance: contribution of species in terms of abundance, biomass and space occupation to the forest communities studied (very low, low, medium, high and very high).
Guettarda scabra and Erithalis fruticosa occurred in shrub formation, mature shrub formation, presylvatic, and young sylvatic forests, in other words, primarily in extrasylvatic forests (Table 7). Randia aculeata appeared more prevalent in shrub formation and mature shrub formation (extrasylvatic forests), while the other species appeared more ecologically restricted, limited to the more advanced stages of our forests (intrasylvatic forests) (Table 7).
Table 8 shows their affinity for various ecosystem types. Erithalis fruticosa and Randia aculeata were more abundant in backshore forests and in xerophilous forests, while Guettarda scabra was particularly abundant in xerophilous forests. Genipa americana, although very poorly represented, was found only in mesophilous forests (Table 8). Ixora ferrea, poorly represented, seemed more associated with mesophilous or even hygrophilous formations, while Randia nitida, also poorly represented, clearly showed a weak affinity for coastal plant formations as well as for hygrophilous forests. Erithalis fruticosa, Guettarda scabra, and Randia aculeata could be found among the dominant species by their population abundance (number of individuals) or by their distribution (spatial distribution of species), while their basal area (phytomasses or biovolumes of plants) was very small. In contrast, for all other species, ecological dominance was clearly negligible.
Table 7. Temperament and ecological profile of the 6 Rubiaceae identified in the stations, according to the qualitative representation model by Professor Philippe JOSEPH [7] [26] [27].
Species |
Erithalis
fruticosa |
Genipa
americana |
Guettarda scabra |
Ixora ferrea |
Randia
aculeata |
Randia nitida (or armata) |
Physiognomy |
Shrub |
Tree |
Tree |
Shrub |
Shrub |
Tree |
PH |
++++ |
|
++++ |
|
+++ |
|
SH |
++ |
+ |
++ |
|
++ |
+ |
SGH |
|
|
++ |
|
|
|
HémSc |
|
|
|
+ |
+ |
|
SPME |
Lower and Middle |
Middle |
Middle and Upper |
Lower |
Lower |
Lower |
O |
NO |
NO |
T |
NO |
P |
P |
S.F |
++++ |
|
+++ |
|
+++ |
|
M.S.F |
+++++ |
|
+++++ |
|
+++ |
|
P.F |
+++ |
|
++++ |
|
+ |
+ |
S.Y.S.F |
+ |
+ |
++ |
+ |
+ |
+ |
S.S.F |
|
|
S |
+ |
+ |
|
PH: Primary heliophile/SH: Secondary heliophile/SGH: Sylvatic gap heliophile/HémSc: Hemisciaphile/SPME: Stratigraphic position at maximum expansion/O: Obsolescence; (T: Total, NO: Not obsolete, P: Partial)/(S.F): Shrub formation/(M.S.F): Mature shrub formation/(P.F): Presylvatic formation/(S.Y.S.F): Structured young sylvatic formation/(S.S.F): Secondary sylvatic formation/S: Seedling/Significance of descriptor: (+) very low, (++) low, (+++) medium, (++++) high, and (+++++) very high.
Table 8. Ecosystem affinity of the 6 Rubiaceae identified in the stations, according to the qualitative representation model by Professor Philippe JOSEPH [7] [26] [27].
|
A.M |
A.P |
F.S.S.T.F.X |
F.T.X.M |
F.S.S.T.T |
F.O.S.S.T |
F.O.S.M.T |
Species |
Ab |
D |
ED |
Ab |
D |
ED |
Ab |
D |
ED |
Ab |
D |
ED |
Ab |
D |
ED |
Ab |
D |
ED |
Ab |
D |
ED |
Erithalis fruticosa |
|
|
|
+++++ |
M |
Lo |
+++ |
M |
Lo |
+ |
EL |
EL |
|
|
|
|
|
|
|
|
|
Genipa americana |
|
|
|
|
|
|
|
|
|
|
|
|
+ |
EF |
EL |
|
|
|
|
|
|
Guettarda scabra |
+ |
EL |
EF |
+ |
Lo |
EL |
++++ |
Lo |
Lo |
++ |
M |
M |
+ |
VL |
VL |
|
|
|
|
|
|
Ixora ferrea |
|
|
|
|
|
|
|
|
|
|
|
|
+ |
EL |
EL |
+ |
VL |
EL |
+ |
VL |
EL |
Randia aculeata |
|
|
|
+++++ |
M |
Lo |
+++ |
Lo |
VL |
++ |
VL |
EL |
+ |
EL |
EL |
|
|
|
|
|
|
Randia nitida |
|
|
|
|
|
|
+ |
VL |
EL |
|
|
|
+ |
Lo |
EL |
|
|
|
|
|
|
Ab: Abundance/D: distribution/ED: Ecological dominance/M: Medium, Lo: Low, VL: Very low, EL: Extremely low /A.M: Back-mangrove floristic units, A.P: Backshore floristic units, F.S.S.T.F.X: Tropical seasonal evergreen and xeric facies plant formations (xerophilous), F.T.X.M: Transitional xero-mesophilous plant formations (tropical seasonal evergreen), F.S.S.T.T: Typical tropical seasonal evergreen plant formations (mesophilous), F.O.S.S.T: Tropical seasonal ombro-evergreen plant formations (mesohygrophilous), F.O.S.M.T: Tropical sub-montane rainforest plant formations (hygrophilous). Significance of descriptor: (+) very low, (++) low, (+++) medium, (++++) high, and (+++++) very high.
4. Discussion
The island context of Martinique is characterized by strong anthropogenic pressure and a rich and extremely vulnerable biodiversity [4] [6] [7] [9] [16] [20]-[22] [25] [26].
Urbanization, agriculture, the introduction of invasive species, and current climate change are increasingly creating major ecological risks, threatening the stability, diversity, and functioning of natural ecosystems [7] [17] [19] [21] [22].
Several in situ conservation measures have been in place in Martinique since the beginning of the 20th century to oversee the preservation of plant biodiversity. Currently, there are several dozen terrestrial and marine areas with various legal statuses, such as strict biological reserves, national or regional nature reserves, regional or marine natural parks, prefectural biotope protection decrees, and nature reserves [30]. The Martinique Regional Natural Park, for example, extends over 2/3 of the island, including forest, agricultural, coastal and marine areas, and its main mission is the protection and enhancement of natural and cultural heritage, the sustainable development of territories, as well as supporting communities in the concerted management of the island’s remarkable and sensitive areas [43].
In addition to protected areas, certain territories without regulatory status play an essential role in the knowledge and inventory of natural heritage, in particular the 24 Natural Zones of Ecological, Faunistic and Floristic Interest (ZNIEFF) spread across the entire island [30].
At the same time, targeted actions are implemented to preserve heritage and threatened species, through tools such as National Action Plans or Conservation Master Plans led in particular by the National Botanical Conservatory of Martinique [44]. These plans aim to coordinate targeted preservation actions for heritage and threatened plant species, by defining priorities, management objectives and concrete conservation measures, generally over periods of 5 to 10 years [44].
It is also worth noting the existence of projects such as those led by public organizations such as INRAE (National Research Institute for Agriculture, Food and the Environment) and CIRAD (Centre for International Cooperation in Agricultural Research for Development), aimed at the conservation and characterization of tropical plant biological resources with the aim of promoting their agronomic, scientific, and heritage value [45].
Municipalities are beginning to integrate these issues into their urban planning documents, although this approach remains poorly coordinated with local scientific actors. The lack of unified governance and synergies between regional actors have long limited the overall effectiveness of all these tools, despite their individual relevance [7] [17] [19] [24].
A stronger integration of scientific approaches is also necessary to strengthen their impact. This is the existential objective of the Ethnobotanical Park of the city of Marin, created in 2017. Its purpose is to strengthen the effectiveness and coherence of conservation and ecological restoration actions carried out on the island, through its scientific role focused on the study of the phenological cycles of species and the conservation of seed banks of threatened native plant species.
It constitutes a strategic asset that is currently little used and can provide essential data for the adaptive management of ecosystems. Our study, conducted between 2016 and 2020 on six threatened or remarkable Rubiaceae conserved at the Marin Ethnobotanical Park, is part of this dynamic. The combined approach based on phenological observations and floristic inventories provides original data on the seasonal cycles, growth rates, and ecological preferences of the species studied, namely Erithalis fruticosa, Genipa americana, Guettarda scabra, Ixora ferrea, Randia aculeata, and Randia nitida [26] [27].
The results demonstrate variability in phenological strategies and different ecological profiles and temperaments, ranging from fast-growing and heliophilous species such as Genipa americana or Guettarda scabra, to others that are slower and shade-tolerant such as Ixora ferrea or Randia nitida. Some exhibit almost continuous flowering (Erithalis fruticosa), while others follow marked seasonal cycles. Finally, their ecological distribution varies according to the formations: some are found in the extra-sylvatic and xerophilous stages (Erithalis fruticosa, Guettarda scabra, Randia aculeata), while others are more restricted to mesophilous or hygrophilous intra-sylvatic forests (Genipa americana, Ixora ferrea, Randia nitida). In terms of in situ conservation, the data collected can be used to refine management strategies for natural populations. For example, species such as Guettarda scabra or Randia aculeata, relatively common in xerophilous or degraded habitats, could be integrated into rehabilitation programs for open or disturbed environments, as indicator species of ecosystem resilience [7]. Others, such as Randia nitida or Genipa americana, which are more ecologically demanding, require specific interventions in mesophilous or transitional habitats, including site protection, regulation of uses and possibly the reconstitution of microhabitats.
Based on these results, several management measures can be proposed to strengthen the conservation of the six Rubiaceae studied. Seed collection could, for example, be carried out opportunistically, during peak fruiting periods, and stored in local seed banks to secure genetic resources for potential reintroduction programs. Restoration efforts should prioritize degraded xero-mesophilic and mesophilic habitats, which represent the natural ecological niches of the rarest species. At the same time, annual monitoring would be relevant and advisable to assess the effectiveness of existing conservation actions. This monitoring would include the assessment of natural regeneration, growth, and survival of young recruits planted in different habitats, and the early detection of restoration failures. This information would allow management strategies to be adapted if necessary, for example, by reinforcing plantations or adjusting restoration sites.
Pending the implementation of these measures, the Marin Ethnobotanical Park represents a major asset for ex situ conservation in Martinique. The phenological and growth data collected in its living collections provide valuable information for refining propagation protocols, consolidating ex situ populations, and supporting future reintroduction or translocation programs.
Strengthening the integration of scientific data generated by the park with the actions of national and regional partners would significantly enhance the coherence of conservation strategies and contribute to the sustainability of threatened or remarkable Rubiaceae in the Lesser Antilles.
5. Conclusions
This study provided a deeper understanding of the phenological rhythms and ecological requirements of six threatened or remarkable Rubiaceae species in the Lesser Antilles, particularly in Martinique, through an interdisciplinary approach combining field observations, floristic inventories, and documentary analysis. Monthly observations conducted between 2016 and 2020 at the Ethnobotanical Park of the town of Marin, a reference site for ex situ conservation, provided valuable data on the seasonal behavior of these species in a controlled environmental context.
These results were enriched by the use of a database of 120 floristic inventories, conducted across Martinique between 2015 and 2020 as part of a doctoral project, allowing for a detailed characterization of the ecological profiles of the species studied. All the information collected contributes to our knowledge of the local dynamics of these Rubiaceae, and provides a useful scientific basis for the implementation of targeted conservation strategies, by identifying the ecological conditions conducive to their maintenance in the natural environment. Furthermore, the results obtained at the park strengthen ex situ conservation capacities, particularly for the safeguarding of genetic material and future reintroduction projects, thus laying the foundations for more coherent and sustainable management of threatened flora in the Lesser Antilles.
Acknowledgements
Monthly observations from 2016 to 2020, at the Ethnobotanical Park of the town of Marin in Martinique, were carried out by Jean-Philippe CLAUDE and Yelji ABATI. The floristic inventories were carried out from 2015 to 2020 with members of the UMR ESPACE DEV-BIORECA laboratory of the University of the Antilles. Several ecoclimatic data were collected from public organisations: Météo-France (French meteorological organisation), the IRD (Institute of Research for Development), the IGN (National Geographic Institute), the CTM (Territorial Collectivity of Martinique), and the DEAL (Department of Environment, Planning and Housing of Martinique).
Data Availability
The raw data from the observations carried out at the Ethnobotanical Park, in the town of Marin in Martinique, are not accessible online to the public. However, the floristic inventories and their analyses can be consulted from the thesis available online: https://theses.fr/2020ANTI0548.
Appendix 1. List of 120 Stations of Floristic Inventory [26]
Station |
Municipality |
Latitude |
Longitude |
1 |
Marin |
1602031 |
729548 |
2 |
Marin |
1602790 |
730343 |
3 |
Marin |
1602804.212 |
730448.953 |
4 |
Marin |
1602635.67 |
729086.75 |
5 |
Marin |
1602702.58 |
729166.36 |
6 |
Marin |
1602832.66 |
729254.34 |
7 |
Marin |
1603153.387 |
729538.061 |
8 |
Marin |
1603128.362 |
729578.997 |
9 |
Marin |
1603232.518 |
729542.695 |
10 |
Marin |
1603228 |
729674 |
11 |
Marin |
1603299 |
729802 |
12 |
Marin |
1602758.07 |
730266.76 |
13 |
Marin |
1602779.81 |
730231.56 |
14 |
Trinité |
1,629,234 |
720,802 |
15 |
Robert |
1,621,000 |
723,455 |
16 |
Trinité |
1,633,603 |
727,300 |
17 |
Trinité |
1,633,665 |
727,750 |
18 |
Sainte-Anne |
1,594,081 |
728,106 |
19 |
Sainte-Marie |
1,632,981 |
709,011 |
20 |
Vauclin |
1,605,426 |
731,836 |
21 |
Vauclin |
1,605,425 |
731,812 |
22 |
Vauclin |
1,605,458 |
731,742 |
23 |
Vauclin |
1,605,573 |
731,777 |
24 |
Vauclin |
1,605,392 |
731,742 |
25 |
Vauclin |
1,605,476 |
731,898 |
26 |
Vauclin |
1,605,603 |
731,905 |
27 |
Vauclin |
1,605,546 |
731,951 |
28 |
Vauclin |
1,605,678 |
731,892 |
29 |
Vauclin |
1,605,489 |
731,943 |
30 |
Vauclin |
1,605,446 |
732,031 |
31 |
Sainte-Anne |
1,596,214 |
728,435 |
32 |
Sainte-Anne |
1,596,267 |
728,688 |
33 |
Sainte-Anne |
1595999.6 |
728,691 |
34 |
Sainte-Anne |
1596055.71 |
728419.08 |
35 |
Sainte-Anne |
1595861.2 |
728400.4 |
36 |
Sainte-Anne |
1595721.1 |
728207.6 |
37 |
Sainte-Anne |
1595767.6 |
728957.4 |
38 |
Sainte-Anne |
1596079.8 |
728883.2 |
39 |
Sainte-Anne |
1596794 |
728901 |
40 |
Sainte-Anne |
1596814.8 |
728864.2 |
41 |
Sainte-Anne |
1597047.7 |
728,891 |
42 |
Sainte-Anne |
1596943.4 |
728911.1 |
43 |
Sainte-Marie |
1,635,425 |
709,662 |
44 |
Sainte-Anne |
1594747.8 |
727796.9 |
45 |
Sainte-Anne |
1594877.7 |
727664.3 |
46 |
Trinité |
1633012.664 |
723263.003 |
47 |
Trinité |
1632970.626 |
723193.05 |
48 |
Sainte-Anne |
1596236.7 |
728530.3 |
49 |
Sainte-Anne |
1,598,801 |
731,246 |
50 |
Sainte-Anne |
1596230.4 |
727970 |
51 |
Marin |
1,601,365 |
730,633 |
52 |
Marin |
1,601,395 |
730,640 |
53 |
Marin |
1,601,582 |
729,986 |
54 |
Marin |
1,601,515 |
729,983 |
55 |
Diamant |
1602644.1 |
710598.1 |
56 |
Diamant |
1602490.7 |
710584.2 |
57 |
Diamant |
1,602,544 |
710568.5 |
58 |
Diamant |
1602803.6 |
710674.1 |
59 |
Diamant |
1602798 |
710624.1 |
60 |
Diamant |
1602519.9 |
710484.1 |
61 |
Diamant |
1602552.8 |
710448.9 |
62 |
Diamant |
1602620 |
710550.8 |
63 |
Diamant |
1602782.7 |
710589.8 |
64 |
Diamant |
1602573.5 |
710500.9 |
65 |
Diamant |
1602653.7 |
710634.6 |
66 |
Diamant |
1602632.3 |
710644.4 |
67 |
Diamant |
1602597.62 |
710624.569 |
68 |
Vauclin |
1606351.026 |
732132.936 |
69 |
Vauclin |
1606278.180 |
732237.385 |
70 |
Vauclin |
1606245.193 |
732278.094 |
71 |
Vauclin |
1606232.594 |
732309.914 |
72 |
Vauclin |
1606316.108 |
732195.638 |
73 |
Vauclin |
1606165.132 |
732544.927 |
74 |
Marin |
1600827.3 |
728519 |
75 |
Marin |
1600805.9 |
728492.3 |
76 |
Marin |
1600789 |
728462.8 |
77 |
Marin |
1600763.6 |
728438.3 |
78 |
Marin |
1600738.6 |
728405.8 |
79 |
Marin |
1600807.7 |
728538.6 |
80 |
Marin |
1600786.3 |
728560.5 |
81 |
Marin |
1600737.7 |
728472.6 |
82 |
Marin |
1600766.7 |
728476.2 |
83 |
Marin |
1598933.353 |
726089.815 |
84 |
Marin |
1598910.04 |
726131.789 |
85 |
Marin |
1598964.295 |
726048.418 |
86 |
Marin |
1599059.894 |
726027.099 |
87 |
Fort de France |
1623701.061 |
705193.128 |
88 |
Fort de France |
1623631.194 |
705126.389 |
89 |
Fort de France |
1623653.201 |
705102.174 |
90 |
Case-Pilote |
1623964.111 |
703308.368 |
91 |
Marin |
1601000.510 |
730073.266 |
92 |
Marin |
1600958.733 |
730097.480 |
93 |
Marin |
1600935.651 |
730078.230 |
94 |
Marin |
1601008.769 |
730099.153 |
95 |
Vauclin |
1604298.099 |
734510.52 |
96 |
Vauclin |
1604458.993 |
734534.568 |
97 |
Sainte-Anne |
1593740.426 |
728079.439 |
98 |
Sainte-Luce |
1603732.913 |
722993.613 |
99 |
Sainte-Luce |
1603655.382 |
722823.402 |
100 |
Case-Pilote |
1620776.101 |
699786.154 |
101 |
Case-Pilote |
1620872.739 |
699733.764 |
102 |
Schoelcher |
1620854.731 |
704085.829 |
103 |
Schoelcher |
1620652.683 |
704241.155 |
104 |
Case-Pilote |
1623076.7 |
701880.338 |
105 |
Sainte-Marie |
1633707.432 |
710027.972 |
106 |
Fort de France |
1,623,610 |
704,779 |
107 |
Marin |
1,599,548 |
726,211 |
108 |
Diamant |
1,604,706 |
712,505 |
109 |
Schoelcher |
1,617,852 |
705,223 |
110 |
Schoelcher |
1,617,795 |
705,287 |
111 |
Schoelcher |
1,617,949 |
705,237 |
112 |
Schoelcher |
1,617,974 |
705,287 |
113 |
Schoelcher |
1,618,098 |
705,376 |
114 |
Schoelcher |
1,618,423 |
705,380 |
115 |
Schoelcher |
1,618,466 |
705,455 |
116 |
Schoelcher |
1,618,569 |
705,636 |
117 |
Schoelcher |
1,618,747 |
705,711 |
118 |
Schoelcher |
1,619,300 |
705,344 |
119 |
Schoelcher |
1,618,566 |
705,426 |
120 |
Schoelcher |
1,618,687 |
705,451 |
Appendix 2. Characteristics of the 120 Inventory Stations of the Thesis [26]
S |
SE |
Bioclimate or mesoclimate |
Tmin |
Tmax |
Taverage |
Rain |
Eva |
Ins |
H |
1 |
SYSF |
ecotone
(xero-mesophile) |
23.2 |
28.9 |
26.1 |
1854 |
1675 |
2800 |
51 |
2 |
SYSF |
ecotone
(xero-mesophile) |
23.1 |
28.4 |
25.8 |
1853 |
1675 |
2800 |
64 |
3 |
SYSF |
ecotone
(xero-mesophile) |
23.1 |
28.4 |
25.8 |
1853 |
1675 |
2800 |
66 |
4 |
PF |
subhumid dry |
23.2 |
28.9 |
26.1 |
1854 |
1675 |
2800 |
45 |
5 |
PF |
subhumid dry |
23.2 |
28.9 |
26.1 |
1854 |
1675 |
2800 |
49 |
6 |
PF |
subhumid dry |
23.2 |
28.9 |
26.1 |
1854 |
1675 |
2800 |
82 |
7 |
PF |
subhumid dry |
22.4 |
27.9 |
25.2 |
1900 |
1675 |
2800 |
84 |
8 |
PF |
subhumid dry |
22.4 |
27.9 |
25.2 |
1900 |
1675 |
2800 |
69 |
9 |
PF |
subhumid dry |
22.4 |
27.9 |
25.2 |
1900 |
1675 |
2800 |
108 |
10 |
SYSF |
subhumid dry |
22.4 |
27.9 |
25.2 |
1900 |
1675 |
2800 |
56 |
11 |
SYSF |
subhumid dry |
22.4 |
27.9 |
25.2 |
1900 |
1675 |
2800 |
72 |
12 |
PF |
subhumid dry |
23.1 |
28.4 |
25.8 |
1853 |
1675 |
2800 |
69 |
13 |
MSF |
subhumid dry |
23.1 |
28.4 |
25.8 |
1853 |
1675 |
2800 |
71 |
14 |
Mangrove |
subhumid dry |
23.8 |
30.4 |
27.1 |
2093 |
1625 |
2700 |
3 |
15 |
Mangrove |
subhumid dry |
24.6 |
30 |
27.3 |
1572 |
1625 |
2700 |
1 |
16 |
Mangrove |
subhumid dry |
23.4 |
30.8 |
27.1 |
1783 |
1650 |
2700 |
5 |
17 |
Mangrove |
subhumid dry |
23.4 |
30.8 |
27.1 |
1783 |
1650 |
2700 |
1 |
18 |
Mangrove |
subhumid dry |
24 |
29.5 |
26.8 |
1393 |
1625 |
2900 |
3 |
19 |
SSF |
humid |
21 |
28.1 |
24.6 |
3870 |
1575 |
2400 |
252 |
20 |
SYSF |
subhumid humid |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
152 |
21 |
SYSF |
subhumid dry |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
143 |
22 |
SYSF |
subhumid humid |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
131 |
23 |
SYSF |
subhumid humid |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
140 |
24 |
SYSF |
subhumid dry |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
130 |
25 |
SYSF |
subhumid dry |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
194 |
26 |
PF |
subhumid dry |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
182 |
27 |
SYSF |
subhumid dry |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
196 |
28 |
SYSF |
Subhumide humid |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
145 |
29 |
PF |
subhumid humide |
23.3 |
28.4 |
25.9 |
1575 |
1675 |
2700 |
201 |
30 |
SYSF |
subhumid dry |
23.5 |
28.9 |
26.2 |
1489 |
1675 |
2700 |
166 |
31 |
PF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
71 |
32 |
SYSF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
65 |
33 |
SYSF |
subhumid dry |
23.9 |
29.3 |
26.6 |
1421 |
1625 |
2900 |
89 |
34 |
SYSF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
80 |
35 |
SYSF |
subhumid dry |
23.9 |
29.3 |
26.6 |
1421 |
1625 |
2900 |
123 |
36 |
PF |
subhumid dry |
23.9 |
29.3 |
26.6 |
1421 |
1625 |
2900 |
30 |
37 |
SYSF |
subhumid dry |
23.9 |
29.3 |
26.6 |
1421 |
1625 |
2900 |
107 |
38 |
PF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
38 |
39 |
PF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
63 |
40 |
SYSF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
51 |
41 |
SYSF |
subhumid dry |
24.6 |
29.9 |
27.3 |
1476 |
1625 |
2900 |
71 |
42 |
SYSF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
82 |
43 |
SSF |
humid |
21.7 |
28.7 |
25.2 |
3123 |
1575 |
2400 |
177 |
44 |
SF |
subhumid dry |
24.3 |
29.6 |
27.0 |
1397 |
1625 |
2900 |
3 |
45 |
MSF |
subhumid dry |
24.3 |
29.6 |
27.0 |
1397 |
1625 |
2900 |
7 |
46 |
PF |
subhumid humid |
23.9 |
30.8 |
27.4 |
1951 |
1625 |
2700 |
14 |
47 |
PF |
subhumid humid |
23.9 |
30.8 |
27.4 |
1951 |
1625 |
2700 |
19 |
48 |
PF |
subhumid dry |
24.2 |
29.7 |
27.0 |
1441 |
1625 |
2900 |
54 |
49 |
SYSF |
subhumid dry |
23.6 |
28.9 |
26.3 |
1625 |
1675 |
2900 |
85 |
50 |
SYSF |
subhumid dry |
24.4 |
29.6 |
27.0 |
1438 |
1625 |
2900 |
13 |
51 |
SSF |
subhumid humid |
23.2 |
28.6 |
25.9 |
1814 |
1625 |
2800 |
30 |
52 |
SYSF |
subhumid dry |
23.2 |
28.6 |
25.9 |
1814 |
1625 |
2800 |
30 |
53 |
SYSF |
ecotone
(xero-mesophile) |
23.5 |
28.7 |
26.1 |
1810 |
1625 |
2800 |
80 |
54 |
SSF |
ecotone
(xero-mesophile) |
23.5 |
28.7 |
26.1 |
1810 |
1625 |
2800 |
80 |
55 |
SF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
108 |
56 |
MSF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
136 |
57 |
SF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
127 |
58 |
SYSF |
ecotone
(xero-mesophile) |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
97 |
59 |
SYSF |
ecotone
(xero-mesophile) |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
103 |
60 |
SF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
130 |
61 |
PF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
124 |
62 |
SF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
108 |
63 |
SYSF |
ecotone
(xero-mesophile) |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
103 |
64 |
SF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
116 |
65 |
PF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
108 |
66 |
SF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
115 |
67 |
MSF |
subhumid dry |
22.8 |
30.9 |
26.9 |
1609 |
1575 |
2700 |
115 |
68 |
SF |
subhumid dry |
24 |
29 |
26.5 |
1393 |
1675 |
2700 |
33 |
69 |
SF |
subhumid dry |
24 |
29 |
26.5 |
1393 |
1675 |
2700 |
67 |
70 |
SF |
subhumid dry |
24 |
29 |
26.5 |
1393 |
1675 |
2700 |
79 |
71 |
PF |
subhumid dry |
24 |
29 |
26.5 |
1393 |
1675 |
2700 |
80 |
72 |
PF |
subhumid dry |
24 |
29 |
26.5 |
1393 |
1675 |
2700 |
61 |
73 |
PF |
subhumid dry |
24 |
29 |
26.5 |
1393 |
1675 |
2700 |
80 |
74 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
75 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
76 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
77 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
78 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
79 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
80 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
81 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
82 |
E.C.D |
subhumid humid |
23.2 |
29.1 |
26.2 |
1751 |
1625 |
2800 |
4 |
83 |
PF |
subhumid dry |
23.4 |
29 |
26.2 |
1592 |
1625 |
2800 |
2 |
84 |
SF |
subhumid dry |
23.4 |
29 |
26.2 |
1592 |
1625 |
2800 |
2 |
85 |
PF |
subhumid dry |
23.4 |
29 |
26.2 |
1592 |
1625 |
2800 |
8 |
86 |
SF |
subhumid dry |
22.7 |
28.4 |
25.6 |
1698 |
1625 |
2800 |
26 |
87 |
SSF |
Humid |
20.2 |
26.7 |
23.5 |
2613 |
1650 |
2500 |
443 |
88 |
SSF |
Humid |
20.2 |
26.7 |
23.5 |
2613 |
1650 |
2500 |
387 |
89 |
SSF |
Humid |
20.2 |
26.7 |
23.5 |
2613 |
1650 |
2500 |
400 |
90 |
SSF |
Humid |
19.7 |
26.7 |
23.2 |
2479 |
1675 |
2500 |
592 |
91 |
SYSF |
subhumid dry |
23.8 |
28.7 |
26.3 |
1814 |
1650 |
2800 |
33 |
92 |
SYSF |
subhumid dry |
23.8 |
29.2 |
26.5 |
1741 |
1650 |
2800 |
37 |
93 |
PF |
subhumid dry |
23.8 |
29.2 |
26.5 |
1741 |
1650 |
2800 |
41 |
94 |
SYSF |
subhumid dry |
23.2 |
28.7 |
26.0 |
1814 |
1650 |
2800 |
27 |
95 |
SYSF |
subhumid dry |
24.2 |
29.2 |
26.7 |
1513 |
1700 |
2800 |
8 |
96 |
MSF |
subhumid dry |
24.2 |
29.2 |
26.7 |
1513 |
1700 |
2800 |
14 |
97 |
PF |
subhumid dry |
24.3 |
29.5 |
26.9 |
1387 |
1625 |
2900 |
2 |
98 |
SSF |
ecotone
(meso-hygrophilic) |
22.3 |
28.3 |
25.3 |
1965 |
1600 |
2700 |
294 |
99 |
SSF |
ecotone
(meso-hygrophilic) |
22.3 |
28.3 |
25.3 |
1965 |
1600 |
2700 |
257 |
100 |
SYSF |
subhumid dry |
23.4 |
31.5 |
27.5 |
1748 |
1775 |
2700 |
101 |
101 |
PF |
subhumid dry |
23.4 |
31.5 |
27.5 |
1748 |
1775 |
2700 |
62 |
102 |
SSF |
subhumid humid |
21.3 |
29.1 |
25.2 |
1749 |
1725 |
2700 |
180 |
103 |
SSF |
subhumid humid |
21.3 |
29.1 |
25.2 |
1749 |
1725 |
2700 |
130 |
104 |
SSF |
ecotone
(meso-hygrophilic) |
20.7 |
28.3 |
24.5 |
2319 |
1675 |
2600 |
611 |
105 |
SSF |
ecotone
(meso-hygrophilic) |
21 |
28.4 |
24.7 |
3443 |
1575 |
2400 |
447 |
106 |
SSF |
Humid |
19.6 |
26.6 |
23.1 |
2507 |
1650 |
2500 |
500 |
107 |
SYSF |
ecotone
(meso-hygrophilic) |
22.7 |
28.4 |
25.6 |
1698 |
1625 |
2800 |
259 |
108 |
SSF |
subhumid humid |
21.3 |
29 |
25.2 |
1757 |
1575 |
2700 |
363 |
109 |
SSF |
subhumid humid |
23.4 |
30.7 |
27.1 |
1708 |
1750 |
2700 |
20 |
110 |
SSF |
subhumid humid |
23.4 |
30.7 |
27.1 |
1708 |
1750 |
2700 |
21 |
111 |
SYSF |
subhumid dry |
23.4 |
30.7 |
27.1 |
1708 |
1750 |
2700 |
34 |
112 |
PF |
subhumid dry |
23.4 |
30.7 |
27.1 |
1708 |
1750 |
2700 |
42 |
113 |
PF |
subhumid dry |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
58 |
114 |
FVTD |
subhumidhumid |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
35 |
115 |
FVTD |
subhumid humid |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
33 |
116 |
FVTD |
ecotone
(xero-mesophil) |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
109 |
117 |
FVTD |
subhumid humid |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
80 |
118 |
SYSF |
ecotone
(xero-mesophil) |
22.4 |
29.7 |
26.1 |
1738 |
1750 |
2700 |
195 |
119 |
FVTD |
subhumid humid |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
39 |
120 |
SSF |
subhumid humid |
23 |
30.2 |
26.6 |
1705 |
1750 |
2700 |
45 |
S = station; SE = Stage of evolution; Tmin = Minimum temperatures (˚C); Tmax = Maximum temperatures (˚C); Taverage = Average temperatures (˚C); Rain = Rainfall; Eva = Evapotranspiration; Ins = Insolation; H = Height; SE = Stage of evolution: (S.F): Shrub formation/(M.S.F): Mature shrub formation/(P.F): Presylvatic formation/(S.Y.S.F): Structured young sylvatic formation/(S.S.F): Secondary sylvatic formation/(E.C.D): Degraded back-mangrove plant eco-unit/(FVTD): Very degraded plant formation.