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Identificacion de especies de arboles en peligro o amenazadas de Costa Rica basada en la anatomia de la madera y fluorescencia.

Identification of endangered or threatened Costa Rican tree species by wood anatomy and fluorescence activity

For its size (51 000[km.sup.2]) Costa Rica is one of the most biodiverse countries in the world. Of its 10 000 plant taxa, 1 300 are endemic and more than 25% of these are considered rare (Burger 1980). Nonetheless, intensive deforestation began at the start of the twentieth century, and reached levels in excess of 50 000ha per year in the 1970s (Alvarez 1986). Natural forest decreased to only 24.4% by 1987 (Sanchez et al. 2001).

Tree timber species receive the highest degree of protection, and two actions are used to protect them: the promulgation of laws and government decrees to stop the cutting of 18 commercial species (Costa Rica 1997) protected by international conventions such as CITES, and actions limiting harvesting in certain areas of the country (Costa Rica 1996a, 1996b). A number of important timber-producing species have been identified by some experts as threatened species due to their rarity (Jimenez 1999). Identification of these timber-producing species using floral and tree characteristics has been widely described (Jimenez 1999). In contrast, identification of these species based on their wood anatomy is limited to only a few species: examples are the species included in CITES Appendices I and II. The CITES Identification Guide-Tropical Wood (CITES 2002) has a general identification key with illustrations of the transverse sections of each species. The book is intended for non-experts (Gasson et al. 2010) and its use is limited. More recently, the International Association of Wood Anatomists (IAWA) published several wood descriptions of tree timbers included in CITES (Gasson 2011, Gasson et al. 2011).

Recently, wood anatomical identification has been accompanied by other analyses that together with special methods, can achieve precise identification that are useful in identification of endangered or threatened species. Some of these techniques are relatively easy to implement, for example, wood density, surface fluorescence, or fluorescence of water or ethanol extracts. (Miller & Wiemann 2006, Guzman et al. 2008, Wiemann & Ruffinatto 2012). Studies of anatomical features accompanied with complex statistical procedures, such as the use of multivariate analysis, helps to identify CITES species (Gasson et al. 2010, MacLanchlan & Gasson 2010).

The objective of the present study was to describe and compare wood anatomy, fluorescence, and density of endangered or threatened species of Costa Rica, and to use these characteristics to build an identification key. The key will allow monitoring of the use and commercialization of these tree species (endangered or threatened) in Costa Rica, and perhaps will be also useful to other countries in the region.


Wood samples and permanent slides: A total 45 timber species growing in Costa Rica were analyzed from March to December of 2010, 22 of which are considered as endangered species by the Costa Rican Government (Costa Rica 1996a and 1996b, Costa Rica 1997) (Table 1). Another 23 timber species studied are considered to be in threat of extinction in this country (Jimenez 1999) (Table 2). Wood samples of all the species were obtained

from the Forest Products Laboratory (FPL), Wisconsin-USA collection (MADw and SJRw) and the Instituto Tecnologico de Costa Rica (ITCR) collection (TECw). Their sample numbers are detailed in tables 1 and 2. In some cases only one sample and slide was available, so it was necessary to prepare additional permanent slides of these species. To do this, a block (1[cm.sup.3]) was obtained from a wood sample and was softened in hot water. Tangential, radial and transverse sections were cut (12-15Lim thick). These sections were stained with safranin and dehydrated with a series of alcohol (5 minutes each in 50, 70 and 95%); finally, sections were rinsed and mounted on microscope slides. Furthermore, a small piece was cut from each wood block to prepare macerated wood using Franklin's method (Ruzin 1999).

Wood anatomical description: The IAWA list (IAWA 1989) was used as the basis for choosing identification characteristics, with some modifications to allow for increased accuracy and subsequent species level separation. The quantitative anatomical features that were measured were: length and diameter of the fibers, lumen diameter, cell wall thickness, vessel length, diameter and frequency of pores, solitary pore frequency, diameter of intervessel pits, and height and width of rays. Fiber dimensions and vessel lengths were measured on macerated wood. Permanent slides were used for measurement of the other anatomical characters. Qualitative anatomical features were also determined using the IAWA List as a guide (IAWA 1989).

Additional information: Other important information about endangered or threatened species used was: endemic category if the species grows only in Costa Rica, wood density, traditional uses of species, and fluorescence. Wood density (weight/volume) was measured in the air-dry condition. The four traditional uses considered were light or heavy construction, flooring, furniture, and handicrafts. Heartwood fluorescence was observed directly the surface, in water extract, and in ethanol extract, as described in the IAWA List (IAWA 1989).

Surface fluorescence was determined in a darkroom from freshly prepared (planed or scraped) transverse and/or longitudinal surfaces exposed to a low intensity, long wave ultraviolet light (around 365nm). The specimens were recorded as either fluorescente (noting color and intensity) or not fluorescente. Only specimens which exhibited a definite yellow, green, orange or blue fluorescence were recorded as fluorescente. For water and ethanol extracts, color and intensity of fluorescence were scored. Froth tests were conducted and were scored as positive (high intensity) if one minute after shaking vigorously, froth was present and covered the entire surface of the solution, negative if all froth had disappeared, and variable if froth was still present around the edge of the test tube but did not extend over the entire surface of the liquid column.


General aspects: 22% (ten species) were Papilionaceae, 18% (eight species) were Caesalpiniaceae, 13% (six species) were Meliaceae, 9% (four species) were Podocarpaceae, 7% (three species) were Lecythidaceae, 4% (two species) each were Caryocaraceae or Humiriaceae; other families represented the 27% (12 species) of endangered or threatened timber species endemic to Costa Rica. Most of these species are used in heavy or light construction or handicrafts; however, species in the Meliaceae are utilized for furniture manufacturing. High wood density was a characteristic of most of the species, and the main use of the lumber was construction (Table 1 and 2).

Fluorescence test: Surface fluorescence was positive in 11 species. This represented 24% of the total number of species. Fluorescence in Caryocar costaricense, Copaifera aromatica, Astronium graveolens, Mora oleifera and Myroxylon balsamum was green, greenish, or weak green, whereas in Dussia macroprophyllata, Lecythis ampla, Hymenolobium mesoamericanum, Peltogyne purpurea and Tachigali versicolor it was yellow, yellowish or weak yellow (Table 3).

Water extract fluoresced in only nine species (Table 3). The fluorescence was yellow in Copaifera species, H. mesoamericanum and P. purpurea and green in Cordia gerascanthus, A. graveolens, M. balsamum and Platymiscium species (Table 3). C. gerascanthus and Platymiscium species did not show surface fluorescence, but they did show water extract fluorescence (Table 3).

Ethanol extract fluorescence was found in 31 of the species. The fluorescence was greenish blue or yellow, weak or light green, purple, yellowish or bluish (Table 3). Several species (Cedrela odorata, Cedrela fissilis, Cedrela tonduzii, Couratari guianensis, Couratari scottmorii, Dalbergia retusa, Dipteryx panamensis, Guaiacum sanctum, Oreomunnea pterocarpa, Parkia pendula, Prioria copaifera, Qualea paraensis, Swietenia humilis, Swietenia macrophylla, Tabebuia guayacan and Vantanea barborii) had positive ethanol extract fluorescence, but negative surface or water extract fluorescence (Table 3).

General features of endangered or threatened species: Porosity: Diffuse porosity was common in endangered or threatened species, being found in 39 species. Three species were both diffuse and semi-ring porous (C. odorata, C. fissilis and C. gerascanthus) and two species were semi-ring porous (Cedrela salvadorensis and C. tonduzii). Sideroxylon capari was unique with a radial or diagonal pore pattern (Fig. 1a). Almost all of the species had solitary pores and pore multiples. However, the percentage of solitary pores was higher than 85% in G. sanctum and V. barbourii, so these species can be classified as pores exclusively solitary. Higher frequency of multiple pores (>15 pores/[mm.sub.2]) was measured in three species: C. guianensis, P. purpurea and T. guayacan. Pores frequency was the highest in Guaiacum sanctum, and M. balsamo. Lower pore frequencies (<2 pores/[mm.sup.2]) were observed in C. tonduzii, D. retusa, D. macrophylla, H. mesoamericanum, L. ampla and Platymiscium curuense.

Vessel lengths for these species varied from 93 to 1 160mm (Table 3). Vessel length was shortest in G. sanctum and longest in H. guianensis, M. guianensis and V. barbourii. Small diameter vessels (from 50-100mm) were found G. sanctum, M. guianensis, M. balsamo, S. capari and T. versicolor. Large diameter vessels were found only in O. pterocarpa. Vessel diameters from 100-300 mm were found in the other species.

Simple perforation plates were found in almost all species (Table 3). However, H. guianensis (with 10-20 bars), M. guianensis (with <=10 bars) and V. barbourii (with <=20 bars) had scalariform perforations.

Tyloses, deposits or gum were found in the vessels of almost all of the angiosperm species (Table 3). Gum was the most common substance in vessel lumina. Nevertheless, neither substance was observed in three of the angiosperm species: D. macroprophyllata, Sclerolobium costaricense and V. barbourii.

All angiosperm species had alternate polygonal pits. Their diameter was minute (<4[micro]m) in 14 species and large (>=10[micro]m) in five species. Pits in the other species were from 4 to 10[im in diameter (Table 3).

Vestured pits were found in all species of Fabaceae (Caesalpixiaceae, Mimosaceae, Papilionaceae) and in V. barbourii (Table 3). Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape throughout the ray cell, were common. They were found in almost all species. Other pits shapes were found in C. guianensis (Table 3). Vessel-ray pits were restricted to marginal rows in C. costaricense. Vessel-ray pits with much reduced borders to apparently simple, with pits rounded or angular and horizontal (scalariform, gashlike) to vertical (palisade) were found in A. graveolens and Caryodaphnopsis. burgeri (Fig. 1c), C. guianensis, and O. pterocarpa. Vesselray pits were restricted to marginal rows in V. barbourii. L. ampla had vessel-ray pits with much reduced borders to apparently simple: the pits were horizontal (scalariform, gash-like) to vertical (palisade).

Fibers: as expected, gymnosperm species tracheids were longer than most of the angiosperm fibers, being almost 2mm length. However, the fiber length of V. barbourii was the longest of any species at 2.45mm. Fiber length varied from 1.50 to 1.95mm in C. costaricense, C. burgeri, C. gerascanthus, D. macroprophyllata, H. mesoamericanum, H. guianensis, M. guianensis and M. balsamum. The shortest fiber lengths were found in G. sanctum, although C. odorata, C. salvadorensis, D. retusa, O. pterocarpa and S. capari also had short fibers (Table 4). Fiber lumen diameter was widest in D. macroprophyllata, O. pterocarpa, H. guianensis, P. macrostachys and V. barbourii, and narrowest in T. guayacan. Narrow fibers (<15[micro]m) were also found in C. fissilis, C. guianensis, G. sanctum, Paramachaerium gruberi and S. costaricense. The lumen diameter varied from 15 to 55[micro]m in other species (Table 4). Fibers with very thick-walls (>7[micro]m) were found in D. retusa, H. guianensis, M. guianensis, M. balsamum, S. capari and V. barbourii. Thin-walled fibers were found in species of Cedrela and in C. guianensis, P. copaifera, S. costaricense and S. humilis. Cell wall thickness of other endangered and threatened species varied from 3 to 7Lm (Table 4). Septate fibers were observed only in A. graveolens, G. sanctum and Swietenia species (Table 4). In the angiosperms, fibers with simple to minutely bordered pits were most common in H. guianensis, L. ampla and V. barbourii. Distinctly bordered pits were common in both radial and tangential fiber walls. The tracheids of species of the Podocarpaceae had, as expected, bordered pits in radial walls. Many of the angiosperm species had storied fibers (Table 4). Crystals were only found in the fibers of P. gruberi.

Ray parenchyma: Ray height over 1mm was observed in five species (C. costaricense, C. gerascanthus, M. guianensis, P. purpurea and Q. paraensis), representing 11% of the species. Rays were exclusively uniseriate in eight species (Fig. 1b): four Podocarpaceae (Fig. 1d and 1e) and four angiosperms (G. sanctum and Platymiscium species). Rays 1-3 cell in width were the most common, represented by 23 species (51% of total species) (Table 4). Large rays (over 8 cells in width) were observed in eight species (18% of total species) (Table 4). The rays of C. aromatica, C. gerascanthus and P. purpurea were 4-10 seriate. Ray frequency varied from 2 to 20 rays per mm (Table 4). The lowest frequencies were found in C. burgeri and L. ampla and the highest ones (>14 rays/mm) were in Cynometra hemitomophylla, G. sanctum and H. guianensis. Rays composed mostly of procumbent cells were observed in 17 species (38% of total species) and heterogeneous rays were in 19 species (42% of total species). Nine species had both homogeneous and heterogeneous rays (Table 4). Storied rays were found in 14 species (Table 4). Two heights were found in D. macrophylla, P. purpurea and Swietenia species. Crystals were observed in ray cells of 27 species (60% of total species); in 11 species (24%) crystals were found in the marginal ray cells only (Table 4). Silica bodies were present in Anthodiscus chocoensis, Couratari species, L. ampla, Q. paraensis and T. versicolor. Sheath cells were only observed in C. gerascanthus.

Axial parenchyma: Apotraqueal parenchyma was present in 21 species (48% of total species) (Table 5). Parenchyma diffuse and diffuse-in-aggregates was seen in D. retusa, G. sanctum, M. guianensis and S. capari, and it was the only parenchyma type in C. costaricense. Almost all the species of angiosperms had paratracheal parenchyma (Table 5). Scanty paratracheal parenchyma was present in sixteen species, and vasicentric parenchyma was found in A. graveolens, C. odorata, C. fissilis, C. tonduzii, C. gerascanthus and T. versicolor. Three or more different paratracheal types were observed in 17 species (38% of total species). Aliform and confluent parenchyma was very common in several species. Banded parenchyma was observed in 30 species (68 of total species; 73% of angiosperm species). Parenchyma marginal or in wide bands (more than 3 cells wide) were the most common banded parenchyma (Table 5). Axial parenchyma was reticulate in Couratari species, C. hemitomophylla, L. ampla and O. pterocarpa. Axial parenchyma was storied in 13 species (Table 5). Fusiform cells were found in Platymiscium species, Paramachaerium grugeri, M. balsamo, P. copaifera, T. guayacan and T. versicolor. Crystals were present in chambered axial parenchyma in almost all gymnosperm species (Table 5). They were found enlarged in P. gruberi. A. chocoensis, A. graveolens, C. gerascanthus and Podocarpus species, T. guayacan did not have any crystals. Silica bodies were present in Couratari species.

Other anatomical features: some species had other distinctive anatomical features, which are detailed in table 5. They could be used to facilitate wood identification. For example, radial or axial canals are found in A. chocoensis, C. costaricense, C. aromatica and P. copaifera. Traumatic canals were observed in C. camibar, H. guianensis and Platymiscium pinnatum. Pores with two distinct diameters are found in D. retusa and Q. paraensis. C. gerascanthus was a unique species with sheath cell in the rays. Finally, helical thickenings were observed in some vessels elements of S. costaricense.

Specific species: Major differences among species groups are included in the next few paragraphs.

Meliaceae species: Two genera of Meliaceae were analyzed: four species of Cedrela (C. odorata, C. salvadorensis, C. fissilis and C. tonduzii), and two species of Swietenia (S. macrophylla and S. humilis). The wood anatomy of these species, especially C. odorata, C. fissilis and S. macrophylla, has been described by several authors (Panshin 1933, White & Gasson 2008). Four species of Meliaceae (C. salvadorensis, C. fissilis, S. macrophylla and S. humilis) are considered as endangered and their cutting has been prohibited in the natural forest in Costa Rica (Costa Rica 1996a). Felling of the other species, C. odorata and C. tonduzii, is permitted. Therefore, it is important to have a method to separate the species. The species of Swietenia are easily separated from species of Cedrela by anatomical features such as storied rays (Fig. 2g, h, i) and diffuse porosity in Swietenia but not in Cedrela (Fig. 2a-2i).

Platymiscium species: some differences were found among species of Platymiscium include presence of traumatic canals in Platymiscium parviflorum (Fig. 3a) but not in the other species (Fig. 3b). Platymiscium pinnatum var. polystachyum had irregularly storied rays (Fig. 4c) whereas in the other species storing was well-defined (Fig. 4a, 4b and 4d). The parenchyma was paratracheal in P. pinnatum var. polystachyum but not in other Platymiscium species. It was scanty, unilateral paratracheal, winged-aliform in P. pinnatum var polystachyum (Fig. 3b), but lozenge-aliform in the other species. We also found marginal parenchyma in P. pinnatum var polystachyum (Fig. 3b).

Couratari species: The two Couratari species can be distinguished by several differences. C. scottmorii has narrow reticulate parenchyma bands that are two cells wide (Fig. 5b), whereas reticulate bands in C. guianensis are up to four cells wide (Fig. 5a). Rays are 1-3 seriate in C. scottmorii (Fig. 5d), but up to 5-seriate in C. guianensis (Fig. 5c). The frequency of silica bodies is different among species, with, the highest frequency observed in C. guianensis. The vessels-ray pits in C. scottmorii have much reduced borders and the pits are rounded or angular, unlike those of C. guianensis which are horizontal (scalariform, gash-like) to vertical (palisade).

Copaifera species: The two Copaifera species differed mainly by paratracheal parenchyma and ray dimensions. Paratracheal parenchyma is more abundant in C. aromatica than in C. camibar. It is vasicentric 2-3 cells in width in Copaifetra aromatica (Fig. 6a), but scanty paratracheal or vasicentric 1-2 cells in C. camibar (Fig. 6b). The rays are 1- 3 cells wide and high in C. camibar (Fig. 6d), but were commonly 4-10 seriate and low in C. aromatica (Fig. 6c). There is no difference in the resin canals of the species.

Podocarpaceae species: We looked at two genera: three species of Podocarpus (P. costaricensis, P. guatemalensis and P. macrostachys), and one species of Prumnopitys (P. standleyi). Axial parenchyma is present in P. macrostachys and P. costaricensis (Fig. 7a-7b) and is scanty in P. guatemalensis and Prumnopitys standleyi (Fig. 7c-7d). The highest proportion of axial parenchyma was observed in P. macrostachys (Fig. 7a), it was rare to moderately abundant in P. costaricensis (Fig. 7b) and scanty in P. guatemalensis (Fig. 7d). Another important difference between Podocarpus and Prumnopitys is ray height. Rays were highest in P. macrostachys (5 to 10 cells) (Fig. 8a), but they were only 2-4 cells high in P. costaricensis (Fig. 8b). Ray frequency is highest in P. guatemalensis (Fig. 7c). The rays of Prumnopitys standleyi are similar in shape and frequency to the rays of P. macrostachys. No differences were seen among species in crossfield pit apertures.


Forty-five Costa Rican timber species are considered endangered or threatened. The Costa Rican government has decreed that 51% of these species are endangered and 49% of them are considered to be threatened. The CITES Appendices includes only eight of these species (S. humilis, S. macrophylla, D. panamensis, G. sanctum, C. costaricense, O. pterocarpa, C. odorata and D. retusa) (CITES 2002). All gymnosperms growing in the Costa Rican tropics are cataloged as endangered or threatened. However, most of the gymnosperm species in tropical areas around the world are in same situation; they are in decline or are restricted to isolated areas (Farjon et al. 1993).

The development of identification keys, like the one presented in this study, requires knowledge of wood characteristics and structure, as described in IAWA standards (IAWA 1989). Fluorescence provides a quick test for wood identification and it had been utilized by several authors for species separation. For example, Miller & Wiemann (2006) found differences in water and ethanol fluorescence between Dalbergia nigra and D. spruceana. Guzman et al. (2008) found fluorescence species in the Anacardiaceae, Leguminose and Rubiaceae from Brazil and South Africa. Fluorescence is one of the important distinguishing characteristics of endangered or threatened

timber species, although surface and water extract fluorescence can separate only six and eight timber species, respectively. Although, ethanol extract fluorescence was present in many timber species, the color of fluorescence was sometimes the same. However, three species were atypical in ethanol extract fluorescence color, the purple color found in G. sanctum, yellow in P. purpurea and bluish in T. versicolor, make them easy to identify. Guzman et al. (2008) established that for Mexican timber identification it is necessary to use mixture of fluorescence tests and other characteristics, such as color or anatomical features. Likewise, identification of the endangered or threatened timber species from Costa Rica also requires anatomical studies. Species of Cedrela are easily separated from each other based on pore arrangement and axial parenchyma types. According to White & Gasson (2008), C. odo rata is more ring-porous with more aliform parenchyma than C. fissilis. Cedrela tonduzii is similar to C. odorata and C. fissilis, although C. tonduzii is less ring porous. Aliform parenchyma is well defined in C. tonduzii, but not in C. odorata, C. salvadorensis or C. fissilis. C. odorata is considered to be easy to identify by its reddish color and distinct odor. Another important difference among C. tonduzii and other Cedrela species is that C. tonduzii has lower wood density than the other four species. C. salvadorensis has distinctive anatomical features that facilitate its identification; its rays are larger, commonly 4-10 seriate, than the rays of other Cedrela species, which are 1-3 seriate in width. Furthermore, its rays are heterogeneous and homogenous, but the rays of the other Cedrela species are not. However, Bonilla et al. (2004) reported that C. salvadorensis has 1-3 seriate rays, similar to those of other Cedrela. Therefore, we attribute the wider rays that we found in C. salvadorensis in Costa Rica to regional differences.

In species of Platymiscium, traumatic canals were seen in P. parviflorum, but not in other species. Traumatic canals have not been previously reported in Platymiscium (Espinoza & Leon 2002). Storied rays have been reported for most Platymiscium, for example P. lasiucarpium, P. duckei, P. pinnatum and P. yucatanum (Perez 1993, Espinoza & Leon 2002). However, Detienne & Jacquet (1983) reported that rays are irregularly storied in P. ulei, similar to P. pinnatum var polystachyum.

The differences found in Couratari species agreed with previous research, which has also reported differences. For example Leon (2008) separated C. guianensis from C. multiflora by ray width, and Richter (1982) maintained that parenchyma distribution as well as type and configuration of inorganic contents can be employed for separating species of Lecythidaceae.

Canessa (1989) agreed with our results in Copaifera species; he found that the axial parenchyma in C. camibar was different than that of C. officinalis and C. pubiflora. Parenchyma is reported to be vasicentric, aliform to confluent, and aliform of the lozenge type in these species (Melandri & Espinoza de Pernia 2009). The ray dimensions and axial parenchyma of some Copaifera species are characterized by high anatomy variation (Regina et al. 2002), and our results confirm that.

On the other hand, Patel (1967) found similar results in Podocarpaceae species when he evaluated axial parenchyma. They mentioned that numerous species of the genus Podocarpus are characterized by diffuse axial parenchyma and a considerable variation of trached crossfield pit apertures (size, form, number per cross-field). Scanty axial parenchyma in P. guatemalensis can be used for separating this species from other Podocarpaceae in Costa Rica. This is in agreements with P. spicatus growing in New Zealand, in which the lack of axial parenchyma is the main feature to separating it from other Podocarpus species. Abundant axial parenchyma also separates P. macrostachys from P. costaricensis and P. guatemalensis. Patel (1967) agreed with this result: he found that P. dacrydioides can be separated from P. totara, P. hallii and P. acutifolius by the abundance of axial parenchyma. Bauch et al. (2006) mentioned that P. costarricensis has more axial parenchyma than the commercially important species P. salignus growing in Chile and Argentina.

Many of the Costa Rican species included in this study, are also present in other tropical regions, and our results, both as wood descriptions and the identification key, and are applicable to the wider region. Species conservation is a goal for many countries, especially in countries where deforestation of natural forests has increased in the last few years. There is a strong interest to protect timber species which have been over-exploited for many years. This identification key and wood descriptions will assist in the protection of species categorized as endangered or threatened, and will promote reliable conservation plans.

Key for identification

The following identification key was developed to distinguish among
the 45 timber species considered to be endangered or threatened in
Costa Rica. This key use microscopic wood features as well and
fluorescence and wood density.

1.      Gymnosperm                                                 3
2.      Angiosperm                                                 6
3a.     Parenchyma present and rays 2-4                            4
          cells in height
3b.     Parenchyma abundant sparse and                             5
          rays 2-10 cells in height
4a.     Parenchyma abundant                  Podocarpus macrostachys
4b.     Parenchyma scanty                   Podocarpus guatemalensis
5a.     Parenchyma sparse                   Podocarpus costaricensis
5b.     Axial Parenchyma absent                Prumnopitys standleyi
6a.     Negative fluorescence                                      7
6b.     Positive fluorescence                                      14
7a.     Pores solitarys and in groups                              8
7b.     Pores solitary or pores in                                 13
          dendritic or diagonal and/or
          radial pattern.
8a.     Scalariform perforations; ray          Minquartia guianensis
          height over 1 mm, axial
          parenchyma diffuse and
          paratracheal scanty;
          vasicentric tracheids
8b.     Simple perforations                                        9
9a.     Semi-ring porous, parenchyma           Cedrela salvadorensis
          diffuse, paratracheal scanty,
          vasicentric and in marginal
          or in seemingly marginal
9b.     Diffuse porous                                             10
10a.    Rays not storied                                           11
10b.    Ray storied, axial parenchyma         Paramachaerium gruberi
          apotraqueal diffuse, scanty,
          lozenge-aliform, winged-
          aliform, and banded in
          narrow bands or lines up to
          three cells wide
11a.    Ray 1-3 cells in width                                    12
11b.    Axial parenchyma apotracheal           Carydaphnopsis burger
          diffuse, scanty, vasicentric,
          and in narrow bands or lines
          up to three cells wide, with
          3-5 or 5-8 cells per
          parenchyma strand
12a.    Paratraqueal parenchyma with          Anthodiscus chocoensis
          over eight cells per
          parenchyma strand
12b.    Parenchyma in marginal or in
          seemingly marginal bands,
          narrow banded or reticulate;
          rays and axial elements
          irregularly storied, ray          Cynometra hemitomophylla
          frequency high
13a.    Pores solitary, scalariform          Humiriastrum guianensis
          perforation plates, axial
          parenchyma vasicentric,
          lozange-aliform, winged-
          aliform and confluent and
          traumatic canals
13b.    Vessels in dendritic or                   Sideroxylon capari
          diagonal and/or radial
          pattern, rays heterogeneous,
          1-3 seriate and parenchyma
          diffuse and diffuse-in-
14a.    Fluorescence in surface only                               15
14b.    Fluorescence in ethanol                                    16
          extract only
14c.    Fluorescence in surface and                                31
          ethanol extract
14d.    Fluorescence in water and                                  35
          ethanol extract
14e.    Fluorescence in surface, water                             37
          and ethanol extract.
15a.    Parenchyma vasicentric,                        Mora oleifera
          lozenge-aliform, winged-
          aliform, confluent and in
          marginal or in seemingly
          marginal bands, with 3-4 and
          5-8 cells per parenchyma
15b.    Ray height over 1 mm,                  Caryocar costaricense
          parenchyma diffuse,
          paratracheal scanty and in
          marginal or in seemingly
          marginal bands and radial
16a.    Pores solitarys                                            17
16b.    Pores solitarys and in groups                              18
17a.    Ray frequency high, fibers                  Guaiacum sanctum
          septate and storied,
          apotraqueal parenchyma
          diffuse and diffuse-in-
17b.    Perforation plates scalariform,           Vantanea barbourii
          axial parenchyma diffuse,
          scanty partracheal and
18a.    Semi-ring porous                                           19
18b.    Diffuse porous                                             21
19a.    Semi-ring porous, parenchyma                 Cedrela odorata
          vasicentric, rays
          heterogeneous ray and 1-3
          seriates in width
19b.    Semi-ring to diffuse porous                                20
20a.    Parenchyma aliform, rays                    Cedrela fissilis
          heterogeneous and 1-3 seriate
20b.    Parenchyma diffuse,                         Cedrela tonduzii
          paratracheal scanty and
          vasicentric and in marginal
          or in seemingly marginal
21a.    Rays uniseriate, pits vestured,                 Sclerolobium
          axial parenchyma vasicentric,                 costaricense
          vessels with helical
21b.    Rays multiseriate                                          22
22a.    Rays 1-10 cells wide,                  Swietenia macrophylla
          irregularly storied or                or Swietenia humilis
          storied, parenchyma
          apotracheal diffuse, scanty
          paratracheal and in marginal
          or in seemingly marginal
22b.    Rays 1 to 3 seriate                                        23
23a.    Axial canals, parenchyma                   Prioria copaifera
          vasicentric, lozange-aliform;
          confluent, and wide band
          banded with 2 and 3-4 cells
          per parenchyma strand
23b.    Axial canals absent                                        24
24a.    Rays storied                                               25
24b.    Ray not storied                                            27
25a.    Ray homogeneous                                            26
25b.    Ray heterogeneous, 1-3 cells
          wide, parenchyma apotracheal
          diffuse and diffuse-in-
          aggregates, scanty,
          vasicentric, in narrow bands              Dalbergia retusa
          or lines up to three cells
          wide and in marginal or in
          seemingly marginal bands
26a.    Parenchyma lozenge-aliform,              Dipteryx panamensis
          winged-aliform and unilateral
          parenchyma, and prismatic
          crystals in chambered cells
26b.    Parenchyma vasicentric,                    Tabebuia guayacan
          lozenge-aliform, unilateral,
          and in marginal or in
          seemingly marginal bands
27a.    Ray heterogeneous                                          28
27b.    Ray homogeneous, parenchyma                   Parkia pendula
          lozenge-aliform and confluent
          with 3-5 cells per parenchyma
28a.    Ray 1-3 seriate                                            29
28b.    Ray 1-10 seriate, fibers pits           Couratari guianensis
          common on both radial and
          tangential walls, two distinct
          sizes or types in the same ray
          cell; parenchyma bands more
          than three cells wide,
          reticulate and in marinal or
          in seemingly marginal bands.
29a.    Apotracheal parenchyma absent .                            30
29b.    Apotracheal parenchyma diffuse,             Qualea paraensis
          vasicentric, lozenge-aliform
          winged-aliform and confluent,
          vessels with 2 distinct
          diameters, ray height over 1mm
30a.    Rays homogeneous and                    Couratari scottmorii
          heterogeneous, parenchyma in
          bands 2 cells wide, reticulate
          and in marginal or in
          seemingly marginal bands,
          reticulate bands 2 cells in
30b.    Parenchyma in marginal or in           Oreomunnea pterocarpa
          seemingly marginal bands,
          narrow banded or reticulate
          with 5-8 cells per
          parenchyma strand
31 a.   Ray storied                                                32
31b     Rays not storied                                           34
32a.    Rays 1 to 3 seriate                                        33
32b.    Larger rays commonly 4-10                 Peltogyne purpurea
          seriate, ray height over 1mm,
          pits vestured; parenchyma
          lozenge-aliform, confluent,
          unilateral, in marginal or in
          seemingly marginal bands
33a.    Parenchyma confluent and             Dussia macroprophyllata
          banded, more than three
          cells wide
33b.    Parenchyma scanty paratracheal,           Myroxylon balsamum
          confluent vasicentric and
34a.    Pits vestured, axial                    Tachigali versicolor
          paratracheal scanty and
          vasicentric with 2 and 3-5
          cells per parenchyma strand
34b.    Fibers with distinctly bordered               Lecythis ampla
          pits, parenchyma diffuse and
          in bands miore than three
          cells wide, reticulate and in
          marginal or in seemingly
          marginal bands
35a.    Rays exclusively uniseriate;                               36
          pits vestured
35b.    Rays 4-10 seriate, semi-ring or          Cordia gerascanthus
          diffuse porous, parenchyma
          apotracheal difuse, rays
          heterogeneous with sheath
          cells, higher than 1mm
36a.    Rays irregularly storied,              Platymiscium pinnatum
          parenchyma scanty, unilateral             var polystachyum
          paratracheal, winged-aliform,
          marginal banded
36b.    Rays storied, parenchyma               Platymiscium pinnatum
          lozenge-aliform with long
          wings, and confluent
36c.    Parenchyma lozenge-aliform             Platymiscium curuense
          and confluent
36d.    Parenchyma lozenge-aliform and                  Platymiscium
          confluent, traumatic canals                    parviflorum
          sometimes present
37a.    Axial canals present                                       38
37b.    Axial canals absent                                        39

38a.    Parenchyma vasicentric,                    Copaifera camibar
          lozenge-aliform, confluent
          and in seemingly marginal
38b.    Parenchyma vasicentric 1-2               Copaifera aromatica
          cells in width, and in
          seemingly marginal bands
39a.    Fibers septate; parenchyma              Astronium graveolens
          diffuse, paratracheal scanty;
          vasicentric and in marginal or
          in seemingly marginal bands
39b.    Ray 1-10 cells in width,                        Hymenolobium
          storied, axial parenchyma                   mesoamericanum
          lozenge-aliform, confluent,
          vasicentric and in bands
          more than three cells wide


The authors wish to thank The Council for International Exchange of Scholars of the Department of Scholar and Professional Programs of USA, Premios Ford de Conservacion of Ford Motor Company, and Vicerrectoria de Investigacion y Extension del Instituto Tecnologico de Costa Rica (ITCR) for financial support of this research.


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46: 59-71.

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Miller, R. & M. Wiemann. 2006. Separation of Dalbergia nigra from Dalbergia spruceana. Department of Agriculture, Research Paper FPL-RP-632. Forest Service, Forest Products Laboratory, Madison, Wisconsin, USA.

Panshin, A.J. 1933. Comparative Anatomy of the Woods of the Meliaceae, Sub-Family Swietenioideae. Am. J. Bot. 20: 638-668.

Patel, R.N. 1967. Wood anatomy of Podocarpaceae indigenous to New Zealand. NZ. J. Bot. 5: 307-321.

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Regina, C., V. Angyalossy-Alfonso & L. Benetati. 2002. Anatomia comparada do lenho de Copaifera langsdorffii Desf. (Leguminosae-Caesalpinoideae) de floresta e cerradao. Rev. Bras. Bot. 24: 311-320.

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Ruzin, S.E. 1999. Plant microtechnique and Microscopy. Oxford University, Oxford, England.

Sanchez, G.A., R.C. Harris & D. Skole. 2001. Deforestation in Costa Rica: a quantitative analysis using remote sensing imagery. Biotropica 33: 378-384.

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Roger Moya1*, Michael C. Wiemann2 & Carlos Olivares3

1. Instituto Tecnologico de Costa Rica, Escuela de Ingenieria Forestal, Apartado 159-7050, Cartago, Costa Rica;

2. Forest Products Laboratory, USDA Forest Service, One Gifford Pinchot Drive Madison, Wisconsin 53726-2398, USA;

3. Instituto Tecnologico de Costa Rica, Escuela de Ingenieria Forestal, Apartado 159-7050, Cartago, Costa Rica;

* Corresponding author

Received 10-I-2012.

Corrected 20-X-2012.

Accepted 12-XI-2012.


Endangered timber species in Costa Rica (decreed by government)

Botanical Family   Species                Wood species   Endemic

Boraginaceae       Cordia                  Angiosperm      No
                     gerascanthus L.
Caesalpiniaceae    Sclerolobium            Angiosperm      Yes
                     Zamora & Poveda
Caryocaraceae      Anthodiscus             Angiosperm      Yes
                     chocoensis Prance
Lauraceae          Caryodaphnopsis         Angiosperm      Yes
                     burgeri N. Zamora
                     & Poveda
Lecythidaceae      Couratari               Angiosperm      Yes
                     scottmorii Prance
Meliaceae          Cedrela fissilis        Angiosperm      No
                   Cedrela                 Angiosperm      No
                   Swietenia humilis       Angiosperm      No
                   Swietenia               Angiosperm      No
                     macrophylla King
Mimosaceae         Parkia pendula          Angiosperm      No
Papilionaceae      Dipteryx                Angiosperm      No
                     (Pittier) Record &
                   Hymenolobium            Angiosperm      No
                     H.C. Lima
                   Myroxylon balsamum      Angiosperm      No
                     (L.) Harms
                   Paramachaerium          Angiosperm      Yes
                     gruberi Briz.
                   Platymiscium            Angiosperm      Yes
                     curuense N. Zamora
                     & Kiitgaard
                   Platymiscium            Angiosperm      No
                     parviflorum Benth.
                   Platymiscium            Angiosperm      Yes
                     pinnatum var.
                     (Jacq.) Dugand
                   Platymiscium            Angiosperm      No
                     pinnatum (Jacq.)
Podocarpaceae      Podocarpus              Gymnosperm      Yes
                     costaricensis de
                   Podocarpus              Gymnosperm      No
                   Podocarpus              Gymnosperm      No
                     macrostachys Parl.
Zygophyllaceae     Guaiacum sanctum        Angiosperm      No

Botanical Family   Species                Traditional
                                          used of wood

Boraginaceae       Cordia                   Fu & LC
                     gerascanthus L.
Caesalpiniaceae    Sclerolobium             HC & Fl
                     Zamora & Poveda
Caryocaraceae      Anthodiscus                 HC
                     chocoensis Prance
Lauraceae          Caryodaphnopsis             HC
                     burgeri N. Zamora
                     & Poveda
Lecythidaceae      Couratari                   HC
                     scottmorii Prance
Meliaceae          Cedrela fissilis          Fu & H
                   Cedrela                   Fu & H
                   Swietenia humilis         Fu & H
                   Swietenia                 Fu & H
                     macrophylla King
Mimosaceae         Parkia pendula              LC
Papilionaceae      Dipteryx                 HC & Fl
                     (Pittier) Record &
                   Hymenolobium                LC
                     H.C. Lima
                   Myroxylon balsamum       HC & Fl
                     (L.) Harms
                   Paramachaerium              HC
                     gruberi Briz.
                   Platymiscium            Fu, Fl & H
                     curuense N. Zamora
                     & Kiitgaard
                   Platymiscium            Fu, Fl & H
                     parviflorum Benth.
                   Platymiscium            Fu, Fl & H
                     pinnatum var.
                     (Jacq.) Dugand
                   Platymiscium            Fu, Fl & H
                     pinnatum (Jacq.)
Podocarpaceae      Podocarpus                  Fu
                     costaricensis de
                   Podocarpus                  Fu
                   Podocarpus                  Fu
                     macrostachys Parl.
Zygophyllaceae     Guaiacum sanctum        HC, Fl & H

Botanical Family   Species                Wood density *

Boraginaceae       Cordia                      0.72
                     gerascanthus L.
Caesalpiniaceae    Sclerolobium                0.74
                     Zamora & Poveda
Caryocaraceae      Anthodiscus                 0.88
                     chocoensis Prance
Lauraceae          Caryodaphnopsis             0.78
                     burgeri N. Zamora
                     & Poveda
Lecythidaceae      Couratari                   0.79
                     scottmorii Prance
Meliaceae          Cedrela fissilis            0.54
                   Cedrela                     0.56
                   Swietenia humilis           0.75
                   Swietenia                   0.60
                     macrophylla King
Mimosaceae         Parkia pendula              0.59
Papilionaceae      Dipteryx                    0.85
                     (Pittier) Record &
                   Hymenolobium                0.70
                     H.C. Lima
                   Myroxylon balsamum          0.91
                     (L.) Harms
                   Paramachaerium              0.82
                     gruberi Briz.
                   Platymiscium                0.78
                     curuense N. Zamora
                     & Kiitgaard
                   Platymiscium                0.79
                     parviflorum Benth.
                   Platymiscium                0.80
                     pinnatum var.
                     (Jacq.) Dugand
                   Platymiscium                0.78
                     pinnatum (Jacq.)
Podocarpaceae      Podocarpus                  0.57
                     costaricensis de
                   Podocarpus                  0.51
                   Podocarpus                  0.55
                     macrostachys Parl.
Zygophyllaceae     Guaiacum sanctum            1.06

Botanical Family   Species                Wood samples   Permanent
                                            observed      slides

Boraginaceae       Cordia                      6             6
                     gerascanthus L.
Caesalpiniaceae    Sclerolobium                1             1
                     Zamora & Poveda
Caryocaraceae      Anthodiscus                 1             1
                     chocoensis Prance
Lauraceae          Caryodaphnopsis             1             1
                     burgeri N. Zamora
                     & Poveda
Lecythidaceae      Couratari                   2             1
                     scottmorii Prance
Meliaceae          Cedrela fissilis            7             7
                   Cedrela                     2             2
                   Swietenia humilis           6             6
                   Swietenia                   7            12
                     macrophylla King
Mimosaceae         Parkia pendula              7             5
Papilionaceae      Dipteryx                    1             3
                     (Pittier) Record &
                   Hymenolobium                1             1
                     H.C. Lima
                   Myroxylon balsamum          7             7
                     (L.) Harms
                   Paramachaerium              2             2
                     gruberi Briz.
                   Platymiscium                1             1
                     curuense N. Zamora
                     & Kiitgaard
                   Platymiscium                1             1
                     parviflorum Benth.
                   Platymiscium                1             1
                     pinnatum var.
                     (Jacq.) Dugand
                   Platymiscium                5             2
                     pinnatum (Jacq.)
Podocarpaceae      Podocarpus                  2             1
                     costaricensis de
                   Podocarpus                  7             3
                   Podocarpus                  6             7
                     macrostachys Parl.
Zygophyllaceae     Guaiacum sanctum            7             9

Source: Law decree No. 23700-MINAE, No. 25167-MINAE and No.

Legend * Wood density was determined at 12% of moisture content.

LC=Light construction, HC=Heavy construction, Fl=flooring,
Fu=furniture and H=handicrafts.


Timber species considered to be in threat of extinction in Costa Rica

Botanical         Species                    Wood species   Endemic

Anacardiaceae     Astronium graveolens       Angiosperm       No
Bignoniaceae      Tabebuia guayacan          Angiosperm       No
                    (Seem.) Hemsl.
Caesalpiniaceae   Copaifera aromatica        Angiosperm       No
                  Copaifera camibar          Angiosperm       Yes
                    Poveda, N. Zamora
                    & P.E. Sanchez
                  Cynometra hemitomophylla   Angiosperm       No
                    (Donn. Sm.) Britton
                    & Rose
                  Mora oleifera              Angiosperm       No
                    (Triana) Ducke
                  Peltogyne                  Angiosperm       No
                    purpurea Pittier
                  Prioria copaifera          Angiosperm       No
                  Tachigali versicolor       Angiosperm       No
                    & L.O. Williams
Caryocaraceae     Caryocar costaricense      Angiosperm       No
                    Donn. Sm.
Humiriaceae       Humiriastrum               Angiosperm       No
                    guianensis Cuatrec.
                  Vantanea barbourii         Angiosperm       No
Juglandaceae      Oreomunnea pterocarpa      Angiosperm       Yes
Lecythidaceae     Couratari guianensis       Angiosperm       No
                  Lecythis ampla Miers.      Angiosperm       No
Meliaceae         Cedrela odorata L.         Angiosperm       No
                  Cedrela tonduzii C.DC.     Angiosperm       No
Olacaceae         Minquartia                 Angiosperm       No
                    guianensis Aubl.
Papilionaceae     Dalbergia retusa           Angiosperm       No
                  Dussia macroprophyllata    Angiosperm       Yes
                    (Donn. Sm.) Harms
Podocarpaceae     Prumnopitys standleyi      Gymnosperm       Yes
                    (Wild.) Ladd.
Sapotaceae        Sideroxylon capari         Angiosperm       No
                    (A. DC.) Pittier
Vochysiaceae      Qualea paraensis           Angiosperm       No

Botanical         Traditional used   Wood Density *
Family                of wood        (g/[cm.sup.3])

Anacardiaceae        HC, Fl & H           0.95
Bignoniaceae          HC & Fl             1.01
Caesalpiniaceae      LC, Fl & H           0.68
                    HC, Fl & Ha           0.78
                         LC               0.78
                      LC & Fl             0.75
                     HC, Fl & H           0.98
                      LC & Fl             0.48
                      LC & Fl             0.64
Caryocaraceae            HC               0.72
Humiriaceae              HC               0.66
                      HC & Fl             0.91
Juglandaceae             LC               0.53
Lecythidaceae            HC               0.69
                         HC               0.78
Meliaceae              FU & H             0.44
                       FU & H             0.44
Olacaceae             HC & Fl             0.89
Papilionaceae        HC, Fl & H           1.01
                         LC               0.59
Podocarpaceae            Fu               0.63
Sapotaceae            HC & Fl             0.79
Vochysiaceae          LC & Fl             0.37

Botanical         Wood samples      Permanent
Family              observed     slides observed

Anacardiaceae          8                9
Bignoniaceae           7                5
Caesalpiniaceae        3                2
                       1                1
                       2                2
                       1                1
                       7                3
                       6                9
                       1                2
Caryocaraceae          6                2
Humiriaceae            2                1
                       6                1
Juglandaceae           3                8
Lecythidaceae          5                1
                       5                1
Meliaceae              7                9
                       9                2
Olacaceae              1                6
Papilionaceae          7                6
                       1                3
Podocarpaceae          3                1
Sapotaceae             1                1
Vochysiaceae           2                8

Source: Jimenez 1999.

Legend: * Wood density was determined at 12% of moisture content.

LC=Light construction, HC=Heavy construction, Fl=flooring,
Fu=furniture and H=handicrafts.


Results of fluorescence test and vessels characteristics
of endangered and threat of extinction species in Costa Rica

Species                 Heartwood fluorescence

                             SF             WEF             EEF

Anthodiscus                  --             --              --
Astronium                 greenish      weak green     greenish blue
Caryocar                 weak green         --              --
Caryodaphnopsis              --             --              --
Cedrela odorata              --             --          light green
Cedrela fissilis             --             --          weak green
Cedrela salvadorensis        --             --              --
Cedrela tonduzii             --             --           variable
Copaifera aromatica       greenish      weak yellow      yellowish
Copaifera camibar         greenish      weak yellow      yellowish
Cordia gerascanthus          --         weak green    greenish yellow
Couratari guianensis         --             --          weak green
Couratari scottmorii         --             --          weak green
Cynometra                    --             --              --
Dalbergia retusa             --             --        yellowish green
Dipteryx panamensis          --             --          weak yellow
Dussia                   weak yellow        --         greenish blue
Guaiacum sanctum             --             --            purple
H. mesoamericanum        weak yellow    weak yellow      greenish
Humiriastrum                 --             --              --
Lecythis ampla            yellowish         --          weak green
Minquartia                   --             --              --
Mora oleifera           greenish blue       --              --
Myroxylon                 greenish          --         greenish blue
Oreomunnea                   --             --          light green

Paramachaerium               --             --              --
Parkia pendula               --             --          weak green
Peltogyne purpurea           --             --            yellow
Platymiscium                 --         light green      greenish
Platymiscium                 --         light green      greenish
Platymiscium                 --         light green      greenish
P. pinnatum var.             --         light green      greenish
Podocarpus                   --             --              --
Podocarpus                   --             --              --
Podocarpus                   --             --              --
Prioria copaifera            --             --          weak green
Prumnopitys                  --             --              --
Qualea paraensis             --             --          weak green
Sclerolobium                 --             --          light green
Sideroxylon                  --             --              --
Swietenia                    --             --          light green
Swietenia                    --             --          weak green
Tabebuia                     --             --           yellowish
Tachigali                 yellowish         --            bluish
Vantanea                     --             --          light green

Species                             Vessels elements

                        Porosity and   PSP    PF

Anthodiscus                  D         66    3.6
Astronium                    D         87    4.7
Caryocar                     D         56    4.3
Caryodaphnopsis              D         34    5.1
Cedrela odorata             D-S        79    3.7
Cedrela fissilis            D-S        77    2.1
Cedrela salvadorensis        S         75    3.3
Cedrela tonduzii             S         66    1.8
Copaifera aromatica          D         72    3.9
Copaifera camibar            D         86    5.8
Cordia gerascanthus         D-S        75    16.3
Couratari guianensis         D         27    2.2
Couratari scottmorii         D         61    3.7
Cynometra                    D         69    2.0
Dalbergia retusa             D         93    1.8
Dipteryx panamensis          D         65    9.9
Dussia                       D         70    1.3
Guaiacum sanctum             D         99    23.2
H. mesoamericanum            D         81    1.3
Humiriastrum                 D         96    10.8
Lecythis ampla               D         62    1.8
Minquartia                   D         43    12.7
Mora oleifera                D         79    3.6
Myroxylon                    D         44    19.4
Oreomunnea                   D         56    3.8
Paramachaerium               D         85    5.9
Parkia pendula               D         92    3.0
Peltogyne purpurea           D         29    7.2
Platymiscium                 D         81    1.6
Platymiscium                 D         79    3.1
Platymiscium                 D         75    2.8
P. pinnatum var.             D         41    2.4
Podocarpus                   --        --     --
Podocarpus                   --        --     --
Podocarpus                   --        --     --
Prioria copaifera            D         59    2.9
Prumnopitys                  --        --     --
Qualea paraensis             D         77    10.6
Sclerolobium                 D         62    3.8
Sideroxylon                D-DE-R      39    8.4
Swietenia                    D         64    5.2
Swietenia                    D         70    13.3
Tabebuia                     D         31    6.8
Tachigali                    D         69    3.9
Vantanea                     D         95    4.5

Species                        Vessels elements

                         LV    DV    PP

Anthodiscus             357    174   S
Astronium               428    129   S
Caryocar                267    152   S
Caryodaphnopsis         462    188   S
Cedrela odorata         240    105   S
Cedrela fissilis        369    140   S
Cedrela salvadorensis   267    150   S
Cedrela tonduzii        332    210   S
Copaifera aromatica     374    144   S
Copaifera camibar       311    118   S
Cordia gerascanthus     202    114   S
Couratari guianensis    244    116   S
Couratari scottmorii    366    164   S
Cynometra               352    140   S
Dalbergia retusa        153    201   S
Dipteryx panamensis     225    206   S
Dussia                  450    261   S
Guaiacum sanctum         93    58    S
H. mesoamericanum       463    243   S
Humiriastrum            1018   101   Sc
Lecythis ampla          352    204   S
Minquartia              930    89    Sc
Mora oleifera           287    102   S
Myroxylon               441    66    S
Oreomunnea              408    318   S
Paramachaerium          263    106   S
Parkia pendula          344    230   S
Peltogyne purpurea      308    149   S
Platymiscium            285    168   S
Platymiscium            172    172   S
Platymiscium            226    177   S
P. pinnatum var.        240    170   S
Podocarpus               --    --    --
Podocarpus               --    --    --
Podocarpus               --    --    --
Prioria copaifera       249    167   S
Prumnopitys              --    --    --
Qualea paraensis        306    166   S
Sclerolobium            371    189   S
Sideroxylon             240    86    S
Swietenia               349    118   S
Swietenia               150    137   S
Tabebuia                313    172   S
Tachigali               315    51    S
Vantanea                1160   211   Sc

Species                         Vessels elements

                        Deposits   DIP

Anthodiscus               G-T      4.2
Astronium                 G-T      8.7
Caryocar                   T       4.0
Caryodaphnopsis            T       6.2
Cedrela odorata            G       2.7
Cedrela fissilis           G       2.5
Cedrela salvadorensis      G       3.0
Cedrela tonduzii           G       5.6
Copaifera aromatica        G       5.8
Copaifera camibar          G       7.1
Cordia gerascanthus       G-T      5.4
Couratari guianensis       T       4.2
Couratari scottmorii       T       10.4
Cynometra                  G       6.9
Dalbergia retusa           G       2.8
Dipteryx panamensis        G       12.0
Dussia                     A       11.4
Guaiacum sanctum           G       2.6
H. mesoamericanum          G       9.0
Humiriastrum               G       2.0
Lecythis ampla             T       3.8
Minquartia                 T       12.3
Mora oleifera              G       5.4
Myroxylon                  G       1.6
Oreomunnea                 T       9.4
Paramachaerium             G       6.0
Parkia pendula             G       1.7
Peltogyne purpurea         G       7.7
Platymiscium               G       9.1
Platymiscium               G       6.4
Platymiscium               G       6.6
P. pinnatum var.           G       6.1
Podocarpus                 --       --
Podocarpus                 --       --
Podocarpus                 --       --
Prioria copaifera          G       2.3
Prumnopitys                --       --
Qualea paraensis           T       6.1
Sclerolobium               A       5.5
Sideroxylon                T       2.5
Swietenia                  G       1.6
Swietenia                  G       2.8
Tabebuia                   G       3.6
Tachigali                  G       1.3
Vantanea                   A       11.0


                        Vestured      VRP *

Anthodiscus                -           30
Astronium                  -          30-32
Caryocar                   -          30-35
Caryodaphnopsis            -          30-32
Cedrela odorata            -           30
Cedrela fissilis           -           30
Cedrela salvadorensis      -           30
Cedrela tonduzii           -           30
Copaifera aromatica        +           30
Copaifera camibar          +           30
Cordia gerascanthus        -           30
Couratari guianensis       -       30-31-32-33
Couratari scottmorii       -           30
Cynometra                  +           30
Dalbergia retusa           +           30
Dipteryx panamensis        +           30
Dussia                     +           30
Guaiacum sanctum           -           30
H. mesoamericanum          +           30
Humiriastrum               -           30
Lecythis ampla             -           30
Minquartia                 -           32
Mora oleifera              +           30
Myroxylon                  +           30
Oreomunnea                 -        30-31-32
Paramachaerium             +           30
Parkia pendula             +           30
Peltogyne purpurea         +           30
Platymiscium               +           30
Platymiscium               +           30
Platymiscium               +           30
P. pinnatum var.           +           30
Podocarpus                 -           --
Podocarpus                 -           --
Podocarpus                 -           --
Prioria copaifera          +           30
Prumnopitys                            --
Qualea paraensis           +           30
Sclerolobium               +           30
Sideroxylon                -           30
Swietenia                  -           30
Swietenia                  -           30
Tabebuia                   -           30

Tachigali                  +           30
Vantanea                   -        30-32-35

Legend: SF=surface fluorescence, WEF=Water extract fluorescence,
EEF: Ethanol extract fluorescence. D=diffuse-porous, S=semi-ring
porous, DE=Vessels in dendritic pattern, R=Vessels in radial or
diagonal pattern, PSP=Percentage of solitary pores, PF=Pores
frequency (pores/[mm.sup.2]), LV=Length of vessels ([micro]m),
DV=Diameter of vessels ([micro]m), PP=Perforation plates,
S=simple perforations plate, Sc=scalariform perforations plate,
Deposits: T=tyloses, G=gums, A=absent, DIP=diameter of intervascular
pits ([micro]m), VRP=Vessels-ray pitting according to IAWA class:
30: Vessel-ray pits with distinct borders; similar to intervessel
pits in size and shape throughout the ray cell, 31: Vessel-ray pits
with much reduced borders to apparently simple: pits rounded or
angular, 32: Vessel-ray pits with much reduced borders to
apparently simple: pits horizontal (scalariform, gash-like)
to vertical, 33: Vessel-ray pits of two distinct sizes or types
in the same ray cell, 35: Vessel-ray pits restricted to marginal
ows (IAWA, 1989). "-" anatomical feature absent, "+"
anatomical feature present, "?" unknown feature.


Fiber or tracheid and ray parenchyma characteristics of
species that are endangered or considered to be in threat
of extinction in Costa Rica

Spcies                             Fibers or tracheids

                                 FL     LD    WCT   FC

Anthodiscus chocoensis           1.5   17.4   6.5   -
Astronium graveolens             1.2   22.1   3.6   -
Caryocar costaricense            1.7   24.1   5.3   -
Caryodaphnopsis burgeri          1.8   21.3   6.6   -
Cedrela odorata                  0.9   24.4   3.3   -
Cedrela fissilis                 1.4   15.0   2.1   -
Cedrela salvadorensis            0.9   17.8   2.5   -
Cedrela tonduzii                 1.2   20.1   2.7   -
Copaifera aromatica              1.2   17.0   3.1   -
Copaifera camibar                1.2   19.1   3.4   -
Cordia gerascanthus              1.6   22.6   5.1   -
Couratari guianensis             1.3   13.1   2.5   -
Couratari scottmorii             1.4   16.7   4.2   -
Cynometra hemitomophylla         1.3   23.7   4.6   -
Dalbergia retusa                 0.6   19.2   7.7   -
Dipteryx panamensis              1.0   16.3   6.3   -
Dussia macroprophyllata          1.7   32.7   6.4   -
Guaiacum sanctum                 0.5   11.6   3.6   -
H. mesoamericanum                1.8   22.8   6.3   -
Humiriastrum guianensis          1.6   25.8   5.5   -
Lecythis ampla                   1.3   15.5   4.0   -
Minquartia guianensis            1.7   18.2   7.5   -
Mora oleifera                    1.3   14.4   6.2   -
Myroxylon balsamum               1.7   17.3   7.0   -
Oreomunnea pterocarpa            0.8   29.3   6.9   -
Paramachaerium gruberi           0.9   14.2   3.9   +
Parkia pendula                   1.1   19.2   4.6   -
Peltogyne purpurea               0.9   15.3   5.3   -
Platymiscium curuense            1.1   19.4   4.3   -
Platymiscium parviflorum         1.2   21.5   3.6   -
Platymiscium pinnatum            1.0   18.4   4.4   -
P. pinnatum var. polystachyum    1.1   19.9   4.9   -
Podocarpus costaricensis         2.4   31.5   6.3   -
Podocarpus guatemalensis         3.2   35.4   3.1   -
Podocarpus macrostachys          2.9   26.5   4.6   -
Prioria copaifera                1.0   21.8   2.9   -
Prumnopitys standleyi            2.5   31.4   5.7   -
Qualea paraensis                 1.1   18.3   4.4   -
Sclerolobium costaricense        1.1   14.6   2.7   -
Sideroxylon capari               0.8   17.0   8.9   -
Swietenia humilis                1.0   18.8   2.9   -
Swietenia macrophylla            0.8   21.0   3.6   -
Tabebuia guayacan                0.9   6.2    3.8   -
Tachigali versicolor             0.9   17.7   3.1   -
Vantanea barbourii               1.6   33.1   6.8   -

Spcies                            Fibers or tracheids

                                 SF    GTF   PFRT   FS

Anthodiscus chocoensis            -    SBD    -     -
Astronium graveolens              +    SBD    -     -
Caryocar costaricense             -    SBD    -     -
Caryodaphnopsis burgeri           -    SBD    -     -
Cedrela odorata                   -    SBD    -     -
Cedrela fissilis                  -    SBD    -     -
Cedrela salvadorensis             -    SBD    -     -
Cedrela tonduzii                  -    SBD    -     -
Copaifera aromatica               -    SBD    -     -
Copaifera camibar                 -    SBD    -     -
Cordia gerascanthus               -    SBD    -     -
Couratari guianensis              -    SBD    +?    -
Couratari scottmorii              -    SBD    -     -
Cynometra hemitomophylla          -    SBD    -     -
Dalbergia retusa                  -    SBD    -     +
Dipteryx panamensis               -    SBD    -     +
Dussia macroprophyllata           -    SBD    -     -+
Guaiacum sanctum                 -+    SBD    -     +
H. mesoamericanum                 -    SBD    -     +
Humiriastrum guianensis           -    BD     +     -
Lecythis ampla                    -    BD     +     -
Minquartia guianensis             -    SBD    -     -
Mora oleifera                     -    SBD    -     -
Myroxylon balsamum                -    SBD    -     +
Oreomunnea pterocarpa             -    SBD    -     -
Paramachaerium gruberi            -    SBD    -     +
Parkia pendula                    -    SBD    -     -
Peltogyne purpurea                -    SBD    -     -
Platymiscium curuense             -    SBD    -     -+
Platymiscium parviflorum          -    SBD    -     +
Platymiscium pinnatum             -    SBD    -     -+
P. pinnatum var. polystachyum     -    SBD    -     -+
Podocarpus costaricensis          -     T     -     +
Podocarpus guatemalensis          -     T     -     +
Podocarpus macrostachys           -     T     -     +
Prioria copaifera                 -    BD     -     -
Prumnopitys standleyi             -     T     -     -
Qualea paraensis                  -    SBD    -     -
Sclerolobium costaricense         -    SBD    -     -
Sideroxylon capari                -    SBD    -     -
Swietenia humilis                 +    SBD    -     -+
Swietenia macrophylla            - +   SBD    -     -+
Tabebuia guayacan                 -    SBD    -     +
Tachigali versicolor              -    SBD    -     -
Vantanea barbourii                -    BD     +     -

Spcies                                     Ray parenchyma

                                   RH          RW        RF

Anthodiscus chocoensis             590     1-3, U (>)   6.1
Astronium graveolens               850     1-3, 4-10    3.2
Caryocar costaricense              >1         1-3       7.8
Caryodaphnopsis burgeri            580        3-10      2.0
Cedrela odorata                    250        1-3       3.5
Cedrela fissilis                   335        1-3       3.6
Cedrela salvadorensis              342     1-3, 4-10    4.8
Cedrela tonduzii                   390        1-3       3.7
Copaifera aromatica                360        4-10      6.7
Copaifera camibar                  292        1-3       6.4
Cordia gerascanthus               290>1       4-10      6.8
Couratari guianensis               440     1-3, 4-10    6.2
Couratari scottmorii               380        1-3       6.8
Cynometra hemitomophylla           306        1-3       14.9
Dalbergia retusa                   134        1-3       16.6
Dipteryx panamensis                194        1-3       6.6
Dussia macroprophyllata            323        1-3       6.6
Guaiacum sanctum                   67          1        14.4
H. mesoamericanum                  360     1-3, 4-10    5.9
Humiriastrum guianensis            400     1-3, U (>)   19.7
Lecythis ampla                     480        1-3       2.8
Minquartia guianensis              >1      1-3, U (>)   7.0
Mora oleifera                      330        1-3       10.0
Myroxylon balsamum                 170        1-3       11.6
Oreomunnea pterocarpa              535        1-3       9.5
Paramachaerium gruberi             190        1-3       12.4
Parkia pendula                     224        1-3       5.8
Peltogyne purpurea                 >1         4-10      5.7
Platymiscium curuense              202         1        10.4
Platymiscium parviflorum           170         1        8.9
Platymiscium pinnatum              160         1        6.0
P. pinnatum var. polystachyum      160         1        9.6
Podocarpus costaricensis           71          1        4.2
Podocarpus guatemalensis           55          1        5.6
Podocarpus macrostachys            111         1        7.7
Prioria copaifera                  550        1-3       5.9
Prumnopitys standleyi              90          1        4.7
Qualea paraensis                 260, >1      1-3       4.1
Sclerolobium costaricense          250         1        11.0
Sideroxylon capari                 224        1-3       12.1
Swietenia humilis                  330     1-3, 4-10    7.0
Swietenia macrophylla              147     1-3, 4-10    8.1
Tabebuia guayacan                  190        1-3       9.5
Tachigali versicolor               227        1-3       8.2
Vantanea barbourii                 650        1-3       11.1

Spcies                                   Ray parenchyma

                                  RT     RS    R (C)

Anthodiscus chocoensis            Ht     -     S(+)
Astronium graveolens              Ht     -    U+ P(-)
Caryocar costaricense             Ht     -      -
Caryodaphnopsis burgeri           Ht     -      -
Cedrela odorata                  Ho-Ht   -    U+P(-)
Cedrela fissilis                  Ht     -    U+ P(-)
Cedrela salvadorensis             Ht     -    U+ P(-)
Cedrela tonduzii                  Ht     -    U+ P(-)
Copaifera aromatica              Ho-Ht   -    U+ P(-)
Copaifera camibar                Ho-Ht   -      -
Cordia gerascanthus               Ht     -    U+ P(+)
Couratari guianensis             Ho-Ht   -     S(+)
Couratari scottmorii             Ho-Ht   -     S(+)
Cynometra hemitomophylla          Ht     -      -
Dalbergia retusa                 Ho-Ht   +      -
Dipteryx panamensis               Ho     +      -
Dussia macroprophyllata          Ho-Ht   +-     -
Guaiacum sanctum                  Ho     +      -
H. mesoamericanum                 Ht     +      -
Humiriastrum guianensis           Ht     -      -
Lecythis ampla                   Ho-Ht   -     S(+)
Minquartia guianensis             Ht     -    U+ P(-)
Mora oleifera                     Ho     -      -
Myroxylon balsamum                Ht     +    U+ P(-)
Oreomunnea pterocarpa             Ht     -      -
Paramachaerium gruberi            Ho     +      -
Parkia pendula                    Ho     -      -
Peltogyne purpurea                Ho     +-     -
Platymiscium curuense             Ho     +      -
Platymiscium parviflorum          Ho     +      -
Platymiscium pinnatum             Ho     +      -
P. pinnatum var. polystachyum     Ho     +      -
Podocarpus costaricensis          Ho     -      -
Podocarpus guatemalensis          Ho     -      -
Podocarpus macrostachys           Ho     -      -
Prioria copaifera                 Ht     -      -
Prumnopitys standleyi             Ho     -      -
Qualea paraensis                 Ho-Ht   -     S(+)v
Sclerolobium costaricense         Ho     -      -
Sideroxylon capari                Ht     -    U+ P(-)
Swietenia humilis                 Ht     +-   U+ P(-)
Swietenia macrophylla             Ht     +-   U+ (-)
Tabebuia guayacan                 Ho     +      -
Tachigali versicolor              Ho     -     S(+)
Vantanea barbourii                Ht     -      -

Fiber: FL=Fiber or tracheids length (mm), LD=lumen or tracheids
diameter (um), WCT=wall cell thick (um), FC=Fibers crystals,
SF=Septate fibers, GTF=ground tissue fiber, SBD=simple to minutely
bordered pits, BD=Fibers with distinctly bordered pit, T=tracheids,
PFRT=Fiber pits common in both radial and tangential, FS=Fibers
stored. Ray parenchyma: RH=ray height (um), RW=ray width cell
(um), U(>)=mainly uniseriate, RF=ray frequency (ray/mm), RT=ray
type, Ht=heterogeneous, Ho=homogenous, RS=ray stored, R(C)=crystals
presence in ray, U=crystals in upright cells, P=procumbent cells,
S=Silica bodies presence, "-" anatomical feature absent, "+"
anatomical feature present.


Fiber, ray parenchyma and axial parenchyma characteristics of
endangered and threatened species in Costa Rica

                                    Axial parenchyma

                            AP     Type      Parenchyma arrangement

Anthodiscus chocoensis      -       --                 Vc
Astronium graveolens        +      Diff             Sca & Vc
Caryocar costaricense       +    Diff-Agg              Sca
Caryodaphnopsis burgeri     +      Diff                Sca
Cedrela odorata             +      Diff              Sca-Vc
Cedrela fissilis            +      Diff            Sca-Vc-Loz
Cedrela salvadorensis       +      Diff              Vc-Con
Cedrela tonduzii            +      Diff             Sca & Vc
Copaifera aromatica         -       --                 Vc

Copaifera camibar           -       --         Vc, Loz-alif & Con

Cordia gerascanthus         +      Diff           Sca, Vc & Con
Couratari guianensis        -       --                 --
Couratari scott-morii       -       --                 --

Cynometra hemitomophylla    -       --                 --
Dalbergia retusa            +    Diff-Agg           Sca & Vc
Dipteryx panamensis         -       --      Loz-alif, win-alif & Uni
Dussia macroprophyllata     -       --                 Con
Guaiacum sanctum            +    Diff-Agg              Sca
H. mesoamericanum           -       --          Loz-alif, Con, Vc
Humiriastrum guianensis     -       --            Vc, Loz-alif,
                                                 Win-alif & Uni
Lecythis ampla.             +      Diff                --
Minquartia guianensis       +    Diff-Agg              --
Mora oleifera               -       --            Vc, Loz-alif,
                                                 win-alif & Con
Myroxylon balsamum          -       --         Sca, Vc, Con & Uni
Oreomunnea pterocarpa       -       --
Paramachaerium gruberi      +      Diff     Sca, Loz-alif & wing-alif
Parkia pendula              -       --           Loz-alif & Con
Peltogyne purpurea          -       --        Loz-alif & Con & Uni
Platymiscium curuense       -       --           Loz-alif & Con
Platymiscium parviflorum.   -       --           Loz-alif & Con
Platymiscium pinnatum       -       --           Loz-alif & Con
P. pinnatum var.            -       --           Loz-alif & Con
Podocarpus costaricensis    +      Diff                --
Podocarpus guatemalensis    +      Sca                 --
Podocarpus macrostachys     +      Diff                --
Prioria copaifera           -       --         Vc, Loz-alif & Con
Prumnopitys standleyi       +      Sca                 --
Qualea paraensis            +      Diff           Vc, Loz-alif,
                                                 win-alif & Con
Sclerolobium costaricense   -       --                 Vc

Sideroxylon capari          +    Diff-Agg              --
Swietenia humilis           +      Diff                Sca
Swietenia macrophylla       +      Diff                Sca
Tabebuia guayacan           -       --         Vc, Loz-alif & Uni
Tachigali versicolor        -       --               Sca-Vc
Vantanea barbourii          +      Diff              Sca-Uni

                                      Axial parenchyma

                                 Banded        Stored      Cell

Anthodiscus chocoensis             --            --         >8
Astronium graveolens              Mar            --         3-8
Caryocar costaricense              --            --         3-8
Caryodaphnopsis burgeri           Nar            --         3-8
Cedrela odorata                   Mar            --         3-8
Cedrela fissilis                  Mar            --         3-8
Cedrela salvadorensis             Mar            --         3-8
Cedrela tonduzii                  Mar            --         3-8
Copaifera aromatica               Mar            --         3-8

Copaifera camibar                 Wid            --         3-8

Cordia gerascanthus               Mar            --       2 & 3-4
Couratari guianensis        Wid-Nar-Ret-Scal     --         3-8
Couratari scott-morii         Nar-Ret-Scal       --         3-8

Cynometra hemitomophylla      Nar-Ret-Mar      + ves        5-8
Dalbergia retusa                Nar-Mar        + ves      2 & 3-4
Dipteryx panamensis               Mar          + wae      2 & 3-4
Dussia macroprophyllata           Wid          + wae     3-8 & >8
Guaiacum sanctum                  Nar          + ves         2
H. mesoamericanum                 Wid          + wae      2 & 3-4
Humiriastrum guianensis            --            --      3-8 & >8

Lecythis ampla.               Wid-Ret-Mar        --         3-8
Minquartia guianensis              --            --         5-8
Mora oleifera                     Mar            --         3-5

Myroxylon balsamum                 --          +- wae     2, 3-4
Oreomunnea pterocarpa         Nar-Ret-Mar        --         5-8
Paramachaerium gruberi            Nar          + wae         2
Parkia pendula                                   --         3-5
Peltogyne purpurea                Mar          + ves        3-4
Platymiscium curuense             Nar            --          2
Platymiscium parviflorum.         Nar          + ves         2
Platymiscium pinnatum             Nar          + ves         2
P. pinnatum var.                  Nar          + ves         2
Podocarpus costaricensis           --            --         <4
Podocarpus guatemalensis           --            --         <4
Podocarpus macrostachys            --            --     <4 and 5-15
Prioria copaifera               Wid-Nar        + ves      2 & 3-4
Prumnopitys standleyi              --            --         <4
Qualea paraensis                   --            --         3-8

Sclerolobium costaricense          --            --         3-8

Sideroxylon capari                 --            --         3-4
Swietenia humilis                 Mar            --         5-8
Swietenia macrophylla             Mar            --         5-8
Tabebuia guayacan                 Mar          + wae      2 & 3-4
Tachigali versicolor               --            --       2 & 3-4
Vantanea barbourii                 --            --      5-8 & 8>

                              Axial      Others anatomical features

                              AP (C)

Anthodiscus chocoensis          --             Radial canals
Astronium graveolens            --
Caryocar costaricense         + (+)            Radial canals
Caryodaphnopsis burgeri       + (+)
Cedrela odorata               + (-+)
Cedrela fissilis              + (-+)
Cedrela salvadorensis         + (-+)
Cedrela tonduzii              + (-+)
Copaifera aromatica           + (+)         Axial & ray canals &
                                                sheath cells
Copaifera camibar             + (+)         Axial & ray canals &
                                                sheath cells
Cordia gerascanthus
Couratari guianensis          + (+)
Couratari scott-morii         + (+),
Cynometra hemitomophylla      + (+)
Dalbergia retusa              + (+)
Dipteryx panamensis           + (+)
Dussia macroprophyllata       + (+)
Guaiacum sanctum              + (+)
H. mesoamericanum             + (+)
Humiriastrum guianensis       + (+)           Traumatic canals

Lecythis ampla.               + (+)
Minquartia guianensis         + (+)        Vasicentric tracheids
Mora oleifera                 + (+)

Myroxylon balsamum            + (+)
Oreomunnea pterocarpa         + (+)
Paramachaerium gruberi        + (-+)     Crystals in enlarged cells
Parkia pendula                + (-+)
Peltogyne purpurea            + (+)
Platymiscium curuense         + (+)
Platymiscium parviflorum.     + (+)
Platymiscium pinnatum         + (+)           Traumatic canals
P. pinnatum var.              + (+)
Podocarpus costaricensis        --
Podocarpus guatemalensis        --
Podocarpus macrostachys         --
Prioria copaifera             + (+)         Axial canals diffuse
Prumnopitys standleyi           --
Qualea paraensis              + (-+)            Vessels of 2
                                             distinct diameter
Sclerolobium costaricense     + (+)          Helical thickening
                                                 in vessels
Sideroxylon capari            + (-+)
Swietenia humilis             + (-+)
Swietenia macrophylla         + (-+)
Tabebuia guayacan               -
Tachigali versicolor          + (+)
Vantanea barbourii            + (+-)

AP=apotraqueal parenchyma, Diff=diffuse. Diff-Agg=diffuse in
aggregates, Sca=scanty, Con=confluent, Vc=vasicentric,
Loz-alif=lozenge-aliform: Loz-w=lozenge-winged, uni=unilateral,
Mar=marginal, Nar=narrow band, Wid=wide band, ret=reticulate,
Scal=Scalarifom band, S=Silica bodies present, AP(C)=crystals
present in axial parenchyma (chambered present), ves=vessel
elements storied, ae=axial elements storied.
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Title Annotation:articulo en ingles
Author:Moya, Roger; Wiemann, Michael C.; Olivares, Carlos
Publication:Revista de Biologia Tropical
Date:Sep 1, 2013
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