Identificacion de especies de arboles en peligro o amenazadas de Costa Rica basada en la anatomia de la madera y fluorescencia.
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.
MATERIAL AND METHODS
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 diffuse-in-aggregates, 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 bands 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- aggregates 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 strand 15b. Ray height over 1 mm, Caryocar costaricense parenchyma diffuse, paratracheal scanty and in marginal or in seemingly marginal bands and radial canals 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- aggregates 17b. Perforation plates scalariform, Vantanea barbourii axial parenchyma diffuse, scanty partracheal and unilateral 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 bands 21a. Rays uniseriate, pits vestured, Sclerolobium axial parenchyma vasicentric, costaricense vessels with helical thickenings 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 bands 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 strand 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 width 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 unilateral 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 bands 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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Roger Moya1*, Michael C. Wiemann2 & Carlos Olivares3
1. Instituto Tecnologico de Costa Rica, Escuela de Ingenieria Forestal, Apartado 159-7050, Cartago, Costa Rica; firstname.lastname@example.org
2. Forest Products Laboratory, USDA Forest Service, One Gifford Pinchot Drive Madison, Wisconsin 53726-2398, USA; email@example.com
3. Instituto Tecnologico de Costa Rica, Escuela de Ingenieria Forestal, Apartado 159-7050, Cartago, Costa Rica; firstname.lastname@example.org
* Corresponding author
TABLE 1 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 costaricense 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 Vell. Cedrela Angiosperm No salvadorensis Standl. Swietenia humilis Angiosperm No Zucc. Swietenia Angiosperm No macrophylla King Mimosaceae Parkia pendula Angiosperm No Benth. Papilionaceae Dipteryx Angiosperm No panamensis (Pittier) Record & Mell Hymenolobium Angiosperm No mesoamericanum 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. polystachyum (Jacq.) Dugand Platymiscium Angiosperm No pinnatum (Jacq.) Dugand Podocarpaceae Podocarpus Gymnosperm Yes costaricensis de Laub. Podocarpus Gymnosperm No guatemalensis Standl. Podocarpus Gymnosperm No macrostachys Parl. Zygophyllaceae Guaiacum sanctum Angiosperm No L. Botanical Family Species Traditional used of wood Boraginaceae Cordia Fu & LC gerascanthus L. Caesalpiniaceae Sclerolobium HC & Fl costaricense Zamora & Poveda Caryocaraceae Anthodiscus HC chocoensis Prance Lauraceae Caryodaphnopsis HC burgeri N. Zamora & Poveda Lecythidaceae Couratari HC scottmorii Prance Meliaceae Cedrela fissilis Fu & H Vell. Cedrela Fu & H salvadorensis Standl. Swietenia humilis Fu & H Zucc. Swietenia Fu & H macrophylla King Mimosaceae Parkia pendula LC Benth. Papilionaceae Dipteryx HC & Fl panamensis (Pittier) Record & Mell Hymenolobium LC mesoamericanum 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. polystachyum (Jacq.) Dugand Platymiscium Fu, Fl & H pinnatum (Jacq.) Dugand Podocarpaceae Podocarpus Fu costaricensis de Laub. Podocarpus Fu guatemalensis Standl. Podocarpus Fu macrostachys Parl. Zygophyllaceae Guaiacum sanctum HC, Fl & H L. Botanical Family Species Wood density * (g/[cm.sup.3]) Boraginaceae Cordia 0.72 gerascanthus L. Caesalpiniaceae Sclerolobium 0.74 costaricense 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 Vell. Cedrela 0.56 salvadorensis Standl. Swietenia humilis 0.75 Zucc. Swietenia 0.60 macrophylla King Mimosaceae Parkia pendula 0.59 Benth. Papilionaceae Dipteryx 0.85 panamensis (Pittier) Record & Mell Hymenolobium 0.70 mesoamericanum 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. polystachyum (Jacq.) Dugand Platymiscium 0.78 pinnatum (Jacq.) Dugand Podocarpaceae Podocarpus 0.57 costaricensis de Laub. Podocarpus 0.51 guatemalensis Standl. Podocarpus 0.55 macrostachys Parl. Zygophyllaceae Guaiacum sanctum 1.06 L. Botanical Family Species Wood samples Permanent observed slides observed Boraginaceae Cordia 6 6 gerascanthus L. Caesalpiniaceae Sclerolobium 1 1 costaricense 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 Vell. Cedrela 2 2 salvadorensis Standl. Swietenia humilis 6 6 Zucc. Swietenia 7 12 macrophylla King Mimosaceae Parkia pendula 7 5 Benth. Papilionaceae Dipteryx 1 3 panamensis (Pittier) Record & Mell Hymenolobium 1 1 mesoamericanum 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. polystachyum (Jacq.) Dugand Platymiscium 5 2 pinnatum (Jacq.) Dugand Podocarpaceae Podocarpus 2 1 costaricensis de Laub. Podocarpus 7 3 guatemalensis Standl. Podocarpus 6 7 macrostachys Parl. Zygophyllaceae Guaiacum sanctum 7 9 L. Source: Law decree No. 23700-MINAE, No. 25167-MINAE and No. 25663-MINAE. Legend * Wood density was determined at 12% of moisture content. LC=Light construction, HC=Heavy construction, Fl=flooring, Fu=furniture and H=handicrafts. TABLE 2 Timber species considered to be in threat of extinction in Costa Rica Botanical Species Wood species Endemic Family Anacardiaceae Astronium graveolens Angiosperm No Jacq. Bignoniaceae Tabebuia guayacan Angiosperm No (Seem.) Hemsl. Caesalpiniaceae Copaifera aromatica Angiosperm No Dwyer 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 Griseb. Tachigali versicolor Angiosperm No Standl. & L.O. Williams Caryocaraceae Caryocar costaricense Angiosperm No Donn. Sm. Humiriaceae Humiriastrum Angiosperm No guianensis Cuatrec. Vantanea barbourii Angiosperm No Standl. Juglandaceae Oreomunnea pterocarpa Angiosperm Yes Oerst. Lecythidaceae Couratari guianensis Angiosperm No Aubl. 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 Hemsl. 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 Ducke 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. TABLE 3 Results of fluorescence test and vessels characteristics of endangered and threat of extinction species in Costa Rica Species Heartwood fluorescence SF WEF EEF Anthodiscus -- -- -- chocoensis Astronium greenish weak green greenish blue graveolens Caryocar weak green -- -- costaricense Caryodaphnopsis -- -- -- burgeri 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 -- -- -- hemitomophylla Dalbergia retusa -- -- yellowish green Dipteryx panamensis -- -- weak yellow Dussia weak yellow -- greenish blue macroprophyllata Guaiacum sanctum -- -- purple H. mesoamericanum weak yellow weak yellow greenish Humiriastrum -- -- -- guianensis Lecythis ampla yellowish -- weak green Minquartia -- -- -- guianensis Mora oleifera greenish blue -- -- Myroxylon greenish -- greenish blue balsamum Oreomunnea -- -- light green pterocarpa Paramachaerium -- -- -- gruberi Parkia pendula -- -- weak green Peltogyne purpurea -- -- yellow Platymiscium -- light green greenish curuense Platymiscium -- light green greenish parviflorum. Platymiscium -- light green greenish pinnatum P. pinnatum var. -- light green greenish polystachyum Podocarpus -- -- -- costaricensis Podocarpus -- -- -- guatemalensis Podocarpus -- -- -- macrostachys Prioria copaifera -- -- weak green Prumnopitys -- -- -- standleyi Qualea paraensis -- -- weak green Sclerolobium -- -- light green costaricense Sideroxylon -- -- -- capari Swietenia -- -- light green humilis Swietenia -- -- weak green macrophylla Tabebuia -- -- yellowish guayacan Tachigali yellowish -- bluish versicolor Vantanea -- -- light green barbourii Species Vessels elements characteristics Porosity and PSP PF arrangement Anthodiscus D 66 3.6 chocoensis Astronium D 87 4.7 graveolens Caryocar D 56 4.3 costaricense Caryodaphnopsis D 34 5.1 burgeri 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 hemitomophylla Dalbergia retusa D 93 1.8 Dipteryx panamensis D 65 9.9 Dussia D 70 1.3 macroprophyllata Guaiacum sanctum D 99 23.2 H. mesoamericanum D 81 1.3 Humiriastrum D 96 10.8 guianensis Lecythis ampla D 62 1.8 Minquartia D 43 12.7 guianensis Mora oleifera D 79 3.6 Myroxylon D 44 19.4 balsamum Oreomunnea D 56 3.8 pterocarpa Paramachaerium D 85 5.9 gruberi Parkia pendula D 92 3.0 Peltogyne purpurea D 29 7.2 Platymiscium D 81 1.6 curuense Platymiscium D 79 3.1 parviflorum. Platymiscium D 75 2.8 pinnatum P. pinnatum var. D 41 2.4 polystachyum Podocarpus -- -- -- costaricensis Podocarpus -- -- -- guatemalensis Podocarpus -- -- -- macrostachys Prioria copaifera D 59 2.9 Prumnopitys -- -- -- standleyi Qualea paraensis D 77 10.6 Sclerolobium D 62 3.8 costaricense Sideroxylon D-DE-R 39 8.4 capari Swietenia D 64 5.2 humilis Swietenia D 70 13.3 macrophylla Tabebuia D 31 6.8 guayacan Tachigali D 69 3.9 versicolor Vantanea D 95 4.5 barbourii Species Vessels elements characteristics LV DV PP Anthodiscus 357 174 S chocoensis Astronium 428 129 S graveolens Caryocar 267 152 S costaricense Caryodaphnopsis 462 188 S burgeri 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 hemitomophylla Dalbergia retusa 153 201 S Dipteryx panamensis 225 206 S Dussia 450 261 S macroprophyllata Guaiacum sanctum 93 58 S H. mesoamericanum 463 243 S Humiriastrum 1018 101 Sc guianensis Lecythis ampla 352 204 S Minquartia 930 89 Sc guianensis Mora oleifera 287 102 S Myroxylon 441 66 S balsamum Oreomunnea 408 318 S pterocarpa Paramachaerium 263 106 S gruberi Parkia pendula 344 230 S Peltogyne purpurea 308 149 S Platymiscium 285 168 S curuense Platymiscium 172 172 S parviflorum. Platymiscium 226 177 S pinnatum P. pinnatum var. 240 170 S polystachyum Podocarpus -- -- -- costaricensis Podocarpus -- -- -- guatemalensis Podocarpus -- -- -- macrostachys Prioria copaifera 249 167 S Prumnopitys -- -- -- standleyi Qualea paraensis 306 166 S Sclerolobium 371 189 S costaricense Sideroxylon 240 86 S capari Swietenia 349 118 S humilis Swietenia 150 137 S macrophylla Tabebuia 313 172 S guayacan Tachigali 315 51 S versicolor Vantanea 1160 211 Sc barbourii Species Vessels elements characteristics Deposits DIP Anthodiscus G-T 4.2 chocoensis Astronium G-T 8.7 graveolens Caryocar T 4.0 costaricense Caryodaphnopsis T 6.2 burgeri 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 hemitomophylla Dalbergia retusa G 2.8 Dipteryx panamensis G 12.0 Dussia A 11.4 macroprophyllata Guaiacum sanctum G 2.6 H. mesoamericanum G 9.0 Humiriastrum G 2.0 guianensis Lecythis ampla T 3.8 Minquartia T 12.3 guianensis Mora oleifera G 5.4 Myroxylon G 1.6 balsamum Oreomunnea T 9.4 pterocarpa Paramachaerium G 6.0 gruberi Parkia pendula G 1.7 Peltogyne purpurea G 7.7 Platymiscium G 9.1 curuense Platymiscium G 6.4 parviflorum. Platymiscium G 6.6 pinnatum P. pinnatum var. G 6.1 polystachyum Podocarpus -- -- costaricensis Podocarpus -- -- guatemalensis Podocarpus -- -- macrostachys Prioria copaifera G 2.3 Prumnopitys -- -- standleyi Qualea paraensis T 6.1 Sclerolobium A 5.5 costaricense Sideroxylon T 2.5 capari Swietenia G 1.6 humilis Swietenia G 2.8 macrophylla Tabebuia G 3.6 guayacan Tachigali G 1.3 versicolor Vantanea A 11.0 barbourii Species Vestured VRP * Anthodiscus - 30 chocoensis Astronium - 30-32 graveolens Caryocar - 30-35 costaricense Caryodaphnopsis - 30-32 burgeri 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 hemitomophylla Dalbergia retusa + 30 Dipteryx panamensis + 30 Dussia + 30 macroprophyllata Guaiacum sanctum - 30 H. mesoamericanum + 30 Humiriastrum - 30 guianensis Lecythis ampla - 30 Minquartia - 32 guianensis Mora oleifera + 30 Myroxylon + 30 balsamum Oreomunnea - 30-31-32 pterocarpa Paramachaerium + 30 gruberi Parkia pendula + 30 Peltogyne purpurea + 30 Platymiscium + 30 curuense Platymiscium + 30 parviflorum. Platymiscium + 30 pinnatum P. pinnatum var. + 30 polystachyum Podocarpus - -- costaricensis Podocarpus - -- guatemalensis Podocarpus - -- macrostachys Prioria copaifera + 30 Prumnopitys -- standleyi Qualea paraensis + 30 Sclerolobium + 30 costaricense Sideroxylon - 30 capari Swietenia - 30 humilis Swietenia - 30 macrophylla Tabebuia - 30 guayacan Tachigali + 30 versicolor Vantanea - 30-32-35 barbourii 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. TABLE 4 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. TABLE 5 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 polystachyum 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 Parenchyma 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 polystachyum 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 parenchyma 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 + (+), S(+) 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. + (+) polystachyum 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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