Biodiversity and Conservation 7, 1563±1575 (1998)
Biodiversity and Conservation 7, 1563±1575 (1998)
Vascular plant species count in a wet forest in the ChocoÂ area on the Paci®c coast of Colombia
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, BogotaÂ, Colombia
STELLA SUAÂ REZ
Apartado 265216, BogotaÂ, Colombia
Department of Systematic Botany, Institute of Biological Sciences, Building 137, University of Aarhus, DK-8000 Aarhus C, Denmark
Received 22 February 1998; revised and accepted 13 May 1998
The total number of vascular plant species was counted and growth form distribution was studied in the ChocoÂ area on the Paci®c coast of Colombia, in two transects 400 ́ 10 m and ten transects 2 ́ 50 m, for a total sampled area of 0.9 ha. The species count of the ten transects (442 species in 0.1 ha) appears to be the highest number of species recorded with this methodology. There were 970 species for the total area (0.9 ha). Ninety to ninety-®ve percent of the species were under 10 cm dbh and 70±86% under 2.5 cm dbh, epiphytes and small trees and treelets £10 cm dbh being the most diverse growth forms. The most species-rich families and genera were those represented by herba- ceous plants and treelets. Individuals were counted only in the ten 2 ́ 50 m transects (0.1 ha), where 4459 individuals were found. Palms and ferns were the most abundant growth forms. Arguments are presented against the way diversity is usually measured. Recommendations are made to include other growth forms besides trees when assessing alpha diversity for conservation purposes.
Keywords: species richness; growth forms; wet forest; ChocoÂ ; Colombia.Introduction
The bulk of information on structure, composition and diversity of neotropical forests is based upon the study of the woody component of the forest. Customarily, quantitative inventories include only trees 310cm dbh and less often they include all woody plants32.5 cm dbh (Gentry, 1982a, 1986, 1988, and see review in Gentry, 1995), and very few studies include plants 31 cm dbh (e.g. Franco-Rosselli et al., 1997). The few quantitative studies of neotropical forests that have considered all the species or have included other non-woody life-forms, have shown the importance of non-tree species for the structure, composition and species richness of tropical forests (Smith, 1970; Whitmore et al., 1985; Gentry and Dodson, 1987; Poulsen and Balslev, 1991; Duivenvoorden, 1994; Balsev et al., in press). The available information, however, is still very scarce. The total sampling of vascular plant alpha diversity faces the diculty of collecting in the forest canopy and of identifying sterile material (Whitmore et al., 1985). Thus, most inventories in tropical forests continue to include only individuals 310 cm dbh.
*To whom correspondence should be addressed. 0960-3115 Ó 1998 Chapman & Hall
1564 Galeano et al.
The ChocoÂ biogeographic region (Gentry, 1982b) is believed to hold about 8000 vas- cular plant species, including a high number of endemic species, and it is considered to include some of the most diverse forests on earth (Lellinger, 1975; Gentry, 1986, 1993; Forero and Gentry, 1989; McNeeley et al., 1990). Quantitative studies on the region have dealt mostly with trees 310 cm dbh or with individuals 32.5 cm dbh (Gentry, 1986; Faber- Langendoen and Gentry, 1991). The only published study dealing with a species count and including all growth forms and size classes in the ChocoÂ region is that of Gentry and Dodson (1987) at RõÂo Palenque, Ecuador, where they recorded a total of 365 vascular plant species in ten 0.01 transects. That locality, however, has a considerably drier climate (2980 mm of annual precipitation) than the site of the present study.
The structure, composition and richness of the forests at the Gulf of TribugaÂ , on the Paci®c coast of ChocoÂ , Colombia, were previously studied based on the individuals 35 cm and 310 cm dbh (Galeano et al., in press; Galeano, unpubl. data). In this paper we report the total number of vascular plant species counted in two 0.4 ha and ten 0.01 ha transects. This is the ®rst total species count ever recorded for a site with more than 5000 mm of rainfall, and it is the second one recorded for the ChocoÂ region. On the basis of these counts, we analyse the importance of dierent growth forms and taxonomic groups to forest structure, composition, and richness, and we make comparisons with similar studies in the ChocoÂ region and elsewhere in the Neotropics.
The study was carried out at the Gulf of TribugaÂ , on the Paci®c Coast of the ChocoÂ Department, Colombia (5° 29¢±6° 08¢N, 77° 10¢±77° 32¢W). The land around the gulf consists of hills lower than 300 m, with steep slopes of 10±50°, mostly covered by undis- turbed tropical wet forest according to Holdridge’s life-zone system (IGAC, 1977). Soil is derived from basaltic rocks, with a moderately developed structure, nutrient poor and acidic (CorteÂ s, 1981). Annual precipitation ranges from 5100 mm (average for 20 years) as recorded in a weather station about 50 km to the north, and 7150 mm (average for 5 years) as recorded in E1 Amargal Biological Station, near the southmost point of the gulf. More information about the study area, the sites, and about the structure, composition and richness of the woody species can be found in Galeano et al. (in press).
Two 0.4 ha transects (400 ́ 10 m) about 10 km from each other, were made at NuquõÂ and CoquõÂ. The third sampling unit was established at E1 Amargal Biological Station, ca 27 and 30km west of the NuquõÂ and CoquõÂ transects, respectively. Here, 10 transects of 2 ́ 50 m, mostly parallel among them and separated by 10±15 m were established. All transects were subjectively placed to include the range of topography common in the area (hillsides, ridges and some ̄at areas). In each transect, all vascular plant species were recorded and collected. Dbh and height were measured on all plant individuals. Trees whose stems straddled the transect border were included in the sample if the midpoint of their trunks fell within the range. Epiphytes and lianas were located with binoculars and then sampled. The sample was enhanced by bringing down one to two large tree branches in each of 20 subtransects of 10 ́ 20 m at NuquõÂ and CoquõÂ and in each of the ten 2 ́ 50 m transects at E1 Amargal, and searching them for epiphytes. At E1 Amargal the
Plant species count in ChocoÂ, Colombia 1565
number of individuals of each species was counted; epiphytes were counted with binoc- ulars; clones of rhizomatous plants were counted as single individuals only when we were certain that all ramets belonged to the same plant. Woody plants shorter than 1.3 m were not sampled at NuquõÂ and CoquõÂ , and saplings less than 50 cm tall were not sampled at E1 Amargal. Given the constrains of the last points mentioned, the samplings of both species and individuals were neither exact nor total. Vouchers were identi®ed at COL, or sent to specialists and duplicates were deposited at INDERENA, CHOCO, and AAU.
For analysis, each species (or morphospecies when identi®cation was not possible) was assigned to one of the following growth forms (some of them modi®ed from Gentry, 1985):
- Trees (woody plants 35 cm dbh, excluding palms, lianas, and hemiepiphytes).
- Treelets (woody plants <5 cm dbh, excluding palms, lianas and hemiepiphytes).
- Palms (members of the family Palmae and Zamia obliqua, Zamiaceae).
- Lianas (woody climbers; begining life as terrestrial seedlings).
- Vines (herbaceous or subwoody climbers; begining life as terrestrial seedlings).
- Hemiepiphytes (climbers; begining life as epiphytes; sometimes with woody stemsappressed to supporting trees).
- Terrestrial herbs (including saprophytes).
- Epiphytes (both erect and climbing epiphytes).
Species richness and composition
The total number of vascular plant species found at the three sites is shown in Table 1. The species±area curve for the cumulative species in all sampling units (Fig. 1) showed no indication of levelling o, indicating that more species would have appeared if a larger area had been sampled. For the three sites pooled (an area of 0.9 ha) there were 970 species of vascular plants, distributed as follows: 890 species of ̄owering plants (dicots ac- counting for 70% of the species, and 80% of the families), 78 species of Pteridophyta, and two species of gymnosperms (Zamia obliqua A. Braun and Gnetum leyboldii Tul.). Family concept for the angiosperms follows Cronquist (1981), but the Fabaceae are treated as one large family including Caesalpiniaceae and Mimosaceae; the Pteridophyta are treated as a large group following Mabberley (1989). Only 38% of the species have been satisfactorily identi®ed so far. The ̄owering plants were distributed among 90 families and, at least, 295 genera. The most species-rich families are shown in Fig. 2. These 15 families (17% of all families found) included 63% of the total number of species. The species-rich families are roughly the same for the three transects, although ranked dierently. Figure 3 shows the species-richest genera. These 12 genera accounted for 23.4% of all the species; most of them are herbaceous*.
Density of individuals
The total number of individuals in ten 0.01 transects at E1 Amargal was 4459. Table 2 shows the number of individuals and their relative proportions in each growth form; Table 3 shows the most abundant species. Terrestrial herbs (most of them represented by ferns) and palms were the most dense groups.
*A complete list of species found can be obtained from the ®rst author.
1566 Galeano et al.
number of species and proportion in parentheses. Species may overlap between size classes but not between growth
El Amargalb0.1 ha
Erect epiphytes Climbing epiphytes Palms <10 cm dbh Lianas <10 cm dbh Vines
Hemiepiphytes <10 cm dbh
96 (19.7) 63 (12.9) 48 (9.8)
53 (24.5) 61 32 (14.8) 115 28 (13.0) 59
(12.5) 28 (23.5) 79 (12.1) 40 (4.1) 14 (4.1) 7 (1.0) 4 (1.8) 5
(10.3) (29.2) (14.8) (5.2) (2.6) (1.5) (1.8)
63 (14.2) 32 (7.2) 69 (15.6) 11 (2.5) 22 (5.0)
Small trees and treelets <10 cm dbh (including saplings >50 cm tall) Trees 310 cm dbh Palms 310 cm dbh Lianas 310 cm dbh Hemiepiphytes 310 cm dbh
9 (1.8) 125 (25.6)
58 (27) 155
488 216 489
ha at these sites come from the ®rst ®ve subtransects.
6 (2.8) 43 (8.8) 17 (3.5)
1 (0.2) 20 13 (6.0) 20 3 (1.4) 5 4 (1.9) 9
8 (1.8) 13 (2.9)
4 (0.8) 2 (0.9) 5 2 (0.4) 0 0 2 (0.4) 1 (0.5) 0
(16.4) 29 (1.0) 2 0 0
34 (7.7) 3 (0.7) 0
a NuquõÂ and CoquõÂ were established using contiguous 200 m2 subtransects; data for 0.1b The 0.1 ha at E1 Amargal was the sum of ten 2 ́ 50 m separated transects.
22 (10.2) 80
Plant species count in ChocoÂ, Colombia 1567
Figure 1. Species±area curve for all vascular plant species in the cumulative area sampled at the Paci®c coast of ChocoÂ , Colombia.
Figure 2. Largest families of vascular plants in three plots totalling 0.9 ha, at the Paci®c coast of ChocoÂ , Colombia. Ferns are treated as one family. Figures at the end of the bars represent per- centages of the total.
Size and growth form distribution
Table 1 shows the growth form distribution at the three sites (including the ®rst 0.1 ha at NuquõÂ and CoquõÂ ). Epiphytes were the most diverse growth form in NuquõÂ and CoquõÂ and the second most diverse at E1 Amargal (Fig. 4). Epiphytes (including erect epiphytes, climbing epiphytes, and hemiepiphytes) accounted for 25±46% of the total species number, being more abundant at CoquõÂ . Small trees and treelets (<10 cm dbh, including their saplings), were also very species-rich, accounting for 26±46% of all species in all
1568 Galeano et al.
Figure 3. Largest genera (with 10 or more species) of vascular plants in three plots totalling 0.9 ha, at the Paci®c coast of ChocoÂ , Colombia. In parentheses growth form represented in order of occurrence (t, trees; tr, treelets; h, terrestrial herbs; e, epiphytes + hemiepiphytes; l, lianas).
counts (Table 1). At E1 Amargal, treelets <2.5 cm dbh were remarkably species-rich (Fig. 4). Ninety to ninety-®ve percent of the species in the transects were under 10 cm dbh and 70±86% were under 2.5 cm dbh (Fig. 5). The non-woody components (terrestrial herbs, epiphytes, vines, and herbaceous hemiepiphytes) accounted for about 50% of the total species number (Fig. 5).
The total of 442 species of vascular plants found in the ten 2 ́ 50 m transects at E1 Amargal is, to our knowledge, the highest ®gure ever recorded for a total species count in 0.1 ha. The second highest number of species has been recorded for the RõÂ o Palenque area, Ecuador (365 species), employing the same methodology used at E1 Amargal but in- cluding saplings less than 50 cm high (Gentry and Dodson, 1987). If those size classes had been included at E1 Amargal, the number of the species would have been probably higher. Also, because we did not employ destructive sampling, it is possible that some species were missed, especially in the high canopy.
The species counts for NuquõÂ and CoquõÂ and for their respective 0.1 ha subtransects are similar to each other. However, the same area (0.1 ha) sampled at E1 Amargal had 1.6±2.0 times as many species as the 0.1 ha transects at NuquõÂ or CoquõÂ , and about as many species as either of the two 0.4 ha transects (Table 1). This remarkable dierence seems to be related to the dierent sampling methodology. First, the dierent size limit for sampling in NuquõÂ and CoquõÂ (1.3 m high) compared to that at E1 Amargal (50 cm high) accounts for a dierence of 69 species. Second, the exploded quadrat method (Gentry, 1982a) used at E1 Amargal seems to yield many more species than the contiguous areas used in NuquõÂ and CoquõÂ. Faber-Langendoen and Gentry (1991) using contiguous plots, recorded
Plant species count in ChocoÂ, Colombia 1569Table 2. Distribution of the total number of individuals among growth forms in 0.1 ha at E1
Amargal, Paci®c coast of ChocoÂ , Colombia
Growth forms No. individuals Percentage
Terrestrial herbs 1704 38.2 Epiphytes 492 11.0 Hemiepiphytes 144 3.2 Lianas 75 1.7 Vines 21 0.5 Palms 1177 26.4
Treelets <2.5 cm dbh (incl. Saplings)aSmall trees and treelets 2.5 £ dbh <10 cmaTrees 310 cm dbh
690 15.5 106 2.4 50 1.1
Total 4459 100a Treelets were divided into two size classes for easier comparison.
1.6±2.4 times fewer species as Gentry (1986) found with the exploded quadrat method; although both sites were extremely close to each other (Duivenvoorden, 1994). Finally, the exploded quadrat method involves a higher possibility of sampling error (Peters, 1996).
Compared to other forests from the ChocoÂ biogeographic region (Table 4), particularly in terms of species 310 cm and 32.5 cm dbh, the forest at the Gulf of TribugaÂ is less rich, and seems to be structurally more similar to the drier site at RõÂ o Palenque, Ecuador than to other Colombian sites with similar or higher rainfall.
Most of the diversity at the study transects is due to the herbaceous component, es- pecially to epiphytes. This overall high importance of epiphytes is similar for some Andean forests as well as for some Central American forests, particularly La Selva (Hammel, 1990), and is one of the most notable dierences between these regions and the Amazon region (see Gentry, 1990; Duivenvoorden, 1994). The ChocoÂ sites studied here, compar- atively poor in tree species 310 cm dbh but with a high richness in vascular plant species, represent an extreme in the dichotomy of growth form distribution between Amazonian and Central American (and now Chocoan) forests (Gentry, 1990). Whereas Amazonia,
Table 3. The most abundant ChocoÂ , Colombia
Wel®a regia H. Wendl.
(O.F. Cook and Doyle) Burret
Calathea elegans KennedyTrichomanes elegans Rich.Philodendron inaequilaterum Liebm.
species in ten 0.01 ha transects at
E1 Amargal Biological Station,
Marantaceae Fern Araceae
palm terrestrial herb
terrestrial herb terrestrial herb hemiepiphyte 93
2.8 2.2 2.1
1570 Galeano et al.
Figure 4. Percentage of species per growth form in three 0.1 ha plots at the Paci®c coast of ChocoÂ , Colombia. At E1 Amargal, treelets £2.5 cm dbh include saplings higher than 50 cm. Some species may overlap between trees and treelets.
Figure 5. Contribution of size classes and growth forms groups to the total number of species in three 0.1 ha plots at the Paci®c coast of ChocoÂ , Colombia. Nonwoody species include terrestrial herbs, epiphytes, hemiepiphytes and vines.
Plant species count in ChocoÂ, Colombia 1571
Table 4. Number of species 310 cm and 32.5 cm recorded for 0.1 ha plots in the ChocoÂ biogeographic region, and at La Selva
Bajo Calima 1 Bajo Calima 2 Tutunendo RõÂo Palenque NuquõÂ
E1 Amargal 1 E1 Amargal 2
La Selva, Costa Rica
310 cm dbh
77 42±48 55 32 24 31 37 21±40 43
32.5 cm dbh
265 108±166 258 119
77 102 90 130
Faber-Langendoen and Gentry (1991) Gentry (1986)
Galeano et al. (in press)
particularly the western part, seems to be the region of highest tree species richness (Gentry, 1990; Duivenvoorden, 1994; Valencia et al., 1994), the ChocoÂ region appears to be the one with the highest species richness of non-woody plants. If this generality proves true, one would expect that a total species count at any ChocoÂ site as rich in woody species as Bajo Calima or Tutunendo (Table 4) would result in a number of species higher than the one recorded here.
The low representation of lianas is noteworthy. The number of liana species 32.5 cm dbh per 0.1 ha at the three 0.1 ha transects in the Gulf of TribugaÂ (2±6) is more similar to that recorded in the temperate zone or in dry tropical forest (x 12 species) than to the number recorded at moist (x 25 species), or wet tropical forests (x 32 species) (Gentry, 1982a). A low density of lianas has also been recorded for other forests in the ChocoÂ region (Gentry, 1986; Faber-Lagendoen and Gentry, 1991). To explain this unusual feature for a neotropical forest, Gentry (1986) suggested that in the ChocoÂ forest the lianas are, in an ecological sense, replaced by hemiepiphytes. Nevertheless, our data do not support this view because hemiepiphytes are also scarce and even less important than lianas at any scale. Gentry (1982a) discussed the existence of an apparent positive cor- relation between liana density and stand maturity. Nevertheless, the structure of the transects in the present study showed that the forest is a mature one (Galeano et al., in press). It may be possible, then, that epiphytes (instead of hemiepiphytes) replace (in an ecological sense) lianas at the study site, as was suggested by Hartshorn and Hammel (1994) for the forest at La Selva, which has also few lianas and abundant epiphytes. The site with fewest liana species, the 0.1 ha at CoquõÂ (2.6% of the species) was the same with the largest number of epiphytes (46% of the species).
Floristic composition, in terms of families and genera, re ̄ects growth form represen- tation. The species-richest families and genera at the Gulf of TribugaÂ were those of her- baceous plants and treelets (Fig. 2). A similar situation has been recorded at La Selva and, to a lesser extent, at Barro Colorado Island (Gentry, 1990). This is in strong contrast to the situation recorded in the Amazon, where families and genera of trees and treelets prevail (Foster, 1990; Gentry, 1990; Prance, 1990; Duivenvoorden, 1994; but see Balslev et al., in press). The Araceae, Piperaceae, ferns, and Cyclanthaceae are particularly important at our study sites. The substantial local diversity of these taxa is due to the coexistence of many sympatric species (Fig. 3); 74% of the species of Anthurium and 72% of the species
1572 Galeano et al.
of Philodendron are epiphytes; this supports Gentry’s (1982a) observation that infrageneric diversi®cation is most common among epiphytes. The diversity of Araceae recorded here (100 species in 0.9 ha) seems to be the highest ever recorded for such a small area. Croat (1992) stated that the aroid ̄ora of Colombia is the highest of any region in the world and that most of the species must be concentrated in western Colombia. Even La Selva, in Costa Rica, well known as rich in Araceae, has an aroid ̄ora as rich as the one recorded here (99 species), although the area of natural forest at La Selva is about 1000 times as large as our sample (Hartshorn and Hammel, 1994). The number of species of Cyclanthaceae found in our study (30 species in 0.9 ha) is probably also a record for this family; this supports the view that this family has its main center of diversi®cation in the ChocoÂ (Harling, 1958).
The number of individuals in 0.1 ha at E1 Amargal was lower than the existing records for wet and dry forests in western Ecuador, 7210 and 5428 individuals per 0.1 ha, respec- tively (Gentry and Dodson, 1987). The main dierences between these sites and E1 Amargal were the strikingly higher density of epiphytes in the wet forest at RõÂ o Palenque (although the species richness of this growth form was similar), and the higher density of herbs in the dry forest. Interestingly, the number of trees 310cm dbh and its relative density at E1 Amargal were almost the same as at RõÂ o Palenque (Gentry and Dodson, 1987). The density of terrestrial herbs at E1 Amargal (1704 individuals per 0.1 ha) was, however, higher than that recorded by Gentry and Dodson (1987) at RõÂ o Palenque (1220 individuals in 0.1 ha) and by Poulsen and Balslev (1991) in Amazonian Ecuador (813±1322 individuals per 0.1 ha). The contribution of palms to forest structure at the study area is apparent from their relative density (26.4% of all individuals). Palms not only characterize the woody com- ponent (Galeano et al., in press; Galeano, unpubl. data) but they are also dominant across the vertical gradient, especially in the understory.
The total number of species in the study area is higher than recorded in the small transects as shown by the species±area curve (Fig. 1). Considering the small area sampled (0.9 ha), the total number of species (970 species) is high compared with the number of vascular species recorded for larger areas, e.g. Gorgona island (in the same ChocoÂ region) with an area of 15.7 km2, has a known ̄ora of 512 species (Barbosa, 1986; Murillo and Lozano, 1989; Rangel and Aguirre, 1990); Barro Colorado Island, 15.6 km2, has 966 forest species (Croat, 1978; Foster and Hubbell, 1990); La Selva, Costa Rica, in about 14 km2has 1458 native species (Hartshorn and Hammel, 1994; McDade and Hartshorn, 1994). Most strikingly, the 970 species found in our sample represent about 25% of all species recorded in the checklist of the ChocoÂ Department (Forero and Gentry, 1989) which encompasses an area of more than 5 million times as large as our sample. This stresses the need of many more collections and ̄oras from the ChocoÂ region. Another count, in the Amazon, recorded a similar richness (1223 species) in ten 0.1 ha plots (Duivenvoorden, 1994). Those plots, however, were located farther from each other and on very dierent soil types, which adds to the possibility of ®nding dierent species.
Comparing the few data available, the forests at the Gulf of TribugaÂ seem to be more similar in terms of growth form distribution, tree species richness and density, and ̄oristic composition at the family level, to the forest at La Selva, Costa Rica (Hartshorn and Hammel, 1994), and at RõÂ o Palenque, Ecuador (Gentry 1986; Gentry and Dodson, 1987), than to other sites so far studied in the Colombian ChocoÂ (Tutunendo and Bajo Calima: Gentry, 1986; Faber-Langendoen and Gentry, 1991). A similarity in ̄oristic composition between La Selva and RõÂo Palenque was pointed out by Gentry (1993).
Plant species count in ChocoÂ, Colombia 1573
The fact that most of the species were under 10 cm dbh, and even under 2.5 cm dbh (Fig. 5), supports the view that tree diversity and treelet or herb diversity are not always necessarily correlated (Gentry, 1982a), and that the earth’s richest plant community is the understory of the tropical rain forest (Gentry and Dodson, 1987; Hartshorn and Hammel, 1994). It is, then, clear that a classical forest inventory, including only individuals 310 cm dbh (and less often individuals 32.5 cm dbh) is not at all the appropriate means to study total composition and species richness. Gentry and Dodson (1987) stated that tree species310 cm dbh constitute only 15±20% of the complete ̄oras of many neotropical sites. Even for many Amazonian forests where tree species prevail, trees 310 cm dbh scarcely account for 10±30% of the total number of species (Duivenvoorden, 1994). Inventories attempting to assess alpha diversity for conservation goals should, of necessity, also consider growth forms other than large trees. If this is not done, by far the greatest part of the species richness will not be taken into consideration.
This paper is part of G. Galeano’s Ph.D. dissertation at the Department of Systematic Botany, Biological Institute, Aarhus University, directed by Henrik Balslev. Financial support was given by DANIDA (Grant no. 104.DAN.8.4/201), Fondo FEN-Colombia, The MacArthur Foundation, INDERENA, Universidad Nacional de Colombia, and FundacioÂn InguedeÂ. We thank L. GarcõÂa, J. Grueso, C. HernaÂndez, O. Hurtado, P. Hurtado, A. Moreno, G. Moreno, J. Moreno, N. Moreno, I. Perea, L. Perea, D. Tuberquia, F. Valencia and E. Ayazo for ®eld assistance. Logistic support was given by Instituto de Ciencias Naturales, Universidad Nacional de Colombia, the Department of Systematic Botany and the herbarium at Aarhus University, Denmark. L. Allorge, M. Amaya, L. Andersson, C.C. Berg, J. Betancur, R. Callejas, L. Clark, J.L. FernaÂ ndez, A. GoÂmez, P. Maas, R.T. Pennington, J. Sarmiento, and C. Taylor helped with identi®cations. We thank A. Barfod, R. Bernal, A. Cleef, J. Duivenvoorden, R. Foster, and P. Hall for reviewing the manuscript.
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