BIOLOGICAL CONSERVATION 136 (2007) 372– 387

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Local illegal trade reveals unknown diversity and involves a high richness of wild vascular epiphytes

Alejandro Flores-Palacios*, Susana Valencia-Dı´az

CEAMISH, University Autonomous of Morelos State, Av. Universidad 1001, Col. Chamilpa, Cuernavaca, 62209 Morelos,

ARTICLE INFO ABSTRACT

Article history: Traffic of and animals of wild origin is a major threat to biological diversity creating Received 31 May 2006 a need for legal protection of traded species. Epiphytic vascular plants are a diverse guild Received in revised form which provides the horticultural market with many species. Many epiphytes are under pro- 18 October 2006 tection intended to stop illegal trade at international and local levels. To measure the rich- Accepted 8 December 2006 ness and volume of the epiphytes that are traded illegally on a local level, we monitored an Available online 12 February 2007 illegal sale point in Xalapa, Veracruz, Mexico for 85 weeks. We recorded 27 traders, 207 spe- cies and 7598 plants. Nineteen species were known for Mexico but not native to Veracruz; Keywords: two others had been collected in Veracruz but had never been reported before for Mexico. About 25% of epiphyte richness in Veracruz and 47% of the State’s epiphytic orchids are Harvest traded illegally. Most of the species traded could have been harvested in lower montane Non timber forest products cloud forest (105 species), however all vegetation types of central Veracruz are potential epiphyte sources. Twenty-seven species protected by Mexican law were traded, along with Traffic 41 species endemic to Mexico and six endemic to Veracruz. We concluded that on a local level, the richness and volume of epiphytes illegally traded are high; in fact, the volume of orchids traded equals the annual average volume of legal Mexican orchid exports. Illegal epiphyte trade occurs worldwide, making necessary crucial increases of protective mea- sures and the development of sustainable ways of harvesting epiphytes. Ó 2007 Elsevier Ltd. All rights reserved.

1. Introduction pets and in plants with medicinal or horticultural use (e.g. Vovides et al., 1997; McMahan and Walter, 1989; SEMARNAP, Legal protection and regulation of biodiversity is intended to 1997; Herna´ndez and Go´ mez-Hinostrosa, 2002); however, we stop unsustainable forms of exploitation by restricting the lack of data to understand the impact and magnitude of ille- extraction of protected species and certifying those plants gal trade on any scale (Blundell and Mascia, 2005). and animals raised or propagated legally for market. Interna- In Mexico there has been an academic effort to develop tional treaties and national laws have been adopted by many strategies to protect biodiversity. This effort has supported countries to stop illegal trade at local, regional and interna- the development of natural protected areas; especially impor- tional scales (Walter and Gillett, 1998; De Grammont and tant was the development of Biosphere reserves, in which the Cuaro´ n, 2006). Even with these legal measures, illegal trade reserve inhabitants are involved in conservation activities of wild animals and plants remains frequent worldwide and and management practices (Halffter, 1984). Moreover, the spe- is considered one of the most profitable illegal activities for cialists have developed and constantly updated threatened some countries (McMahan and Walter, 1989; SEMARNAP, lists (e.g. Vovides, 1981; Vovides et al., 1997). As a partial 1997). Illegal trade is especially intense in animals used as result of this academic initiatives, Mexico signed the CITES

* Corresponding author: Tel./fax: +52 777 329 7019. E-mail addresses: alejandro.fl[email protected] (A. Flores-Palacios), [email protected] (S. Valencia-Dı´az). 0006-3207/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2006.12.017 BIOLOGICAL CONSERVATION 136 (2007) 372– 387 373

convention in 1991, and more recently Mexican authorities Table 1 – Number of animal and plant species protected developed an official list of threatened species (SEMARNAT, by the Mexican law (SEMARNAT, 2002) and the number of 2002). However, illegal trade is still recognized as a major species in each group in Mexico threat to biodiversity (SEMARNAP, 1997), indicating that these Group Number of Number of Percent initial protection measures have been insufficient and a bet- species species in protected ter understanding of the magnitude of illegal trade is urgently protected Mexico needed. a Epiphytic plants are a diverse guild characteristic of tropi- Fish 186 2122 8.8 b cal forests and represent a substantial proportion of the spe- Amphibians 197 285 69.1 Reptilians 466 693b 67.2 cies richness in those ecosystems (Wolf and Flamenco, 2003). Birds 371 1076c 34.5 Many epiphytic species are important in the international Mammals 295 525d 56.2 horticultural market, especially species from Araceae, Brome- Plants 980 24 601e 4.0 liaceae, Orchidaceae and ferns. Consequently, tropical forest Cactaceae 285 570f 50.0 canopies are sources of valuable non-timber forest products Orchidaceae 181 1150g 15.7 h (NTFP) (Verhoeven and Beckers, 1999). As with other groups, Palmae 64 100 64.0 Zamiaceae 43 48i 89.6 eradication or extinction of epiphytic species are affected by Agavaceae 39 198j 19.7 a combination of deforestation, fragmentation, and illegal k Pinaceae 36 60 60.0 collecting (Koopowitz, 1992; Turner et al., 1994; Williams-Line- Bromeliaceae 21 326l 6.4 ra et al., 1995; Olmsted and Jua´rez, 1996; Sosa and Platas, 1998; Another 96 311 21 873e 1.4 Pickens et al., 2003). In Singapore, for example, epiphytes ac- plant families counted for the largest number of plant extinctions or eradi- The number of Mexican species was taken from: cations (Turner et al., 1994). In Yucatan State, Mexico, 22 a Pe´rez et al. (1998). epiphytic orchid species have been eradicated (Olmsted and b Flores-Villela (1998). Jua´rez, 1996), while in Veracruz, six epiphytic orchid species c Ceballos et al. (2000). have been eradicated and one is proposed as extinct (Sosa d Ceballos and Oliva (2005). and Platas, 1998). e Espejo-Serna et al. (2004a). f Herna´ndez and Go´ mez-Hinostrosa (2002). Some epiphytes can be collected when populations are g Espejo-Serna and Lo´ pez-Ferrari (1997, 1998). large enough (Wolf and Konings, 2001; Pickens et al., 2003). h Quero (1994). Unfortunately, the methods used to collect NTFP are aimed i Vovides (2000). at the quickest profit, depleting the resources. Exception are j Garcı´a-Mendoza (1995). cases in which the people involved depend heavily on them, k Narave-Flores and Taylor (1997). follow traditional management practices, or have ecological l Espejo-Serna et al. (2004b). knowledge (Ticktin and Johns, 2002; Shaanker et al., 2004). The only study that documents changes in harvested vascu- lar epiphyte populations is that of the fern Platycerium 2. Methods stemaria (P. Beauv.) Desv. (Polypodiaceae); where commercial harvest reduced plant frequency on trees and modified 2.1. Study area plant distribution in cocoa plantations (Porembski and Bie- dinger, 2001), suggesting that collection pressure reduced The research took place in the Xalapa-Coatepec area in cen- abundance and restricted sites where the species can tral Veracruz, Mexico (Fig. 1). The region features intense hor- survive. ticultural activity, with wild plants sold legally and illegally. Epiphytes, especially orchids, are protected plants, both The zone is mountainous (Fig. 1), and lies within an hour’s nationally and internationally. In Mexico, the national list of drive of the coast. Therefore, many different vegetation types protected plants covers 980 species (SEMARNAT, 2002), of are easily accessible from Xalapa-Coatepec, including pine which at least 19% are epiphytes; Orchidaceae has the second forest, lower montane cloud forest, oak forest, and tropical highest number of protected species among plant families dry forest. The coastal area of central Veracruz contains man- (Table 1). Despite this legal protection, wild epiphyte traffic groves, tropical evergreen forest, and sand dune vegetation is common in markets and streets (e.g., Aguilar, 2001; Peralta (Rzedowski, 1978). and Torres, 2004); ironically, some epiphyte species have been A partial flora for the state of Veracruz exits (Sosa and reported for some regions on the basis of this activity (e.g., Go´ mez-Pompa, 1994) and can be complemented by more re- Cha´vez and Arenas, 1996). In fact, some protected species cent lists of Orchidaceae (Espejo-Serna and Lo´ pez-Ferrari, (e.g., the orchid Laelia speciosa (Kunth.) Schltr.) are subject to 1997, 1998), Bromeliaceae (Espejo-Serna et al., 2004b), Pterido- systematic collection. No studies have focused on local epi- phyta (Palacios-Rı´os, 1992, Melhtreter unpub. data), and a phyte traffic, which makes it impossible to assess the effec- partial guide to the epiphytes of Veracruz (Hietz and Hietz- tiveness of protection measures or know whether the only Seifert, 1994); according to these sources, Veracruz features species affected are those of high horticultural value. We at least 604 epiphytic plant species. The most diverse groups hypothesized that if legal protection is effective, then the ille- of epiphytes are Orchidaceae (262 species), Pteridophyta (168 gal trade of wild harvested epiphytes will be low, infrequent at species), and Bromeliaceae (68 species). In different localities any scale and that those plants harvested will be offered of central Veracruz the ecology of epiphyte communities has rooted to conceal any signs of wild origin. been studied (e.g. Hietz and Hietz-Seifert, 1995; Garcı´a-Franco, 374 BIOLOGICAL CONSERVATION 136 (2007) 372– 387

Fig. 1 – Central area of Veracruz, Mexico. The area surrounding Xalapa-Coatepec is mountainous with altitudes between 1200 and 1400 m asl; however, the Gulf of Mexico is ca. 60 km east of Xalapa, and 20 km west of Xalapa the zone reaches elevations of up to 4200 m asl.

1996; Flores-Palacios, 2003), and it has been estimated that the individual plants are homogeneous in size and are clean these plants constitute up to 32% of the flora in at least one and potted. Thus, we concluded that a specimen had been ta- municipality (Flores-Palacios, 2003). The results of these stud- ken from the wild when it was not planted, had accompanying ies, along with personal observations, suggest that there are flora (e.g. bryophytes, epiphylls, ferns, lichens, seedlings), at least 297 epiphyte species in central Veracruz, 133 of which scars caused by herbivores, and if the cohort was composed are Orchidaceae, 63 Pteridophyta, and 42 Bromeliaceae. of plants of different sizes (e.g. seedlings, adults). In the case of potted orchids, we concluded that a specimen had been 2.2. Data collection taken from the wild if the older pseudobulbs were not rooted to the substrate, were disoriented from the pot (or even bur- A ‘‘tianguis’’ is a temporal market place. From July 6th, 2001 to ied), showed signs of herbivory, or had accompanying flora. April 18th, 2003, we visited the Leyzegui tianguis (hereafter the We recorded only ornamental species and ignored epiphyte tianguis) every Friday around 09:00–11:00 h. The tianguis has species sold for food (e.g. the cacti Hylocereus undatus operated every Friday for the last 30 years, and according to (Haw.) Britton and Rose), and the bromeliad usneoides market managers and our own experience, wild flora has been (L.) L., which is sold in huge quantities during Christmas sold since 1988. The first eight weeks were allocated approach- season. ing and familiarizing ourselves with the traders. Following When we could not identify plants at the marketplace, we this period, we established informal conversation with the obtained flowers or entire individuals for subsequent identifi- traders to obtain their names, town of residence, whether they cation. Such specimens were deposited at the Instituto de Eco- collected the plants they offered, the harvest localities and the logı´a A. C. herbarium (XAL), or at that of the Universidad plant names (scientific and common). During each visit, we Auto´noma del Estado de Morelos (HUMO), or the Universidad Auto´- identified and recorded the plants offered, the number of indi- noma Metropolitana-Iztapalapa (UAMIZ). For most species, we viduals per species, prices (in Mexican pesos), the presence of made a photographic record. Additional data were obtained reproductive structures, if plants were offered whole or in from an anonymous collector who allowed us to examine the pieces (cuttings), and whether they were cultivated. A cohort plants he purchased every week and informed us of their of artificially propagated plants can be recognized because prices. BIOLOGICAL CONSERVATION 136 (2007) 372– 387 375

We determined the habit (epiphyte, terrestrial, or saxico- mal sites and five regular sale points. There were only four lous), level of endemism (Mexico or Veracruz) and the primary informal sites and two regular sale points where they did type of vegetation where each species grows. During the not participate; in most cases these infrequent sites involved study period, the mean exchange rate was 9.7 ± 0.6 Mx pesos peddlers who occasionally sold an epiphyte, except for one for 1 US dollar (minimum 9.0 and maximum 11.2; Bank of site in front of the Guadalupe Church in Coatepec, where Mexico, http://www.banxico.gob.mx). the trader knew the bromeliads and orchid species. We identified those species currently protected in Mexico Plant prices varied widely (Appendix) and depended on (SEMARNAT, 2002). Mexican law recognizes four protection species, size, and the presence of flowers. The average price categories: probably extinct in the wild (Ex), in danger of was $23 ± 41 per plant (n = 716; minimum $1, maximum extinction (DE), endangered (EE), and subject to special protec- $500); however, certain species could be sold in small groups tion (Pr). These categories are partially equivalent to those of (all orchids of the genera Lepanthes, Pleurothallis, and Trichosal- the IUCN 1994 red list of threatened species; Ex is similar to ex- pinx) while others were very valued, such as the white forms tinct in the wild; DE with critically endangered and endan- of the orchid Laelia anceps Lindl. ($500) and species not native gered; EE with vulnerable; and Pr with at lower risk (Walter to Veracruz (Appendix). and Gillett, 1998; SEMARNAT, 2002). In Mexico, it is a requisite to obtain a government authorization for a commercial exploi- 3.2. Species richness tation of wild species (General Law of Ecological Equilibrium). Exploitation of protected species is forbidden, but the govern- On average 19 species of plants were sold each week (Table 3). ment could confer an authorization when there is a manage- Orchids accounted for the greatest richness every week (Table ment program; which would warrant the reproduction and the 3). The highest number of species sold was in June (Fig. 2b). development of wild populations. We asked to the traders if There was a positive correlation between the number of spe- they had already this authorization. cies sold and the number of traders (r = 0.76, P < 0.001). To ascertain other epiphyte marketplaces in the region, we We recorded a total of 207 species (Table 4, Appendix). interviewed private collectors and visited the towns of Coate- Nineteen were epiphytic orchid species non-native to Vera- pec, Xico, and Teocelo (Fig. 1). We were thus able to determine cruz and collected from other regions of Mexico (Appendix). whether or not merchants in the tianguis were also working Two species were collected in Veracruz but not currently re- in other areas. Species names follow the nomenclature of ported for Mexico (Pleurothallis cf. dodsoni Luer and P. cf. herre- Sosa and Go´mez-Pompa (1994); Espejo-Serna and Lo´ pez-Fer- rae Luer) (Fig. 4). These two species were collected and sold by rari (1997, 1998), and Espejo-Serna et al. (2004b). a Cosautla´n vendor who collected her own merchandise and assured us that they came from the municipality of Cosa- 3. Results utla´n, Veracruz. We found 27 species that are protected in Mexico, all of 3.1. Vendors, markets, and prices them epiphytes (Appendix); 17 are considered in danger of extinction (one Bromeliaceae and 16 Orchidaceae) and 10 are No one vendor had a government authorization to harvest subject to special protection (one Bromeliaceae and nine and sell wild flora. Vendors were grouped according to their Orchidaceae). Forty-seven species were endemic (Appendix); place of origin into (a) Chiltoyac, (b) Coatepec, (c) Cosautla´n 41 endemic to Mexico (36 epiphytic orchids, four epiphytic bro- and (d) other regions or independent vendors (see Table 2 meliads, and one terrestrial bromeliad) and six endemic to for group characteristics). Each week, 6 ± 3 vendors (mean ± 1 Veracruz (one terrestrial bromeliad and one terrestrial cactus, SD; n = 78, minimum 0, maximum 14) sold wild vascular epi- one epiphytic bromeliad and three epiphytic orchids). phytic flora at the tianguis. The number of vendors reached Orchidaceae comprised 81% of the species (Table 4); the its peak in June (Fig. 2a); only on Good Friday (March 29, rest belonged to six other families. Of all species, 84% were 2002) they were absent (Fig. 2a). epiphytes (mainly orchids, 70%), and the rest were saxicolous Apart from the tianguis, we identified 15 places where wild (2%) or terrestrial plants (14%). Although no vendor revealed epiphytes were sold in the Xalapa-Coatepec region (specific the exact collection sites, most species were probably from list can be requested from the authors). Of the 15 wild epi- the lower montane cloud forest (Fig. 3), but species from all phyte sale points, tianguis traders participated in four infor- vegetation types of central Veracruz were sold.

Table 2 – Characteristics of the groups of epiphytes traders registered in a tianguis of Xalapa, Mexico

Chiltoyac Coatepec Cosautla´n Independent

Number of persons 4 4 4 15 Frequency of attendance at the tianguis 15% 92% 30–70% 49–73% Do they know scientific names? Yes Yes No Not the majority Do they collect the plants in the field? Yes Yes Yes Yes Do they pay others to collect the plants? No Yes No No Do they buy plants at the tianguis? No Yes No No Do they sell artificially propagated plants, too? No Yes No No Do they have a nursery? No Yes No No 376 BIOLOGICAL CONSERVATION 136 (2007) 372– 387

a 15 Table 3 – Mean number of species and plants illegally traded in a tianguis of Xalapa, Mexico, from July 6th, 2001 to April 18th, 2003 12 Mean ± SD n Min.–max.

Number of species 9 Araceae – 82 0–1 Bromeliaceae 1.1 ± 1.5 82 0–7 Cactaceae 0.3 ± 0.6 82 0–3 6 Crassulaceae 0.1 ± 0.3 82 0–2 Lentibulariaceae 0.1 ± 0.3 82 0–1 Number of Vendors Lycopodiaceae 0.6 ± 0.8 82 0–3 3 Orchidaceae 16.7 ± 10.6 82 0–45 All families 18.9 ± 12.3 82 0–51

0 Number of plant or plant divisions 7/28/01 11/5/01 2/13/02 5/24/02 9/1/02 12/10/02 3/20/03 Araceae – 80 0–2 Bromeliaceae 2.6 ± 5.2 80 0–1 b 55 Cactaceae 1.0 ± 2.8 80 0–20 50 Crassulaceae 0.2 ± 0.7 80 0–4 Lentibulariaceae 1.0 ± 4.1 80 0–29 45 Lycopodiaceae 1.5 ± 2.5 80 0–12 40 Orchidaceae 88.4 ± 79.0 80 0–354 All families 95.7 ± 82.3 80 0–382 35

30 The n value refers to the number of weeks from which the mean was obtained. 25

20

Number of species 15

10

5 Table 4 – Number of species illegally traded in a tianguis of Xalapa, Mexico, from July 6th, 2001 to April 18th, 2003 0 7/28/01 11/5/01 2/13/02 5/24/02 9/1/02 12/10/02 3/20/03 Epiphytes Saxicolous Terrestrial All habits c 400 Number of species 350 Araceae 1 – – 1 Bromeliaceae 20 2 3 25 300 Cactaceae 5 – 3 8 Crassulaceae 1 – 1 2 250 Lentibulariaceae – – 1 1 200 Lycopodiaceae 3 – – 3 Orchidaceae 144 3 20 167 150 All families 174 5 28 207 Number of plants 100 Number of plants or plant divisions Araceae 2 – – 2 50 Bromeliaceae 180 6 22 208 Cactaceae 13 – 67 80 0 Crassulaceae 9 – 3 12 7/28/01 11/5/01 2/13/02 5/24/02 9/1/02 12/10/02 3/20/03 Lentibulariaceae – – 80 80 Date Lycopodiaceae 116 – – 116 Orchidaceae 6763 167 170 7100 Fig. 2 – Number of vendors (a) species (b) and plants (c) sold All families 7083 173 342 7598 in a tianguis of Xalapa, Mexico, each Friday during July 6th, 2001 to April 18th, 2003. The arrows point out the two consecutive dates when confiscations took place by the Mexican Federal Commission on Environmental Protection. number of plants was positively correlated to the number of species (r = 0.80, P < 0.0001) and vendors (r = 0.71, P < 0.001). 3.3. Species abundance Although all the species could be sold either complete or as cuttings, most (186) were sold as entire plants, with only On average over 90 plants or cuttings were sold each week 21 species sold as cuttings. Nearly all (99.6%) of the plants (Table 3). As with richness, the weekly average volume of were not planted; their roots were visible and there were clear the plants offered was primarily orchids (Table 3). The high- signs that they had been removed from the wild. Only some est number of plants was in June (Fig. 2c). Each week, the individuals of 10 orchid species not native to Veracruz were BIOLOGICAL CONSERVATION 136 (2007) 372– 387 377

Sand dune concentrated there. The total abundance of trafficked orchid plants found in our survey is equal to the average annual Mangrove (1975–2004) quantity of orchid plants exported by Mexico TEF (7193 ± 7211 plants/year) (UNEP-WCMC CITES trade data- Pine base, http://sea-swift.unep-wcmc.org) and is similar to the pooled 1994–1996 cacti confiscations at the airports of Mex- Not know ico, Germany, France and the United States (Herna´ndez and TDF Go´ mez-Hinostrosa, 2002). However, our data of numbers of Oak traders, sale points, richness and quantity of plants offered, should be considered as the minimum because we did not LMCF visit established greenhouses near or within the zone. 0 10 20 30 40 50 60 70 80 90 100 110 Number of species 4.1. Species richness

Fig. 3 – Number of species, grouped by vegetation type, sold Although the species richness and abundance of plants re- in a tianguis of Xalapa, Mexico, during July 6th, 2001 to April corded was high; it underestimates the real value and are 18th, 2003. LMCF, lower montane cloud forest; Mangrove, probably the minimum estimates. Our study had two sources mangrove forest; Oak, oak forest; Pine, pine forest; of error that could lead to underestimates of the richness and Sand-dune, sand-dune vegetation; TDF, tropical dry forest abundance: both the time of our visits and taxonomical and TEF, tropical evergreen forest. uncertainties. Before our arrival, specialized collectors and intermediary traffickers had already made purchases, partic- ularly with rare species. The more substantial source of potted (Cuitlauzina pendula Lex, Encyclia denocaula (Lex.) Schltr., underestimation was taxonomic uncertainty. Many species E. spatella (Rchb. f.) Schltr., L. crawshayana Rchb. f., L. eyerman- in the region are similar in vegetative structure, making iden- iana Rchb. f., L. rubescens Lindl., O. unguicullatum Lindl., Osmo- tification difficult without flowers. At least six names on our glosum pulchellum (Bateman ex Lind.) Schltr., Prosthechea list reflect similar groups of species that we preferred to treat linkiana (Kl.) W.E. Higgins, and P. maculosa (Ames, FT Hubb. as one species in order to decrease errors when recording and Schweinf.) W.E. Higgins); however, evidence of wild origin data at the market. However, only nine species could be was present, indicating they were likely leftovers from previ- occluded due to this policy. ous collection efforts. Harvest efforts by traffickers have revealed an orchid In total, we counted 7598 plants or cuttings (Table 4). Of diversity of Veracruz that has been either poorly docu- these, 93% were epiphytes, mostly Orchidaceae (89%). Three mented (e.g. Amparoa beloglossa (Rchb. f.) Schltr.), recently re- species were sold in quantities above 300 plants (Appendix), ported (e.g. Pleurothallis lateritia Endres and Rchb. f.), reported while over 200 plants were sold of four species, and over as extinct (Encyclia flabellate (Lindl.) B. Thurst. and W. 100 plants were sold of 13 species. Less than 100 plants or cut- Thurst.), or new to Mexico (Pleurothallis cf. dodsoni and P. cf. tings were sold of the remaining species (Appendix). herrerae)(Fig. 4). Amparoa beloglossa has been reported for On two consecutive Fridays (July 26th and August 2nd, the state of Veracruz based on one specimen without data 2002), there were confiscations by the Mexican Federal Com- (Jime´nez-Machorro, 2001), and P. lateritia was reported for mission on Environmental Protection. Confiscations reduced the first time for both Mexico and Veracruz in 2002 (Soto, both the number of vendors (especially those from other re- 2002); however, both species were sold abundantly and fre- gions and from Cosautla´n) and plant quantities (Fig. 2c) but quently during the study period. We recorded only two not the richness (Fig. 2b). In the weeks following the confis- plants of Encyclia flabellata, an orchid species endemic to cation, sellers from Coatepec and Cosautla´n continued offer- central Veracruz, reported as possibly extinct (Sosa and Pla- ing plants, but in smaller quantities and furtively, hiding tas, 1998); our results suggest that the species is rare but not plants only to show them to habitual visitors or customers extinct. they trusted. Four weeks after the confiscation, those who Orchids, constituted the main group sold illegally. The had stopped selling were back and offering their plants trafficked richness we recorded accounts for 25% of epiphyte again. richness for all Veracruz and 59% for the central part of this state. For epiphytic orchids, 47% of the species richness for 4. Discussion the entire state was recorded and 93% of the species rich- ness was recorded for the central part of Veracruz. However, Despite legal protection, epiphytes are sold illegally, regularly, of the total orchid species recorded, only 17 are protected in plain daylight, without being rooted and with clear signs of species. Although Mexican law protects 16% of all Mexican wild origin. Collection pressure is exerted not only on species orchid species (SEMARNAT, 2002), our data show that field of high horticultural value but also on species of interest only extraction includes a substantial proportion of species both to collectors (e.g. Pleurothallidinae). Not only is the quantity at the state and regional level; clearly, legal protection of plants sold high, but trafficking is revealing unknown should cover more species. Furthermore, many of the spe- biodiversity. cies sold are plants without high horticultural value and The tianguis can be considered one of the major market- have small flowers that only specialized collectors appreci- places in the region due to the great diversity and volume ate (e.g. Pleurothallidinae). 378 BIOLOGICAL CONSERVATION 136 (2007) 372– 387

Fig. 4 – (A) Amparoa beloglossa is a Mexican endemic orchid already known for the state of Veracruz but with unknown distribution inside Veracruz (scale bar = 1 cm); (B) Pleurothallis cf. herrerae an orchid species illegally traded but unrecorded for Mexico (scale bar = 1.75 cm); (C) Encyclia candollei (right) and E. flabellata (left), the last orchid species was proposed as extinct, but our data show that the species is not extinct and is illegally traded; E. flabellata can be distinguish from E. candollei because it has a fanlike obscurely lobed lip, while E. candollei has a three-lobed lip (scale bar = 1.5 cm ); (D) Pleurothallis cf. dodsoni a species illegally traded but unknown for Mexico (scale bar = 1.4 cm); (E) The main arch decorating the main church of Coatepec during the 2001 San Jero´ nimo festivities, this arch measured 3.5 m wide and 10 m height and contained 1623 inflorescences of the bromeliad Tillandsia multicaulis.

4.2. Species abundance dors’ preference for certain species from the species abun- dance in the wild. For example, we know that some species Although only 17 orchid species were seen sold at large-scale are highly coveted in the region, but we recorded few plants (more than 100 plants/species), the abundance data is difficult sold (e.g. Cuitlauzina pendula). The low abundance of such spe- to understand because we can not distinguish between ven- cies relates to the fact that during that particular year, the BIOLOGICAL CONSERVATION 136 (2007) 372– 387 379

plants sold may have been leftovers from previous harvest ef- Konings, 2000) and includes more species than just epiphytes; forts. Other species are widely distributed and very common for example, for cycads Donaldson et al. (2003) give a global in Veracruz, but were registered as low abundance, possibly overview of trade and sustainable use, Vovides et al. (2002) indicating species that are not popular (e.g. Prosthechea ochra- outline sustainable use of cycads in Mexico. In Me´xico illegal ceae (Lindl.) W.E. Higgins). We recorded large numbers of other traffic has been reported for many cities such as Oaxaca species for which distribution data do not exist, or that are (Mijangos and Gil, 2004), Mexico City, Puebla (E. Aguirre Leo´n, widely distributed but usually in small populations (e.g. Acin- pers. obs.), Morelia, Michoaca´n(Peralta and Torres, 2004), and eta barkeri (Bateman) Lindl.). To better interpret the abun- Cuernavaca, Morelos (A. Flores-Palacios unpublished data). dance of illegally-sold plants, we need to update species Extensive documentation is therefore needed on how such distribution maps and document abundance in Veracruz. De- activities are affecting diversity and the survival of the har- spite our underestimation, the total abundance of some vested populations, especially since government confisca- traded species in the tianguis is higher than the annual aver- tions underestimates the real volume of plants illegally age exportation (1975–2004) by Mexico (e.g. Tillandsia spp., traded. We suggest that the following actions will help to con- Acineta barkeri, Stanhopea tigrina) (UNEP-WCMC CITES trade trol illegal trade: database, http://sea-swift.unep-wcmc.org). An alarming fact is that all species are trafficked when 1. Increases the list of protected species. Although there are blooming, and the large majority of specimens were not potted presently many protected species, it is necessary preven- or rooted in a substrate. This practice could have two important tive actions that allow the protection of more species. consequences: it diminishes the likelihood of a plant’s survival The list of protected flora in Mexico only represents 4% outside its habitat, especially if the buyer is not a skilled of the country’s entire flora, while the percentage of pro- grower; and second, it leads to a reduction in reproductive tected land vertebrates is higher, ranging between 34% plants in the harvested population. The growth rate of epiphyte and 69% (Table 1). Among plant families, those with the populations is strongly influenced by adult survival (Zotz et al., most protected species are Cactaceae and Orchidaceae 2005; Zotz, 2005; Zotz and Schmidt, 2006), clonal growth (Mond- (Vovides et al., 1997; SEMARNAT, 2002), but relative to the rago´ n et al., 2004), and the minimum number of seeds pro- total number of species per family they have, they do not duced can limit new recruitment (Ackerman et al., 1996). For qualify as the most protected (Table 1). Maybe one action some species, the blooming rate in wild populations is low or is to recognize that those species listed in CITES should highly variable; for example, in central Veracruz from 33% to be considered protected inside countries as Mexico and 52% of all plants bloom each season in populations of the not only for international trade. non-autogamous orchid Ryncholaelia glauca (Lindl.) Schltr., 2. Increase the surveillance in local markets. Surveillance and they reach a fruit set of 6–14% (Flores-Palacios and Gar- and confiscation are temporary measures for controlling cı´a-Franco, 2003). Fruit set in orchids tends to be low (Neiland illegal traffic. Permanent surveillance is needed along with and Wilcock, 1998), and if the abundance of flowers is reduced, stricter punishments for plant traffickers to instigate fruit production could drop (Schemske, 1980; Zimmerman and reduction or stop the selling of wild plants. Aide, 1989; Calvo, 1990). Although specific research is needed to 3. Develop management strategies for wild populations. It is confirm our suspicions, our data suggest that illegal trafficking necessary to develop management strategies that allow could be exerting pressure on adult survival, clonal growth, the sustainable use of epiphytes as have been done with and reproduction, with possible negative demographic conse- bromeliads (Ticktin and Johns, 2002; Pickens et al., 2003; quences for the harvested populations. Wolf et al., 2006) and other groups of plants, such as cycads Although most of this plant traffic was orchid related, the (Vovides and Iglesias, 1994; Vovides et al., 2002). These study area is characterized by other sources of harvest pres- strategies should incorporate the owners of the resource sure for epiphytes, especially for some bromeliads. For exam- (Halffter, 1984; Vovides et al., 2002; Wolf et al., 2006), pro- ple, in October 2001, during the San Jero´ nimo festivities in viding them with harvest estimations (Wolf and Konings, Coatepec, the main church was decorated with three arches, 2001), horticultural and propagation protocols (Pickens in which we counted 2438 inflorescences of Tillandsia multi- et al., 2003; Wolf et al., 2006), while the species under man- caulis Steudel, 109 of T. punctulata Schltdl. and Cham., and agement are monitored demographically (see Vovides 12 of T. imperialis E. Morren ex Roezl (Fig. 4d). The construction et al., 2002; Wolf et al., 2006). A propagation program could of such arches is a common practice in the region. Although help to sustain the harvest of wild flora such has been our data indicated that many bromeliads suffer little pressure achieved for cycads by Vovides and Iglesias (1994) and from horticultural collection, the former scenarios may threa- Vovides et al. (2002), through the introduction of artificially ten them, also affecting both clonal growth and sexual repro- propagated plants into the wild populations (increasing duction and this practice occur in others regions of the rate of population growth), lowering the rate of harvest Mesoamerica (Pickens et al., 2003). and reintroducing plants to areas where the species were removed; however, few such experiences have been under- 4.3. Conservation measurements and research priorities taken with epiphytes (e.g. Pickens et al., 2003; Yam and Thame, 2005; Wolf et al., 2006). Illegal traffic of wild flora is a phenomenon that occurs world- 4. To promote the artificial propagation of species with low wide (McMahan and Walter, 1989; Ferna´ndez, 2005; Rao, 2005; population densities. As has been pointed out by Wolf Shakya and Bajracharya, 2005; Vovides et al., 1997; Wolf and et al. (2006), the harvest of many epiphyte species is not 380 BIOLOGICAL CONSERVATION 136 (2007) 372– 387

possible because of their low population densities. For this come is that a close look at illegal trafficking revealed the species the development of artificial propagation methods insufficiency of diversity data. It is imperative to intensify is the only option. botanical research in order to ascertain the distribution and 5. Research the impact of illegal collection on wild popula- abundance of traded species. Also, it is important to find tions. While we do not know the extent to which illegal molecular markers that permit us to grasp the phylogeogra- trade is endangering the wild populations of epiphytes, phy of these species in order to reintroduce confiscated spec- we suspect that it has a negative effect because unman- imens into appropriate wild populations (Morrison et al., aged harvesting of wild resources often leads to depletion 2005). Otherwise, confiscation is nothing more than an action (e.g. Shaanker et al., 2004; Tang et al., 2005). Extinction that discourages illegal sales; unless the traffickers reveal operates through the elimination of single populations, their harvest sites, it would be impossible to reintegrate con- until the species disappear. The uncontrolled harvest of fiscated plants into their natural habitat; instead the confis- epiphytes could be promoting the eradications of popula- cating authorities send the plants to botanic gardens that tions, but further investigations are needed (but see Pic- act as custodians. kens et al., 2003). However, a seller told us he was unable to find more populations of the widespread orchid Pleuro- Acknowledgements thallis ornata Rchb. f. This is not an isolated example of the possible eradication of a population due to overhar- J.M. Jasso Parada lent valuable assistance and additional vesting; populations of many species, including plants observations during fieldwork. J.G. Garcı´a-Franco and R. Land- from the horticultural market (e.g. Withner, 1998; Herna´n- grave made the map. K. Mehltreter provided a list of epiphytic dez and Go´ mez-Hinostrosa, 2002), have suffered declines Pteridophytes for Veracruz. A. Espejo-Serna, A. Rosa Lo´ pez- or disappeared due to illegal collection, according to the Ferrari (Bromeliaceae), R. Palestina (Pteridophyte ), J. Garcı´a- reports of taxonomic authorities. Cruz, M.A. Soto-Arenas, and V. Sosa (Orchidaceae) helped with species identification. J.G. Garcı´a-Franco, A. Espejo-Serna, Vascular epiphytes are a vital, yet little known component I. Haeckel, I. Marquez, G. Williams-Linera, R. Primack and four of tropical forests. However, our data show that they are being anonymous reviewers provided comments and constructive subject to intense illegal collection. The most surprising out- criticism.

Appendix List of wild plants illegally trade in a tianguis of Xalapa, Veracruz

Species Habit Con. Endemic Price (Mexican pesos) Unit Number of Number of status to units weeks (x Æ SE) Min.–max.

Araceae Anthurium schlechtendallii Kunth Ep – – – – P 2 1

Bromeliaceae lueddemanniana Sax – – – 30.0 P 1 1 (K. Koch) Mez A. nudicaulis (L.) Griseb. Ep – – – 10.0 C 7 4 A. sp. Sax – – – 20.0 P 5 1 Catopsis sessiliflora (Ruı´z and Pav.) Mez Ep – – – 3.0–5.0 P 2 1 heterophylla (Lindl.) Beer Te – – – 10.0 P 7 2 P. ringens Klotzsch Te – Mexico – 25.0 P 2 2 P. schiedeana Baker Te – Veracruz 9.3 ± 0.7 8.0–10.0 P 13 7 Tillandsia brachycaulos Schltdl. Ep – – – 5.0–10.0 P 1 1 T. deppeana Steudel Ep – Mexico – 20.0 P 1 1 T. fasciculata Sw. Ep – – – 5.0 P 4 4 T. filifolia Schltdl. and Cham. Ep – – – 5.0 P 6 3 T gymnobotrya Baker Ep – Mexico 8.8 ± 2.4 5.0–15.0 P 11 10 T heterophylla E. Morren Ep – Mexico – 15.0–20.0 P 1 1 T. imperialis E. Morren ex Roezl Ep Pr Mexico – 20.0 P 6 5 T. ionantha Planch. Ep – – – 5.0 P 38 3 T. leiboldiana Schltdl. Ep – – 12.5 ± 2.5 10.0–20.0 P 12 9 T. lucida E. Morren ex Baker Ep – – – 5.0–20.0 P 2 2 T. multicaulis Steudel Ep – – 8.6 ± 1.0 5.0–10.0 P 26 12 T. palmasolana Matuda Ep – Veracruz – 5.0–10.0 P 6 4 T. polystachia (L.) L. Ep – – – 5.0–10.0 P 1 1 T. punctulata Schltdl. and Cham. Ep – – – 5.0–10.0 P 7 3 T. schiedeana Steud. Ep – – – 5.0 P 21 2 T. sp. Ep – – – 10.0 P 2 1 T. streptophylla Scheidw. ex E. Morren Ep – – 18.3 ± 8.8 5.0–35.0 P 25 12 T. tricolor Schltdl. and Cham. Ep EE – – 5.0 P 1 1 BIOLOGICAL CONSERVATION 136 (2007) 372– 387 381

Appendix – continued Species Habit Con. Endemic Price (Mexican pesos) Unit Number of Number of status to units weeks (x Æ SE) Min.–max.

Cactaceae Coryphantha sp. Te – – – 29.0 P 20 4 Epiphyllum oxypetalum (DC) Haw. Ep – – – 10.0 C 2 2 E. sp.1 Ep – – – 20.0 C 3 2 E. sp.2 Ep – – – 15.0–20.0 C 5 3 E. sp.3 Ep – – – 10.0 C 1 1 Hylocereus sp. Ep – – – 10.0 C 2 1 Mammillaria eriacantha Link Te – Mexico – 20.0–30.0 P 45 11 and Otto ex Pfeiff. Unknown columnar cacti Te – – – 10.0–30.0 P 2 1 Crassulaceae Echeveria rosea Lindl. Ep – – – 5.0 P 9 5 E. sp. Te – – – 5.0 P 3 1 Lentibulariaceae Pinguicula moranensis Kunth Te – – – 2.0–10.0 P 80 8 Lycopodiaceae Hupertzia myrsinites (Lam.) Trevis. Ep – – 7.5 ± 1.4 5.0–10.0 P 12 8 H. pringlei (Underw. and Ep – – 7.0 ± 1.2 5.0–10.0 P 6 6 F. E. Lloyd) Holub. H. taxifolia (Sw.) Trevis. Ep – – 10.3 ± 0.9 5.0–20.0 P 98 33 Orchidaceae Acineta barkeri (Bateman) Lindl. Ep EE Mexico 26.5 ± 4.7 10.0–50.0 P 109 29 Amparoa beloglossa (Rchb. f.) Ep EE Mexico 6.0 ± 2.1 3.0–10.0 P 33 10 Schltr Arpophyllum medium Rchb. f. Ep – – 8.3 ± 1.7 5.0–10.0 C 135 6 Barkeria scandens (La Llave Ep Pr Mexico – 50.0 P 5 3 and Lex.) Dressler and Halb. Beloglottis costaricensis (Rchb. f.) Te – – – 5.0 P 3 2 Schltr Bletia campanulata Lex. Te – – – 15.0–20.0 P 1 1 B. neglecta Sosa Te – Mexico – 10.0 P 44 5 B. purpurata A. Rich. and Galeotti Te – – – – P 4 1 B. purpurea (Lam.) DC Te – – – 15.0–20.0 P 8 2 B.sp. Te – – – – P 1 1 B. spp.a Te – – – 3.0 P 11 1 Brassavolla cucullata (L.) R. Br. Ep – – 10.0 ± 2.0 5.0–15.0 C 56 3 B. nodosa (L.) Lindl. Ep – – – – C 1 1 verrucosa Bateman ex Lindl. Ep – – 11.7 ± 1.1 5.0–20.0 C 110 21 Calanthe calanthoides (A. rich. and Te – – – – P 1 1 Galeotti) Hamer and Garay Campylocentrum schiedei (Rchb. f.) Ep – – – – P 2 2 Benth. ex Hemsl. Catasetum intergerrimum Hook. Ep – – 31.4 ± 9.3 10.0–60.0 P 22 10 Cattleya aurantiaca (Bateman Ep – – – 10.0 P 15 2 ex Lindl.) P.N. Don Chysis leavis Lindl. Ep – – 23.3 ± 4.4 15.0–30.0 P 6 5 Coelia macrostachya Lindl. Ep – – – 10.0 P 8 4 C. triptera (Sm.) G. Don ex steud. Ep – – 17.5 ± 4.3 10.0–30.0 P 13 6 Cohniella cebolleta (Jacq.) Sw. Ep – – – – P 5 5 Comparettia falcata Poepp. and Ep – – 3.0 ± 0.7 2.0–5.0 P 62 8 Endl. Cuitlauzina pendula Lex. Ep EE Mexico 86.7 ± 13.3 60.0–100.0 P 15 7 Cyrtopodium macrobulbon (Lex.) Ep – – – 20.0 P 15 5 G. Romero and Carnevali Deiregyne pyramidalis (Lindl.) Te – – – 5.0–10.0 P 2 2 Burns–Bal Dichaea glauca (Sw.) Lindl. Ep – – 13.8 ± 1.6 10.0–20.0 P 30 13 Dichaea intermedia Ames and Ep – – – 5.0 P 2 2 Correll D. muricatoides Hamer and Garay Ep – – – 10.0 P 5 4 D. neglecta Schltr. Ep – – 7.5 ± 2.5 5.0–15.0 P 13 7 Elleanthus cynarocephallus Ep – – – 10.0–20.0 P 21 4 (Rchb. f.) Rchb. f. (continued on next page) 382 BIOLOGICAL CONSERVATION 136 (2007) 372– 387

Appendix – continued Species Habit Con. Endemic Price (Mexican pesos) Unit Number of Number of status to units weeks (x Æ SE) Min.–max.

Encyclia adenocaula (Lex.) Schltr. Ep Pr Mexico – 100.0 P 3 2 E. atrorubens (Rolfe) Schltr. Ep Pr Mexico – – P 1 1 E. bractescens (Lindl.) Hoehme Ep – – – 5.0–15.0 P 14 1 E. candollei (Lindl.) Schltr. Ep – Mexico 9.0 ± 1.0 5.0–10.0 P 26 9 E. dickinsoniana (Withner) Ep – – 10.0 ± 2.0 5.0–15.0 P 37 4 Withner E. flabellata (Lindl.) B. Thurst. Ep – Veracruz – 5.0 P 2 2 and W. Thurst. E. gravida (Lindl.) Schltr. Ep – – – 10.0 P 7 4 E. luteorosea (A. Rich. and Ep – – – 5.0–10.0 P 2 1 Galeotti) Dressler E. parviflora (Regel) Withner Ep – Mexico 11.7 ± 1.7 10.0–15.0 P 14 6 E. polybulbon (Sw.) Dressler Ep – – 10.0 ± 1.6 5.0–15.0 P 77 27 E. spatella (Rchb. f.) Schltr. Ep – Mexico – 50.0 P 1 1 E. spp.a Ep – – 16.7 ± 1.7 15.0–20.0 P 22 14 Epidendrum ciliare L. Ep – – – 25.0 P 16 10 E. diffusum Sw. Ep – – – 8.0–15.0 P 7 5 E. galeottianum A. Rich. and Ep – – – 5.0–10.0 P 3 1 Galeotti E. laucheanum Bohnhof ex Rolfe Ep – – – 20.0 P 1 1 E. longipetalum A. Rich. and Ep – Mexico 6.9 ± 0.9 5.0–10.0 P 44 16 Galeotti E. parkinsonianum Hook. Ep – – 15.0 ± 2.0 10.0–20.0 P 23 8 E. polyanthum Lindl. Ep – – 14.3 ± 1.3 10.0–20.0 P 70 20 E. radicans Pav. Ex Lindl. Ep – – – 5.0 P 9 5 E. raniferum Lindl. Ep – – 19.0 ± 4.1 5.0–50.0 P 55 15 E. repens Cogn. Ep – – – 5.0 P 4 3 E. spp.a Ep – – – – P 5 3 E. spp.b Ep – – 20.8 ± 7.8 5.0–50.0 P 56 16 E. spp.c Ep – – 7.0 ± 1.2 5.0–10.0 P 20 9 E. spp.d Ep – – – 15.0 P 3 3 E. veroscriptum Ha´gsater Ep – – – – P 3 2 Euliche citrina (La Llave and Ep Pr Mexico – 60.0 P 65 5 Lexarza) Withner E. mariae (Ames) Withner Ep EE Mexico 64.8 ± 9.1 40.0–100.0 P 34 6 Gongora galeata (Lindl.) Rchb. f. Ep – Mexico 18.3 ± 3.3 15.0–25.0 P 17 12 Govenia mutica Rchb. f. Te – – – 10.0–15.0 P 1 1 G. superba (Lex.) Lodd. Te – – – 15.0–20.0 P 11 1 Habenaria floribunda Lindl. Te – – 3.8 ± 0.5 3.0–5.0 P 9 3 Isochilus spp. Ep – – 7.3 ± 2.7 2.0–10.0 P 39 18 Jacquiniella equitantifolia (Ames) Ep – – – 5.0–10.0 P 5 2 Dressler J. leucomelana (Rchb. f.) Schltr. Ep – Mexico – 5.0 P 2 2 J. teretifolia (Sw.) Britton and Ep – – – 5.0 P 3 3 P. Wilson Laelia albida Bateman ex Lindl. Ep – Mexico 66.7 ± 17.6 40.0–100.0 C 7 2 L. anceps Lindl. Ep – – 57.0 ± 28.3 5.0–250.0 P 310 34 L. autumnalis (Lex.) Lindl. Ep – Mexico 41.7 ± 4.4 35.0–50.0 C 33 5 L. crawshayana Rchb. f. Ep – Mexico – 70.0 C 2 1 L. eyermaniana Rchb. f. Ep – Mexico – – C 1 1 L. rubescens Lindl. Ep – – – – C 1 1 L. speciosa (Kunth) Schltr. Ep Pr Mexico – 30.0–35.0 C 207 1 Leochilus carinatus (Knowles Ep – Mexico – 2.0–5.0 P 46 11 and Wesct.) Lindl. L. oncidioides Knowles and Westc. Ep – – 4.4 ± 1.4 3.0–10.0 P 42 13 Lepanthes acuminata Schltr. Ep – – – 3.0 P – 1 var. acuminata D. neglecta Schltr. Ep – – 7.5 ± 2.5 5.0–15.0 P 13 7 Elleanthus cynarocephallus Ep – – – 10.0–20.0 P 21 4 (Rchb. f.) Rchb. f. L. avis Rchb. f. Ep – Mexico – 3.0 P – 1 L. schiedei Rchb. f. Ep – Mexico – 2.0 P 47 2 BIOLOGICAL CONSERVATION 136 (2007) 372– 387 383

Appendix – continued Species Habit Con. Endemic Price (Mexican pesos) Unit Number of Number of status to units weeks (x Æ SE) Min.–max.

Lophiaris cavendishiana Ep – – 22.9 ± 3.7 10.0–45.0 P 114 30 (Bateman) Braem L. cosymbephorum C. Morren Ep – – 23.3 ± 6.0 15.0–35.0 P 10 7 L. spp.a Ep – – 8.3 ± 1.7 5.0–10.0 P 24 8 L. straminea (Bateman ex Lindl.) Ep EE Veracruz 11.3 ± 1.2 8.0–15.0 P 151 24 Braem Lycaste aromatica (Graham) Lindl. Ep – – 9.3 ± 1.1 5.0–20.0 P 169 21 L. deppei (Lodd.) Lindl. Ep – – 12.9 ± 2.0 5.0–25.0 P 98 12 Masdesvallia floribunda Lindl. Ep – – 6.5 ± 1.6 3.0–10.0 P 93 17 Maxillaria cucullata Lindl. Ep – – 15.0 ± 6.3 5.0–40.0 P 84 11 M. densa Lindl. Ep – – 7.5 ± 2.5 5.0–15.0 P 43 17 M. elatior (Rchb. f.) Rchb. f. Ep – – 16.0 ± 4.8 5.0–30.0 C 238 24 M. meleagris Lindl. Ep – – – 10.0 P 1 1 M. moralesii Carnevalli and Ep – – 8.4 ± 2.9 5.0–20.0 P 26 9 J. T. Atwood M. tenuifolia Lindl. Ep – – 14.0 ± 1.9 10.0–20.0 C 70 14 M. variabilis Bateman ex Lindl. Ep – – 9.4 ± 1.2 5.0–15.0 P 151 43 Mormodes maculata (Klotzsch) Ep – Mexico 52.0 ± 16.2 15.0–100.0 P 11 5 L. O. Williams maculata M. maculata (Klotzsch) Ep EE Mexico 32.5 ± 5.7 10.0–50.0 P 63 22 L.O. Williams unicolor (Hook.) Myrmecophila grandiflora (Lindl.) Ep – Mexico 21.7 ± 1.7 20.0–25.0 P 34 7 Carnevalli, Tapia Mun˜ oz and I. Ramı´rez Nageliella purpurea (Lindl.) Ep – – 10.0 ± 2.9 5.0–15.0 P 45 13 L. O. Williams Nidema boothii (Lindl.) Schltr. Ep – – 6.3 ± 1.2 5.0–10.0 P 40 13 Notylia barkeri Lindl. Ep – – 10.0 ± 5.0 5.0–25.0 P 47 12 Oncidium incurvum Barker Ep EE – 15.0 ± 5.8 5.0–25.0 P 13 11 ex Lindl. O. maculatum (Lindl.) Lindl. Ep – – 22.5 ± 4.2 10.0–40.0 P 94 18 O. ornithorrynchum Kunth Ep – – – – P 9 7 O. reichenheimii (Linden and Ep – Mexico 38.3 ± 6.0 30.0–50.0 P 25 5 Rchb. f.) Garay and Stacy O. sp. Ep – – – 15.0 P 4 1 O. sphacelatum Lindl. Ep – – 10.5 ± 1.2 4.0–15.0 C 126 16 O. stenoglossum (Schltr.) Dressler Ep – – – 15.0 P 10 5 O. tigrinum Lex. Ep EE Mexico – – P 4 4 O. unguiculatum Lindl. Ep EE Mexico – – P 4 3 Ornithocephalus spp.e Ep – – – 3.0–5.0 P 3 2 Osmoglosum pulchellum Ep – – – – P 1 1 (Bateman ex Lindl.) Schltr. Papperitzia leiboldii (Rchb. f.) Ep Pr Mexico – 5.0 P 53 5 Rchb. f. Platystele repens (Ames) Garay Ep Pr – 3.8 ± 1.4 2.0–8.0 P 31 5 P. stenostachya (Rchb. f.) Garay Ep – – – 0.5–5.0 P 45 8 Pleurothallis angustifolia Lindl. Ep – – – 10.0 P 1 1 P. cardiothallis Rchb. f. Ep – – – 20.0 P 75 12 P. cf. dodsoni Luer Ep – – – 5.0 P 5 1 P. cf. herrerae Luer Te – – – 10.0 P 5 2 P. circumplexa Lindl. Ep – – – 1.0 P 1 1 P. digitale Luer Ep EE Mexico 1.5 ± 0.5 0.5–2.5 P 56 8 P. immersa Linden and Rchb. f. Ep – – 7.6 ± 1.6 3.0–15.0 P 25 10 P. lateritia Endres and Rchb. f. Ep – – 2.9 ± 0.5 1.0–5.0 P 92 9 P. microphylla A. Rich. and Ep – – – 2.0 P 5 1 Galeotti P. ornata Rchb. f. Ep – Mexico 5.3 ± 0.9 1.0–10.0 P 113 18 P. platystylis Schltr. Ep – – – 5.0–10.0 P 44 2 P. pubescens Lindl. Ep – – 8.3 ± 1.7 5.0–10.0 P 43 12 P. sertularioides (Sw.) Spreng. Ep – – 5.3 ± 1.5 3.0–8.0 C 3 1 P. spp.a Ep – – – 5.0–10.0 P 9 3 P. tribuloides (Sw.) Lindl. Ep – – 2.6 ± 0.9 1.0–5.0 P 75 14 (continued on next page) 384 BIOLOGICAL CONSERVATION 136 (2007) 372– 387

Appendix – continued Species Habit Con. Endemic Price (Mexican pesos) Unit Number of Number of status to units weeks (x Æ SE) Min.–max.

P. tubata (Lodd.) Steud. Ep – – 7.0 ± 1.0 5.0–8.0 P 19 8 Ponera sp. Ep – – – 5.0–50.0 C 25 5 Prescottia stachyodes (Sw.) Lindl. Te – – – 8.0–10.0 P 6 2 Prosthechea baculus (Rchb. f.) Ep – – – 100.0 P 1 1 W. E. Higgins P. cochleata (L.) W. E. Higgins Ep – – – 5.0–40.0 P 119 41 P. chondylobulbon (A. Rich. and Ep – – – 5.0 P 22 6 Galeotti) W. E. Higgins P. linkiana (Kl.) W. E. Higgins Ep – Mexico – 30.0 P 1 1 P. livida (Lindl.) W. E. Higgins Ep – – – 10.0 P 7 3 P. maculosa (Ames, FT Hubb. and Ep – – – 30.0 P 1 1 Schweinf.) W. E. Higgins P. michuacana (Lex ) W. E. Higgins Ep – – – 15.0–25.0 P 4 1 P. ochracea (Lindl.) W. E. Higgins Ep – – 6.5 ± 1.5 5.0 – 8.0 P 42 16 P. pseudopygmaea (Finet) Ep – – – 15.0 P 2 2 W. E. Higgins P. radiata (Lindl.) W. E. Higgins Ep – – 11.1 ± 0.7 10.0–5.0 P 180 20 P. rhynchophora (A. Rich. and Ep – – 8.3 ± 1.7 5.0–10.0 P 53 5 Galeotti) W. E. Higgins P. varicosa (Bateman ex Lindl.) Ep – – 25.7 ± 7.4 5.0–60.0 P 39 20 W. E. Higgins P. vitellina (Lindl.) W. E. Higgins Ep Pr – 13.0 ± 2.9 4.0–25.0 P 335 20 Restrepiella ophiocephala (Lindl.) Ep – – – 15.0–20.0 P 31 14 Garay and Dunst. Rhyncholaelia glauca (Lindl.) Ep – – 20.0 ± 3.5 10.0–40.0 P 207 23 Schltr. Rhynchostele aptera (Lex.) Soto Ep – Mexico – 20.0 P 3 1 Arenas and Salazar R. bictoniensis (Bateman) Soto Ep – – 18.3 ± 1.7 15.0–20.0 P 52 5 Arenas and Salazar R. cervantessii (La Llave and Lex.) Ep EE Mexico – 30.0 P 2 2 Soto Arenas and Salazar R. cordata (Lindl.) Soto Arenas Ep EE – 7.5 ± 1.7 5.0–15.0 P 117 12 and Salazar R. ehrenbergii (Link, Klotzch and Ep EE Mexico – 5.0–10.0 P 6 3 Otto) Soto Arenas and Salazar R. maculata (La Llave and Lex.) Ep – – 16.4 ± 4.0 10.0–40.0 P 57 7 Soto Arenas and Salazar R. rossii (Lindl.) Soto Arenas and Ep EE – 13.5 ± 1.5 5.0–30.0 P 366 29 Salazar Sarcoglottis cerina (Lindl.) Te Pr – – 10.0 P 1 1 P. N. Don S. sceptrodes (Rchb. f.) Schltr. Te – – – 5.0 P 4 2 Scaphyglottis lindeniana (A. Rich. Ep – – – 10.0–20.0 P 3 1 and Galeotti) L. O. Williams S. livida (Lindl.) Schltr. Ep – – – 10.0 P 10 7 Sobralia macrantha Lindl. Sax – – 15.0 ± 1.9 5.0–20.0 P 157 26 S. sp. Sax – – – 500.0 P 1 1 S. xantholeuca B. S. Williams Sax – – – 150.0 P 9 5 Stanhopea oculata (Lodd.) Lindl. Ep EE – 47.1 ± 14.1 15.0–120.0 P 44 11 S. sp. Ep – – – 80.0 P 1 1 S. spp.a Ep – – 21.4 ± 4.2 5.0–40.0 P 67 26 S. tigrina Bateman ex Lindl. Ep EE Mexico 36.0 ± 8.2 10.0–180.0 P 231 8 Stelis spp.f Te – – 7.1 ± 0.7 3.0–10.0 P 62 18 Stenorrhynchos aurantiacus (Lex.) Te – – – 15.0–25.0 P 4 1 Lindl. S. lanceolatus (Aubl.) Rich. Te – – – 25.0–30.0 P 1 1 ex Spreng. S. sp.1 Te – – – 25.0–30.0 P 1 1 S. sp.2 Te – – – 25.0–30.0 P 1 1 S. speciosus (Jacq.) Rich. ex Ep – – 12.2 ± 2.1 5.0–25.0 P 73 21 Spreng. BIOLOGICAL CONSERVATION 136 (2007) 372– 387 385

Appendix – continued Species Habit Con. Endemic Price (Mexican pesos) Unit Number of Number of status to units weeks (x Æ SE) Min.–max.

Trichosalpinx spp.g Ep – – 3.5 ± 0.7 1.0–5.0 P 79 15 Xylobium foveatum (Lindl.) Ep – – – 10.0 P 3 4 G. Nicholson

Plant species were recorded during a survey from July 2001 to April 2003. The conservation status (Con. Status) corresponds to that of the Mexican government (SEMARNAT, 2002). During the study period the mean exchange rate was 9.7 ± 0.6 Mx pesos for 1 US dollar (Bank of Mexico, http://www.banxico.gob.mx). EE, endangered of extinction; C, cutting; Ep, epiphyte; P, whole plant; Pr, subject to special protection; Sax, Saxicolous; Te, terrestrial. a Plants recorded without flowers that could not be identified. Probably belonged to one already recorded species and did not increase the species list. b This group included Epidendrum chlorocorymbos Schltr. and E. melistagum Ha´gsater. We grouped this species because both species were recorded but also infertile plants that could belong to any of the two species. c This group included Epidendrum cardiophorum Schltr. and E. nitens Rchb. f. We grouped this species because both species were recorded but also infertile plants that could belong to any of the two species. d This group included Epidendrum propinquum A. Rich and Galeoti, E. pseudoramosum Schltr. and E. ramosum Jacq. We recorded E. propinquum and E. pseudoramosum, and we grouped this two species with E. ramosum because infertile plants that could belong to any of the three species were recorded too, and the three species had been recorded for central Veracruz (Garcı´a-Cruz and Saldan˜ a, 1999). e We recorded two clearly different species of Ornithocephalus, O. inflexus Lindl. and O. tripterus Schltr., but we grouped this species because we recorded infertile plants that could belong to any of the two species. f We recorded four different species of Stelis, Stelis endresii Rchb. f., S. purpurascens A. Rich. and Galeotti, S. rubens Schltr. and S. veracrucencis Solano, we grouped all this species because was very difficult to identified this plants in the field and we also recorded infertile plants that could belong to any of the species. g We recorded Trichosalpinx blaisdellii (S. Watson) Luer and T. ciliaris (Lindl.) Luer, but we grouped this species because we recorded infertile plants that could belong to any of the two species.

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