A systematic revision of the Encyclia adenocarpos complex (Orchidaceae: Laeliinae) from Megamexico, including two new species from Mexico
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The Encyclia adenocarpos complex is restricted to the Pacific slopes of Megamexico, from Sonora, Mexico, to northern Nicaragua. It is characterized by pyriform to suborbicular pseudobulbs, (1-)2-3(4) thickly coriaceous, narrow leaves (> 15 times longer than wide), thin, verruculose inflorescences, and a verruculose ovary. The column is broad and wingless, fused to the labellum at its proximal 1/4. The flowers of the Encyclia adenocarpos complex feature what is here dubbed the Tupperware© labellum-column morphology, where the columnar ventral face is concave and fits tightly around the rim or upper margins of the callus, similar to the seal of this plasticware. All species grow on dry forests at low elevations from sea level to ca. 1500 m; most populations occurring below 500 m. The complex consists of six species, two of which are newly described herein: E. acapulcensis sp. nov., from southern Guerrero and E. enriquearcilae sp. nov., from western Oaxaca and neighboring Guerrero, Mexico. Encyclia rodolfoi is the correct name for populations of the Tehuantepec Isthmus whereas E. × nizandensis is here regarded as a natural hybrid between E. rodolfoi and E. parviflora; the evidence is presented and discussed. Encyclia schaeferi is treated as a morph of E. papillosa and included in its synonymy. A neotype is selected for Epidendrum adenocarpos and Epidendrum crispatum is rejected as a synonym thereof. All species are illustrated and discussed, and a key to the species is presented. Furthermore, the conservation status of each species is analyzed under the IUCN methodology. Other aspects of the biology of this complex are also discussed.
KeywordsComments of biogeography Diversity patterns IUCN Red List Criteria Megamexico Tupperware© labellum-column morphology
Encyclia Hooker is a Neotropical genus with more than 150 species concentrated in two centers of diversity, Megamexico II (as defined by Rzedowski 1991) and SE Brazil, with ca. 50 and 45 species, respectively (van den Berg and Carnevali 2005). Secondary centers of diversity exist in the West Indies (particularly Cuba), where ca. 25 species are found (Ackerman 2014), and in the periphery of the Amazonian Basin, particularly the eastern Andean foothills and the Guianas (Carnevali et al. 1994). Encyclia species are usually found in elevations below 1500 m, in forests with a marked dry season, although some species inhabit evergreen forests. Most of the species are strictly epiphytes, but a few are true lithophytes (e.g., E. granitica (Lindl.) Schltr. and E. leucantha Schltr.) or subterrestrials (e.g., E. tarumana Schltr. and, in ocasions, E. guatemalensis (Klotzsch) Dressler and G.E.Pollard). Other aspects of the biology and distribution of the genus can be found in Higgins (1997), Carnevali and Ramírez (2003, 2004), van den Berg and Carnevali (2005), Ackerman (2014), and Leopardi et al. (2016).
The genus consists of several clades, supported and identified by molecular and structural evidence. These clades seem to be distinctly correlated geographically (Leopardi et al. 2016), several of which are restricted to Megamexico II. The phylogenetic position and circumscriptions of these Megamexican clades have been discussed elsewhere (Leopardi et al. 2016), and the phylogenetic structure of the genus will thus be only briefly discussed herein in the context of the group treated.
Several clades of Encyclia have radiated in or colonized Megamexico, some restricted in their distribution, whereas others are shared with other Neotropical areas, particularly with the remainder of the Mesoamerican Isthmus (MI henceforth, as defined by Ossenbach et al. 2007) and the West Indies. The most recent phylogeny of the genus (Leopardi et al. 2016) suggests that ancestral Encyclia were related to E. bractescens (Lindl.) Hoehne and to the species phylogenetically close to E. microbulbon (Hook.) Schltr.
In this contribution, we provide a taxonomical and nomenclatural synopsis of this species complex. In addition, two new species are proposed. The conservation status of each taxon is evaluated using the IUCN Red List Criteria (IUCN 2010) and the GeoCAT platform (Bachman et al. 2011).
Materials and methods
The descriptions were prepared from live, herbarium, and/or pickled material, sometimes complemented with available iconography. Live plants were cultivated in Mérida, Yucatán at the Jardín Botánico Regional “Roger Orellana” at the Centro de Investigación Científica de Yucatán, A.C. or at several private orchid collections; plants were flowered, and flowering portions were eventually vouchered and, in most instances, flowers were preserved in a 70:25:5 ethanol/water/glycerine solution. Flowers from herbarium material were rehydrated by boiling and then soaking them in a soapy solution or, else, by soaking them in concentrated ammonium hydroxide for about 1 min, then rinsing in water; all material was examined under a dissecting microscope. Flowers thus pretreated were temporarily preserved by pickling (as indicated above) for further study and eventually returned to herbarium sheets.
Materials from the following herbaria were consulted: AMES, AMO, BIGU, CICY, ENCB, FCME, IBUG, M, MEXU, MO, NY, SEL, SERO, UAMIZ, and W (acronyms according to Thiers 2017). Specimens cited were in flower unless otherwise indicated. All specimens consulted were cited and annotated. Specimens were cited by countries, ordered south to north. Within countries, they were ordered alphabetically by major (states, provinces, departments) and subordinate (departments, municipalities) administrative subdivisions. Citations are as in the labels, but in some cases geographical coordinates and elevations were corrected or added (indicated by the abbreviation “approx.”) wherever information on localities was precise enough as to allow for this. Ecological data from the labels were omitted for brevity but were relevant to assess and inform our discussions.
Relevant bibliography and iconography was consulted and cited to correct errors in the literature. Whenever errors in published iconography were detected, they were corrected in the appropriate section of each species entry.
Pictures of live flowers were taken with a variety of digital cameras. Further images were captured at several resolutions, ranging from 600 to 1200 dpi with an Epson Expression 1640 XL scanner. Some measurements were performed or corroborated with ImageJ (Rasband 1997–2000). Cartography was produced by plotting the locality on an image data “shaded and colored SRTM elevation model” (NASA/JPL/NIMA 2002) using ArcView 3.2 (ESRI 1999). Google Earth 22.214.171.12436 (Google Inc. 2017) was employed for georeferencing and geographical assessments of distributions and distances between localities.
The conservation status of all the species of the Encyclia adenocarpos clade was assessed using the IUCN Red List Criteria (IUCN 2010). Because population data of these species were generally not available, we relied mostly on the B criteria, geographical distribution assessed both as B1 (Extent of Occurrence) or B2 (Area of Occupancy), both as implemented in GeoCAT (Bachman et al. 2011). We complemented these assessments with our own field experience, information and opinions provided by experts, published data and iconography, whenever available.
Results and discussion
Vegetative and floral morphology
In the Encyclia adenocarpos complex, the labellum is fused to the column for 1/4–1/3 of its length, a character state shared within the genus by only a few, distantly related taxa. The relatively small to minute lateral lobes are widely spreading, more or less flat, and they do not enclose the column. The massive callus, wider than the short claw on which it develops, is composed of two broad plates that widen toward the apex, with only a thin, linear to narrowly elliptic, shallow fovea (as opposed to most other members of the genus where the fovea is elliptic and relatively deep). The column is relatively thin in texture, yet broad, and devoid of true auricles or stelidia. However, the margins of the column are membranous, wide, and concave, broadening distally.
In most species of Encyclia, there is a pair of downpointing “wings,” “auricles,” or “stelidia” near the distal end of the column, on each side of the rostellum and the stigmatic surface. The homologies of these structures are unknown at this time but are here hypothesized to be staminodial remnants. Depending on the species, these stelidia often embrace the labellum at the claw of the midlobe, and/or the lateral lobes of the labellum tightly clasp the column (van den Berg and Carnevali 2005). Thus, the central lobe of the labellum is rigidly kept in place by both the stelidia and/or the lateral lobes. Hence, potential pollinators that try to penetrate Encyclia flowers have to force their way in, searching for the deceptive nectary by forcing the central lobe downward until it becomes dislodged from the embrace of the stelidia and/or the clasp of the lateral lobes, thereby excluding insects that cannot exert this force (Romero-González et al. in prep.). As we have observed for E. diurna (Jacq.) Schltr. and a few other species (Carnevali and Ramírez 1986), once the pollinator retreats with the pollinia attached to their back, the lateral lobes close down under the column, potentially inhabilitating the flower from receiving further visits by pollinators. In others, instead, the lateral lobes return to their original position, but a full survey of the genus on this respect is lacking at this time. A legitimate visitor, most likely a bee, will dislodge the anther upon withdrawal, allowing for the four pollinia to become attached to the back of the bee’s thorax. The colors of flowers that have had their pollinia removed soon turn drab and nectar guides become indistinct within a short period of time. Presumably, this mechanism works in optimizing the rare potential pollinator visits by flagging flowers that have already been visited.
Biogeography and diversity patterns
All species of this complex grow in dry forests at elevations ranging from sea level to 1000 m (a few collections of Encyclia papillosa (Lindl.) Aguirre.-Olav. up to 1400 m); most known populations occurring below 500 m. The northernmost range of the complex occurs in SW Chihuahua and Sonora at 27°–28° N for E. adenocarpos (making this species the most northerly in the genus), whereas toward the south, the complex ranges (E. papillosa) to circa 12°N at the Nicaragua Lakes in the Nueva Segovia and Jinotega Departments.
Most species of the Encyclia adenocarpos complex appear to be closely related, differing not by distinctive apomorphies but by unique combinations of character states in combination, most remarkably, with geographical distributions or the association to particular ecological conditions. The species can be distinguished by the contrasting inflorescence length and density, labellum shape (mostly the shape and relative sizes of the lobes), and floral color combinations, particularly of the labellum and column. Furthermore, the species are mostly allopatric or presumably parapatric. Wherever or whenever two species occur in close proximity, such as happens presently with, e.g., E. adenocarpos and E. trachycarpa (Lindl.) Schltr. in Jalisco and Colima, the two species involved feature strikingly different color patterns or floral sizes. In this particular case, one of the species displays a white or pale yellow labellum midlobe with a few purple or red–purple short veins, or these lacking altogether (E. adenocarpos), whereas the other features larger flowers and longer, bolder stripes or veins arranged in a ± regular pattern suggestive of nectar guides on a bright white background. All of the species flower more or less contemporaneously (May–June), and these color patterns are likely to play a role in species isolation.
Encyclia × nizandensis Pérez-García & Hágsater, found in the vicinity of Nizanda, in southern Oaxaca, is apparently a natural hybrid between E. rodolfoi and the locally rare E. parviflora (Regel) Withner. Pérez-García and Hágsater (2012) reported another natural hybrid involving E. rodolfoi, in this case with E. hanburyi (Lindl.) Schltr., Encyclia × nizanburyi Pérez-García & Hágsater (see further discussion under E. rodolfoi).
Four species of the Encyclia adenocarpos complex are currently known on the NW side of the Tehuantepec Isthmus. Starting from the isthmus and along the northwesternward narrow strip of lowland tropical dry forests by the Pacific drainage, we first encounter E. enriquearcilae. This taxon, proposed herein, is well-documented from two localities in coastal Oaxaca and two from SE Guerrero. Although labellum morphology is distinctive (yet clearly reminiscent of E. adenocarpos), color patterns in the flowers suggest those in E. papillosa, to which this species may be closely related. Further NW, in Guerrero, E. acapulcensis occurs, known currently from a single set of populations. This species resembles E. rodolfoi, but colors are distinctive. The fact that these two species (E. enriquearcilae and E. rodolfoi) are most similar to two transisthmian counterparts, suggests two vicariant events at each side of the Tehuantepec Isthmus.
Two species are found from mid-Guerrero up to southern Sonora and Chihuahua. Encyclia trachycarpa is restricted to a relatively small area extending from SW Jalisco state to SW Nayarit state. It is nested within the distribution of the other species, E. adenocarpos, but is florally distinctive with larger blossoms and strikingly different color patterns. The second, Encyclia adenocarpos, has an extensive range along the coastal Pacific drainage, but it also gets farther inland than any other taxon of the complex. It is well-documented as occurring in the Balsas River Basin in the states of Guerrero, México, and Morelos. It also follows the coastal strip along the Pacific from central Guerrero up to southern Sonora reaching its northernmost range near the village of Álamos, the northernmost outpost of tropical deciduous forests in the boreal hemisphere. It has been also collected on the eastern slopes of the Sierra Madre Occidental in Jalisco (e.g., near the Tacotán Dam, Ramírez 2018 sub Carnevali 8030) at elevations of ca. 1300 m.a.s.l.
Very little is known about the reproductive biology of Encyclia in general (van den Berg and Carnevali 2005) and of the E. adenocarpos clade in particular. As in most member of Encyclia, reproductive structures and phenology suggest deceptive pollination by diurnal bees and wasps (or, more rarely, flies) of the appropriate size. Flowers are long lasting (3–4 weeks) and offer no apparent reward. With a few exceptions, very few fruits are produced under natural conditions. Particular populations of E. papillosa are exceptional in producing many fruits (ca. 40% of flowers in a given inflorescence yield fruits, at least under cultivation in Mérida, Yucatán). However, visit to natural populations of these encyclias and analysis of herbarium specimens suggest this high level of fertility is not common in nature. The flowers of the Encyclia adenocarpos complex feature what is here dubbed the Tupperware© labellum-column morphology, where the ventral face of the column is concave and fits tightly around the rim or upper margins of the callus, working like the seal in a piece of Tupperware© container. The pollinator has to forcefully work its way into the flower to accede to the deceptive nectary (see above) by pushing down the central lobe of the labellum, dislodging the “seal.” Flowers that have been pollinated soon change colors, usually with the central lobe of the labellum turning yellow and the purple nectar guides becoming paler or indistinct.
The flowers of the members of this species complex produce little fragrance if at all, at least not perceptible to the human nose. However, using gas chromatography and mass spectrography chemical analysis of the fragrances of some members of Encyclia and Prosthechea Knowles and Westc. from Chiapas, Del Mazo and Damon (2006), reported the fragrance of a member of this species complex to be composed of hexadecene, E-nerilidol, and phenylformide. These authors identified their Chiapas plants as E. adenocarpos (as “E. adenocarpa”), but since this taxon is absent from Chiapas, we assume Del Mazo and Damon’s plants must be referrable to the more widespread and locally common E. papillosa. Kaiser (1993) had reported benzyl and butyl caproate, hydroquinone dimethyl ether, and ß-ionone for E. adenocarpos, compounds that were absent from Del Mazo and Damon’s samples. Kaiser’s citation refers to a non-vouchered plant (and the picture provided along with the text is of E. aspera (Lindl.) Schltr., a totally unrelated Andean species). Thus, because evidence allowing for accurate determinations of the taxa analyzed by these two authors is unavailable at this time, and most taxa of the complex have been identified as E. adenocarpos at one time or another, we assume they must refer to two different species. The relevance, if any, of these differences in fragrance composition for the pollination biology of Encyclia adenocarpos cannot be assessed at this time.
We have observed males of Euglossa viridissima Friese or of the newly described E. dilemma Bembé & Eltz. (Apidae: Euglossini) visiting flowers of Encyclia trachycarpa in cultivation in Mérida (Yucatán; Carnevali, pers.-obs.). Bees “scratched” repeatedly the apices of the petals and sepals as well as the labellum underside, as if collecting fragrances. However, the bees never approached the ventral face of the labellum or the column, and thus could not possibly pollinate the flowers. However, E. trachycarpa occurs naturally on the Pacific drainages of Mexico and it may be visited there by another species of Euglossini that could serve as pollinator. Regardless, the interaction observed in the cultivated plants suggests an evolutionary pathway potentially leading to the androeuglossinophilous pollination syndrome.
Key to the species of the Encyclia adenocarpos species complex
To properly use this key, good floral material is required, including color descriptions; good photographs and pickled flowers can be excellent aids for determinations. Locality data are also very important because of the clear biogeographical patterns in the distribution of the species in the complex. In most cases, a description of color patterns of the flowers coupled with geographical data should suffice to confidently identify the species of the complex. It is important to understand that all of these species are variable and variation patterns often overlap, which is the reason why the key is polythetic, and thus several characters have to be evaluated to arrive at a positive identification.
Petals and sepals narrowly elliptic, sharply acute, concave in natural position, dark brown, with narrow, pale green margins; anther white; labellum with triangular oblong, acute lateral lobes; central and lateral lobes dirty white or pale rose with numerous purple, raised veins (giving the overall impression of a purple labellum); plants from coastal Guerrero state, Mexico … E. acapulcensis
Petals and sepals usually narrowly spatulate, obtuse, rounded to subacute, flat or with recurved margins in natural position, color variable, usually pale brown, greenish or yellowish, variously tinged in purple or darker brown but never dark brown with pale green margins; anther in several hues of yellow, rarely whitish; labellum with variable lateral lobes, only rarely with triangular oblong, acute lateral lobes; central lobe white to yellow with few dark purple veins but lateral lobes never white or pale pink with many purple veins; plants from throughout the range of the species complex but never from the central portion of coastal Guerrero, Mexico … 2
Widest point of the labellum across the midlobe; lateral lobes very reduced (less than half the width of the midlobe); flowers large, nutant, with pedicellate ovary 22–25 mm long, dorsal sepal 20–25 mm long and labellar midlobe 14–16 mm wide; perianth segments usually not widely spreading; anther bright yellow, sharply contrasting against the apex of the clear white column … E. trachycarpa
Widest point of the labellum across the spread lateral lobes; lateral lobes usually more conspicuous (usually longer than the width of the midlobe, always longer than half the width of the midlobe); flowers smaller, patent, with pedicellate ovary 14–16 mm long, dorsal sepal 15–20 mm and labellar midlobe 7–11 mm wide; perianth segments usually widely spreading; anther pale yellow to bright yellow, not contrasting sharply against the apex of the dull yellow, purplish-striped, or dirty white column … 3
Lateral lobes of the labellum large and conspicuous, at least three times longer than wide, obovate, oblong obovate, linear to linear oblong or narrowly triangular; column basally white, very pale yellow or pale green, apically bright yellow, the apex always sharply contrasting in color with the base; anther bright yellow; lateral lobes white to yellow, almost always without colored nerves; central lobe white to pale yellow, margin concolor or bright yellow; callus obovate, 1.5 times longer than wide, white with yellow apex, the yellow color often extending to the margins; at flowering time, turgent pseudobulbs suborbicular … 4
Lateral lobes smaller and less conspicuous, at most two times longer than wide, often as long as wide, triangular, acute to obtuse; column dirty white to pale yellow or yellow, concolor, never the apex sharply contrasting in color with column base; anther white to very pale yellow; lateral lobes pale yellow, with or without purple nerving; central lobe color variable, usually same colored as column apex, with or without purple nerving, never with a contrasting yellow margin; callus elliptic, at least twice as long as broad, entirely white or white with purple veins; at flowering time, turgent pseudobulbs pyriform … 5
Lateral lobes of the labellum oblong to obovate-oblong, apically rounded to broadly obtuse, relatively broad, 3–4 mm wide at midlength; central lobe concolorous white (pale yellow upon aging), rarely with a yellow margin; plants from the Central Depression of Chiapas southeasternward to northern Nicaragua, along the Pacific drainage … E. papillosa
Lateral lobes of the labellum narrowly triangular to narrowly oblong, apically acute, narrow, 1.2–1.5 mm wide at midlength; central lobe white at base with a conspicuous yellow band in the center and the margin, or totally yellow; plants from the coastal plain of southwestern Oaxaca and Guerrero, Mexico … E. enriquearcilae
Lateral lobes of the labellum variable but always reduced as compared to other taxa of the complex, 2.0–3.5 mm long, always much shorter that central lobe, shorter than the callus, less than 1/5 the length of the central lobe, ranging from broadly triangular, acute to oblong, apically rounded, colored as central lobe or pale yellow; central lobe concolorous white or pale yellow, with or without a few short purple veins, lateral veins pale yellow, lacking colored veins; sepals usually flat in natural position; plants from Guerrero northward up to southern Sonora, particularly common in the Balsas River Basin and in Michoacán, Jalisco, Colima, and Nayarit states, Mexico … E. adenocarpos
Lateral lobes of the labellum longer, at least 5 mm long and about as long as the central lobe or slightly shorter, at least as long as 3/4 of the length of the callus, as long or slightly shorter than central lobe, triangular or oblong triangular, apically obtuse to acute, intense to medium yellow with fine but conspicuous purple nerving; central lobe yellow with conspicuous purple nerving, often with pale pink suffusion; sepals often somewhat recurved in natural position; plants from the Tehuantepec Isthmus area, where common, into the Central Depression of Chiapas, Mexico, where rare, southward to Nicaragua along the Pacific drainage … E. rodolfoi
Etymology: The specific epithet refers to Acapulco, the touristic city of Guerrero, Mexico, where several populations of this species are known to occur.
Description: Epiphytic or lithophytic herbs, 21–27 cm tall without inflorescences, up to 95 cm counting inflorescences. Rhizome, short, woody. Pseudobulbs (3.2–)4.9–7.0 × 2.2–3.2 cm, aggregate, ovoid to more rarely subglobose, apically (1-)2-3 leaved, green and smooth when young, slightly wrinkled upon aging; clothed with 1-2 papery sheaths, defibrating upon maturity. Leaves 18.0–26.6 × 0.8–1.4 cm, rigidly coriaceous, linear oblong, acute, slightly falcate, dark green. Inflorescence 52–65.5 cm long, terminal, emerging from the mature pseudobulb, suberect to arching, 5–15-flowered, racemose or paniculate with a lateral branch bearing up to 3 flowers; peduncle and rachis verruculose, thin but strong and elastic, intense green, the rachis often red-tinged; peduncle bracts 0.5–1.4 × 0.05–0.60 cm, tubulose, conspicuous, oblong, acute, decreasing progressively in shape toward the apex; floral bracts 0.1–0.5 × 0.05–0.60 cm, similar in shape to the bracts of the peduncle but smaller and inconspicuous. Flowers 2.5–3.0 cm diameter across the spread tips of the petals, 3.1–3.2 cm across the apex of dorsal sepal and apex of labellum, resupinate, showy, petals and sepals green basally, slightly tinged red or dark brown in the middle section and apex, margins narrowly delineated in pale green; labellum ground color pale green, suffused with pale to dark purple, the central lobe with 5–10 darker colored veins; lateral lobes with fine dark purple veins, denser distally; callus ground color white with pale to dark purple veins, fovea almost white; column dorsally pale green at base, distally white or pale purple with darker purple veins, almost solid red toward the clinandrium, ventrally pale green proximally, white or pale purple distally with dark maroon or red-brown veins; anther white or extremely pale yellow; ovary with pedicel 1.4–1.6 cm long, cylindrical, verruculose; sepals elliptic-lanceolate, acute, dorsal 1.5–2.0 cm × 0.4–0.5 cm, laterals 1.7–2.0 × 0.4 cm, slightly oblique, longitudinally concave; petals 1.5–2.0 × 0.3–0.4 cm, linear elliptic, acute, attenuate in the proximal fourth. Labellum 1.67–2.00 × 1.05–1.40 cm, fused to the ventral face of the column at the basal 1/4, then free, 3-lobed; central lobe 0.95–1.00 × 0.80–0.87 cm, suborbicular to oblong elliptic or oblong obovate, apically truncate; lateral lobes 0.58–0.70 × 0.22–0.28 cm, free, spreading and thus not enclosing the column, subtriangular, apex obtuse-rounded or truncate; callus subelliptic consisting of two keels divided by a conspicuous narrow depression, the keels converging distally into a single keel that reaches the central lobe apex. Column 0.98–1.4 cm × 0.49–0.54 cm, straight, subrhombic in ventral view, narrowing proximally, lacking stelidia; anther white, cordate in ventral view; pollinia 0.10–0.21 × 0.07–0.12 mm, yellow, in two subequal pairs; stigmatic surface obcordate, 1.2 × 2.0 mm.
Diagnosis: A species related to Encyclia rodolfoi Archila, Chirón, and Véliz but with pale to dark purple veins on the labellum over a pale purple or pink background (vs very dark purple or maroon veins over a yellow-white or yellow background and a yellow margin); a purple column (vs dorsally white or yellowish with a few dark purple veins), acute petals and sepals (vs obtuse or broadly acute), and patent, concave, almost conduplicate lateral sepals at the distal half (vs somewhat reflexed, longitudinally flat to very slightly concave).
Phenology: Flowering material of this species has been collected in June and July.
Distribution area and habitats: Encyclia acapulcensis is restricted to a narrow band of seasonal dry forest in coastal Guerrero, where it grows at 75–360 m. Plants of the species are usually lithophytic, with the roots entering the cracks of the rocks where some litter accumulates; some plants have been seen growing as epiphytes. This is an area with a hot climate, a yearly average temperature of 27 °C, and strongly seasonal precipitation on average 1200 mm.
Relationships and diagnostic characteristics: Although vegetatively very similar to the other species of the complex, the flowers of this species are most distinctive and the pseudobulbs often present a reddish cast. Petals and sepals are narrowly elliptic, acute, whereas the labellum central lobe is subquadrate to subquadrate oblong, apically rounded to truncate. The color of the petals and sepals is basally green, slightly tinged red or dark brown in the middle section and apex, margins narrowly delineated in pale green; the sepals, particularly the laterals, are concave. The labellum ground color is pale purple and has many, relatively thick purple veins that reach the margins of the lobes; the lateral lobes are pale yellow with many purple veins, thinner than those of the central lobe. The column and callus are white with many fine purple veins. The general shape of the flower and the color pattern in the flower (although colors are different) suggest a relationship with E. rodolfoi from further south in Mexico in Oaxaca to El Salvador, which also often grows on rocks close to the sea.
Variation range: Encyclia acapulcensis is only known from a few localities, and little is known of its natural variation. We have seen live flowers from the Acapulco populations, and the only variation we can detect is in the shape of the central lobe of the labellum which ranges from almost rounded or subquadrate to subquadrate oblong.
Taxonomic notes: Encyclia acapulcensis, as with most members of this complex of closely related species, has been confused with E. adenocarpos or else included in its circumscription (e.g., Hágsater et al. 2005). Pressed specimens of both species look deceptively similar, but the shape of the sepals and the color description of the flower should aid in its diagnosis.
Iconography: Hágsater et al. (2015), page 124., Figure 248, color photograph (as “E. aff. adenocarpa de la costa de Guerrero”).
Paratypes: MEXICO. Guerrero: Municipio Acapulco de Juárez, Parque el Veladero, selva baja caducifolia sobre la cañada entre la Universidad Loyola del Pacífico y el Fraccionamiento las Brisas, 270 m a. s. l., 16°51′27.80″N, 99°50′23.69″W, flowering in cultivation in Mérida, Yucatán, from plants of the original collection by J. Viccon and U. Muñoz, 18 Jun 2014, G. Carnevali 7714 (AMES, CICY); same locality, 15 Jun 2016, G. Carnevali 7966 (AMES, AMO, CICY); same locality, 20 Jul 2017, G. Carnevali 8101 (AMES, CICY); estación de microondas en Las Cumbres de Llano Largo, en una pequeña desviación de la carretera Escénica a Puerto Marqués, 360 m a. s. l., 16°49′32″N, 99°50′39″W, Francke s.n. (AMO).
IUCN conservation status: According to the IUCN (2010), Encyclia acapulcensis would be considered an Endangered (EN) species both because it occurs over a reduced surface area (“Extent of Occurrence”) of 204 km2 and covers an extremely small Area of Occupancy (16 km2, cell width of 2 km). The species is known from five records, all coming from the same general locality, a series of small hills surrounding the SE extreme of the Acapulco Bay, along the Carretera Escénica which leads from downtown Acapulco to Puerto Marqués through a series of luxurious suburbs. These localities are within the Parque Nacional “El Veladero” that affords some protection to them. However, even within this national park, the vegetation where Encyclia acapulcesis grows is subject to demographic pressure and its populations are severely threatened (Vargas-Márquez 1984). Nearby Acapulco, it grows as a lithophyte on volcanic outcrops along with Tillandsia sp. (Bromeliaceae), Philodendrum sp. (Araceae), and columnar cacti.
All collections come from seasonally dry forests at elevations of 75–360 m a. s. l. (altough we have vouched records only from above 270 m a. s. l.). This kind of vegetation has been severely disturbed for agricultural (mostly slash-and-burn-agriculture, “milpas”) or housing (mostly beach tourism) developments. The right type of habitats for the growth of this species is dispersed in a mosaic of other types of habitats, and if we consider that at least ¼ of the Extent of Occurrence is covered by unsuitable habitats (e.g., mangroves, marshland, or other types of vegetation associated with the sea) or remnants of several types of vegetation converted into pasturelands, we must assume that the species is severely endangered.
Etymology: From the Greek adeno, gland or glandular, and carpus, fruit, referring to the texture of the fruit surface.
Description: Epiphytic or lithophytic herbs, 9–35 cm tall without inflorescences, up to 100 cm counting inflorescences. Rhizome, short, woody. Pseudobulbs 3.6–5.6 × 1.1–1.8 cm, aggregated, conical to pyriform, apically (1–)2–3-leaved, green and smooth when young, eventually wrinkled upon maturity, clothed by papyraceous, eventually defibrating sheaths. Leaves 9.57–28.4 × 0.38–1 cm, basally conduplicate, oblong-linear, acute, blades often partially falcate, rigidly coriaceous, dark green, in some individuals and/or populations, slightly to heavily tinged in purple, midnerve carinate mainly in the lower half. Inflorescences 20–86 cm, terminal, originating from the mature pseudobulb, erect, suberect, or arching, with 4–15(–25) flowers, racemose or, more commonly, paniculate with 1–3(–5) lateral branches of 2–5 flowers; peduncle thin and wiry, deep green or purple tinged; peduncle bracts 3.5–13.1 × 0.4–4.5 mm, inconspicuous, triangular oblong, acute; rachis verruculose, green, often purple tinged; floral bracts 0.1–0.3 × 0.3–0.5 mm, inconspicuous, papyraceous, ovate-triangular, acute; ovary with pedicel 1.4–1.7 long, cylindric, densely verruculose. Flowers 26–37 mm across the spread apices of the petals, resupinate, more or less showy, petals and sepals pale green to yellow green, variously tinged in hues of brown on the distal half; labellum with a central lobe white or cream white, often with yellow tinges or the yellow restricted to the margins, distally often with a slight purplish or brownish tone, veins concolorous or several (3–)5–9(–11) of them purple-colored along midsection, lateral veins pale yellow or pale green, callus white or white tinged pale yellow, often with a few purple veins; column dorsally pale green at the proximal half, distally white or pale yellow, ventrally white or pale yellow, expanded margins often brown tinged; sepals: dorsal 13.4–20 × 3.2–3.9 mm, narrowly oblanceolate to narrowly spatulate, obtuse to subacute; laterals 12.4–18.1 × 3.7–4.1 mm, similar to dorsal but slightly oblique, longitudinally concave; petals 13.2–16.2 × 1.9–4.1 mm, linear oblanceolate to linear spatulate, apices rounded to subacute, narrowed in the proximal half; labellum 26.9–31.0 mm, fused with the column at about the basal 1/5, then entirely free, 3-lobed; central lobe 4.5–5.7 × 5.6–7.9 mm, suborbicular to oblong elliptic, truncate to rounded, emarginate, with 3 or 5 straight or sinuous keel-like veins protruding form the callus; lateral lobes 2.4–4.8 × 0.7–1.1 mm, broadly triangular to oblong elliptic, obtuse to acute, spreading and thus not clasping the column; callus 4.5–6.5 × 3.1–4.9 mm, fleshy, massive, broadly elliptic to sub-rhomboid, occupying all of the isthmus and entering the central lobe, consisting of two thick, reniform-elliptic keels or ridges separated by a narrow sulcus or a shallow, elliptic fovea, the keels converging distally and ending in 1 or 3 thick teeth that may or may not extend themselves as veins entering the central lobe. Column 7.0–10.5 × 3.7–4.9 mm, straight, oblanceolate-spathulate, depressed-ellipsoid in transversal section, stellidial remnants porrect, acute; anther 2–2.1 × 2–2.1 mm, broadly obovate-orbicular in dorsal view, cordate, white to pale yellow, always same colored as column apex; pollinia 0.9 × 1.0 mm, bright yellow, in two subequal pairs; stigmatic surface obcordate-subquadrate. Capsules densely verruculose.
Notes: Encyclia adenocarpos is the most variable species of the complex, often exhibiting conspicuous variation even within populations. Local or regional populations can also be distinctive in shape and coloration pattern. In particular, the species is variable in coloration with perianth segments pale or deep green variously tinged in brown or brown-maroon, often to the extent that in some populations in Jalisco and Nayarit dark chestnut-colored individuals are found. The labellum is also variable in color with the central lobe almost pure white to pale yellow, often with a slight purplish or brownish tone and the veins concolorous or several (3–)5–9(–11) of them purple-colored along midsection of the central lobe. The lateral lobes of the labellum are usually of the same color as the central lobe margin. These lobes are also variable in shape and size relative to the central lobe. In some populations (e.g., Carnevali 6967), the lateral lobes are very small (ca. 1.9 mm long) and broadly triangular, whereas in others they are oblong elliptic to oblong triangular, acute to rounded. However, we have not been able to identify clear patterns in this variation, and thus a single species is recognized for this aggregate of forms featuring small lateral lobes and mostly white, fundamentally elliptical calli.
It is possible that Encyclia adenocarpos is a paraphyletic taxon, with at least one of the other Mexican species of the complex, E. trachycarpa, that occurs within the range of E. adenocarpos, having most likely originated from populations we would refer to E. adenocarpos. Our ongoing and future phylogenetic and phylogeographical analyses of the genus Encyclia particularly of the E. adenocarpos complex, should eventually allow us to address this possibility.
Distribution area and habitats: Encyclia adenocarpos has the most extensive distributional range of the species complex, which is ca. 1300 km in a straight line, ranging from coastal Guerrero into the Balsas River Basin with localities recorded from the States of México, Michoacán, and Morelos. Then, along the coast, it extends as far north as Álamos, Sonora at 27°N, thus becoming the most northerly Encyclia species of all. In general, it can be stated that its distribution follows that of the tropical dry forests on the western side of Megamexico II (e.g., De Nova et al. 2012) as several other taxa with similar requirements do. Encyclia adenocarpos grows commonly as an epiphyte, but plants have often been collected growing as lithophytes, usually on northern or eastern exposures. Populations have been recorded at elevations ranging from sea level to up to 1250 m (rarely somewhat higher), but most collections have been made below 500 m. At this time, there is an important gap in our knowledge of the distribution and variation of the species of this complex in coastal Guerrero and Michoacán, an area that may harbor more populations referable to this taxon or to additional (and at this time unknown) species or even of populations bridging the variation gap between known species.
Additional specimens examined: MEXICO. No state specified: received from R. Oberg (supposedly probably from Chiapas), Garay 3727 (AMES, USF). Chihuahua. Morelos: Real Morelos y Real La Dura, 600 m a. s. l., 26°37′N, 107°48′W, W.P. Hewitt 380 (AMES). Colima: Rancho La Salida, 400 m a. s. l., 3 May 2002, Contreras s.n (IBUG); Ixtlahuacán, 19 May 2000, Contreras and González-Tamayo s.n. (IBUG). Guerrero: Alpoyeca: 2 km al SO de Tecoyo, bosque tropical caducifolio, 1200 m a. s. l., 17°35.081′N, 98°30.500′W, 17 May 1995, J. Calónico 1812b (MEXU); 9 km al NE de Tlapa, camino Tlapa-Huamuxtitlán, Puente El Salado, 990 m a. s. l., approx. 17°36′23.02″N, 98°30′58.72″W, 16 Sep 1982, Martínez et al. 2620 (MEXU, MO). Coyuca: Coyuca, near Chacamerito, ca. 350 m a. s. l., 18°19′9.40″N, 100°35′17.15″W, 5 Jul 1934, Hinton et al. 6015 (AMES, GBH, NY, P, US). Iguala: near Iguala, 731 m a. s. l., 18°19′N, 99°34′W, 11 May 1933, Ignacio P. 2297 (AMES, F). Huamuxtitlán: 1 km al O de Jilotepec, bosque tropical caducifolio, 1260 m a. s. l., 17°57.711′N, 98°29.522′W, 8 Aug 1993, E. Moreno and J. Calónico 271 (AMO). Mochitlán: region of Chilpancingo, on oaks on Mt. El Terrero, 1 Jul 1942, H.D. Sawyer 636 (F); Iguala y Buenavista, Cañón de la Mano, entre Los Amates y el Naranjo, 10 km al N de Iguala por el ferrocarril, 900–1000 m a. s. l., approx. 18°26′1.46″N, 99°33′40.35″W, cañón angosto con acantilados y afloramientos de roca caliza, bosque tropical caducifolio, 2 May 1987, C. Catalán et al. 617 (MEXU). San Jerónimo Técpan: 2 m a. s. l., 17°05′N, 100°35′W, 2 Jun 1933, O. Nagel 1818 (AMO). Técpan de Galeana, Técpan-El Reparo, 75 m a. s. l., 4 Sep 1939, G. Hinton 14122 (AMES, AMO, F, MO, NY). Tlapa: ca. del puente sobre El Río Salado, 8 km al N de Tlapa, road to Huamuxtitlan, 1060 m a. s. l., 17°34′35″N, 98°31′24″W, 7 May 1997, A. Garcia-Mendoza 6515 (B, MEXU). Xochihuehuetlán: 0.6 km NW de Jilotepec, faldas del Xilotzin, bosque tropical caducifolio, 1260 m a. s. l., 17°57′57"N, 98°29′15"W, 8 May 1993, E. Moreno and J. Jiménez 271 (MEXU). Jalisco. La Huerta: Estación Biológica Chamela, Sendero Calandria, El Mirador, 90–95 m a. s. l., 19°15′30"N, 105°02′8"W, selva baja caducifolia con epífitas como Tillandsia spp., Encyclia ssp. y Hechtia jaliscana L.B. Smith, 2 Jun 2015, W. Cetzal and G. Carnevali 431(CICY); Vereda Tejón, 19°30′N, 105°03′W, 28 May 1982, Emily J. Lott 1068 (AMO); Loma Alta, 40 km. de La Huerta hacia Barra de Navidad, 10 m a. s. l., 19°13′30.89″N, 104°41′10.69″W, selva mediana subcaducifolia con elementos de selva baja caducifolia en muy buen estado de conservación con árboles como Cordia elaeagnoides DC. y Bursera simaruba Sarg. con alta diversidad de epífitas xeromorfas tales como Tillandsia caput-medusae E. Morren y Myrmecophila galeottiana (Reichb. f.) Rolfe, 15 Jun 1998, G. Carnevali and G. A. Salazar 5133 (CICY). Magdalena: no precise locality, 1200 m a. s. l., S. Rosillo de Velasco 12 (AMO). Pihuamo: carretera Pihuamo-Colima, 9 Jun 1997, Contreras s.n. (IBUG). Tecolotlán: 4 km S de Tecolotlán, por la vía a Juchitán, 1169 m a. s. l., 20°09′21.48"N, 104°03′01.39"W, Soltero and Contreras s.n. (IBUG). Tenamaxtlán: Miraplanes, 1467 a. s. l., approx. 20°8′51.27"N, 104°10′17.05"W, Contreras s.n (IBUG); 7 km E Rancho Miraplanes, 1200 m a. s. l., approx. 20°07′53.96"N, 104°06′38.35"W, May 1985, Soltero and Contreras 144 (IBUG). Tolimán: Camino a Chachahuatlan, 2 km Paso Real, bosque tropical caducifolio, 700 m a. s. l., 19°37′5.18″N, 103°58′11.02″W, 26 May 1990, A. Rodríguez-Contreras et al. 2029 (IBUG, MEXU). Tomatlán: Laguna Agua Dulce, 70 m a. s. l., 20°03′00N, 105°30′30"W, 11 Mar 1993, G. Castillo et al. 11055 (XAL); 13 km ESE from Tomatlán on road to Talpa de Allende, 2 km SSE de Puentecillas, 140 m a. s. l., 19°51′37.51″N, 105°09′54.58″W, H. Iltis s.n. (IBUG), same locality, 14 Jan 1979, Iltis and Nee 1658 (drawing, IBUG). Unión de Tula: 3 km delante de Tacotán, rumbo a Sta. Rosalía, selva baja caducifolia, 1275 m a. s. l., 20°2′3.99″N, 104°19′39.38″W, 16 Jun 1988, A. Espejo et al. 3275 (UAMIZ). México. Temascaltepec: 1000 m a. s. l., 24 Apr 1933, G.B. Hinton 3528 (AMES); Barranca de Tonatico, 1200 m a. s. l., 15 Mar 1954, E. Matuda et al. 32158 (F, MEXU); Ixtapan, 1000 m a. s. l., 19 Mar 1933, G. B. Hinton 3828 (AMES); same locality, 2 Apr 1937, G.B. Hinton 3909 (AMES); Ixtapan, 1000 m a. s. l., 27 Apr 1937, G.B. Hinton 3909 (AMES). Michoacán. Aquila: Km 120 Playa Azul-Manzanillo, afloramiento volcánico, 110 m a. s. l., 18°18′12.93"N, 103°14′25.22"W, 10 Jan 1987, J. Lomelí 19920 (IBUG), same locality, 150 m a. s. l., 10 Jan 1987, J. Lomelí 20492 (IBUG); km 35 carretera Tecomán-Coahuayana, 16 m a. s. l., 18°40′43.90′’N, 103°41′12.28′’W, 30 Jun 1984, Pérez de la Rosa s.n. (IBUG). Coahuayana de Hidalgo: 22 km después de los límites de los estados de Colima y Michoacán, carretera Manzanillo-Cd. Lázaro Cárdenas, 120 m a. s. l., 18°33′48.83″N, 103°38′23.09″W, 17 Jun 1988, A. Flores Castorena et al. 884 (UAMIZ); El Mirador 5 km al SE de San Juan de Alima, bosque tropical caducifolio, 150 m a. s. l., 24 May 1982, J. Rzedowski 37774 b (F, MO, NY); derrame de lava ca. de un mirador sobre la carretera, ca. San Juan de Alima, carretera Zihuatanejo– Manzanillo, unos 16.5 km al SE del límite con el estado de Colima, 174 m a. s. l., 18°33′48"N, 103°38′50"W, 1 Jun 2010, G. Carnevali 6967 (AMES, CICY, NY). Morelos: Jojutla, A. Espejo 818 (UAMIZ); A. Espejo et al. 2528 (UAMIZ); Temixco, J. Vázquez S. 1977 (MEXU). Tlaquiltenango, J. Ceja et al. 841 (UAMIZ); J. Ceja et al. 847 (UAMIZ); 2 km al S de la desviación a San José de Pala, sobre el camino a Huautla, selva mediana caducifolia, 1150 m a. s. l., 18°32′28"N, 99°00′38"W, 7 Nov 1996, A. Espejo et al. 5607 (UAMIZ); A. Espejo et al. 5609 (UAMIZ); J. García-Cruz and R. Jiménez 1004 (UAMIZ); camino a Huautla, selva mediana caducifolia, 1150 m a. s. l., 18°27′42"N, 99°00′10"W, 16 May 1997, A.R. López-Ferrari et al. (CHIS, UAMIZ, XAL); B. Maldonado sub MORE 5353 (UAMIZ); 2.5 km al SW de Ajuchitlan, 1061 m a. s. l., 18°26′43.95″N, 98°59′19.50″W, 18 May 1995, R. Ramírez Rodríguez 1556 (MEXU); 1 km sobre el camino a Ajuchitlán a partir del camino a Huautla, selva mediana caducifolia, 1160 m a. s. l., 18°27′53"N, 99°00′00"W, 8 Nov 1996, L. Sanchez-Saldaña et al. 384 (AMO); km 7 camino de El Higuerón-Xicatlacotla, selva baja caducifolia, 910 m a. s. l., 18°31′53.81″N, 99°11′36.09″W, A. Espejo 818 (AMO); Puente de Ixtla, 1 km después de La Tigra rumbo a El Zapote, a partir del camino Tehuixtla-Tizapotla, vegetación riparia, 1110 m a. s. l., 18°30′58″N, 99°19′50″W, 22 Jun 1996, J. García Cruz 832 (AMO); 2 km sobre la desviación a Ajuchitlan a partir del camino a Huautla, 1050 m, 18°27′42″N, 99°00′10″W, 16 May 1997, A.R. López-Ferrari et al. 2457 (AMES, UAMIZ). Nayarit: Acaponeta: Acaponeta, approx. 22°29′2.51″N, 105°21′43.94″W, 20 Mar 1897, J.N. Rose 1533 (AMES). Ahuacatlán, Malpaís del Volcán Ceboruco, carretera entre Ahuacatlan y Tepic cerca de Capopales, bosque tropical caducifolio, 1200 m a. s. l., 21°6′6.59″N, 104°32′36.01″W, 14 Jun 1992, M. Cházaro et al. 6957 (XAL); lavas del Ceboruco, lava bed about 8 km west of Ahuacatlan, 20 Jun 1961, approx. 21°5′27.72″N, 104°34′17.73″W, R.L. Dressler 2636 (US); same locality, 10 Jun 1979, F. Castañón sub E. Hágsater 2400 (AMO); Volcán Ceboruco, Malpaís al NW de Ahuacatlán, 1100 m a. s. l., 21°06′05.03″N, 104°32′58.68″W, 9 Jun 1997, González-Tamayo and Harber s.n (IBUG 3-sheets), same locality, 1000 m a. s. l., 21°3′41.96″N, 104°30′46.28″W, 3 Jun 1997, Harker et al. s.n (IBUG). Nayar: Islote 193 al W del ejido de La Palmita, approx. 6 km al E de La Cortina, embalse del P.H. Aguamilpa, 180 m a. s. l., approx. 21°50′13.50″N, 104°48′7.38″W, bosque tropical caducifolio, 20 Aug 1993, J.I. Calzada et al. 18620 (MEXU, MO, XAL); Arroyo de los Bueyes, bosque tropical caducifolio, 90 m a. s. l., approx. 20°50′48.8″N,105°25′24.3″W, 3 Jun 1991, A. Benites Paredes 3121 (MEXU). Ruíz: cafetales afuera del Zopilote, orilla del rio, selva mediana subcaducifolia, 200 m a. s. l., approx. 21°57′22.54″N, 104°56′35.58″W, 5 May 1986, M.A. Soto and G.A. Salazar 2428 (AMO). San Blas: near San Blas, 5–10 m a. s. l., approx. 21°32′47.89″N, 105°15′44.20″W, 10 Jul 1961, R.L. Dressler 2697 (US); old lava flows 2 miles NW Ahuacatlán; among broken rocks on steep wooded slopes, 900 m a. s. l., 19 Jun 1957, R. Mcvaugh 14908 (AMES). Santiago Ixcuintla, ca. Estación Yago, 250 m a. s. l., approx. 21°50′N, 105°04′W, 22 Aug 1936, Otto Nagel 5126 (US); Volcán Ceboruco, 1 Feb 1972, F. Castañón 2400 (AMO). Tepic: km 54.3 del camino de la carretera Tepic-Mazatlán a Jesús María, adelante del poblado del Naranjo, 320 m a. s. l., 22°01′46″N, 104°50′40.2″W, 25 Jun 1999, M.A. Soto and E. Huerta 8676 (AMO). Sinaloa: Culiacán, a 5 km al SE de la presa Sanalona, selva baja caducifolia de Acacia, con algunas Bombacaceas, 24°47.045′N, 107°25.155′W, 6 Jun 1980, R. Vega Aviña 750 (MEXU); camino hacia El Salado hacía tierra adentro, selva baja caducifolia, 50 m a. s. l., 24°29.942′N, 107°10.236′W, 25 Apr 1986, G. Salazar 1952 (AMO); Sierra de Álamos, 26°56.157′N, 108°53.747′W, 17 Mar 1910, J.N. Rose 13062 (US); 4 Apr 1897, J.N. Rose 1533 (US). Mocorito, 5 mi. NO of Pericos, rocky thorn forest, 25°7.147′N, 107°40.890′W, 22 Dec 1949, R.L. Dressler 980 (US). Sinaloa: about 10 km south of Mazatlán, approx. 23°7′29.05″N, 106°18′32.42″W, 21 Jun 1961, R.L. Dressler 2642 (NY, US). Sonora: Álamos, Sierra de Álamos, ladera O de Los Picacho, Rancho La Sierra, selva baja caducifolia, 920 m a. s. l., 26°58′05.3′N, 108°57′17.5′W, 24 Jun 2008, A. García-Mendoza et al. 9162 (MEXU); Sierra de Álamos, 26°56.157′N, 108° 53.747′W, 20 Jun 1993, M.A. Dimmitt s.n. (AMES); 30 miles NE of Álamos, past San Pedro, on rock, 27°10.805′N 109°24.581′W, 1 Apr 1967, Foster 540 (MEXU); Álamos, in the vicinity of Álamos, 27°1.331′N, 108°56.723′W, 18 Mar 1910, Rose, Standley and Russel 13042 (NY); Álamos, obtenida en el patio de una casa dentro del pueblo, 27°1′13.68″N, 108°56′9.66″W, floreciendo en cultivo en Mérida Yucatán, México, 8 Jun 2016, G. Carnevali and A. Can Sulú 7955 (CICY); in the vicinity of Álamos, 27°1′19.86″N, 108°56′43.38″W, 18 Mar 1910, Rose et al. 13042 (NY).
Iconography: Dressler and Pollard (1974): plate 55, color photograph (as Encyclia adenocarpon). Withner (2000): plate 4, color photograph (as E. adenocarpa). Espejo-Serna et al. (2002): 76, line drawing, p. 77, habitat photograph, plate 10, photograph p. 32. Szeszko (2011): 154–155, color photograph, p. 155 (as E. adenocarpa). Hágsater et al. (2005), p. 124, Figure 246 (as E. adenocarpa de Morelos); IOSP site, http://www.orchidspecies.com/encadenocarpa.htm (as E. adenocarpa, the image corresponds to a population from Sonora).
IUCN conservation status: Due to its large distribution area and many known populations, Encyclia adenocarpos is here treated as a LC (least concern) species. As with many species of the genus and particularly members of this complex, E. adenocarpos tends to occur as localized populations, often associated with rocky outcrops, which are easily perturbed by anthropogenic activities. However, the sheer geographical area that covers and the fact that there should be many more intervening populations awaiting to be recorded between those already known (especially in remote places of Sinaloa, Sonora, and Nayarit) suggest that the species as a whole is not severely threatened.
Taxonomic notes: The species was first described as Epidendrum adenocarpon by the Michoacán born botanical team of Antonio de La Llave and Juan José Martínez de Lejarza (“Lexarza”). It was eventually transferred to Encyclia as E. adenocarpa by the German orchidologist Rudolf Schlechter. According to Dr. Kanchi Gahndi at HUH, the basionym epithet adenocarpon is in Greek neuter form; Schlechter erred in citing it as “adenocarpum” (Latin neuter) and changing it to “adenocarpa” (Latin feminine) in transferring the species to Encyclia; it is corrected to adenocarpos (Greek feminine).
Epidendrum adenocarpon var. rosei was described by the Harvard-based team of Oakes Ames, F. Tracy Hubbard, and Charles Schweinfurth from material cultivated at Washington D.C., originally collected by Joseph Nelson Rose and collaborators in the vicinity of Álamos, Sonora. This locality is at the extreme north of the distribution of the species. However, the type and subsequent collections from this and neighboring areas fail to display any significant differences with populations from further south. In fact, they are indistinguishable from some plants collected in Jalisco and Nayarit. Thus, this variety is here treated as a synonym of Encyclia adenocarpos.
Several taxonomic treatments purportedly of this species in floras are actually composites of several species. Examples are those of Ames and Correll (1985: 299) and McVaugh (1985). The first is a mixed description of E. papillosa and E. rodolfoi, the second one of E. adenocarpos, E. acapulcensis, and E. enriquearcilae. That of Dressler and Pollard (1974) includes E. papillosa, and true E. adenocarpos; to our knowledge, E. enriquearcilae and E. acapulcensis had not been collected at that time.
The name Epidendrum crispatum Knowles & Westcott (Floral Cabinet 2: 2 [No. XVII]: 79. [July] 1838. non G.Forst. 1786 (= Dockrilia crispata (G.Forst.) Rauschert), (TYPE: Mexico, ex Hort. G. Barker s.n., not located) has been referred to the synonymy of Encyclia adenocarpos (CONABIO 2009). Although we have not seen the type, the protologue of the name clearly states that “… the side portions are wrapped around the column …”. The “side portions” referred to in the description are undoubtedly the lateral lobes of the labellum. No species of the Encyclia adenocarpos complex features such a labellum. Thus, the name is rejected as applicable to a potential member of the complex.
Etymology: This species is named after Enrique Arcila Arcila, from Mérida, Yucatán, who grew a small plant of this species for several years that became a specimen large enough to provide cuttings to prepare the type material.
Description: Epiphytic herbs, 40–90 cm with the inflorescences. Rhizome short, tough, woody. Pseudobulbs 3.0–7.0(–8.0) × 1.0–3.0 cm, clustered but shortly creeping, pyriform, apically 2-3-leaved, green and smooth when young and then clothed by 1-2 thin membranous, papyraceous, non-persistent sheaths, which are eventually deciduous. Leaves 29–35 × (0.5–)0.7–0.9(–1.1) cm, linear elliptic, acute thick, rigid, midnerve carinate mainly in the lower half, blades coriaceous. Inflorescences 45–80 cm long, terminal, erect, a simply pinnate panicule bearing 2–4(–6) lateral branches, (4–)8–15 cm long, each bearing 2-5 flowers for a total of 5–25 flowers per inflorescence; peduncle 20–30 cm long, lightly verruculose, thin but fairly strong clothed by 4–8 remote, tubular sheaths 1.0–1.9 × 0.3–0.45 cm, oblong elliptic, acute, internodes 4.5–7 cm long, naked in the upper ¾; rachis lightly verruculose, bracts subtending the branches 0.5–0.6 mm long, similar to those of the peduncle, all of the inflorescence is light green. Ovary with pedicel 1.9–25 cm long, surface laxly but conspicuously verruculose, cylindrical, thicker at apex. Flowers resupinate, relatively showy, fairly lax on the inflorescences, 24–26 mm across the spread apices of the petals; perianth segments sub-fleshy; sepals and petals pale green or yellow green, the apical portion heavily overlaid with dull purplish maroon, the veins showing faintly; the labellum ground color pale yellow-white to pale yellow, the central portion paler, the midlobe with 5-7 bright purple, short (> 0.5 cm) stripes, lateral lobes pale yellow, the callus basally white, the apical half pale yellow, darker toward the apex; column dorsally white at base, apically yellow, the ventral surface white in the proximal half, the distal half pale bronzy yellow-orange, the stigmatic surface pale yellow; sepals subsimilar but the lateral somewhat oblique, dorsal sepal (1.3–)1.5–1.8 × 3.5–4.5 cm, linear obovate to narrowly obovate, broadly acute to obtuse; lateral sepals 1.6–1.7 × 0.5 cm; petals 1.38–1.70 × 0.40–0.44 cm, linear obovate to linear spatulate, broadly acute to obtuse, attenuated basally to 1–1.5 mm wide; labellum 3-lobed, free from column except at base where fused for 2.5–3 mm to the ventral face of the column, the lateral lobes free to base, total labellum length 1.19–1.3 cm, 0.9–1.1 cm across spread lateral lobes; the central lobe 0.65–0.9 × 0.81–1.0 cm, subquadrate to suborbicular, rounded to subtruncate at apex and base, apex emarginate, the margins coarsely undulate, the blade with raised veins, particularly toward the base; lateral lobes irregularly and obliquely triangular to oblong triangular, perpendicular to labellum main axis, acute to broadly acute, the anterior (shorter) margin 0.34–0.44 cm, the posterior (longer) margin 0.5–0.55 cm, basal width (upon merging with labellum disk) 0.25–0.35 cm, width at midlength 0.12–0.15 cm; callus 0.45–0.49 × 0.41–0.43 cm, rhomboid obovate, apically subacute to rounded, massive, consisting of two obliquely elliptic, flat, smooth plates 0.20–0.21 cm wide running from the base of the free portion of the labellum to the base of the central lobe, from its apex emerge 5–7 thick veins, 0.10–0.12 cm long and thus reaching the lower third of the central lobe. Column 0.79–0.85 × 0.43–0.47 cm, narrowing to a 0.17–0.18 cm wide base, broadly obovate, lengthwise thicker along the central portion, the margins much thinner, wing-like particularly on the distal half where it is produced into a pair of flat, broadly triangular, porrect auricles, these 0.6 mm long; stigmatic surface obcordate, 0.9 × 1.1 cm, clinandrium shallowly 3-dentate; anther 0.2 mm wide and thick, pollinia 0.08 mm along the longer axis. Capsule 3.2–3.5 × 1.5–1.6 cm, ellipsoid, heavily verruculose.
Diagnosis: An Encyclia species phenetically related to E. papillosa with similar suborbicular flowering pseudobulbs and color pattern but narrower petals and sepals, with smaller, narrowly oblong triangular or narrowly triangular, acute lateral lobes to the labellum (vs obovate, rounded to obtuse) that are also perpendicular to labellum main axis (vs lateral lobes at an 40–45°angle); also, the inflorescences are longer and laxer.
Notes: Encyclia enriquearcilae is phenetically closest to Encyclia papillosa from which it is allopatric. Both share a generalized habit (pseudobulbs tend to be subspheroid when turgid) and flower morphology, particularly the massive rhomboid or trapezoidal callus that covers much of the labellum disk combined with the bright yellow column apex and lateral lobes to the labellum. However, E. papillosa features broader (3.8–4.4 mm), longer (6–6.6 mm along the main axis) lateral lobes to the labellum; furthermore, these are obovate, rounded apically and are partially porrect (at a 41°–46° angle to main labellum axis). In addition, the inflorescences in E. enriquearcilae are longer (to 80 cm) and laxer than in E. papillosa. Encyclia papillosa occurs from Nicaragua north of the lakes northwesternward to the Tehuantepec Isthmus. No records of E. papillosa exist from west of the Tehuantepec Isthmus. Encyclia enriquearcilae, instead, grows along the Oaxacan coast from Lagunas de Chacahua northward just across the Guerrero border at elevations from near sea level to ca. 150 m. There are ca. 500 km in straight line from the Laguna de Chacahua locality to the nearest known populations of Encyclia papillosa in SW Chiapas.
Paratypes: MEXICO. Oaxaca: Municipio Juquila, 4.5 km al SE de Charco Redondo camino a Las Salinas, Parque Nacional Lagunas de Chacahua, 1–3 m a. s. l., selva baja caducifolia, 20 May 1980, M. Martínez 2231 (MEXU!); same locality and date, M. Martínez 2232 (MEXU!); same locality of type, collected June 11, 2012, along with J. P. Pinzón, C. Jiménez Nah and A. Castillo; grown at Mérida, Yucatán, Mexico, 23 May 2013, G. Carnevali 7694 (AMES!, AMO!, CICY!, MO!, OAX!). Guerrero: Municipio Ometepec, Zacualpan, orilla del río Mazapa, cerca del límite entre Oaxaca y Guerrero, ca. 70 m a. s. l., colectada originalmente por Imelda Santiago Ramírez en el 2012, E.A. Pérez-García 2617 (AMO!, CICY!). San Luis Acatlán: Jolotichán, 251 m a. s. l., 16°44′47"N, 98°54′59"W, on tropical dry forest, A. Cano Nava s.n., (photograph).
IUCN conservation status: EN (Endangered). Encyclia enriquearcilae meets criteria B1a and B2a of the IUCN (2010). It is currently known from only four localities, and its Extent of Occurrence is of 2.725.05 km2, whereas its Area of Occupancy is of 16.00 km2 (with a cell size of 2 km). One of the populations resides inside the Parque Nacional Lagunas de Chacahua where it is afforded some degree of protection, whereas the other three inhabit areas where the tropical dry forest has been severely fragmented due to cattle ranching and “milpas”, thus restricting the species to isolated forest remnants where small populations survive. Encyclia enriquearcilae is rare in nature, apparently occurring as isolated populations of few individuals, but there are several plants of the species under cultivation in private collections and botanical gardens (including a recent batch of seed-raised plants) thus ensuring the long-term survival of the species ex situ.
Etymology: The epithet refers to the “papillae” (actually warts) of the pedicel and ovary of the species.
Description: Epiphytic, more rarely lithophytic herbs,16–31 cm tall, 23–71 cm counting the inflorescence. Rhizome thick, fibrous. Pseudobulbs 2.8–6.0 × 1.5–2.0 cm, aggregated, pyriform to typically subspheroid, apically (1-)2-3-foliate, enveloped with papyraceous, eventually evanescent sheaths, plump and smooth when young, wrinkled upon maturation. Leaves 16–31 × 0.41–2 cm, linear oblong, acute, often slightly falcate, thickly fleshy coriaceous, dark green. Inflorescences 23–71 cm long, terminal and emerging from the mature pseudobulb, suberect, patent or arching, (3–)8–15(–25)-flowered, peduncle and rachis verruculose, but the rachis less so, racemose or, more commonly, paniculate with (1-)2-3(–6) lateral branches, the lateral branches bearing (2-)3–5(–7) flowers; peduncle wiry (relatively thin, strong but flexible), deep green; peduncle bracts 3–12 × 0.4–5 mm, inconspicuous, oblong-ovate, acute, floral bracts 1–3 × 3–5 mm, similar to the bracts of peduncle, successively smaller distally, inconspicuous, thinly fleshy when fresh, papyraceous when dry; ovary with pedicel 13–17 mm long, verruculose, green or green with red tinges. Flowers 21.3–23.6 mm across the spread petal tips, resupinate, showy as compared with the flowers of other members of this complex; petals and sepals yellow green to pale green, brown or pale maroon tinged, particularly on the distal half, the color usually more conspicuous on the veins, often showing reticulations; labellum usually bright white, the base of the lobes bright yellow, more rarely the whole labellum pale yellow (not to be confused with the general yellowing of the entire labellum as the flowers ages); the central lobe with (1–)3–5(–9) purple veins, rarely colored veins lacking; lateral lobes white, the proximal half yellow or green-yellow, often the entire lobe yellow; callus white, distally tinged yellow, the yellow color often extending to the margins of the callus; column proximally white or more commonly pale yellow, bright yellow on the distal half; anther white or very pale yellow; sepals elliptic-spatulate to narrowly elliptic-spatulate, the laterals somewhat oblique, rounded-obtuse to broadly acute, more rarely acute; dorsal 15–17 × 3.5–3.9 mm, laterals 14.6–16.1 × 4.3–5.2 mm, distally concave with flat to somewhat reflexed margins; petals 14.1–15.5 × 2.0–2.4 mm, linear spatulate to narrowly spatulate, rounded to broadly acute, strongly narrowed proximally; labellum 14.1–15.2 mm long, fused to the column at the proximal 1/5, then free, sharply 3-lobed, 14.7–15.5 mm across the spread apices of the lateral lobes; central lobe 7.5–8.7 × 9.1–10.5 mm, suborbicular, subquadrate to broadly ovate-elliptic, rounded to truncate, more rarely very broadly obtuse, margins and surface ondulate, disk with 3-5 thick veins arising from the callus and reaching the proximal third; lateral lobes 5.5–6.5 × 3.4–4.1 mm, widely spreading, flat to slightly suberect, obovate, elliptic-obovate to oblong elliptic, rounded to broadly ovate; callus 4.9–5.6 × 3.9–4.2 mm, broadly obovate-elliptic, apex broadly obtuse, composed of two flat, broad keels bisected by a narrow, linear fovea, deeper in the proximal half, the two basal endings slightly bilobed, the apex of the callus is thick and merges into the several veins that enter the central lobe. Column 8.3–9.3 × 3.8–4.5 mm, straight, depressed-hemicylindric, obovate, basally clavate attenuate, no obvious stelidia, dorsally with a thick keel; anther 1.7–1.9 mm long, ovate-subcordate in dorsal view; clinandrium smooth, with no lateral teeth, pollinia 0.8–1.1 × 0.4–0.5 mm, yellow, in two subequal pairs; stigmatic surface 2.4–2.6 × 2.3 mm, obcordate, proximally acute, distally truncate. Capsule 34–38 × 7–8 mm, densely verruculose, ellipsoid.
Distribution area and habitats: Encyclia papillosa has one of the largest distributional ranges of the E. adenocarpos clade, occurring from the Lago de Nicaragua to the south up to the southern section of the central Depression of Chiapas, approximately 800 km in straight line, always along the Pacific drainage. Along this huge distributional area, the species is known to occur in several types of seasonally dry ecosystems. It has been reported both as an epiphyte or a lithophyte. We have also collected it as a subterrestrial on steep slopes in Chiapas. Elevations range from sea level to ca. 1300 m, but most often occurs below 700 m. It usually grows on exposed positions, more rarely in the shade. It has often been collected growing as an epiphyte over Crescentia cujete L., the jicara or jicaro of Central American countries.
Notes: Encyclia papillosa has a great deal of internal variation, and this variation is somewhat geographically correlated, as opposed to the variation found in E. adenocarpos. Populations from E. papillosa from Nicaragua through southern Guatemala have narrower lateral lobes to the labellum than those from Mexican populations. Some Nicaraguan populations have particularly narrow, acute, lateral lobes and in that somewhat resemble E. enriquearcilae. However, in this latter species, the lateral lobes are even narrower and are totally flat, whereas the narrow-lobed populations of Nicaragua display somewhat erect (or even slightly porrect) lateral lobes.
Encyclia papillosa is easily distinguishable from similar species of the clade on account of its relatively large, broad, lateral lobes of the labellum, which is white with just a few (or none) purple veins. However, it is particularly noteworthy because of its bright yellow column and bases of the lateral lobes. The related E. enriquearcilae has much narrower lateral lobes which are totally yellow. Both species have orbicular pseudobulbs when turgid, as opposed to the most commonly ovoid or pyriform pseudobulbs of other members of the clade.
Additional specimens examined: EL SALVADOR. Chalatenango: Near the Lempa, approx. 13°58′22.64″N, 89°0′1.05″W, epífita, 30 May 1924, Calderón 30 (AMES); La Libertad: Comasagua, 900 m a. s. l., approx. 13°38′22.28″N, 89°22′44.38″W, Apr 1941, M.L. von Séveren s.n. (AMES); Morazán: Before Osicala, Puente Torola, 575 m a. s. l., approx. 13°50′25.06″N, 88°8′45.89″W, epífita, 23 Mar 1967, F. Hamer 27A (AMES); San Salvador: Between Apopa and Toma de Aguilares, 400 m a. s. l., approx. 13°48′50.05″N, 89°14′32.67″W, sobre árboles de Morro, 3 Apr 1963, F. Hamer 27 (AMES, F, MO, SEL); Garden of María Reyes Guerra, 4 May 1939, S. Calderón s.n. (AMES); Jardín Botánico La Laguna, zona 26 N, 800–850 m a. s. l., (approx. 13°40′10.81″N, 89°14′49.57″W), 17 May 1986, J. Flores 67 (MO). Santa Ana: ca. Finca El Jardín, ± 5.5 km de Santa Ana, 600 m a. s. l., approx. 14°2′24.09″N, 89°31′14.84″W, 27 Apr 1994, J.L. Linares and C.A. Martínez 1328 (EAP, MO). Sonsonate: Pedregal de San Isidro, approx. 13°48′39.57″N, 89°35′09.42″W, 800 m a. s. l., F. Hamer s/n (SEL). GUATEMALA. Baja Verapaz: Purulha, 11 Jul 1978, J. D. Ackerman 1223 (F, SEL). Chiquimula: Chiquimula, 350–400 m a. s. l., approx. 14°47′05.55″N, 89°32′01.04″W, 1 Apr 2003, M. Véliz and M. Pérez 13256 (BIGU); 300–400 m a. s. l., 14°51′03″N, 89°31′02″W, 6 May 2003, M. Véliz and M. García 13448 (BIGU). El Progreso: Sanarate, 742 m a. s. l., 14°50′14.20″N, 90°19′40.20″W, 10 Apr 2011, M. Véliz and 23004? (BIGU-57843); 4 km East of El Rancho, on Highway 4, 440 m a. s. l., approx. 14°53′45.46″N, 89°59′1.55″W, 16 Jun 1971, W. E. Harmon and J. A. Fuentes 5670 (ENCB); Rancho, 275 m a. s. l., approx. 14°54′31.99″N, 89°59′56.94″W, on tree on mountain, 8 Jun 1909, C.C. Deam 6249b (AMES, US); El Rancho, 275 m a. s. l., 14°54′31.99″N, 89°59′56.94″W, 23 Jul 1906, W.R. Maxon 3772 (US); at miles 136.12 on main line IRCA (International Railways of Central America), 275 m a. s. l., approx. 14°54′2.52″N, 89°59′34.28″W, Jul 1937, M.W. Lewis 121 (AMES); Morazán, 200–300 m a. s. l., 14°55′33″N, 90°00′12″W, 27 May 2003, M. Véliz and M. García 13506 (BIGU); San Agustín Acasaguastlán, Aldea El Rancho, 300–400 m a. s. l., 14°54′46.80″N, 90°02′06.00″W, 9 May 2003, A. Cóbar and F. Ramírez 352 (BIGU); 200–300 m a. s. l., 14°53′28″N, 90°01′17″W, 7 May 2003, F. Ramírez and A. Cóbar 638 (BIGU); San Agustín Acasaguastlán, 200–300 m a. s. l., 14°53′46″N, 89°57′34″W, 8 May 2003, A. Cóbar and F. Ramírez 319 (BIGU); 300–400 m a. s. l., 14°54′43″N, 89°55′52″W, 6 May 2003, M. García and M. Véliz 559 (BIGU); 14°55′33″N, 90°00′12″W, 7 May 2003, M. Véliz and M. García 13517 (BIGU); 14°54′42″N, 90°02′08″W, 4 Apr 2003, M. Véliz and M. Pérez 13422 (BIGU); 200–300 m, 14°54′43″N, 89°55′52″W, 6 May 2003, M. García and M. Véliz 543 (BIGU); Sanarate, Finca San Miguel, 773 m a. s. l., 14°51′03.38″N, 90°16′30.18″W, 23 Aug 2010, J. Valdez and F. Gomes JVQ1 456 (BIGU); 1041 m a. s. l., 14°50′09.23″N, 90°18′25.51″W, 23 Aug 2010, J. Valdez and F. Gomes JVT2 407 (BIGU); 784 m a. s. l., 14°49′04.99″N, 90°17′36.26″W, 25 Apr 2011, J. Valdez and C. Reyes JVQ3 1049 (BIGU); 1041 m a. s. l., 14°50′09.23″N, 90°18′25.51″W, 25 Apr 2011, J. Valdez and C. Reyes JVT2 1074 (BIGU); 773 m a. s. l., 14°51′03.38″N, 90°16′30.18″W, 25 Apr 2011, J. Valdez and C. Reyes JVQ1 1035 (BIGU); 25 Mar 2011, J. Valdez and C. Reyes JVQ1 999 (BIGU); 912 m a. s. l., 14°50′09.83″ N, 90°18′43.52″ W, 23 Aug 2010, J. Valdez and F. Gomes JVT4 436 (BIGU); Peña de la Virgen, 881 m a. s. l., 14°48′59″N, 90°18′44.10″W, 10 Apr 2011, M. Véliz 23089 (BIGU); Paso de los Jalapa, 300–400 m a. s. l., 14°52′33″N, 89°58′24″W, 8 May 2003, M. Véliz and M. García 13525 (BIGU). Guatemala: Carretera a Puerto Barrios, km 25 a Ciudad de Guatemala, 1000 m a. s. l., 14°46′59″N, 90°13′48″W, 16 May 1963, A. Molina R. and A. R. Molina 12339 (F); Garden of Don Mariano Pacheco, 1650 m a. s. l., (originally collected at Depto. Santa Rosa), 4 May 1942, J.A. Steyermark 46404 (F). Huehuetenango: Jacaltenango, Aldea Cuatro Caminos, 600 m a. s. l., approx. 15°33′48.81″N, 91°44′38.94″W, 13 Mar 2002, M. Véliz and 12407b (BIGU). Jutiapa: Plains and swampy ground along railroad, between Agua Blanca and Amatillo, 950–990 m a. s. l., approx. 14°30′24.95″N, 89°36′34.26″W, 24 Oct 1939, J.A. Steyermark 30431 (F). Zacapa: Cabañas, 200–300 m a. s. l., 14°54′51″N, 89°47′01″W, 6 May 2003, M. García and M. Véliz 588 (BIGU); camino hacia Chiquimula, 200–300 m a. s. l., approx. 14°53′29.48″N, 89°31′16.79″W, 8 Mar 2003, M. García and F. Ramírez 264 (BIGU); Km 94 camino a San Cristóbal Acasaguastlán, approx. 14°58′11.33″N, 89°46′41.92″W, 9 Mar 2003, A. Cóbar and M. Véliz 195 (BIGU); Gualán, Carretera de terracería a Zacapa, 236 m a. s. l., approx. 15°05′08.50″N, 89°02′ 53.20″W, 22 May 2008, M. Véliz and 20237 (BIGU); Río Hondo, 200–300 m a. s. l., 15°01′32″N, 89°35′06″W, 6 May 2003, M. García and M. Véliz 527 (BIGU); Zacapa, 200–300 m a. s. l., 14°56′39″N, 89°31′43″W, 6 May 2003, A. Cóbar and F. Ramírez 312 (BIGU); 300–400 m a. s. l., 14°56′24″N, 89°32′19″W, 6 May 2003, A. Cóbar and F. Ramírez 293 (BIGU). HONDURAS. s.loc., s.d., A. Zimmerman 2604 (TEFH); no department mentioned but probably Comayagua: 40 km from Tegucigalpa, north road near monument, flowering in cultivation at the Marie Selby Botanical Gardens, 29 May 1974, Nepple s.n. (SEL 000327). Choluteca: 8 km Southeast of Choluteca, 200 m a. s. l., approx. 13°14′47.13″N, 87° 9′19.00″W, epífita, 2 Jun 1965, R.W. Lent 596 (F); Near San Antonio de Flores, 100 m a. s. l., approx. 13°39′37.28″N, 87°21′44.71″W, epífita, 3 May 1948, L.O. Williams 14350 (F); Carretera 5, San Antonio de Flores, 75 km S of Tegucigalpa, 90–140 m a. s. l., 13°40′N, 87°23′W, 2 May 1996, T. Hawkins 1005 (EAP, MO). Comayagua: Valle between Comayagua and Villa de San Antonio, 570 m a. s. l., approx. 14°24′55.65″N, 87°38′54.56″W, epífita, 1 May 1947, L.O. Williams and A. Molina R. 12608 (F); Quebrada a lo largo de la carretera Villa San Antonio a Flores, 600 m a. s. l., approx. 14°18′10.96″N, 87°34′47.02″W, epífita, 18 Apr 1957, A. Molina R. 8281 (F, NY); Ajuterique, 750 m a. s. l., approx. 14°22′48.01″N, 87°43′3.85″W, 31 Mar 1945, J.V. Rodríguez 2607 (F). Francisco Morazán: Campus of Escuela Agrícola Panamericana, Zamorano, 800 a. s. l. approx. 14°0′32.84″N, 87°0′33.60″W, 800 m, “on Crescentia cujete”, 29 May 1947, A. Molina R. 2 (AMES, F); El Zamorano, 32 km E de Tegucigalpa, 800 m a. s. l., approx. 14°0′34.62″N, 87°0′39.28″W, 10 Aug 1986, C. Sosa 7 (MO, UNAH; fruiting); 30 km NE Tegucigalpa, Montaña La Tigra, 1700 m a. s. l., approx. 14°11′38.21″N, 87°7′29.05″W, bosque montano húmedo, 16 Feb 1972, A. Clewell 3033 (TEFH); Río Yegüare (Campus El Zamorano), 800 m a. s. l., approx. 14°0′7.47″N, 86°59′16.15″W, Apr 1944, J.V. Rodríguez 2112 (AMES); Cerro San Pedro, Maraita, 45 km E of Tegucigalpa, 640 m a. s. l., 13°51′N, 87°06′W, 9 May 2011, K. Barahona 32 (TEFH). Intibucá: Valle de Jesús Otoro, matorrales cerca del puente sobre el Rio Grande, 600 m a. s. l., approx. 14°28′7.32″N, 87°59′59.46″W, epífita sobre Bursera simaruba, 25 May 1964, A. Molina R. and A.R. Molina 14113 (F). Valle de Otoro, ca. 10 km W of Jesús de Otoro, vic. Río Sirima, 800 m a. s. l., 14°33′N, 88°06′W, 3 Jun 1991, G. Davidse 34950 (EAP, MO, TEFH). Valle: Isla de Zacate Grande, La Guayaba, Andino 24a (TEFH); same locality, 0 m a. s. l., 13°19′8.14″N, 87°34′56.27″W, 9 May 1991, R. Varela 348 (TEFH). MEXICO. Chiapas: Comitán de Domínguez: ca. 3.2 km SW of Berlin, 2.95 km ENE of Francisco J. Mújica, along the road from Comitán to the Belisario Domínguez Dam, 640–650 m, 16° 3′24.05″N, 92°11′20.79″W, flowering in cultivation in Mérida, Yucatan, 14 Jun 2017, G. Carnevali 8060 (AMES, CICY). Frontera Comalapa: ca. El Recuerdo, 620 m a. s. l., 15°46′24.57″N, 91°58′46.45″W, flowering in cultivation, 15 Jun 2017, G. Carnevali 8092 (photovoucher, CICY). Motozintla: ca. Motozintla, 1290–1300 m a. s. l., approx. 15°22′15″N, 92°14′54″W, flowering in cultivation, 19 May 2008, G. Carnevali and I.M. Ramírez 7339 (CICY). Yucatán (doubtful): km 35 Kinchil-Celestún, 10 m a. s. l., approx. 20°51′14.88″N, 90°16′20.16″W, 2 Jul 1989, J.M. Andrews 745 sub/E Hágsater 7924 (AMO). NICARAGUA. Boaco: San José de los Remates, “La Majada”, 380–400 m a. s. l., 12°37′N, 85°51′W, 10 May 1982, P.P. Moreno 16292 (AMO, MO, SEL), 4 km S de Boaquito, “San Antonio”, 12°26′N, 135°44′W, 200 m a. s. l., 12 Oct 1982, A. Moreno 18017 (MO); Boaco: cerca de Camoapa, 500–550 m a. s. l., approx. 12°23′21.41″N, 85°29′52.12″W, 10 May 1975, J.A. Atwood A148 (MO). Chinandega: municipio de Cinco Pinos, comunidad Las Pozas, quebrada Las Pozas, 275 m a. s. l., 13°11′N, 86°51′W, 4 May 2004, I. Coronado and J. Soriano 641 (HULE, MO, NY); La Pavona, 6 km al N de Somotillo, camino a Cinco Pinos, 60–80 m a. s. l., 13°05′N, 86°53′W, 28 Sep 1981, P.P. Moreno 11527 (MO); Km 142.5 carretera a Corinto, 50–60 m a. s. l., approx. 12°30′8.05″N, 87°10′52.63″W, 26 April 1982, J.C. Sandino 2601 (MO). Chontales: In plains on road to Camoapa (as “Camuapa”) from Comalapa, approx. 12°19′4.21″N, 85°30′34.07″W, 10 May 1975, J.T. Atwood Jr. A148 (NY); along Hwy. 7 at Comalapa road entrance, 150 m a. s. l., ca. 12°10′N, 85°33′W, 19 May 1980, W.D. Stevens and O.M. Montiel 17168 (MO); 1.8 km N of Comalapa on road to Camoapa, ca. 300 m a. s. l., ca. 12°17′N, 85°30′W, 19 May 1980, W.D. Stevens and O.M. Montiel 17198 (MO); Municipio Comalapa, camino entre la carretera al Rama y el pueblo, 200–300 m a. s. l., approx. 12°15′0.38″N, 85°31′46.01″ W, 10 July 1980, M. Guzmán, D. Castro and A. Montiel 314 (MO); sobre el camino entre Juigalpa y Puerto Diaz, 31–130 m a. s. l., approx. 12° 3′11.04″N, 85°24′37.56″W, 11 Jul 1980, M. Guzmán D. Castro and A. Montiel 369 (MO; fruiting); Hacienda Corpus, 11 km W of Juigalpa, 100 m a. s. l., 12°07′N, 85°28′W, J.S. Miller and M. Nee 1338 (MO, fruiting). Jinotega: Cordillera Dariense, below Cerro Chimborazo, 1200 m a. s. l., approx. 13°4′16.08″N, 85°59′11.44″W, 24 May 1975, J.T. Atwood 242a (HNMN, MO); ca. 0.6 km from Hwy 3 on road to Aranjuez, ca. 1430 m a. s. l., ca. 13°01′N, 85°55′W, 9 Apr 1978, W.D. Stevens 7549 (HNMN, MO). León: Isla Momotombito, NW slope, 40–300 m a. s. l., ca. 12°21′N, 86°28′W, steep slopes of broken basalt, W.D. Stevens and M. Araquistain 13238 (AMES, MO; fruiting); Along Hwy 12, ca 1 km SE of junction with Hwy 28 (first quebrada SE of junction), ca. 30 m a. s. l., ca. 12°15′N, 86°43′W, rocky to muddy stream bed and level rocky savanna, 28 May 1980, W.D. Stevens 17274 (ENCB, SEL); same locality, 17 Dec 1978, W.D. Stevens 11172 (HNMN, MO; fruiting specimen); along Hwy 12 ca. 1 km SE of junction with Hwy 28 (first quebrada SE of junction), ca. 30 m a. s. l., ca. 12°15′N, 86°43′W, 28 May 1980, W.D. Stevens and O.M. Montiel 17274 (MO); carretera León-Managua, cerca Rio Tamarindo, approx. 12°14′33.50″N, 86°42′55.87″W, 24 May 1975, J. Atwood s.n. (HNMN, MO); Rt. 12 north and Rio Tamarindo, ca. 30 m a. s. l., approx. 12°14′33.50″N, 86°42′55.87″W, 24 May 1975, J.T. Atwood A245 (MO; MSC); sobre la carretera vieja a León, 1 km antes del empalme con la carretera nueva, km 64, 60 m a. s. l., 12°16′N, 86°44′W, P.P. Moreno 3553 (MO); carretera vieja a León, ca. Empalme Izapa, 40–60 m a. s. l., 12°16′N, 86°44′W, 12 Sep 1980, M. Gumán 1038 (MO); Municipio de la Paz Centro, Hacienda el Tempate, 100 m a. s. l., 12°17′N, 86°46′W, 2 Jun 1998, R. Rueda 8138 (HULE, MO). Managua: Lava Fields near Managua, collected by Frank Mathews, flowering in cultivation at the Marie Selby Botanical Gardens, 26 May 1978, A. Pridgeon s.n. (SEL 019405); 2 km E de San Francisco Libre, 60-80 m a. s. l., 12°30′N, 86°15′W, 6 Jun 1983, P.P. Moreno 21448 (AMO, MO); Carretera a San Francisco Libre, 1 k despues del Río San Antonio, 50–60 m a. s. l., 12°23′N, 86°09′W, 21 Oct 1980, P.P. Moreno 3805 (MO; fruiting); along road from Hwy 1 (N of San Jacinto entrance) to San Francisco del Carnicero, ca 4.8 km W of Hwy 1, 100 m a. s. l., approx. 12°21′22.19″N, 86° 4′37.65″W, low thorny forest on heavy black (soil?), 9 May 1980, W.D. Stevens 17113 (ENCB, MO, SEL); about 2.4 km SW of Rio Pacora bridge on road between San Francisco del Carnicero and Hwy. 1, ca. 70 m a. s. l., ca. 12°26′N, 86°11′W, 3 Jun 1979, W.D. Stevens 13150 (MO); south shore of Lago de Managua, ca. km 31 on carretera nueva a León near Piedras Azules, 40–50 m a. s. l., ca. 13°15′N, 86°28′W, 3 Jun 1979, W.D. Stevens 13151 (MO); a 3 km de la carretera Managua-Sebaco, camino a San Francisco Libre, 100–120 m a. s. l., approx. 12°29′42.01″N, 86°15′51.43″W, 26 Jan 1981, J.C. Sandino 362 (MO; fruiting); 8 km de carretera Managua-Sébaco, camino a San Francisco Libre, 50–100 m a. s. l., approx. 12°22′48.16″N, 86°7′13.08″W, 26 Jan 1981, J.C. Sandino 380 (MO; fruiting); La Mojarra, 60–80 m a. s. l., 12°35′N, 86°21′W, 6 Jun 1983, P.P. Moreno 21437 (HNMN, MO); 9 km ENE of Tipitapa (as “Piptapa”), 60 m a. s. l., approx. 12°18′51.96″N, 86°0′33.66″W, 22 Sep 1976, Avinoan Danin 76-15-9 (MO; sterile); 20 km NE of Tipitapa, 60 m a. s. l., approx. 12°26′31.32″N, 86°2′20.20″W, 22 Sep 1976, Avinoan Danin 76-16-5 (MO; fruiting); along Hwy. 1, ca. 8 km N of Las Maderas, ca. 340 m a. s. l., ca. 11°29′N, 86°03′W, W.D. Stevens 11132 (MO; fruiting); W of Quebrada Las Ruedas, N of road, NW of El Tránsito, 15–30 m a. s. l., ca. 12°05′N, 86°43′W, 16 Oct 1977, W.D. Stevens 4699 (MO; fruiting), flowering in cultivation from same population, 27 Jun 1978, W.D. Stevens 9057 (MO); same locality, 13 May 1981, W.D. Stevens 20158 (MO); 3 km E de Las Maderas, camino a Aseses, 80 m a. s. l., 12°26′N, 86°04′W, 19 Nov 1982, P.P. Moreno 18747 (HMNM, MO; fruiting). Masaya: Laguna de Masaya, 1.5 km al S de Nindirí, parte NW de la laguna, 140–160 m a. s. l., 11°08′N, 86°08′W, bosque tropical seco, 11 May 1980, P.P. Moreno 387 (ENCB, MO); Laguna de Masaya, 120–160 m a. s. l., 11°59′N, 86°08′W, 30 May 1981, P. Moreno and J. Heinrich 8932 (HMNM, MO; fruiting); 2 km W del valle de Tisma, 50–80 m a. s. l., 12°05′N, 86°02′W, 7 Nov 1980, P.P. Moreno 4370 (MO; fruiting); Parque Nacional Volcán Masaya, ca. 1.7 km from Hwy. on road along W shore of Laguna de Masaya, 135–145 m a. s. l., ca. 11°59′N, 86°07′W, 21 May 1980, W.D. Stevens 17209 (MO); Parque Nacional Volcán Masaya sobre la costa O de la Laguna de Masaya, a 1.7 km de la carretera sobre el camino de tierra, 135–145 m a. s. l., 11°59′07 N, 86°07′W, 16 Apr 1980, M. Guzmán and D. Castro 75 (MO). Matagalpa: SW slopes of Cerro El Pílon and adjacent Laguna Tecomapa, 420–540 m a. s. l., 12°37′N, 86°02′W, 20 Jul 1978, W.D. Stevens 9433 (F, MO; fruiting). Nueva Segovia: 3 km W of Ocotal, 620–630 m, approx. 13°37′56.87″N, 86°30′46.98″W, 20 Dec 1968, J.T. Atwood 3940 (MO; fruiting).
IUCN conservation status: Least Concern (LC). Encyclia papillosa has quite an extensive distributional range (see above), exceeding 36000 Km2 and occurs in a variety of seasonally dry environments. Encyclia papillosa often survives as an epiphyte in calabash trees left standing in extensive pastures, or in mangrove associations. It also occurs as a subterrestrial or lithophyte on road cuts, a demonstration of the resilience and adaptability of the species. Although some collecting for horticultural purposes occurs (as this is possibly the showiest species of the complex), and some of the areas where this taxon occurs have been transformed for extensive cattle ranching, we suspect that the species is not currently threatened.
Iconography: Encyclia papillosa was figured as E. adenocarpos in Dressler and Pollard (1974: plate 55). These authors considered all the members of the clade to belong to only two species, one being E. adenocarpos with relatively small flowers, the other, E. trachycarpa, with larger flowers. The image at the IOSPE site (http://www.orchidspecies.com/encpappilosa.htm) represents a totally unrelated species.
Taxonomic notes: Encyclia schaeferi was described from a plant collected at the San Marcos Department in Guatemala and named after Hanno Schaefer, from the Munich Technical University (Technische Universität München) “… a student of plant biodiversity whom has supported the research work of the species’ authors…” (our translation). The original plant was described as lithophytic in dry forest. We have not been able to locate the type collection or any similar plants among the very large sample of the species we evaluated. One of us (WCI), visited the herbarium BIGU, where the type collection was supposedly deposited but was unable to locate it. Judging from the original description and iconography (a line drawing that is but a caricature of a flower and a poor-quality photograph) we assume this to be an extreme or teratological form of E. papillosa with a narrow central lobe and somewhat reduced lateral lobes. In fact, some specimens from Guatemala (e.g., C. Leopardi 428-01, CICY, from ca. La Virgen) and Nicaragua (e.g., Sune Holt E0318) or from Guatemala (without precise locality) somewhat resemble Archila’s crude drawing and photograph.
Encyclia papillosa has been reported from near Celestún, on the extreme NW of the Yucatan Peninsula (Andrews and Gutiérrez 1988, Carnevali et al. 2001). The record is based upon a single collection, J. M. Andrews 745 sub/E Hágsater 7924 (AMO), also vouchered as Carnevali 6203 (CICY, pickled). The specimen was prepared from a cultivated plant grown at the garden of J. Andrews, of Mérida, Yucatán. When the senior author visited this garden in 1997, the collector had the plant growing on a tree, unlabeled, along with many other orchids, including exotic species. We have explored the reported locality and similar habitats in NW Yucatan State during the last 20 years and have been unable to locate this species again. However, the superficially similar E. guatemalensis is exceedingly common at these places. Because species of the E. adenocarpos clade are only known from the Pacific drainage (as the numerous collections herein reported demonstrate), and the report is based upon an unlabeled cultivated plant, we suspect a confusion from the plant owner and assume E. papillosa to be absent from the Yucatan Peninsula until reliably collected again.
Etymology: Named after Rodolfo Hernández, collector of the species and orchid photographer.
Description: Herbs, epiphytes or more commonly lithophytes, 13.8–40 cm tall without inflorescences, 58–71 cm with them, cespitose or shortly creeping. Rhizome thick, fibrous. Pseudobulbs 3–6 × 1–2.5 cm, aggregate, pyriform, more rarely subspheric, apically (1-)2-3-foliate, green and smooth when young, often reddish-tinged, enveloped by papery sheaths eventually defibrating, somewhat wrinkled upon maturity. Leaves 17–40 × 0.5–1 cm, basally conduplicate, linear oblong, acute, straight of somewhat falcate, rigidly-coriaceous, dark green. Inflorescence 58–71 cm, terminal, borne on the mature pseudobulb, erect to somewhat arcuate, racemose to (more commonly) paniculate, bearing (2–)8–11(–15) flowers, lateral branches with 2–5 flowers; peduncle dark green, laxly verruculose; peduncle bracts 5–14 × 0.5–0.6 mm, conspicuous, linear oblong, acute; rachis verruculose, green, red-tinged; ovary with pedicel 22.4–25.9 mm, cylindrical, thicker at apex, surface laxly but conspicuously verruculose; floral bract 1–4 × 4–5 mm, inconspicuous, papyraceous, acute, narrowing toward the apex. Flowers 2–3.13 cm across the spread petals, resupinate, relatively showy; petals and sepals widely spreading, rarely the petals somewhat incurved distally, pale green to yellowish at the proximal half, pale to dark brown to reddish on the distal half, with many dark brown to reddish fine veins; labellum ground color white or pale yellow to cream with pale yellowish-green margins; central lobe white, greenish or yellowish, or white proximally and a greenish or yellowish distal half, with 4–8 purple or red–purple veins on each half, radially diverging from the apex of the callus, the veins bifurcating on the distal half or simple, reaching the labellum margin or only halfway through; lateral lobes pale green or more commonly yellowish with some reddish tinges and minute spots, with several (3–5) longitudinal purple or red–purple veins; callus white with longitudinal fine dotting of red–purple or purple, the distal third white, three solid red–purple veins at the apex, entering the midlobe; column ground color white to dull yellowish with many, densely arranged fine lines of red–purple dots on both sides, anther white; sepals oblanceolate, obtuse to subacute, minutely apiculate, dorsal 16.3–17.9 cm × 3.6–3.9 mm, 9-nerved, laterals 16.2–16.5 × 3.6–3.9 mm elliptic-oblanceolate, slightly oblique, longitudinally concave; petals 16.1–16.9 × 2.2–2.4 mm, narrowly oblanceolate, subacute to obtuse, minutely apiculate, abruptly attenuated at the proximal ¼. Labellum 13.4–15.5 × 12.5–13.6 mm, fused with the column at the proximal 1/5, then free, 3-lobed, with the relatively small lateral lobes widely spreading and not enclosing the column; central lobe 6.6–7.1 × 6.6–7.0 mm, suborbicular, broadly elliptic to transversely broadly elliptic, apically truncate, shallowly emarginate, margins slightly emarginate; lateral lobes 5.9–6.1 × 2.3–2.6 mm, free, obliquely triangular to triangular elliptic, subacute to broadly obtuse; callus 0.58–0.65 × 0.43–0.46 mm, broadly elliptic, massive but flat, made up of two thick keels with a conspicuous narrow sulcate fovea in between, the keels converging proximally and distally. Column 07.1–8.8 × 3.2–4.7 mm, straight, subrhombic, basally somewhat clavate; anther white, subelliptic; pollinia 0.8–2.2 × 1.5 mm, yellow, in two subequal pairs; stigmatic surface obcordate; ovary with pedicel 22.4–25.9 mm long, cylindric, densely verruculose; capsule verruculose.
Notes: Encyclia rodolfoi is characterized by the color of its flowers, particularly the labellum. The sepals and petals are of a pale green or pale yellow green ground color, but heavily suffused with dark, dull reddish or maroon, resulting in a dull, dark maroon color. This coloration is darker on the distal half whereas the proximal half is almost devoid of color suffusion. The labellum is of a white or pale yellow to cream ground color with pale yellowish-green margins and several (4–8) purple or red–purple veins on each half, radially diverging from the apex of the callus. The lateral lobes are well-spread and yellow, with or without (less often) purple or red–purple veins which may or may not reach the lobe apex. They are also longer than in E. adenocarpos, at least 5 mm long and about as long as the central lobe or slightly shorter. Encyclia acapulcensis is probably the most similar phenetically (including the relatively short-leaved plants), but can be easily diagnosed by the color pattern and by the acute, distally concave sepals and petals. Although variable, E. rodolfoi is easily recognizable by the color pattern of the flowers.
Distribution area and habitats: Encyclia rodolfoi is common in the southern portion of the Tehuantepec Isthmus, where it grows mostly as a lithophyte, more rarely as an epiphyte. Toward the east and the south, in Chiapas, Guatemala, and El Salvador, it has always been collected, to our knowledge, as an epiphyte. Habitats are tropical dry forests (“selva baja caducifolia”). Elevations recorded are always below 600 m. A collection from Guatemala (Ackerman 999, SEL) is reported from an elevation of approximately 1500 m, but we assume this to be a mistake and that the specimen was collected elsewhere, most likely in coastal Pacific Guatemala.
Additional specimens examined: EL SALVADOR. San Salvador. Lós Cóbanos: 42. 5 km along the Carretera del Litoral toward Tamanique, on the Pacific coast, 0–5 m a. s. l., 13°31′55.78″N, 89°48′40.68″W, F. Hamer 35 (AMES, HUH, SEL). GUATEMALA. Baja Verapaz: Purulha?, flowering in cultivation at the Marie Selby Botanical Gardens accesion SEL 26-1976-52-5, 7 Jul 1977, H. Wiehler s.n. sub J.D. Ackerman 999 (SEL). MEXICO. Chiapas. Acapetahua: Cerrito, 11 m a. s. l., approx. 15°12′16.77″N, 92°44′55.14″W, 3 May 1948, “on tree, near beach”, E. Matuda 17712 (F, NY). Chiapa de Corzo: Carretera Tuxtla Gutiérrez- Aeropuerto Internacional Ángel Albino Corzo, unos 6–7 km al SE de Suchiapa; 550–560 m a. s. l., 16°36′28.52″N, 93°2′25.82″W, 6 May 2011, G. Carnevali 7626 (AMES, AMO, CICY). s.loc., 3 May 1948, E. Matuda 17712 (NY). Cintalapa: 14.7 km W of Crucero Las Cruces, 650 m a. s. l., 16°30′14.06″N, 93°57′23.45″W, collected by E. Greenwood, 1967, flowered in cultivation in Oaxaca, 22 Feb 1975, sub Pollard J-186a-55 (AMO). Tonala: Mojarra, no precise locality and date, E. Matuda 17234 (F, MEXU). Oaxaca. Asunción Ixtaltepec: Cerro La Pasión, en el extremo SE del poblado de la Mata, unos 6.4 km en línea recta del poblado de Nizanda, 85 m a. s. l., 16°37′5.05″N, 94°58′23.46″W, 22 Jul 2007, G. Carnevali et al. 7228 (AMES, CICY); same locality and date, R. Balam et al. 95 (CICY); Cerro de la Piedra, 1 km en línea recta al NE (32º) de Nizanda, 200 m, 16º39′53″N, 95º 0′26″W, selva baja caducifolia, 8 Jun 2001, E.A. Pérez-García 2094 (MEXU); same locality, 31 May 1996, E.A. Pérez-García and B. Reyes Ríos 1129 (MEXU, 3 sheets); same locality and date; same locality and date, E.A. Pérez-García and B. Pérez 2092 (MEXU); same locality and date, C. Gallardo Hernández and B. Reyes Ríos 2095 (MEXU); pedrera cercana a la vía del tren Transístmico, a 1.75 km en línea recta al NO (375º) de Nizanda, 16°40′12.50″N, 95°1′37.53″W, 7 Jul 1995, J. Meave and E.A. Pérez-García 1773 (MEXU); same locality and date, J. Meave and E.A. Pérez-García 1759 (MEXU). Ciudad de Juchitán de Zaragoza: ca. 4 km carretera La Ventosa-Santo Domingo, 150 m a. s. l., 16°33′53.68″N, 94°54′16.23″W, selva baja caducifolia, Aug 1995, E.A. Pérez-García s/n (MEXU 1189376). Nejapa de Madero: 5–10 mi. NW of Nejapa, on volcanic rocks in limestone area, ca. 1080 m a. s. l., approx. 16°42′47.80″N, 95°59′7.09″W, 28 Jan 1947, Y. Dawson s.n. (US).
IUCN conservation status: LC (Least Concern) when evaluated by its Extent of Occurrence (50, 941.2 km2) but EN (Endangered) as assessed by its Area of Occupancy of 48.00 km2 (with a cell size of 2 km). Encyclia rodolfoi meets criteria B1a and B2a of the IUCN (2010). It is currently known from a series of widely spread, isolated localities (hence the large Extent of Occurrence) embedded in matrix of suitable but highly perturbed environments. In the general Nizanda area, the species is exceedingly common growing on rocks over several low hills covered with limestone outcrops, and most collections of the species have originated there. Elsewhere, Encyclia rodolfoi is apparently rare and most of the habitats have been transformed to cattle pastures or other anthropogenic environments as it has happened in coastal Chiapas. We have visited some of the old localities documented in herbaria (e.g., Matuda’s), but tree coverage is mostly gone and there are no rocky outcrops capable of supporting populations of this or any other Encyclia species. However, because of its large range and its ability to thrive in localized, inaccessible microhabitats unsuitable for agriculture or husbandry, Encyclia rodolfoi seems to have guaranteed its long-term survival.
Taxonomic notes: The status of the populations of the Encyclia adenocarpos complex in the southern Tehuantepec Isthmus here referred to E. rodolfoi have long been a source of controversy and have been considered by authors the result of gene exchange between a member of the complex and E. parviflora. The first to propose this hypothesis were Dressler and Pollard (1974: 142) who stated “We have one plant, collected by T. McDougall at Nizanda, Oaxaca, which different persons have assigned either to E. adenocarpon (sic) or to E. belizensis ssp. parviflora …”. They applied a simple mathematical formula to calculate the sizes estimated for such a hybrid and concluded that “… it agrees well … except that the petals and sepals are narrower than we would expect.” The size concordance is not surprising since the flowers of both species are very similar in size and the shape differences most likely reflect the fact that the E. adenocarpos complex parent is apparently dominant.
Pérez-García and Hágsater (2003) first proposed a new taxon, Encyclia nizandensis, for the plants around Nizanda. Their discussion of the new taxon affinities centered around the differences with E. papillosa, which grows south and east of these populations, without explicitly stating or discussing the purported hybrid origin of their new taxon. However, the Latin diagnosis clearly states “Planta inter E. papillosam (Batem.) Aguirre-Olav. et E. parvifloram (Rege/) Withner quasi intermedia et verisimiliter ex hybridatione earum genita ….”. Thus, their intention was to propose a hybrid entity, leaving unidentified the parental taxon of E. adenocarpos affinity hypothetically involved in the hybrid.
However, they mentioned that the hybrid had stabilized and stressed the fact that in any case it does not consist of an isolated individual but of populations and that is why they treated it as a distinct species instead of as a hybrid. We have visited the type locality of E. nizandensis several times and have found that, albeit variable in size and leaf shape as well as in flower shape and color, plants from the area are rather homogeneous, which supports the stabilized introgression hypothesis.
Eventually, Archila et al. (2013) proposed Encyclia rodolfoi for what they thought was an entirely unrelated taxon without realizing that they were providing a name for the member of Encyclia adenocarpos complex that grows in the southern Tehuantepec Isthmus and is most likely involved in the origin of E. nizandensis.
Later, Leopardi et al. (2016), in a phylogeny of Northern Hemisphere Encyclia, found several genotypes of Encyclia rodolfoi (as E. nizandensis) to be nested within the other members of the E. adenocarpos complex, in analyzes involving both nuclear (ITS) and plastid (rpl32-trnL + ycf1) DNA regions. However, clonotype material of E. nizandensis behaved differently. In the plastid analyses, it fell, as expected, within the E. adenocarpos clade, along with other the genotypes of E. rodolfoi/nizandensis. However, in the nuclear analyses, this specimen was retrieved nested within the Caribbean clade, along with such species as E. alata (Batem.) Schltr. and E. parviflora. This molecular evidence suggests that the type of E. nizandensis is of hybrid origin, with the putative parentage E. rodolfoi × E. parviflora. Furthermore, it also suggests that the pod parent was a member of the E. adenocarpos clade, in this case most likely E. rodolfoi.
What follows is a discussion of Encyclia nizandensis, E. parviflora, and E. rodolfoi in the context of current cladistic, biogeographical, and morphological evidence.
Vegetative evidence: Members of the Encyclia adenocarpos clade feature concave, narrow leaves (4.0–)4.5–14 mm wide, at least 20–25 times longer than wide, whereas E. parviflora is a larger, more robust plant with flat, broader leaves (8–)20–42 mm wide. The type description of E. nizandensis reports a leaf width of 10–18 mm, which is in fact broader than the usual for the E. adenocarpos clade. Our measurements of a clonotype specimen of E. nizandensis indicate leaves 8–14.2 cm long, 11–15 mm wide, which is in agreement with the diagnosis. A sampling of many genotypes of E. rodolfoi from the southern Tehuantepec area (see section of additional specimens examined), plus many other plants analyzed in the field and under cultivation, clearly reveals a foliar width range of 5–10(–15) mm for the species. The somewhat broader leaf brings some support to the hybrid hypothesis. Plants of E. rodolfoi from east and south of this area have much narrower, relatively longer leaves, similar to those of E. adenocarpos. The relatively broad leaves homogeneously found in the plants from the Tehuantepec area suggest that genes of E. parviflora may have introgressed these populations; the molecular evidence, however, at least for genotypes other than the clonotype of E. nizandensis, appears to contradict this notion.
Biogeographical evidence: Encyclia parviflora is a member of the Caribbean clade (Leopardi et al. 2016), composed of species of which have distinct stelidia on the sides of the column and well-developed foveas in the labellar calli. No species of the clade are known to have substantial populations on the Pacific slope of Megamexico. However, E. parviflora has been collected a few times in the vicinity of the Nizanda area (Cerro Mazagua; 2 km SW of Cienaguita, A. Saynes, A. Sánchez and S. Salas 3203, SERO; same locality, collected by E. Pérez-García s.n., sub Carnevali 8088, CICY) and the possibility of occasional gene exchange between E. parviflora and E. rodolfoi exists. It is noteworthy that E. rodolfoi has already proven to be capable of generating hybrids with other non-related members of Encyclia, as in the above mentioned E. × nizanbury.
Thus, all the available evidence strongly suggests that the type plant of Encyclia nizandenzis is actually a hybrid between E. rodolfoi and E. parviflora. Hence, the name E. nizandensis should in all property only be applied to this particular plant and to any other that features the morphological characteristics of this particular plant. We have visited the Nizanda area several times analyzing the variation of the Encyclia adenocarpos-type plants found in the area and have found them to be extremely variable in floral color and morphological details (see Fig. 9). Plants are also vegetatively variable at the site, particularly leaf relative length, with some plants somewhat resembling the type plant of Encyclia nizandenzis. This variation pattern may suggest the acquisition of E. parviflora genes in the history of these populations. However, we have never found any plants quite like the type. We (Leopardi et al. 2016) have obtained DNA sequences of five genotypes of these plants, including the type of Encyclia × E. nizandensis, that span the morphological variation of the entity. All genotypes other than the type of E. × nizandensis are retrieved from the analyses (both nuclear and plastid) nested within the E. adenocarpos clade, regardless of their morphological characteristics. Thus, albeit morphologically variable suggesting introgression, the critical analysis of all evidence (both molecular and morphological) does not support the hypothesis of a hybrid origin for the Encyclia adenocarpos-type plants found in the Nizanda area and the name Encyclia rodolfoi should be applied to them, as well as for the populations of similar morphology occurring in Chiapas, Guatemala, and El Salvador, whereas the name E. nizandensis should only, then, be applied to the type plant of that name.
Etymology: The specific name refers to Nizanda, Oaxaca, which means “hotspring water” in Zapotec language.
Discussion: As elaborated above, the name is here restricted to plants of Encyclia rodolfoi morphology but with broader leaves (7.5–9.5 times longer than wide in E. × nizandensis vs at least 15 times longer than wide in E. rodolfoi), well-developed stelidia at the column apex, and with a shallow yet well-developed ellipsoid fovea in the labellar calli. So far it is only known for sure from a single genotype. This plant has already been in cultivation for several years and can be found in several orchid collections.
Iconography: An image at the IOSPE site (http://www.orchidspecies.com/encnizandensis.htm) is of the true E. × nizandensis (the type plant). On the other hand, the plate in Hágsater et al. (2015), p. 124., Figure 247 (as E. nizandensis) is Encyclia rodolfoi as circumscribed here.
Etymology: From the Greek trachus = “asperous” and karpos = “fruit”, referring to the rugulose surface of the ovary with pedicel and the capsule.
Description: Epiphytic herbs, densely caespitose and eventually forming large clumps, 19–40(–60) cm without inflorescences, up to 90 cm with them. Rhizome thick, fibrose. Pseudobulbs 3.4–6.7(–8.0) × 1.3–2.3(–3.5) cm, aggregate, pyriform, apically (1-)2-3-leaved, first green and smooth, wrinkled and often red-tinged upon aging; clothed by 2-3 marcescent sheaths that are first membranous, becoming papyraceous and defibrating on age. Leaves 20.0–37.0 × 1.0–1.8(–2.0) cm, proximally conduplicate, v-shaped distally, rigidly fleshy coriaceous, linear oblong, often somewhat falcate, acute, dark green. Inflorescences 19–62(–80) cm, terminal and emerging from the mature pseudobulb, suberect to somewhat arcuate, (3–)5–15(–20)-flowered, racemose or more commonly paniculate with 1-2(-3) lateral branches, these 2–4(–6)-flowered, 3–4.5 cm long, branches appearing midway or above along inflorescence length; peduncle thin and flexible, but strong, surface verrucose, more so distally; peduncle bracts 5–17 mm long, becoming shorter distally, tubular, oblong-ovate when forcibly spread, acute; rachis verruculose, green or green red-tinged; ovary with pedicel 18–22 mm long, verruculose; floral bracts 1–4 × 4–5 mm, inconspicuous, thinly fleshy first, then papyraceous, ovate, acute, decreasing in dimensions distally. Flowers 23.1–41 mm across the spread apices of the petals, resupinate, showy; petals and sepals spreading proximally, slightly porrect in the distal half, thus flowers in some clones appearing somewhat campanulate, green or yellow green, darker proximally, the veins overlayed with dark dull maroon or brown from base to apex; labellum clear white, the margins with a faint to ± intense band of yellow, the central lobe with 10–12 longitudinal purple veins radiating from the callus, the veins simple of branching on the distal half, the central most 4-6 darker and thicker than the others; lateral lobes pale dull maroon with a pale yellow margin (colored as for sepals); callus white with 4-5 relatively thick purple veins running longitudinally, darker and thicker proximally, becoming discontinuous and fainter distally; column bright white, slightly tinged red along middle of the proximal half; anther pale to bright yellow; sepals (15–)16–20(–22) × 4–7 mm, narrowly elliptic, acute to broadly acute, somewhat attenuate and concave proximally, the laterals slightly oblique, and longitudinally concave, petals 17–21 × 5–6 mm, narrowly elliptic to linear spathulate, acute to broadly acute, proximally more attenuate than the sepals; labellum 14–19 mm long, fused to the column at the basal 1/5, then free, sharply 3-lobed; central lobe 8–10(–11) × 9–10(–11) mm, suborbicular, subquadrate to ovate subquadrate, rounded to broadly truncate, flat to slightly ondulate at the margins; lateral lobes 3–4 × 1–1.2 mm, the proximal margin ca. 2 mm long, the distal margin ca. 4.5 mm long, flat and widely spreading (thus not enclosing the column), triangular, obliquely acute; callus 6.5–8.5 × 3.5–4.8 mm, subelliptic, apex broadly obtuse to subacute, composed of two flat, broad keels bisected by a narrowly linear elliptic fovea, deeper and broader in the proximal half, the two basal endings slightly bilobed, the apex of the callus is thick and grades into the several veins that enter the central lobe. Column 8.5–11 × 3.6–5.4 mm, fused with the ventral face of the column by 5 mm, rhombic-obovate, proximally somewhat attenuate, without stelidia; anther 1.5 × 1.8 mm, cordate in distal view; pollinia 1.0 × 0.7–1.2 mm, yellow, in two subequal pairs; stigmatic surface obcordate, 3.1 × 3.7 mm; pedicellate ovary 23.2–25 mm long, cylindrical, densely verruculose.
Distribution area and habitats: Encyclia trachycarpa has a fairly limited distribution in Jalisco and Nayarit, at elevations from sea level up to 400 m but much more commonly below 200 m. As is the rule for species in the E. adenocarpos clade, it grows on tropical dry forests. Its distributional range is totally embedded within that of E. adenocarpos. Furthermore, both species occupy the same general habitats and flower simultaneously, suggesting they are most likely using a different set of pollinators. Wherever sympatric, E. trachycarpa can easily be distinguished by its larger, showier flowers with a much broader central lobe to the labellum (9.5–11 mm wide vs 5.6–7.9 mm) and relatively much smaller lateral lobes. The flowers of E. trachycapa are usually displayed in a more or less nutant fashion. Also, the colored veins in E. trachycarpa are more abundant, thicker, and more colorful. In general, mature plants of E. trachycarpa present larger, more massive pseudobulbs than those of E. adenocarpos, but the ranges overlap and a large plant of the later may reach the dimensions of a small plant of the former. As opposed to E. adenocarpos, which is often found growing on rocks, E. trachycarpa has only been collected as an epiphyte to our knowledge.
Additional specimens examined: MEXICO. Jalisco. “Costa Tropical de Jalisco” cultivated at the Universidad de Guadalajara, 20 Jun 1999, L.M.V. de Puga s.n. (IBUG). Ciahuatlán: ca. 10 km from Barra de Navidad on road to Bahía de Tecanatita, 170–175 m a. s. l., approx. 19°15′54.93″N, 104°43′36.26″W, 5 Jul 1961, R.L. Dressler and M. Wirth 2695 (MO, US); 2 km NW de Cihuatlán, 400 m a. s. l., approx. 19°14′43.72″N, 104°36′1.78″W, 16 Jun 1982, Thurston and E. Hágsater T-2213 (AMO); Río Careyitos 1-2 km. al NW de la entrada a Careyitos, carretera Ciahuatlán-Puerto Vallarta, unos 11–11.5 km en línea recta al SE de Chamela, 14–20 m a. s. l., 19°263′46″N, 105°013′37.7″W, selva baja caducifolia con epífitas como Tillandsia spp. y Encyclia ssp., 19 May 2008, G. Carnevali 7340 (CICY); same locality, 30 Jul 2016, G. Carnevali and I. Ramírez 7963 (CICY). La Huerta: Estación Biológica de Chamela, 150 m a. s. l., 19°31′43.64″N, 105°4′19.80″W, 9 Mar 1973, M. Sousa et al. 3921 (US); same locality, 30 May 1975, C.L. Diaz L. 5857 (GUADA); 1 km al N. de Careyes, Arroyo La Lima, km 55 de la carretera Barra de Navidad-Pto. Vallarta, 19°27′45.21″N, 105°1′58.90″W, selva baja caducifolia, 7 Jun 1984, J.A. Magallanes 4193 (MEXU); La Manzanilla, (10 m a. s. l., approx. 19°17′10.17″N, 104°47′15.00″W), 28 May 2000, I. Contreras V. s.n. (IBUG). Puerto Vallarta: Puerto Vallarta, 50–100 m a. s. l., approx. 20°37′29.98″N, 105°12′25.68″W, 19 Jun 1984, S. Rosillo de Velazco s.n (AMO); 3 km from El Colorado to Las Palmas, 20°47′47.52″N, 105°8′51.17″W, 25 May 2017, A. Zabalgoitia s.n. sub G. Carnevali 8032 (CICY). Tomatlán: ca. Tomatlán, 50 m a. s. l., approx. 19°55′46.69″N, 105°14′44.33″W, S. Rosillo de Velazco s.n (AMO). Nayarit. San Pedro Ixcatán, 8 km N San Pedro Ixcatán, ca. Los Tepetates y la cañada Los Bueyes, 24°08′89″N, 50°61′69″W (sic) (130–135 m a. s. l., approx. 22°6′34.44″N, 104°56′48.93″W,), 14 Jan 2008, A. Castro-Castro and A. Frías 1158 (IBUG).
IUCN conservation status: VU (Vulnerable) Encyclia trachycarpa meets criteria B1a and B2a of the IUCN (2010). Its Extent of Occurrence is of 11.211,6 km2, with an Area of Occupancy of 44 km2, a value which would rank it as EN (endangered). Although we know of only 15 herbarium collections of the species, it is certainly somewhat more common in the area where we have seen it on trees in forest remnants. The coastal zone of Jalisco and Nayarit, where E. trachycarpa is restricted, has been heavily perturbated, and much of the tropical dry forests of the area, the preferred habitat of the species, have been reduced to cow pastures for extensive cattle ranching. Populations of the species, however, occur within the boundaries of the Estación Biológica Chamela, a scientific research station of the UNAM (Universidad Nacional Autónoma de México) where some degree of protection is afforded to the continual survival of the species.
Taxonomic notes: Epidendrum trachycarpum was first collected by George W. Barclay during the trip of the H.M.S. Sulphur on 20 May, 1837 at Manzanillo Bay. The plants resulting from the expedition were studied by G. Bentham (1846) who published them in The Botany of the Voyage of H.M.S. Sulphur, which appeared between 1844–1846. Orchids from this expedition were studied by John Lindley, who described this species as related to Epidendrum adenocarpos but with fleshy flowers (instead as membranous) and extremely rigid leaves. The leaves are in reality very similar in texture to those of most members of this clade, but the fleshy character of the flowers is fairly conspicuous, at least in the live state and they are longer lasting. Although McVaugh (1985) strongly argues that the accounts of the H.M.S. Sulphur’s trip indicates that the “Manzanillo” mentioned in the protologue is the port in what is now Colima, the species has never been collected there nor in any other place in the state of Colima, even after 30 additional years of botanical collecting in the area. Dressler (1964) believed that the type specimen could have been collected near La Manzanilla (at Tenacatita Bay), in Jalisco, ca. 100 km to the west of the Coliman port, a view that we support. However, the right kind of habitats may exist in the vicinity of Manzanillo, Colima. Thus, it is not impossible that Encyclia trachycarpa grew there at the time of the H.M.S. Sulphur’s trip and that it is now gone due to habitat transformation under anthropogenic activities.
Iconography: The species is sufficiently distinct and has thus not been often confused with related species. However, some authors have mistaken populations here referred to E. rodolfoi from Honduras and El Salvador with this species. For example, the photograph in Orchids of Nicaragua (van den Berghe and van den Berghe (2008): “Photo and maps,” plate A49) is correctly identified as Encyclia trachycarpa, but the image must correspond to a Mexican plant because that species does not occur in Nicaragua.
Enrique Arcila Arcila, Eduardo Pérez-García, Denis Szeszko, Claudia Ramírez, Pablo Carrillo, and Alejandro Zavalgoitia provided plant material and photographs that were crucial to the development of this project. Rodrigo Duno de Stefano and Iván Tamayo Cen (CICY), Eduardo Pérez-García (MEXU), and Ramón López (Barquisimeto, Venezuela) contributed discussion and reviews of earlier drafts of the manuscript. Gregorio Amílcar Castillo (CICY) and Juan Pablo Pinzón Esquível (UADY), collaborated in the field work associated with the developement of this project. Jesús García Robles and Cristina Taddei Bringas, from CIAD, Hermosillo, provided logistical support in a collecting trip to the vicinity of Álamos, Sonora in September 2015, for which we are extremely grateful. We are indebted to Felipe Escudero Ganem and Sergio Reynaud for sharing habitat data of E. acapulcensis. Kanchi Gandhi at HUH helped us with nomenclatural matters. Eric Hágsater and colleagues at AMO (Rolando Jiménez Machorro and Luis Sánchez-Saldaña, r.i.p.), helped us with photographs, illustrations, discussion of the species complex, and loaning of critical material. Katya Romero, Edgar Mó, Hermes Vega, and Illiam Rivera, among others contributed photographs of plants in habit that were fundamental to arrive at the species concepts employed here. We thank the curators of the many herbaria that allowed us to study their plant material. This paper would not have been possible without their contribution. This research was partially funded by CONACyT project CB-2011-168640 “Sistemática y Filogenia de Encyclia Hook. (Orchidaceae: Laeliinae), con énfasis en Megaméxico” awarded to the senior author. CL and ACS thank for scholarships granted by CONACyT. The authors acknowledge the American Orchid Society for its support to our Encyclia research through the funding of project “Systematics and evolution of Encyclia Hook. s.s. (Orchidaceae: Laeliinae) with emphasis in Megamexico” awarded to GC. Finally, we would like to acknowledge two anonymous reviewers that helped make this a much better article with their suggestions and comments: we are very grateful for their input.
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