Haseltonia 14: 161–169. 2008 161 and disTribuTion of epiphyTic cacTi in uruguay—noTes Towards a checklisT of cacTaceae of uruguay, parT 3

URS EggLI Sukkulenten-Sammlung Zürich, Mythenquai 88, CH-8002 Zürich, Switzerland; [email protected]

EdUARdO MARCHESI, MAURICIO BONIFACINO Laboratorio de Botánica, Herbario MVFA, Facultad de Agronomía, Casilla de Correos 1238, Montevideo, Uruguay; [email protected]

RETO NyFFELER Institut für systematische Botanik der Universität Zürich, Zollikerstrasse 107, CH-8008 Zürich, Switzerland; [email protected]

Abstract: The most recent treatments of the family Cactaceae for Uruguay are that of Arechavaleta (1905), Osten (1941), and Herter (1953–55). While the globular members of the family (Parodia [including Notocactus], Frailea, and Gymnocalycium) are relatively well known through numerous later individual publications, the epiphytic of and are comparatively poorly-known. Based on herbarium material and recent col- lections, we present data on the geographical distribution and taxonomy for two species of Lepismium and two species of Rhipsalis. and Rhipsalis floccosa are new records for the flora of Uruguay. The occurrence ofEpiphyllum phyllanthus in Uruguay could not be confirmed. Resumen: Los tratamientos mas recientes de la familia Cactaceae para Uruguay son las publicaciones de Arechavaleta (1905), Osten (1941), y Herter (1953–55). En comparación con las especies globulares de la familia (géneros Parodia [incl. Notocactus], Frailea y Gymno- calycium), que son bastante bien conocidas a través de numerosas publicaciones posteriores, las especies epífitas (génerosLepismium y Rhipsalis) carecen de datos detallados. Basado en es- pecimenes de herbario y colectas recientes, presentamos datos sobre distribución y taxonomía de dos especies de Lepismium y dos especies de Rhipsalis. Lepismium cruciforme y Rhipsalis floccosa presentan nuevos registros para Uruguay. La ocurrencia de Epiphyllum phyllanthus en Uruguay no fue confirmada. Key words: conservation, Cactaceae, distribution, Lepismium, Rhipsalis, taxonomy, Uruguay

1. Introduction not a single entry for epiphytic taxa (R Mar- Epiphytic cacti: Species of cacti are an impor- tin, pers. comm.). Epiphytic cacti are tradi- tant element in arid and semi-arid vegetations tionally classified into two distinct taxonomic in North and South America. The vast major- groups (Barthlott and Hunt 1993; Anderson ity of taxa are terrestrial, and most of 2001). The tribe Hylocereeae comprises six these groups are reasonably well known through genera and 64 species (Bauer 2003), which many systematic and/or regional studies. This occur in tropical forests of central and north- is especially true for the small-growing taxa ern South America. It is not considered here with high “collectors’ appeal.” because no representatives are native to Uru- Epiphytically growing cacti are, however, guay. The tribe consists of four comparatively less well known. From 1356 col- genera and 61 species (Barthlott and Taylor lection records for Uruguayan cacti, the vast 1995). The center of species diversity is east- majority (over 1300) refer to the genera Paro- ern , in particular the area of the Mata dia, Gymnocalycium, and Frailea, and there is Atlântica. Only a few species occur in cen- 162 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 163 tral and southern North America. Moreover, cated in a transition area between hot and and Osten (1941) merely cover a selection of relation to the total cactus diversity—only two (J. S. Mueller) Stearn is humid regions to the north and cold and dry taxa. The checklist of cacti in the Uruguayan out of the previously recorded 51 taxa are ep- the only species which is also naturally oc- areas to the south. Annual precipitation var- flora by Herter (1953–55) is mostly based on iphytes! This over-representation of epiphytic curring in Africa, Madagascar, and islands in ies from 1600 mm in the north to 1100 mm a literature survey, with the addition of his cacti could be due to the fact that these com- the Indian Ocean. The Rhipsalis, in the in the south. The Uruguayan vegetation is own (mostly undocumented) observations. paratively less-succulent are easier to dry current circumscription, comprises species largely made up of grasslands (“pampa”). Nat- Hunt (1999) lists 51 taxa (42 accepted and 9 than other cacti. In addition, the holdings of B, with acrotonic branching, while those with ural forests are mostly restricted to river banks provisionally accepted) for Uruguay, and An- US, Z and ZSS were checked for Uruguayan predominantly basitonic branching are cur- and steep ravines in the extensive hill system derson (2005) lists 53 taxa (including infra- material, but none were found. rently included in an expanded genus Lepis- that covers most of the country. Cactaceae specific taxa). Taxonomy and distribution: Herbarium mium (Barthlott and Taylor 1995). However, are found on rocky outcrops mostly associ- Genera of epiphytic cacti reported from specimens were geo-coded (± 1–5 geograph- recent molecular phylogenetic analyses indi- ated with hill tops, as well as on shallow stony Uruguay are Lepismium and Rhipsalis, with ical minutes) whenever the locality data was cate that the genus Lepismium in its broad soils overlying bedrock, but also occur in other one species each, namely Lepismium lumbri- sufficiently detailed to allow localization of the circumscription is polyphyletic and that the xerophytic habitats, such as Prosopis forests in coides (Lemaire) Barthlott and Rhipsalis ce- collection on modern maps. Distribution maps groups presently treated as subgenera Pfei- the west and coastal scrub along the Río de la reuscula Haworth (Arechavaleta 1905; Herter were then generated using ESRI ArcMap (ver- ffera, Acanthorhipsalis, and Lymanbensonia Plata and Atlantic Ocean shorelines. 1953–1955; Barthlott and Taylor 1995, 1999; sion 9.1) software and topographic and other should be classified into a distinct genusPfei - Much of the vegetation is influenced to Anderson 2005). data from the Digital Basemap of the Ameri- ffera (Nyffeler 2000, 2002, unpublished data; a considerable degree by increasingly inten- The occurrence of Rhipsalis linearis cas project (see Bletter and others 2004). The Taylor and Zappi 2004). sive agriculture and sylviculture, as well as by K. Schumann (a synonym of Lepismium diagnostic taxon descriptions were compiled Vegetation of Uruguay: Uruguay is lo- grazing. Severe overgrazing with subsequent warmingianum (K. Schumann) Barthlott from herbarium specimens and published de- colonization by neophytes and native Eryn- according to Barthlott and Taylor 1995) and scriptions (Taylor and Zappi 2004; Anderson gium spp (Apiaceae) has a major impact on R. pentaptera A. Dietrich as communicated 2005), and—for L. lumbricoides—our own col- the naturally occurring biodiversity. The dis- by Britton and Rose (1923) was not con- lections. Variation patterns could only be ex- appearance of natural forests is of major con- firmed in later literature and must be consid- plored to a limited extent due to the relative cern for the continued survival of epiphytes, ered erroneous. paucity of the material available. Generic clas- including epiphytic cacti. Hunt (1999) also lists Epiphyllum phyllan- sification and species synonymy follow Barth- Cacti of Uruguay: In contrast to coun- thus (Linné) Haworth for Uruguay, but no lott and Taylor (1995). tries such as , Brazil, and Chile, the specimens to support this occurrence have been cactus diversity of Uruguay is comparatively located, nor is the species reported by other 3. Results: Descriptions, smaller and less well known. No detailed authors. The species has a wide range, but in classification, and identification study of the family at the national level has the most recent synopsis of the group (Bauer 3.1. Key to genera and species ever been prepared, and Arechavaleta (1905) 2003) Uruguay is not listed in the distribu- tion range. The nearest regions from which 1 Branching basitonic to mesotonic; stems this taxon is known are and north- terete or 3- to 6-angled (or rarely flat)..... Figure 1. Lepismium cruciforme. a. Distribution in ern Argentina. An occurrence in Uruguay is ...... Lepismium,2 b. c, d, e. Uruguay. Tip of young growing stem; Flow- unlikely, but possible. 1 Branching predominantly or exclusively ers; f. Flower bud; g. Fruit. Photographs: M Bonifaci- no, of cultivated material in hort. Marchesi. acrotonic; stems terete ...... Rhipsalis, 3 a 2. Material and methods 2 Stems distinctly angled or rarely flat; are- b c d e Fieldwork: Fieldwork was carried out in Uru- oles 1 cm or more apart...... guay by UE and RN in October/November of ...... Lepismium cruciforme 2004 and 2005. A total of 138 localities scat- 2 Stems terete; areoles < 1 cm apart ...... tered over a large part of Uruguay were visited ...... Lepismiumlumbricoides in order to sample cactus diversity in the form 3 Stems heteromorphic (extension shoots of herbarium vouchers and photographic doc- and second-order and third-order stems umentation. Sampling of cactus diversity was of different lengths), terete...... carried out in a general way, and no special em- ...... Rhipsalis cereuscula f g phasis was placed on epiphytic taxa. Herbarium 3 Stems all similar, terete to slightly angled material (143 specimens) was prepared using ...... Rhipsalis floccosa the method of Eggli and Leuenberger (1996). Sets were deposited at MVJB and ZSS. All col- 3.2. Lepismium lections are geo-coded by using a GPS receiver in the field. 3.2.1. Lepismium cruciforme (Fig 1) Herbarium work: We studied all accessi- Epiphytic or rarely lithophytic; stems creep- ble herbarium material of cacti in the herbaria ing, with few to numerous adventitious roots, of Montevideo (MVFA, MVJB, MVM). Out or pendent, forming chains to 3 m long, seg- of a total of 76 usable (in the sense of having mented, irregularly branched basitonically to sufficient data or being sufficiently complete) mesotonically, more rarely also acrotonically, all specimens, 40 belong to the study group. Epi- similar, medium green, margins often flushed phytic cacti are thus vastly over-represented in dark red, fully exposed plants completely red- 162 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 163 tral and southern North America. Moreover, cated in a transition area between hot and and Osten (1941) merely cover a selection of relation to the total cactus diversity—only two Rhipsalis baccifera (J. S. Mueller) Stearn is humid regions to the north and cold and dry taxa. The checklist of cacti in the Uruguayan out of the previously recorded 51 taxa are ep- the only species which is also naturally oc- areas to the south. Annual precipitation var- flora by Herter (1953–55) is mostly based on iphytes! This over-representation of epiphytic curring in Africa, Madagascar, and islands in ies from 1600 mm in the north to 1100 mm a literature survey, with the addition of his cacti could be due to the fact that these com- the Indian Ocean. The genus Rhipsalis, in the in the south. The Uruguayan vegetation is own (mostly undocumented) observations. paratively less-succulent plants are easier to dry current circumscription, comprises species largely made up of grasslands (“pampa”). Nat- Hunt (1999) lists 51 taxa (42 accepted and 9 than other cacti. In addition, the holdings of B, with acrotonic branching, while those with ural forests are mostly restricted to river banks provisionally accepted) for Uruguay, and An- US, Z and ZSS were checked for Uruguayan predominantly basitonic branching are cur- and steep ravines in the extensive hill system derson (2005) lists 53 taxa (including infra- material, but none were found. rently included in an expanded genus Lepis- that covers most of the country. Cactaceae specific taxa). Taxonomy and distribution: Herbarium mium (Barthlott and Taylor 1995). However, are found on rocky outcrops mostly associ- Genera of epiphytic cacti reported from specimens were geo-coded (± 1–5 geograph- recent molecular phylogenetic analyses indi- ated with hill tops, as well as on shallow stony Uruguay are Lepismium and Rhipsalis, with ical minutes) whenever the locality data was cate that the genus Lepismium in its broad soils overlying bedrock, but also occur in other one species each, namely Lepismium lumbri- sufficiently detailed to allow localization of the circumscription is polyphyletic and that the xerophytic habitats, such as Prosopis forests in coides (Lemaire) Barthlott and Rhipsalis ce- collection on modern maps. Distribution maps groups presently treated as subgenera Pfei- the west and coastal scrub along the Río de la reuscula Haworth (Arechavaleta 1905; Herter were then generated using ESRI ArcMap (ver- ffera, Acanthorhipsalis, and Lymanbensonia Plata and Atlantic Ocean shorelines. 1953–1955; Barthlott and Taylor 1995, 1999; sion 9.1) software and topographic and other should be classified into a distinct genusPfei - Much of the vegetation is influenced to Anderson 2005). data from the Digital Basemap of the Ameri- ffera (Nyffeler 2000, 2002, unpublished data; a considerable degree by increasingly inten- The occurrence of Rhipsalis linearis cas project (see Bletter and others 2004). The Taylor and Zappi 2004). sive agriculture and sylviculture, as well as by K. Schumann (a synonym of Lepismium diagnostic taxon descriptions were compiled Vegetation of Uruguay: Uruguay is lo- grazing. Severe overgrazing with subsequent warmingianum (K. Schumann) Barthlott from herbarium specimens and published de- colonization by neophytes and native Eryn- according to Barthlott and Taylor 1995) and scriptions (Taylor and Zappi 2004; Anderson gium spp (Apiaceae) has a major impact on R. pentaptera A. Dietrich as communicated 2005), and—for L. lumbricoides—our own col- the naturally occurring biodiversity. The dis- by Britton and Rose (1923) was not con- lections. Variation patterns could only be ex- appearance of natural forests is of major con- firmed in later literature and must be consid- plored to a limited extent due to the relative cern for the continued survival of epiphytes, ered erroneous. paucity of the material available. Generic clas- including epiphytic cacti. Hunt (1999) also lists Epiphyllum phyllan- sification and species synonymy follow Barth- Cacti of Uruguay: In contrast to coun- thus (Linné) Haworth for Uruguay, but no lott and Taylor (1995). tries such as Argentina, Brazil, and Chile, the specimens to support this occurrence have been cactus diversity of Uruguay is comparatively located, nor is the species reported by other 3. Results: Descriptions, smaller and less well known. No detailed authors. The species has a wide range, but in classification, and identification study of the family at the national level has the most recent synopsis of the group (Bauer 3.1. Key to genera and species ever been prepared, and Arechavaleta (1905) 2003) Uruguay is not listed in the distribu- tion range. The nearest regions from which 1 Branching basitonic to mesotonic; stems this taxon is known are Paraguay and north- terete or 3- to 6-angled (or rarely flat)..... Figure 1. Lepismium cruciforme. a. Distribution in ern Argentina. An occurrence in Uruguay is ...... Lepismium,2 b. c, d, e. Uruguay. Tip of young growing stem; Flow- unlikely, but possible. 1 Branching predominantly or exclusively ers; f. Flower bud; g. Fruit. Photographs: M Bonifaci- no, of cultivated material in hort. Marchesi. acrotonic; stems terete ...... Rhipsalis, 3 a 2. Material and methods 2 Stems distinctly angled or rarely flat; are- b c d e Fieldwork: Fieldwork was carried out in Uru- oles 1 cm or more apart...... guay by UE and RN in October/November of ...... Lepismium cruciforme 2004 and 2005. A total of 138 localities scat- 2 Stems terete; areoles < 1 cm apart ...... tered over a large part of Uruguay were visited ...... Lepismiumlumbricoides in order to sample cactus diversity in the form 3 Stems heteromorphic (extension shoots of herbarium vouchers and photographic doc- and second-order and third-order stems umentation. Sampling of cactus diversity was of different lengths), terete...... carried out in a general way, and no special em- ...... Rhipsalis cereuscula f g phasis was placed on epiphytic taxa. Herbarium 3 Stems all similar, terete to slightly angled material (143 specimens) was prepared using ...... Rhipsalis floccosa the method of Eggli and Leuenberger (1996). Sets were deposited at MVJB and ZSS. All col- 3.2. Lepismium lections are geo-coded by using a GPS receiver in the field. 3.2.1. Lepismium cruciforme (Fig 1) Herbarium work: We studied all accessi- Epiphytic or rarely lithophytic; stems creep- ble herbarium material of cacti in the herbaria ing, with few to numerous adventitious roots, of Montevideo (MVFA, MVJB, MVM). Out or pendent, forming chains to 3 m long, seg- of a total of 76 usable (in the sense of having mented, irregularly branched basitonically to sufficient data or being sufficiently complete) mesotonically, more rarely also acrotonically, all specimens, 40 belong to the study group. Epi- similar, medium green, margins often flushed phytic cacti are thus vastly over-represented in dark red, fully exposed plants completely red- 164 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 165 brown to brown-green; stem segments 15–30 bristles 5–10 mm long, straight, whitish to scar left by the deciduous perianth conspic- (–40) cm long and 0.6–3 cm wide, frequently brownish. Flowers usually scattered along the uous, 1.5–2 mm wide, fruit wall 2.5 mm 3-ribbed, occasionally flat or with up to six length of the stems, solitary (but the same thick, pulp strongly mucilaginous; seeds nu- angles or ribs, rib margins very shallowly to areole can form another flower later), diur- merous (ca. 30–35), brown. Flowering time: distinctly and often irregularly crenate to un- nal, opening for a single day only, unscented, December to March; fruiting time: February dulate-crenate; areoles small, 1.2–2 cm apart, broadly funnel-shaped at anthesis, 0.9–1.5 cm to September. situated in the crenations along the ribs, be- diam. and 0.7–1 cm long; pericarpel medium Comments: This is the first report ofLepis - fore flowering subtended with a conspicuous, green to pinkish green, rounded to obconi- mium cruciforme in Uruguay, though its pres- triangular, scale-like leaf rudiment ca. 1 mm cal, 3–5 (–7) mm long and 2–5 mm diam., ence could have been expected. It is a very long, later naked, enlarged with age and with without scales; perianth elements 8–9, nar- widespread and variable taxon. Although the a conspicuous tuft of wool and weak bristles, rowly obovate to oblong, 5–7 mm long and synonymy (see Anderson 2005) is extensive to 2 mm wide, delicate, white to pale cream, and the variability pronounced, no infraspe- white flushed with pink, or pale pink, tip cific classification has been proposed so far. The obtuse, recurved; stamens ca. 60, filaments name has a checkered nomenclatural history 3–7 mm long, white, lower filaments placed and is based on the conserved name Cactus against the style; anthers/pollen very pale yel- cruciformis Vellozo 1829 (Brummitt 1996). a 1 low to white; style ca. 5 mm long, lower ⁄3 In earlier literature, this species is often named white, otherwise pinkish; stigma lobes 3–5, Lepismium myosurus (Salm-Dyck ex De Can- spreading, ca. 1 mm long, white to pinkish. dolle) Pfeiffer (based on Cereus myosurus Fruits depressed globose to globose to glo- Salm-Dyck ex De Candolle 1828). bosely pear-shaped, rarely elongate-globose, Identification: Though the plants are vari- first dull green, magenta to dark red at matu- able, they are easy to identify amongst Uru- rity, to 7 mm diam. and 11 mm long, glossy, guayan epiphytic cacti due to the flat or ribbed/ ridged, massive stem segments. Figure 2. Lepismium lumbricoides. a. Distribu- General Distribution: NE to SW Brazil tion in Uruguay. b, c. Lepismium lumbricoides flow- (Pernambuco and southwards), Uruguay, SE ers; d. Flowering stem; e. Tip of young growing stem; Paraguay, NE Argentina (Formosa, Chaco, b f. Habit. Photographs: M Bonifacino, of material col- Corrientes, Misiones). a lected at Buceo, Montevideo. Distribution in Uruguay: Artigas, Cerro Largo. b c d Ecology: There is no data on the specimens available, but at least in Cerro Largo, plants have been observed (EM and MB) as epiphytes on Erythrina crista-galli (Fabaceae). 3.2.2. Lepismium lumbricoides c d (Lemaire) BarthLott (Fig 2) Epiphytic or rarely lithophytic; stems creep- ing on tree branches and then with numer- ous adventitious roots, or irregularly parallel- pendent, irregularly branched basitonically to mesotonically, all similar, dull green to me- dium green, 10–40 cm long, 4–6 mm diam., e f e f terete or slightly longitudinally ridged in the Figure 3. Rhipsalis cereuscula. a. Distribution in dry season or when old with 6–8 indistinct Uruguay. b. Habit; c, d. Flowers; e. Detail of branch- ribs; areoles minute, regularly scattered along ing stems; f. Fruit. Photographs: M Bonifacino, of cul- the length of the stems, 8–14 mm apart, on tivated material in hort. Marchesi. young stems with a minute pointed triangu- lar subtending scale-like leaf-rudiment 0.5– 1 mm wide, later naked and almost invisible, opening for ca. three days; buds ascending at or with a minute tuft of dirty white short 45° angle, greenish; open flowers spreading wool, or more rarely with weak bristly spines; to pendent, campanulate to broadly funnel- spines present on some or all areoles, 1–8 per shaped, 1–1.5 cm diam. and long; pericarpel areole, 2–5 mm long, softly bristly, straight dull dark green, globose or bluntly angled, 2– or irregularly curved, brown to almost black. 3 mm long and ca. 2 mm diam., sometimes Flowers usually scattered along the length with 1–2 small scales; perianth elements (8– of the stems, sometimes more aggregated ) 12–18, oblong to narrowly obovate, 11– towards the stem tips, diurnal, unscented, 13 mm long, 3–4 mm wide, obtuse, delicate, 164 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 165 brown to brown-green; stem segments 15–30 bristles 5–10 mm long, straight, whitish to scar left by the deciduous perianth conspic- (–40) cm long and 0.6–3 cm wide, frequently brownish. Flowers usually scattered along the uous, 1.5–2 mm wide, fruit wall 2.5 mm 3-ribbed, occasionally flat or with up to six length of the stems, solitary (but the same thick, pulp strongly mucilaginous; seeds nu- angles or ribs, rib margins very shallowly to areole can form another flower later), diur- merous (ca. 30–35), brown. Flowering time: distinctly and often irregularly crenate to un- nal, opening for a single day only, unscented, December to March; fruiting time: February dulate-crenate; areoles small, 1.2–2 cm apart, broadly funnel-shaped at anthesis, 0.9–1.5 cm to September. situated in the crenations along the ribs, be- diam. and 0.7–1 cm long; pericarpel medium Comments: This is the first report ofLepis - fore flowering subtended with a conspicuous, green to pinkish green, rounded to obconi- mium cruciforme in Uruguay, though its pres- triangular, scale-like leaf rudiment ca. 1 mm cal, 3–5 (–7) mm long and 2–5 mm diam., ence could have been expected. It is a very long, later naked, enlarged with age and with without scales; perianth elements 8–9, nar- widespread and variable taxon. Although the a conspicuous tuft of wool and weak bristles, rowly obovate to oblong, 5–7 mm long and synonymy (see Anderson 2005) is extensive to 2 mm wide, delicate, white to pale cream, and the variability pronounced, no infraspe- white flushed with pink, or pale pink, tip cific classification has been proposed so far. The obtuse, recurved; stamens ca. 60, filaments name has a checkered nomenclatural history 3–7 mm long, white, lower filaments placed and is based on the conserved name Cactus against the style; anthers/pollen very pale yel- cruciformis Vellozo 1829 (Brummitt 1996). a 1 low to white; style ca. 5 mm long, lower ⁄3 In earlier literature, this species is often named white, otherwise pinkish; stigma lobes 3–5, Lepismium myosurus (Salm-Dyck ex De Can- spreading, ca. 1 mm long, white to pinkish. dolle) Pfeiffer (based on Cereus myosurus Fruits depressed globose to globose to glo- Salm-Dyck ex De Candolle 1828). bosely pear-shaped, rarely elongate-globose, Identification: Though the plants are vari- first dull green, magenta to dark red at matu- able, they are easy to identify amongst Uru- rity, to 7 mm diam. and 11 mm long, glossy, guayan epiphytic cacti due to the flat or ribbed/ ridged, massive stem segments. Figure 2. Lepismium lumbricoides. a. Distribu- General Distribution: NE to SW Brazil tion in Uruguay. b, c. Lepismium lumbricoides flow- (Pernambuco and southwards), Uruguay, SE ers; d. Flowering stem; e. Tip of young growing stem; Paraguay, NE Argentina (Formosa, Chaco, b f. Habit. Photographs: M Bonifacino, of material col- Corrientes, Misiones). a lected at Buceo, Montevideo. Distribution in Uruguay: Artigas, Cerro Largo. b c d Ecology: There is no data on the specimens available, but at least in Cerro Largo, plants have been observed (EM and MB) as epiphytes on Erythrina crista-galli (Fabaceae). 3.2.2. Lepismium lumbricoides c d (Lemaire) BarthLott (Fig 2) Epiphytic or rarely lithophytic; stems creep- ing on tree branches and then with numer- ous adventitious roots, or irregularly parallel- pendent, irregularly branched basitonically to mesotonically, all similar, dull green to me- dium green, 10–40 cm long, 4–6 mm diam., e f e f terete or slightly longitudinally ridged in the Figure 3. Rhipsalis cereuscula. a. Distribution in dry season or when old with 6–8 indistinct Uruguay. b. Habit; c, d. Flowers; e. Detail of branch- ribs; areoles minute, regularly scattered along ing stems; f. Fruit. Photographs: M Bonifacino, of cul- the length of the stems, 8–14 mm apart, on tivated material in hort. Marchesi. young stems with a minute pointed triangu- lar subtending scale-like leaf-rudiment 0.5– 1 mm wide, later naked and almost invisible, opening for ca. three days; buds ascending at or with a minute tuft of dirty white short 45° angle, greenish; open flowers spreading wool, or more rarely with weak bristly spines; to pendent, campanulate to broadly funnel- spines present on some or all areoles, 1–8 per shaped, 1–1.5 cm diam. and long; pericarpel areole, 2–5 mm long, softly bristly, straight dull dark green, globose or bluntly angled, 2– or irregularly curved, brown to almost black. 3 mm long and ca. 2 mm diam., sometimes Flowers usually scattered along the length with 1–2 small scales; perianth elements (8– of the stems, sometimes more aggregated ) 12–18, oblong to narrowly obovate, 11– towards the stem tips, diurnal, unscented, 13 mm long, 3–4 mm wide, obtuse, delicate, 166 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 167 white to greenish white or whitish cream, ceae), Prosopis (Fabaceae). Rarely lithophytic known. Flowering time: October to March; pollen pale yellow to whitish; style white, ca. sometimes almost semi-transparent, outer on somewhat shaded rocks (Arechavaleta 1905; fruiting time: April to October. 7 mm long; stigma lobes (3–) 4 (–6), spread- elements smaller; stamens ca. 30, filaments at Pan de Azucar). Comments: Variability in R. cereuscula is ing, white, 3 mm long. Fruits before maturity whitish to greenish white, 5–7 mm long; an- not very pronounced, apart from some vari- pale green, sometimes flushed dark red, white thers/pollen very pale yellow to white; style 3.3. Rhipsalis ation in fruit color (usually white but “occa- or red when ripe, turbinate, ca. 5 mm diam., white to greenish white, filiform, 9–10 mm sionally red outside Brazil” according to Taylor sometimes with two minute areoles with scant long, longer than the stamens; stigma lobes 3.3.1. Rhipsalis cereuscula haworth and Zappi 2004). The only Uruguayan speci- wool, scar left from the deciduous perianth (3–) 4–5, spreading, 2.5 mm long, white to (Fig 3) men with fruit data (Del Puerto 2061, MVFA) quite conspicuous, ca. 1.5 mm diam., fruit pale greenish. Fruits ca. globose to slightly Epiphytic or lithophytic; stems markedly het- was reported to have white fruits. wall 2 mm thick, translucent white; seeds 20– elongate-globose, ca. 5 mm diam., first green, eromorphic, with acrotonical branching, te- Identification: R. cereuscula is easily rec- 22, black. Flowering time October to Novem- changing to dark red when ripe, dry perianth rete, pale fresh green; extension shoots 7–40 ognizable due to its heteromorphic shoot ber; fruiting time June to October. remains usually remaining attached, brown (–60) cm long, 3–4 mm diam., with numer- organization with usually clavate ultimate Comments: R. floccosa is a widespread and to almost black, pulp strongly mucilaginous; ous adventitious roots, attached to branches segments. variable taxon that is divided into a total of six seeds ca. 20, black. Flowering time: Septem- or tree trunks or erect to irregularly porrect- General distribution: E and S Brazil (from subspecies with mostly allopatric distribution ber to November; fruiting time November spreading and finally pendent, with closely Pernambuco southwards), Uruguay, NE Bo- and little-pronounced differences (Anderson to February (?). set areoles bearing finely bristled, appressed, livia (La Paz), E Paraguay, NE Argentina (Cor- 2005). The total variability of the species needs Comments: Plants are variable, espe- soon-caducous spination; second-order shoots rientes, Entre Ríos, Misiones). further assessment (Taylor and Zappi 2004). cially as to presence or absence of bristly in clusters of 3–4 (–10 or more) at or near Distribution in Uruguay: Artigas, in the This taxon is newly recorded for Uruguay, but spines, and also to some extent with respect the tips of extension shoots, not always pres- drainage of the Río Cuareím but also reported the material at hand (herbarium specimens to growth form and distance between are- ent, 4–10 cm long, spineless; third-order from “Río Negro, Tacuarembó and others” only) does not allow us to assign it with cer- oles, but none of these characters appears shoots in pairs or in dense clusters at the (Arechavaleta 1905, for the synonym R. saglio- tainty to any of the subspecies. to follow a geographical or ecological pat- tip of second-order shoots or in groups of nis (Lemaire) Otto ex Walpers (Herter 1953– Identification: The combination of rubbery tern. Barthlott and Taylor (1995) recognize up to five from the tip of older third-order 1955), but no specimens have been traced to stems of equal architecture with hidden ster- two forms of this species, but it appears un- shoots, 0.5–1.5 cm long, 2–4 mm thick, usu- corroborate these citations. ile areoles and widely opening flowers make certain whether these two taxa (forma lum- ally distinctly clavate, sometimes irregularly Ecology: Forest remains along arroyos, the identification of R. floccosa easy. bricoides; forma aculeatum (F. A. C. Weber) sausage-like, rarely much abbreviated and al- epiphytic. General distribution: , Brazil, Barthlott & N. P. Taylor [= Rhipsalis acu- most globose, slightly 4–5-ribbed when old Uruguay, Paraguay, , Argentina, Bolivia. leata F. A. C. Weber], reported only for NW or desiccated, areoles 2–6 mm apart, minute, 3.3.2. Rhipsalis floccosa SaLm-Dyck ex Distribution in Uruguay: Only known Argentina) can be upheld in view of the gen- without subtending scale, segment tip with a Pfeiffer (Fig 4) from Rivera. eral variation presented by the species over bristly composite areole, bristles 2–4, trans- Epiphytic; stems spreading-pendent, in total Ecology: Epiphytic on Citharexylum mon- its geographical range. The Uruguayan ma- parent, delicate, to 1.5 mm long. Flowers at to 1.5 (–3) m long, segmented, basally becom- tevidense (‘Tarumán’; Verbenaceae), and prob- terial falls within the range of variation that the tip of third-order segments only, pen- ing woody and to 1.5 cm thick, usually with- ably local and rare. Barthlott and Taylor imply for forma lumbri- dent, diurnal, opening for several days, un- out adventitious roots, branched acrotonically coides, although their differential description scented, campanulate to funnel-shaped, ca. with groups of (2–) 3–5 branches, all stems 4. Discussion of forma aculeatum calls for whitish bristles 1.2 cm diam. and 15 mm long; pericarpel similar, to 20 (–30) cm long and 4–5 (rarely In South America, subtropical forests reach (rather than brown to blackish as observed green to pale yellowish green, more or less to 12) mm thick, terete or somewhat irregu- their southern limits in Uruguay and in the in Uruguayan collections). globose to obconical, 4–5 mm diam. and larly angled, very weakly thickened below the Buenos Aires province of Argentina. In Uru- Identification: The combination of ba- long, naked or occasionally with 1–2 scales, rudimentary leaf subtending the areoles, gray- guay most of these forests are associated with sitonic to mesotonic branching and terete scales reddish triangular, 0.2 mm long, with green, dull, somewhat rubbery but firm and the extensive system of hills and accompany- stems with greenish white to whitish cream or without bristles in their axils; perianth el- not flaccid, minutely scabrid; areoles 1.5–4 cm ing rivers, and it is in these habitats where the flowers allow easy identification of the taxon ements 11–15, the inner 7–8 slightly larger distant from one another, immersed into the Uruguayan epiphytic cacti are to be expected. in Uruguay. than the outer, 8–10 mm long and 3 mm stem tissue and largely invisible before produc- L. lumbricoides is widely distributed through- General distribution: Brazil (São Paulo wide, narrowly elliptic, acute, white to very ing a flower, without subtending scale, older out the country. Our knowledge of the distri- and southwards), Uruguay, Paraguay, Argen- pale creamy-white, tip faintly flushed pink- areoles to 4 mm wide, densely woolly-bristly, bution of the other three species of epiphytic tina (Catamarca and northwards), E Bolivia. ish, the outer elements more spreading, the otherwise spineless, producing additional flow- cacti (L. cruciforme, R. floccosa, and R. cereus- This is a widespread taxon. Font (2003) gives inner remaining more erect; stamens ca. 36, ers over the course of time. Flowers scattered cula) is scanty. Considering the long history of more details for the occurrence in Argentina, filaments white with somewhat reddish base, along the length of the stems, diurnal, flatly exploration in the country and its rather small including a list of provinces. 7–10 mm long; anthers/pollen pale yellow; opening, 1.2–2.2 cm diam., opening for 2– area we think this is a case of genuine rarity Distribution in Uruguay: Throughout style white, filiform, 9–10 mm long, longer 3 days, scented, nectariferous disc surround- in the country, rather than overall poor sam- the country. Herter (1953–1955) lists Artigas, than the stamens and usually also surpass- ing the style; pericarpel semi-globose, 3 mm pling. All three taxa reach the fringe of their Cerro Largo, Durazno, Maldonado, Rivera, ing the perianth; stigma lobes 3–5, spread- diam. and long, completely immersed in the southern distribution in Uruguay. It is an- Salto, San José, Soriano, Tacuarembó, and ing to recurved, whitish. Fruits more-or-less woolly areole, cream-colored, without scales; ticipated, however, that more records will be Treinta y Tres; Herter (1930) also gives Co- globose, white or occasionally red, 5–7 mm outer perianth elements 3–5, creamy yellow added for these three rare species when more lonia (for the synonym Rhipsalis aculeata). diam., dried perianth remains persistent, with reddish tips, inner perianth elements 7–8, areas are explored. Ecology: Forests and forest remains along base somewhat immersed into the top of oblong, obtuse and slightly cucullate, 8 mm Uruguay has witnessed rather aggressive de- arroyos, epiphytic on a variety of trees, for in- the fruit, pulp strongly mucilaginous; seeds long and 3–4.5 mm wide, greenish-white to forestation implemented throughout the coun- stance Myrsine (Myrsinaceae), Erythrina crista- black (pers. obs., Ritter 1190 from Paraguay, white, delicately translucent; stamens very nu- try during the last three decades, the effects of galli (‘Ceibo’; Fabaceae), Celtis tala (Ulma- herbarium ZSS 13619, 14677), number not merous (60+), filaments white, 5–7 mm long; which are yet to be determined in terms of the 166 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 167 white to greenish white or whitish cream, ceae), Prosopis (Fabaceae). Rarely lithophytic known. Flowering time: October to March; pollen pale yellow to whitish; style white, ca. sometimes almost semi-transparent, outer on somewhat shaded rocks (Arechavaleta 1905; fruiting time: April to October. 7 mm long; stigma lobes (3–) 4 (–6), spread- elements smaller; stamens ca. 30, filaments at Pan de Azucar). Comments: Variability in R. cereuscula is ing, white, 3 mm long. Fruits before maturity whitish to greenish white, 5–7 mm long; an- not very pronounced, apart from some vari- pale green, sometimes flushed dark red, white thers/pollen very pale yellow to white; style 3.3. Rhipsalis ation in fruit color (usually white but “occa- or red when ripe, turbinate, ca. 5 mm diam., white to greenish white, filiform, 9–10 mm sionally red outside Brazil” according to Taylor sometimes with two minute areoles with scant long, longer than the stamens; stigma lobes 3.3.1. Rhipsalis cereuscula haworth and Zappi 2004). The only Uruguayan speci- wool, scar left from the deciduous perianth (3–) 4–5, spreading, 2.5 mm long, white to (Fig 3) men with fruit data (Del Puerto 2061, MVFA) quite conspicuous, ca. 1.5 mm diam., fruit pale greenish. Fruits ca. globose to slightly Epiphytic or lithophytic; stems markedly het- was reported to have white fruits. wall 2 mm thick, translucent white; seeds 20– elongate-globose, ca. 5 mm diam., first green, eromorphic, with acrotonical branching, te- Identification: R. cereuscula is easily rec- 22, black. Flowering time October to Novem- changing to dark red when ripe, dry perianth rete, pale fresh green; extension shoots 7–40 ognizable due to its heteromorphic shoot ber; fruiting time June to October. remains usually remaining attached, brown (–60) cm long, 3–4 mm diam., with numer- organization with usually clavate ultimate Comments: R. floccosa is a widespread and to almost black, pulp strongly mucilaginous; ous adventitious roots, attached to branches segments. variable taxon that is divided into a total of six seeds ca. 20, black. Flowering time: Septem- or tree trunks or erect to irregularly porrect- General distribution: E and S Brazil (from subspecies with mostly allopatric distribution ber to November; fruiting time November spreading and finally pendent, with closely Pernambuco southwards), Uruguay, NE Bo- and little-pronounced differences (Anderson to February (?). set areoles bearing finely bristled, appressed, livia (La Paz), E Paraguay, NE Argentina (Cor- 2005). The total variability of the species needs Comments: Plants are variable, espe- soon-caducous spination; second-order shoots rientes, Entre Ríos, Misiones). further assessment (Taylor and Zappi 2004). cially as to presence or absence of bristly in clusters of 3–4 (–10 or more) at or near Distribution in Uruguay: Artigas, in the This taxon is newly recorded for Uruguay, but spines, and also to some extent with respect the tips of extension shoots, not always pres- drainage of the Río Cuareím but also reported the material at hand (herbarium specimens to growth form and distance between are- ent, 4–10 cm long, spineless; third-order from “Río Negro, Tacuarembó and others” only) does not allow us to assign it with cer- oles, but none of these characters appears shoots in pairs or in dense clusters at the (Arechavaleta 1905, for the synonym R. saglio- tainty to any of the subspecies. to follow a geographical or ecological pat- tip of second-order shoots or in groups of nis (Lemaire) Otto ex Walpers (Herter 1953– Identification: The combination of rubbery tern. Barthlott and Taylor (1995) recognize up to five from the tip of older third-order 1955), but no specimens have been traced to stems of equal architecture with hidden ster- two forms of this species, but it appears un- shoots, 0.5–1.5 cm long, 2–4 mm thick, usu- corroborate these citations. ile areoles and widely opening flowers make certain whether these two taxa (forma lum- ally distinctly clavate, sometimes irregularly Ecology: Forest remains along arroyos, the identification of R. floccosa easy. bricoides; forma aculeatum (F. A. C. Weber) sausage-like, rarely much abbreviated and al- epiphytic. General distribution: Venezuela, Brazil, Barthlott & N. P. Taylor [= Rhipsalis acu- most globose, slightly 4–5-ribbed when old Uruguay, Paraguay, Peru, Argentina, Bolivia. leata F. A. C. Weber], reported only for NW or desiccated, areoles 2–6 mm apart, minute, 3.3.2. Rhipsalis floccosa SaLm-Dyck ex Distribution in Uruguay: Only known Argentina) can be upheld in view of the gen- without subtending scale, segment tip with a Pfeiffer (Fig 4) from Rivera. eral variation presented by the species over bristly composite areole, bristles 2–4, trans- Epiphytic; stems spreading-pendent, in total Ecology: Epiphytic on Citharexylum mon- its geographical range. The Uruguayan ma- parent, delicate, to 1.5 mm long. Flowers at to 1.5 (–3) m long, segmented, basally becom- tevidense (‘Tarumán’; Verbenaceae), and prob- terial falls within the range of variation that the tip of third-order segments only, pen- ing woody and to 1.5 cm thick, usually with- ably local and rare. Barthlott and Taylor imply for forma lumbri- dent, diurnal, opening for several days, un- out adventitious roots, branched acrotonically coides, although their differential description scented, campanulate to funnel-shaped, ca. with groups of (2–) 3–5 branches, all stems 4. Discussion of forma aculeatum calls for whitish bristles 1.2 cm diam. and 15 mm long; pericarpel similar, to 20 (–30) cm long and 4–5 (rarely In South America, subtropical forests reach (rather than brown to blackish as observed green to pale yellowish green, more or less to 12) mm thick, terete or somewhat irregu- their southern limits in Uruguay and in the in Uruguayan collections). globose to obconical, 4–5 mm diam. and larly angled, very weakly thickened below the Buenos Aires province of Argentina. In Uru- Identification: The combination of ba- long, naked or occasionally with 1–2 scales, rudimentary leaf subtending the areoles, gray- guay most of these forests are associated with sitonic to mesotonic branching and terete scales reddish triangular, 0.2 mm long, with green, dull, somewhat rubbery but firm and the extensive system of hills and accompany- stems with greenish white to whitish cream or without bristles in their axils; perianth el- not flaccid, minutely scabrid; areoles 1.5–4 cm ing rivers, and it is in these habitats where the flowers allow easy identification of the taxon ements 11–15, the inner 7–8 slightly larger distant from one another, immersed into the Uruguayan epiphytic cacti are to be expected. in Uruguay. than the outer, 8–10 mm long and 3 mm stem tissue and largely invisible before produc- L. lumbricoides is widely distributed through- General distribution: Brazil (São Paulo wide, narrowly elliptic, acute, white to very ing a flower, without subtending scale, older out the country. Our knowledge of the distri- and southwards), Uruguay, Paraguay, Argen- pale creamy-white, tip faintly flushed pink- areoles to 4 mm wide, densely woolly-bristly, bution of the other three species of epiphytic tina (Catamarca and northwards), E Bolivia. ish, the outer elements more spreading, the otherwise spineless, producing additional flow- cacti (L. cruciforme, R. floccosa, and R. cereus- This is a widespread taxon. Font (2003) gives inner remaining more erect; stamens ca. 36, ers over the course of time. Flowers scattered cula) is scanty. Considering the long history of more details for the occurrence in Argentina, filaments white with somewhat reddish base, along the length of the stems, diurnal, flatly exploration in the country and its rather small including a list of provinces. 7–10 mm long; anthers/pollen pale yellow; opening, 1.2–2.2 cm diam., opening for 2– area we think this is a case of genuine rarity Distribution in Uruguay: Throughout style white, filiform, 9–10 mm long, longer 3 days, scented, nectariferous disc surround- in the country, rather than overall poor sam- the country. Herter (1953–1955) lists Artigas, than the stamens and usually also surpass- ing the style; pericarpel semi-globose, 3 mm pling. All three taxa reach the fringe of their Cerro Largo, Durazno, Maldonado, Rivera, ing the perianth; stigma lobes 3–5, spread- diam. and long, completely immersed in the southern distribution in Uruguay. It is an- Salto, San José, Soriano, Tacuarembó, and ing to recurved, whitish. Fruits more-or-less woolly areole, cream-colored, without scales; ticipated, however, that more records will be Treinta y Tres; Herter (1930) also gives Co- globose, white or occasionally red, 5–7 mm outer perianth elements 3–5, creamy yellow added for these three rare species when more lonia (for the synonym Rhipsalis aculeata). diam., dried perianth remains persistent, with reddish tips, inner perianth elements 7–8, areas are explored. Ecology: Forests and forest remains along base somewhat immersed into the top of oblong, obtuse and slightly cucullate, 8 mm Uruguay has witnessed rather aggressive de- arroyos, epiphytic on a variety of trees, for in- the fruit, pulp strongly mucilaginous; seeds long and 3–4.5 mm wide, greenish-white to forestation implemented throughout the coun- stance Myrsine (Myrsinaceae), Erythrina crista- black (pers. obs., Ritter 1190 from Paraguay, white, delicately translucent; stamens very nu- try during the last three decades, the effects of galli (‘Ceibo’; Fabaceae), Celtis tala (Ulma- herbarium ZSS 13619, 14677), number not merous (60+), filaments white, 5–7 mm long; which are yet to be determined in terms of the 168 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 169 changes inflicted to native vegetation. However, on the distribution of the three rare species Este with Minas Corrales, 41 km S of Ruta Rhipsalis floccosa the biggest threat to epiphytic cacti is the cut- is needed in order to assess their conservation 27 ( = 35 km S of Amarillo), 21.10.2005, Uruguay. Rivera; en quebrada afluente ar. ting down of native forests, in particular “ser- status with more confidence on the national Nyffeler and Eggli 1615 (MVJB, ZSS); Ar- Lonarejo, ruta 30, km 107, 11.4.1984, Bayce rano” forest (a rather low, 4–6 m high, xero- Uruguayan level. Taylor (2006) lists an over- tigas; some km SE of Artigas on Ruta 30, and al. s.n. (MVFA 17322); Rivera; “Ao. phytic forest associated with hilly areas), for all conservation assessment of “least concern” then 7 km on lateral road to and beyond Pie- Potrero, Po. Ataques”, 12.12.1997, Bonifac- fuel wood and in the reduced and fragmentary for all four taxa. dra Pintada, 23.10.2005, Nyffeler and Eggli ino and al. s.n. (MVFA 27638); Rivera; Ar- nature of the most humid and lush forests lo- 1633 (MVJB, ZSS); Río Negro; Ruta 20, km royo Rubio Chico, 12.5.1995, Brussa and al. cated in the north of the country (EM and MB, 5. Material examined 32, 22.3 km E of Ruta 24, Arroyo Sanchez, s.n. (MVFA 24963). pers. obs.). These northern forests constitute 25.10.2005, Nyffeler and Eggli 1655 (MVJB, the habitat of the three more-rare species. Lepismium cruciforme ZSS); Treinta y Tres; Ruta 18, Arroyo Ceibal, Acknowledgments As a result of the condition of the Uru- Uruguay. Artigas; “Riusa, San Gregorio”, 21.10.1969, Olano and al. s.n. (MVFA 8783); UE and RN are grateful to the Museo y guayan native forests and the threats imposed 30.3.1962, Del Puerto 2060 (MVFA s.n.); Treinta y Tres; “Río Olimar a la altura de T y Jardín Botánico de Montevideo for help by logging, all taxa could be considered slightly Cerro Largo; “en camino sobre Sierra de Tres”, 22.10.1969, Olano and al. s.n. (MVFA provided during two field trips in 2004 and to moderately threatened. More information Ríos, al S de Esc. 25 y caserón, al N Puesto 8815); Rivera; “Tranqueras”, 22.2.1947, Os- 2005. Support for fieldwork was provided Policial”, 21.10.1992, Izaguirre and al. s.n. orio s.n. (MVM 13526); Soriano; “Mercedes, by the Brasilien-Fonds of Rösly and Wer- (MVFA 21094B). Cololó”, 17.2.1892, Osten 2936 (MVM s.n.); ner Uebelmann as well as the British Cac- Figure 4. Rhipsalis floccosa. a. Distribution in b. c. Soriano; “Cololó prope Mercedes”, 29.9.1895, tus and Succulent Society to UE and by the Uruguay. Habit; Flowering and fruiting branch; Lepismium lumbricoides d. Flower; e. Fruit. Photographs: M Bonifacino, of Osten 4190 (MVM s.n.). Claraz-Schenkung to RN. The curators of cultivated material in hort. Marchesi.captino Uruguay. Colonia; “Isla Juncal”, s.a., Anony- MVM, MVFA, and MVJB are thanked for mus s.n. (MVM 17095); San José; “Arazatí”, Rhipsalis cereuscula granting access to their collections. We are 11.1933, Ardao s.n. (MVM 10122); Rocha; Uruguay. Artigas; “Bella Unión”, 23.1.1942, further grateful to the CITES authorities of “Laguna de Castillos, costa N”, 3.12.1995, Anonymus s.n. (MVFA 3743); Artigas; “Cuaró”, Uruguay for the necessary collecting permits, Bayce and al. s.n. (MVFA 26477); Artigas; s.a., Anonymus s.n. (MVM 17328); Artigas; and UE expresses his thanks to the Sukku- “Cuareím”, 9.1901, Berro s.n. (MVFA 4046); “Riusa, San Gregorio”, 30.3.1962, Del Puerto lenten-Sammlung Zürich and Grün Stadt Artigas; “Cuareím, Sta. Rosa”, 1.9.1901, Berro 2061 (MVFA s.n.); Artigas; “Río Cuareím, al Zürich for their permission to participate s.n. (MVFA 2144 pp); Artigas; “Cuareím, Sta. oeste del ar. Yucutujá”/“Estancia El Ombu de in this project. Ralph Martin (Cardiff, GB) Rosa”, 15.9.1901, Berro s.n. (MVFA 4045); Mallo, A. Yucutujá y R. Cuareím”, 12.4.1978, provided statistical information on recorded Artigas; “Cuareím”, 1.9.1901, Berro s.n. Del Puerto and Marchesi s.n. (MVFA 15313); Uruguayan collecting localities. Beat Leuen- (MVFA 2144 pp); aldonado; “Pan de Azu- Artigas; Río Cuareím y ar. Yucutujá, 6.10.1997, berger (Berlin) supplied description informa- car”, 12.1899, Berro s.n. (MVFA 4044); Mal- Marchesi s.n. (MVFA 26950). tion on Rhipsalis floccosa. donado; “Pan de Azucar”, 12.1899, Berro s.n. (MVFA 3038); Maldonado; “Pan de Azucar”, Literature Cited 12.1899, Berro s.n. (MVFA 3088); Soriano; “Vera”, 4.11.1902, Berro s.n. (MVFA 2540); Anderson EF. 2005. Das grosse Kakteenlexikon. Herter G. 1930. Estudios botánicos en la región Treinta y Tres; “Cerro Aspero”, 29.11.1899, Ulmer, Stuttgart ( = augmented and corrected Uruguaya. IV. Florula Uruguayensis. Plantae a Berro s.n. (MVFA 4047); Artigas; “Ar. Yucu- German translation of the original English edi- vasculares. Republica Oriental del Uruguay, tujá, Paso iraponchos, Ruta 30, 26.8.1995, tion of 2001: The cactus family. Timber Press, Montevideo. b c Portland). Herter G. 1953–1955. Flore illustrée de l’Uruguay. Bonifacino s.n. (MVFA 25017); Montevi- Arechavaleta J. 1905. Flora Uruguaya. Vol- Cactaceae. Cactus (Paris) No. 38: 267–276; No. deo; “Cementerio del Buceo”, Bonifacino ume 2. [Cactaceae]. Anales Mus Nac Montevi- 39: 19–24; No. 41: 91–96; No. 42: 119–124; No. s.n. (photographs only); Río Negro; Palmares deo 5: 161–292. 44: 177–179; No. 45: 203–206. Pre-publication de Porrúa, 16.10.1995, Brescia and al. s.n. Barthlott W, Taylor NP. 1995. Notes towards with additional notes of Herter 1956. (MVFA 26810); Tacuarembó; “Rtua 44, Ao. a monograph of Rhipsalideae (Cactaceae). Brad- Herter G. 1956. Flora Ilustrada del Uruguay, Yaguarí”, 7.10.1961, Del Puerto 242 (MVFA leya 13: 43–79. Fasc. 12. Publicado por el autor. Cactaceae s.n.); Rocha; “Parque San Miguel”, 6.10.1965, Bauer R. 2003. A synopsis of the tribe Hylocer- pp581–600. Del Puerto and Marchesi 5270 (MVFA s.n.); eeae F. Buxb. Cactaceae Syst Init 17: 3–63. Hunt DR. 1999. CITES Cactaceae Checklist. Ed. Artigas; “area a inundar por Represa Salto Bletter N and others. 2004. A digital base 2. RBG Kew, Richmond. map for studying the neotropical flora. Taxon Nyffeler R. 2000. Should Pfeiffera be resurrected? Grande al sur del Ao. Guaviyú (Paredón)”, 53(2): 469–477. Cactaceae Syst Init 10: 10–11. 12.1977, Del Puerto-Berreta 14753 (MVFA Britton NL, Rose JN. 1923. The Cactaceae. Nyffeler R. 2002. Phylogenetic relationships in s.n.); Salto; Arapey, 9.1949, Herter [Pl. Urug.] Volume IV. Carnegie Institution, Washing- the Cactus Family (Cactaceae) based on evi- 1130c (B, MO?); Rivera; Ruta 6, 2 km al N ton DC dence from trnK/matK and trnL-trnF sequences. d e de Vichadero”, 20.10.1992, Izaguirre and al. Brumitt RK. 1996. Report of the Committee for Amer. J. Bot. 89: 312–326. s.n. (MVFA 21048B); Rocha; “Arroyo Cha- Spermatophyta, 44. Taxon 45(4): 671–681. Osten C. 1941. Notas sobre Cactáceas. Legrand falote”, s.a., Legrand 3733 (MVM s.n.); Mal- Eggli U, Leuenberger BE. 1996. A quick and D, editor. Museo Nacional de Historia Natu- donado; “Sierra de las Animas”, 15.9.1963, easy method for drying specimens, in- ral, Montevideo. cluding succulents, for the herbarium. Taxon Taylor NP, Zappi DC. 2004. Cacti of eastern Bra- Marchesi 818 (MVFA s.n.); Salto; Campos 45: 259–261. zil. Royal Botanic Garden Kew, Richmond. de Zunini, Río Arapey, 17.9.1976, Marchesi Font F. 2003. Cactáceas de la Provincia de Bue- Taylor NP (compiler). 2006. Conservation assess- s.n. (MVFA 12802); Rivera; unnumbered nos Aires. Rev Circ Colec Cact Crasas Rep Ar- ments; pp. 324–335. In Hunt DR, editor. The road connecting Ruta 27 at Cerro Pelado al gent 2(4): 100–128. new cactus lexicon. dh books, Milborne Port. 168 EGGLI AND OTHERS—EpIpHyTIc cAcTI IN URUGUAy HASELTONIA 14, 2008 169 changes inflicted to native vegetation. However, on the distribution of the three rare species Este with Minas Corrales, 41 km S of Ruta Rhipsalis floccosa the biggest threat to epiphytic cacti is the cut- is needed in order to assess their conservation 27 ( = 35 km S of Amarillo), 21.10.2005, Uruguay. Rivera; en quebrada afluente ar. ting down of native forests, in particular “ser- status with more confidence on the national Nyffeler and Eggli 1615 (MVJB, ZSS); Ar- Lonarejo, ruta 30, km 107, 11.4.1984, Bayce rano” forest (a rather low, 4–6 m high, xero- Uruguayan level. Taylor (2006) lists an over- tigas; some km SE of Artigas on Ruta 30, and al. s.n. (MVFA 17322); Rivera; “Ao. phytic forest associated with hilly areas), for all conservation assessment of “least concern” then 7 km on lateral road to and beyond Pie- Potrero, Po. Ataques”, 12.12.1997, Bonifac- fuel wood and in the reduced and fragmentary for all four taxa. dra Pintada, 23.10.2005, Nyffeler and Eggli ino and al. s.n. (MVFA 27638); Rivera; Ar- nature of the most humid and lush forests lo- 1633 (MVJB, ZSS); Río Negro; Ruta 20, km royo Rubio Chico, 12.5.1995, Brussa and al. cated in the north of the country (EM and MB, 5. Material examined 32, 22.3 km E of Ruta 24, Arroyo Sanchez, s.n. (MVFA 24963). pers. obs.). These northern forests constitute 25.10.2005, Nyffeler and Eggli 1655 (MVJB, the habitat of the three more-rare species. Lepismium cruciforme ZSS); Treinta y Tres; Ruta 18, Arroyo Ceibal, Acknowledgments As a result of the condition of the Uru- Uruguay. Artigas; “Riusa, San Gregorio”, 21.10.1969, Olano and al. s.n. (MVFA 8783); UE and RN are grateful to the Museo y guayan native forests and the threats imposed 30.3.1962, Del Puerto 2060 (MVFA s.n.); Treinta y Tres; “Río Olimar a la altura de T y Jardín Botánico de Montevideo for help by logging, all taxa could be considered slightly Cerro Largo; “en camino sobre Sierra de Tres”, 22.10.1969, Olano and al. s.n. (MVFA provided during two field trips in 2004 and to moderately threatened. More information Ríos, al S de Esc. 25 y caserón, al N Puesto 8815); Rivera; “Tranqueras”, 22.2.1947, Os- 2005. Support for fieldwork was provided Policial”, 21.10.1992, Izaguirre and al. s.n. orio s.n. (MVM 13526); Soriano; “Mercedes, by the Brasilien-Fonds of Rösly and Wer- (MVFA 21094B). Cololó”, 17.2.1892, Osten 2936 (MVM s.n.); ner Uebelmann as well as the British Cac- Figure 4. Rhipsalis floccosa. a. Distribution in b. c. Soriano; “Cololó prope Mercedes”, 29.9.1895, tus and Succulent Society to UE and by the Uruguay. Habit; Flowering and fruiting branch; Lepismium lumbricoides d. Flower; e. Fruit. Photographs: M Bonifacino, of Osten 4190 (MVM s.n.). Claraz-Schenkung to RN. The curators of cultivated material in hort. Marchesi.captino Uruguay. Colonia; “Isla Juncal”, s.a., Anony- MVM, MVFA, and MVJB are thanked for mus s.n. (MVM 17095); San José; “Arazatí”, Rhipsalis cereuscula granting access to their collections. We are 11.1933, Ardao s.n. (MVM 10122); Rocha; Uruguay. Artigas; “Bella Unión”, 23.1.1942, further grateful to the CITES authorities of “Laguna de Castillos, costa N”, 3.12.1995, Anonymus s.n. (MVFA 3743); Artigas; “Cuaró”, Uruguay for the necessary collecting permits, Bayce and al. s.n. (MVFA 26477); Artigas; s.a., Anonymus s.n. (MVM 17328); Artigas; and UE expresses his thanks to the Sukku- “Cuareím”, 9.1901, Berro s.n. (MVFA 4046); “Riusa, San Gregorio”, 30.3.1962, Del Puerto lenten-Sammlung Zürich and Grün Stadt Artigas; “Cuareím, Sta. Rosa”, 1.9.1901, Berro 2061 (MVFA s.n.); Artigas; “Río Cuareím, al Zürich for their permission to participate s.n. (MVFA 2144 pp); Artigas; “Cuareím, Sta. oeste del ar. Yucutujá”/“Estancia El Ombu de in this project. Ralph Martin (Cardiff, GB) Rosa”, 15.9.1901, Berro s.n. (MVFA 4045); Mallo, A. Yucutujá y R. Cuareím”, 12.4.1978, provided statistical information on recorded Artigas; “Cuareím”, 1.9.1901, Berro s.n. Del Puerto and Marchesi s.n. (MVFA 15313); Uruguayan collecting localities. Beat Leuen- (MVFA 2144 pp); aldonado; “Pan de Azu- Artigas; Río Cuareím y ar. Yucutujá, 6.10.1997, berger (Berlin) supplied description informa- car”, 12.1899, Berro s.n. (MVFA 4044); Mal- Marchesi s.n. (MVFA 26950). tion on Rhipsalis floccosa. donado; “Pan de Azucar”, 12.1899, Berro s.n. (MVFA 3038); Maldonado; “Pan de Azucar”, Literature Cited 12.1899, Berro s.n. (MVFA 3088); Soriano; “Vera”, 4.11.1902, Berro s.n. (MVFA 2540); Anderson EF. 2005. Das grosse Kakteenlexikon. Herter G. 1930. Estudios botánicos en la región Treinta y Tres; “Cerro Aspero”, 29.11.1899, Ulmer, Stuttgart ( = augmented and corrected Uruguaya. IV. Florula Uruguayensis. Plantae a Berro s.n. (MVFA 4047); Artigas; “Ar. Yucu- German translation of the original English edi- vasculares. Republica Oriental del Uruguay, tujá, Paso iraponchos, Ruta 30, 26.8.1995, tion of 2001: The cactus family. Timber Press, Montevideo. b c Portland). Herter G. 1953–1955. Flore illustrée de l’Uruguay. Bonifacino s.n. (MVFA 25017); Montevi- Arechavaleta J. 1905. Flora Uruguaya. Vol- Cactaceae. Cactus (Paris) No. 38: 267–276; No. deo; “Cementerio del Buceo”, Bonifacino ume 2. [Cactaceae]. Anales Mus Nac Montevi- 39: 19–24; No. 41: 91–96; No. 42: 119–124; No. s.n. (photographs only); Río Negro; Palmares deo 5: 161–292. 44: 177–179; No. 45: 203–206. Pre-publication de Porrúa, 16.10.1995, Brescia and al. s.n. Barthlott W, Taylor NP. 1995. Notes towards with additional notes of Herter 1956. (MVFA 26810); Tacuarembó; “Rtua 44, Ao. a monograph of Rhipsalideae (Cactaceae). Brad- Herter G. 1956. Flora Ilustrada del Uruguay, Yaguarí”, 7.10.1961, Del Puerto 242 (MVFA leya 13: 43–79. Fasc. 12. Publicado por el autor. Cactaceae s.n.); Rocha; “Parque San Miguel”, 6.10.1965, Bauer R. 2003. A synopsis of the tribe Hylocer- pp581–600. Del Puerto and Marchesi 5270 (MVFA s.n.); eeae F. Buxb. Cactaceae Syst Init 17: 3–63. Hunt DR. 1999. CITES Cactaceae Checklist. Ed. Artigas; “area a inundar por Represa Salto Bletter N and others. 2004. A digital base 2. RBG Kew, Richmond. map for studying the neotropical flora. Taxon Nyffeler R. 2000. Should Pfeiffera be resurrected? Grande al sur del Ao. Guaviyú (Paredón)”, 53(2): 469–477. Cactaceae Syst Init 10: 10–11. 12.1977, Del Puerto-Berreta 14753 (MVFA Britton NL, Rose JN. 1923. The Cactaceae. Nyffeler R. 2002. Phylogenetic relationships in s.n.); Salto; Arapey, 9.1949, Herter [Pl. Urug.] Volume IV. Carnegie Institution, Washing- the Cactus Family (Cactaceae) based on evi- 1130c (B, MO?); Rivera; Ruta 6, 2 km al N ton DC dence from trnK/matK and trnL-trnF sequences. d e de Vichadero”, 20.10.1992, Izaguirre and al. Brumitt RK. 1996. Report of the Committee for Amer. J. Bot. 89: 312–326. s.n. (MVFA 21048B); Rocha; “Arroyo Cha- Spermatophyta, 44. Taxon 45(4): 671–681. Osten C. 1941. Notas sobre Cactáceas. Legrand falote”, s.a., Legrand 3733 (MVM s.n.); Mal- Eggli U, Leuenberger BE. 1996. A quick and D, editor. Museo Nacional de Historia Natu- donado; “Sierra de las Animas”, 15.9.1963, easy method for drying plant specimens, in- ral, Montevideo. cluding succulents, for the herbarium. Taxon Taylor NP, Zappi DC. 2004. Cacti of eastern Bra- Marchesi 818 (MVFA s.n.); Salto; Campos 45: 259–261. zil. Royal Botanic Garden Kew, Richmond. de Zunini, Río Arapey, 17.9.1976, Marchesi Font F. 2003. Cactáceas de la Provincia de Bue- Taylor NP (compiler). 2006. Conservation assess- s.n. (MVFA 12802); Rivera; unnumbered nos Aires. Rev Circ Colec Cact Crasas Rep Ar- ments; pp. 324–335. In Hunt DR, editor. The road connecting Ruta 27 at Cerro Pelado al gent 2(4): 100–128. new cactus lexicon. dh books, Milborne Port.