Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations

1988 Flora and vegetation of the Huascarán National Park, Ancash, Peru: with preliminary taxonomic studies for a manual of the flora David Nelson Smith Iowa State University

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Recommended Citation Smith, David Nelson, "Flora and vegetation of the Huascarán National Park, Ancash, Peru: with preliminary taxonomic studies for a manual of the flora " (1988). Retrospective Theses and Dissertations. 8891. https://lib.dr.iastate.edu/rtd/8891

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Flora and vegetation of the Huascarân National Park, Ancash, Peru, with preliminary taxonomic studies for a manual of the flora

Smith, David Nelson, Ph.D. Iowa State University, 1988

300N.ZeebRd. Ann Aibor, MI 48106

Flora and vegetation of the Huascardn National Park, Ancash, Peru, with preliminary taxonomic studies for a manual of the flora

by

David Nelson Smith

A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY

Department: Botany Major Botany (Taxonomy)

Approved;

Signature was redacted for privacy. In Charge of Major Work

Signature was redacted for privacy. For the Major Department

Signature was redacted for privacy. le Graduate College

Iowa State University Ames, Iowa

1988 11

TABLE OF CONTENTS Page PART I. THE HUASCARAN NATIONAL PARK: ENVIRONMENT, VEGETATION, AND FLORA 1 INTRODUCTION 2 GEOLOGY 6 CLIMATE 17 VEGETATION OF THE HUASCARAN NATIONAL PARK 29 THE FLORA: COMMENTARY AND LIST OF TAXA 45 BIOGEOGRAPHIC RELATIONSHIPS OF THE HUASCARAN FLORA 67 BIOGEOGRAPHIC LISTS 72

PART II. PRELIMINARY STUDIES OF THE FLORA OF THE HUASCARAN NATIONAL PARK: MONOCOT FAMILIES (EXCLUDING POACEAE) 112 TO THE READER 113 MATERIALS AND METHODS 116 GENERAL KEYS TO THE FAMILIES 122 FAMILY TREATMENTS: 126 Agavaceae 126 Alliaceae 129 Alstroemeriaceae 131 Amaryllidaceae 138 Antherlcaceae 141 Bromeliaceae 143 Commelinaceae 164 Cyperaceae 167 Dloscoreaceae 193 Hydrocharitaceae 196 Iridaceae 198 Juncaceae 209 Juncaginaceae 221 Lemnaceae 223 Orchidaceae 225 iii

Potamogetonaceae 253 Zannichelliaceae 2SS LITERATURE CITED 257 ACKNOWLEDGMENTS 279 1

PART I. THE HUASCARAN NATIONAL PARK: ENVIRONMENT, VEGETATION, AND FLORA 2

INTRODUCTION The Huascarân National Park and International Biosphere Reserve is located in the Department of Ancash, Peru, immediately north of the Department of Lima. The Park occupies the major part of the Cordillera Blanca, which is the world's highest tropical mountain range, and has the highest mountain peaks in South America north of Argentina and Chile. The Park (see Map 1) is oriented northwest-southeast, has a length of 158 km, and the width varies from 11 to 39 km. The reserve has an area of 340,000 ha, and includes elevations from 3240 to 6770 meters above sea level, at the peak of Nevado Huascardn Sur. The boundaries of the Park are usually located at 3500 m, or above. The Cordillera Blanca forms part of the continental divide between western, coastal Peru and the Amazon Basin to the east. Water from the western valleys flows to the Rio Santa, and then to the Pacific Ocean, while the water from the eastern valleys flows to the Atlantic Ocean via the Amazon and Maraflon Rivers and their tributaries. The mountain building events that created the Cordillera Blanca occurred rapidly, with great uplift, and the range has been strongly eroded since uplift began (details of the geologic history and references are in the chapter on geology). These combined factors have resulted in a very steep and dissected topography with many river valleys on both the east and west sides of the Cordillera. The rough topography, extensive areas of snow and glaciers, torrential streams, and vegetation give the Park great scenic beauty. Since the last century, visitors have commented on its beauty, and in this century there have been several proposals for the creation of a Natural Park. These came to fruit in 1975, when the Huascarân National Park was established by presidential decree (ORDEZA, no date). At a later date, it was declared an International Biosphere Reserve by the UNESCO, one of the three in Peru. The western side of the Cordillera Blanca is accessible from the many towns and villages in the Rio Santa valley. Huaraz is the principal city of the valley, and the departmental capitol as well. Caraz, Yungay, Carhuaz, Recuay, and Catac are also cities of regional importance. The Rio Santa valley is readily accessible via a paved road from the coast. There is direct bus service from Lima to Huaraz and Caraz, serving all the major towns in the valley. The eastern side of the Cordillera is less readily accessible; there is a gravel road crossing the Cordillera between Catac and Chavin, which continues northward around the Park via Pomabamba, with connection to the Rio Santa valley. More or less regular bus service is available on that route. MAJOR PEAKS ALTITUDE* 1 Champara Este 5750 m 2 Mllluacociia 5480m 3 Pucahirea Noite 6050 m 40 s 4 Santa Cniz Sur 6260 m 5 Caraz 6025m 6 HuandoyNorte 6395m 7 Chacraraju 6075 m 8 Huascaran Sur 6770 m 9 Contraheirbas Central 6040 m 10 Hualcan Norte 6125 m 11 CopaSur 6190 m 12 Palcaralu Este 6275 m w 8*50 S 13 CMnchey 6220m 14 Huantzan 6395m 15 Uruashraju Norte 5720 m 16 Morron]u 5710 m 17 Rarla Norte 5580m 18 Caullaraju 5690m *Altitude rounded to nearest S m. Sources: Bartie (1981) and DIaz Bustos (1984) ry YUNGAY @"10 S

CARHUAZ

9'20"S + Ishinea

*"30 S HUARAZ

CARHUAZ

9®20'S + Ishinea

HUANTAR 4 9*30 s

9*40 s

RECUAY

CATAC 9"50 S HUASCARAN NATIONAL PARK LA UNION ANCASH, PERU

Park Boundary 10*00 S Continental Divide Major Road Departmental Capital o Town ^ Major Mountain Peaks A CONOCOCHA Approximate Scale: CHIQUIAN LIMA 0k—25 km Adapted from ORDEZA (no date) and Diaz Bustos (1984)

Map 1. Huascarân National Park

4

The Park has considerable economic importance. It is the largest Peruvian National Park within easy access of Lima; the pleasant climate of the Rio Santa valley and the spectacular scenery attract a large number of tourists, domestic and foreign, to Huaraz and the adjoining Park, making tourism a major regional industry. The principal activities of the visitors to the Park are sightseeing, nature watching, hiking, and mountain climbing. Expeditions come from all parts of the globe to enjoy the world class climbing afforded by the many glaciers and peaks in the Park, of which 26 have an elevation of over 6000 m. Scientifically, the Reserve offers many possibilities for study in a variety of fields, for example, meteorology, geology, glaciology, botany, limnology, zoology, ecology, and park, range, and wildlife management. The Cordillera Blanca is interesting for a number of factors, including its high elevation, wide range of habitats and microsites, climatic extremes, biogeographical position and connections, and its size. Its possibilities as an outdoor laboratory are immense, and it is representative of much of the tropical Andes in terms of environmental conditions and biotic elements. There are several reasons for embarking on a floristic study of the Huascaràn National Park: — The Andes are the largest continuous mountain range in the tropics, and the only one which is oriented north-south. The continuity of the chain is offers excellent opportunities for plant migration, yet the highest elevation habitats are not physically contiguous. There is a delicate balance between the potential for dispersal and factors of isolation; it is an excellent natural laboratory for evolution. And, the isolating factors may be large scale, such as the Huancabamba depression (Berry, 1982), or small scale, such as the difficulties of gene flow between small populations separated by local topography. — The bulk of studies on high Andean plants have been done in the Andes north of the Equator; the Cordillera Blanca is in the Southern Hemisphere. — The Huascaràn florula is the only complete, recent study of the high elevation vegetation of Peru. It is representative of a large portion of the Peruvian high mountain region, and comparisons will now be possible between the floras of western Cordilleras of Peru and those of the eastern Cordilleras. This florula also offers a unique comparison with earlier floristic studies in the Andes, and with the floristic studies in other equatorial high elevation regions. 5

— Biogeographically the Cordillera Blanca is just between the southernmost extension of vegetation related to the paramos in the western cordilleras of the Andes and the northernmost extension of the puna. — A florula of the Huascarân International Biosphere Reserve has practical value as a basic tool in management and for future scientific research, and it is a logical extension of the Missouri Botanical Gardens' scientific activities in Peru. In late 1984, the Missouri Botanical Garden began a study of the flora of the Huascarân National Park, with the objectives of cataloguing the flora and, as such, creating a data base on the flora of the Park, creating an on-site herbarium as a resource for resource managers, scientists, and other persons interested in the flora, in-field training of Park personnel in field botany, and preparation of a manual to the flora. The field work was completed between December 1984 and August 1986, during which time 3843 specimen sets were collected, forming the base for the study of the Park flora. Herbarium study has been in progress since September 1986. The results of that study are reported here, as well as preliminary taxonomic studies in preparation for the manual of the flora. 6

GEOLOGY The Pacific Ocean is rimmed by a tectonically and orogenically active arc of mountains and volcanic islands (Zeil, 1979). The Andes are composed of several volcano-plutonic chains (Cobbing, 1978; Sillitoe, 1974), which extend along the western edge of South America from northern Venezuela and Colombia to southern Chile and Argentina, and are part of the circumpacific mountain system. This does not imply that the tectonic events and mechanisms are uniform throughout the circumpacific system, nor on a larger, global scale (Cobbing and Pitcher, 1972). At present there are three models for mountain building: collision of continental plates, continental uplift by an underthrusting oceanic plate, and the Andean model (Cobbing and Pitcher, 1972). The Andean model involves convergence of a continental and an oceanic plate and the subduction of the oceanic plate, but due to the relative thicknesses of the two plates, the thin oceanic plate is unable to bend or lift the thick continental plate (Cobbing and Pitcher, 1972; Myers, 1974, 1975). In the process of the Andean orogeny, the oceanic plate pushed against the continental South American plate, which fractured into narrow strips of crust parallel with the plate margin and these reacted tectonically by moving up and down along vertical fractures (Myers, 1974, 1975). The fractures represent major fault lines in the crystalline, Precambrian basement rocks, and possibly shear zones in the crust (Cobbing and Pitcher, 1972; Myers, 1975; Pitcher and Bussell, 1977). These ancient lines have determined the course of sedimentation, deformation, magmatism, and mineralization from the Mesozoic to the present (Myers, 1975; Pitcher and Bussell, 1977). Much is left to study regarding the orogenesis and tectonic history of the Andes, and not all students of the problem are in accord; James (1971, 1973) maintains that an underthrusting oceanic plate is causing the uplift. The Andes system is differentiated longitudinally into a series of parallel cordilleran systems of different ages, lithology, and history, and is divided along its length into distinct tectonic segments recognizable by surface and structural features, vulcanism, geophysical evidence, and tectonic boundaries (Hall and Wood, 1985; Sillitoe, 1974). The Central Andes of Peru between Huancabamba and Pisco are within a single, volcanically quiet tectonic segment (Hall and Wood, 1985; Sillitoe, 1974), and are divided into two Cordillera systems, the younger, western Cordillera Occidental and the older, eastern Cordillera Central, which are separated by the Rio Marafion. In Ancash, the 7

Cordillera Occidental is further divided into the Cordillera Negra, the Rio Santa valley, and the Cordillera Blanca (Wilson et al., 1967). Regional Geologic History This account follows Wilson et al. (1967), except when) noted. The Precambrian and Early Paleozoic strata were formed by marine géosynclinal deposition, which ended in the Early Paleozoic with a period of orogenesis, including intrusion of plutonic rocks and metamorphosis. The Andes have continued to be an active orogenic belt at intervals throughout Phanerozoic time (Cobbing, 1972). The period of mountain building was followed by a long period of erosion. In the Devonian, a series of major sedimentation basins were formed, controlled by the long established underlying fault structure and parallel with the directional trend of the Andes (Pitcher and Bussell, 1977). The Mississippian to early Permian was a period of deposition characterized by alternating marine submergence and continental emergence. The region was emerged and continental throughout the Pennsylvanian, and submerged and marine during the early Permian. Continental masses have been in movement relative to each other throughout the earth's history. At the end of the Palaeozoic Era, the palaeocontinents Laurasia and Gondwana came into contact forming Pangaea. This "supercontinent" lasted from the late Carboniferous to the early Jurassic. It was, however, not a static landmass, but in a process of constant change (Irving, 1977). Pangaea began to separate into North America, Eurasia, and Gondwana in the late Jurassic, and Gondwana began to separate into South America, Africa, Australia, Antarctica, and India about the same time as the separation of Pangaea. Historically, the western margin of West Gondwana, present day South America, was tectonically quiet (James, 1971). Beginning in the middle to late Permian, major tectonic activity began, and by the early Mesozoic Era the eastern Pacific subduction zone was functioning (James, 1971; Zeil, 1979). Cobbing (1976) places the beginnings of the eastern Pacific subduction zone in the early Jurassic. This was followed by elevation and continental deposition in the middle to late Permian. In the early and middle Triassic, the region was emerged slightly above sea level, and suffered little erosional destruction, becoming a stable erosion surface, which subsided to marine deposition in the late Triassic-early Jurassic. In the upper Jurassic, the region was emergent, although little above sea level, and lightly eroded. 8

During the Cretaceous, the major features of northern Peru were the Eastern and Western Peruvian geosynclines separated by the MaraAon geanticline. The Tapacocha fault is the axis and most important tectonic line of the Western Peruvian geosyncline (Myers, 1975), although Cobbing (1978) considers the Cordillera Blanca fault to be the most important, and the Rio Santa basin was a miogeosyncline on the Chavin crustal block between the Cordillera Blanca and Tapacocha faults (Myers, 1975). West of the Tapacocha fault, the underlying blocks subsided faster than the blocks east of the fault and a eugeosyncline formed over the Paramonga and Paracas crustal blocks (Myers, 1974, 1975). The Early Cretaceous was, largely, a period of subsidence (Myers, 1974), during which the western geosyncline was alternately slightly elevated and slightly subsided, a site for shallow water deposition and marine invasion. In the Early Cretaceous, sea floor spreading of the South Atlantic initiated (James, 1973) and West Gondwanaland began to separate into Africa and South America. By the late Cretaceous, the western geosyncline was uplifted, and the region has remained continental to the present. This initial phase of orogenesis completely disrupted the paleotectonic units. With uplift, the region went through a period of deformation, and was strongly folded by decollement (Myers, 1975) with the formation of several thrust fault systems. The deformation is especially evident in the Jurassic and Cretaceous sedimentary series. The mountain building events, which continue to the present, were initiated in the Tertiary with large scale block-faulting, uplift, and volcanic and plutonic igneous activity, including the formation of the batholiths of the western cordillera. The tectonic and magmatic events occured in stages, rather than continuously (Noble et al., 1974). The great Coastal batholith formed early in the period, and the Cordillera Blanca batholith much later (Cobbing, 1978). In the late Miocene-early Pliocene, the region was extensively eroded, and the "puna" erosion surface formed. A renewed period of uplift was initiated in the Pliocene and Pleistocene, and possibly as early as the late Miocene. The "puna" erosion surface was uplifted about 3000 meters to an average elevation of 4200 meters above sea level. This uplift was a two stage event; the first stage elevated the "puna" surface 1000 meters and initiated a period of deep erosion known as the valley stage, whereas the second stage elevated the "puna" surface an additional 1000-2000 meters, and initiated the canyon erosional stage, which was stronger than the valley stage. The effects of these two erosion stages are evident in the regional geomorphology. Since the "puna" erosion surface was extensively faulted, uplift was uneven and many blocks were raised to 9

greater heights. The Cordillera Blanca was raised about 2000 m heigher than the bulk of the "puna" erosion surface. The Andes were extensively glaciated in several events during the Pleistocene and Recent time; this is discussed in the section on glaciology. Sedimentary and Volcanic Stratigraphy The lowermost strata exposed within the Park are Jurassic in age, and the youngest are Quaternary and Recent sediments. The following is an overview of the geologic column, based on the works of Benavides-Câceres (1956), Coney (1970), Wilson (1963), and Wilson et al. (1967). The distribution of the formations is shown on Map 2. Upper Jurassic: Chicama formation (Js). A very thick formation with a complicated structure (regional decollement), composed of dark gray shales with bands of fine, silty sandstones, it was deposited under reducing conditions in a deep marine environment. Both the Jurassic and the Cretaceous were major sedimentary periods, and the weight of the sediments resulted in about 3000 meters of subsidence (Wilson, 1963). The Cretaceous rocks can be divided into a lower group that is basically non- marine, clastic sediments, the Goyllarisquisga Group, and an upper group of basically marine calcareous sediments, represented here by the Parihuanca Formation alone (Wilson, 1963). The clastic materials forming the Cretaceous sedimentary series probably came from the MaraAon geanticline. These sediments were strongly folded during the Tertiary. Lower Cretaceous: Goyllarisquisga Group (Ki). Wilson (1963) named this group, which includes the pre-Albian Cretaceous units from north and central Peru. Cobbing et al. (1981) found the concept useful, although they disagreed with his interpretation of the stratigraphie relationships, including only the Carhuaz and Farrat Formations in the group. This Group has three facies, an eastern, southern, and western, the western is the thickest and most varied. It is represented within the Park by four formations: (1) Chimu Formation: This formation is relatively thin (150-400 meters deep). The lower layers are composed of brown sandstones and quartzites intercalated with bands of dark gray, silty, carbonaceous shale and coal. The sandstone predominates over the shales in a ratio of about 10:1 (Coney, 1970). The coal is often present in thin bands, but can occur in veins up to 2 meters thick. The upper layers are grayish white quartzite. The formation was deposited in delta and floodplain environments. (2) Santa Formation; Also relatively thin, this has a lower layer of dark gray shales and an upper layer of sandy oolytic limestone. Deposition occurred in littoral and near shore marine environments. The strata weather to a brown color. HUASCARAN NATIONAL PARK GEOLOGY

Faults: thrust observed Inferred Approx. Scale: 0 25 Km Adapted from Instituto de GeologIa y MInerIa (1875)

QUATERNARY: O Qc Quaternary Sediments TERTIARY: B TsQv Yungay Formation B Ti Cordillera Blanca Batholith B T-v Callpuy Formation CRETACEOUS: EI KTI-C Huaylas Formation B Kms Parlahuanca Fonnation B M Qoyllarisqulsga Group JURASSIC: B Js Chicama Formation 2. Huascaràn National Park: Geology 11

<3) Carhuaz Formation: This is the dominant formation in the Group; it is up to 1000 meters in depth. At the base, there are bands of oolitic limestone 2-5 meters thick, some of these are gypsum-bearing. The formation is principally clastic, fine brown sandstone and quartzite intercalated with thin bands with black, gray, or yellowish red shales and siltstones. Some of the shales are carboniferous. The limestones are of shallow marine environments, and the clastic members were deposited in coastal swamps. (4) Farrat Formation: A very thin formation, 40-80 meters thick, the strata are fine whitish quartzites with intercalations of red shale. These are from delta and fluvial environments. Wilson (1963) was the first to recognize this as a formation; previously it had been mapped with the Carhuaz Formation. Lower Cretaceous: Pariahuanca Formation (Kms). Relatively thin, it is a fine gray limestone with a few bands of fine dark shale. The sedimentary environment was littoral to shallow marine. Upper Cretaceous: Huaylas Formation (KTi-c). This formation and several others throughout Peru, including the Chota Formation, are part of a group of mostly fresh water clastic strata with varing amounts of volcanic material included, which were deposited in the Late Cretaceous to Early Tertiary (Noble et al., 1974). Locally it can -be up to 350 meters thick, and is composed of gray conglomerate with volcanic fragments and intercalations of reddish sand and siltstones. The volcanic fragments are concentrated in the lower levels. Late Cretaceous-Earlv Tertiarv: Calipuy Group (T-v). Usually a thick accumulation (lOOO meters), this deposit is mixture of flows, brecchias, and tuffs of dacitic, rhyolitic, or porforitic andesites. The lowermost 100 meters are often a conglomerate. The stratigraphy of the group is poorly understood. Late Tertiarv-Early Ouaternarv: Yungay Formation (TsQv). This is a thin formation of light gray to white dacitic tuff of quartz crystals in feldspar matrix. Quaternary: Quaternary sediments (Qc). These deposits are very thick, and variable in nature and composition depending on their parent material and transport process. They include moraines, fluvio-glacial material, usually poorly sorted sands and gravels, alluvial deposits, and colluvial materials, especially mudflows (huaicos) and talus. Lacustrine deposits, glacial varves, are reported from near Vicos. Intrusive Igneous Rocks The outcrop of the Cordillera Blanca batholith (Ti) is 200 km long and 12-15 km wide. It is mostly coarse-grained granodiorite, with amphiboles and quartziferous 12

perfides in the Llanganuco valley. Relatively, the Cordillera Blanca batholith is very young. Intrusion of the batholiths in the Cordillera Occidental began 50-60 million years ago (James, 1973); radiometric dating of the Cordillera Blanca batholith places its age between 12 and 2.7 million years (Farrar and Noble, 1976). The main emplacement occurred earlier; the ages younger than 4 million years are likely cooling or uplifting dates, or more recent intrusive phases. Earlier dating produced age ranges that did not concur with the stratigraphie evidence. The recent age range places the origin of the batholith early enough in the Miocene to account for the truncation of the associated mineralization zone by the erosion environment which produced the "puna" erosion surface. The batholith was intruded along the Cordillera Blanca fault. Metamorphlc Rocks and Mineralization At the contact of the intrusive and sedimentary rocks, there is a zone of alteration about 1 km wide (Cobbing et al., 1981; Ricker, 1977; Wilson et al., 1967). The principal alteration is a hardening of the strata in the Chicama Formation, and closest to the contact metamorphosed rocks are present. Mica schists occupy the 100 meters closest to the contact, with andalusitic slates formed at the periphery. A zone of mineralization is present at the contact zone on the eastern flank of the batholith. Egeler and de Booy (1956) found thermal metamorphosis in the Cretaceous rocks in contact with the intrusive rocks. Economic Geology Within the zone of mineralization, there are deposits of metals: lead, zinc, and silver, and, in smaller quantities, copper, tungsten, molybdenum, and gold (Ricker, 1977; Wilson et al., 1967). These have been exploited since colonial times (Ricker, 1977). Currently the mines are not functioning due to the state of the world metal market and restrictions on mining within the National Park. The mines are located in Carhuazcancha, Honda, Manto Mina, Rurichinchay, and Safuna. Gypsum and limestone have occasionally been mined on a small scale, usually in the Rio Santa valley well outside the Park boundaries. The Chimû Formation contains exploitable quantities of anthracite coal, which is currently being mined within the National Park at Manto Mina, and nearby at Machac. The glaciers of the Cordillera Blanca have been a source of ice for the nearby towns for many years in the past, and continue to be such. The quarrying and cartage of ice is a cottage industry. 13

The most important geologic resource is hydropower. The runoff of the western side of the Cordillera Blanca powers several small hydroplants, as well as the large plant at Huallanca. There are mineral and thermal springs within and adjacent to the Park. Thermal baths are found at Cashapampa, Chances, and Monterrey, and cold-water mineral springs in Quebrada Honda and Rio Pachacoto. Glaciology There have been repeated glacial events in South America since the Late Pliocene; Clapperton (1983) states that van der Hammen and coworkers found evidence of 20-30 glacial events. These events are the glacial advances and interglacial periods of the four major glaciations recorded in the last 3.5 million years (Clapperton, 1983). There are very poor data on the older glaciations, and the evidence has been altered by succeeding glacial events; there may have been more than four, perhaps six. Considering the youth of the Cordillera Blanca, there is disagreement regarding the timing of the maximum glaciation in the range. Lliboutry et al. (1977) are of the opinion that maximum effect of glaciation occurred during the last glaciation, given that the recent uplift of the range would have been inadequate for effective glaciation before the last glaciation. However, the evidence reviewed by Clapperton (1983) indicated that the timing and magnitude of Pleistocene glacial events were synchronous and relatively equivalent throughout South America, and relatively synchronous with Pleistocene glacial events on a global scale. The best documented glaciations are the last and the penultimate, the latter occurring between 170,000-130,000 yr BP. Radiocarbon dates from the Quelccaya ice cap in southern Peru place the last major glaciation between 28,000 and 14,000 yr BP (Mercer and Palacios, 1977). Clapperton (1972) found few glacial features related to that glaciation in the Cordillera Blanca. The most extensive advances occurred early in the period, between 45,000-36,000 yr BP and 28,000-20,000 yr BP. Other advances were recorded at 20,000-19,000 yr BP, 16,000-14,500 yr BP, ca. 13,000 yr BP, and 11,000- 10,000 yr BP (Clapperton, 1983). The maximum of glaciation in the tropics was reached at 15,500-14,000 yr BP. During the Holocene, the dynamism of glacial advance and recession continues. Since the last major glaciation, there have been advances in 8000-5000 BP, 4700-4200 BP, 1300-1100 BP, and in the past 400-500 years. These advances are "Little Ice Age" 14

events, part of a period from ca. 6500-600 BP. The advance of 1300-1100 BP coincides with the alpine advance in the Medieval times. In this century, the glaciers in Peru have been receding since the late 1920s (Clapperton, 1983; Wilson et al., 1967), during which time the snowline rose 100-200 meters in altitude. The rate of recession has declined since the 1960s. Kinzl (1968) found that the fluctuations of the Andean glaciers coincide with the fluctuations of the Alpine glaciers. There is no evidence of the existence of an ice cap in northern Peru. All the northern glaciers have formed in cirques (Clapperton, 1983), and these flowed out of the high valleys of the Cordillera into the major river systems to the east and west. The lowest altitudes reached by glacial tongues are marked by terminal moraines at 2700- 2900 m in the west and 3200-3600 m in the east (Wilson et al., 1967). Given the very steep altitudinal gradient on the western flank of the Cordillera Blanca, the major erosive power is on that side (Wilson et al., 1967), hence the unusual phenomenon that the cirques of the major glaciers flowing to the west are located to the east of the line of summits on the batholith. Wilson et al. (1967) observe that the path of the glaciers followed the major fracture lines in the batholith. Despite the general recession of glaciers, there are still many active glaciers in the Cordillera Blanca. The snouts of several of these are completely covered by rock and debris, and function somewhat like rock glaciers (Lliboutry, 1977). Throughout the Park glacial and periglacial structures are prominent. In the northern part where the intrusives and the altered sedimentary rocks are resistant, there are many deep, U-shaped glacial valleys: Llanganuco, Parôn, and Santa Cruz being outstanding examples. Several of the higher peaks are glacial horns, and throughout the park there are many areas of scoured rock surfaces. Moraines of many different sizes and several different glacial advances can be seen in all parts of the Park, as can massive fluvio-glacial deposits. The lakes of any size are either formed in abandoned cirques or behind terminal moraines. Given the extent of glaciation in the Park, it is probable that there was permafrost in the immediate periglacial environment; the only evidence of this is patterned ground that can be seen in the bottoms of some valleys in areas now covered by bogs or marshy ground. Soils To my knowlege, there are no detailed studies of the high Andean soils. Drosdoff et al. (1960) describe the soils as loams of varying depth, usually with 15 abundant organic matter, or as having rudimentary, if any, development. Seek and Bramao (1968) found the high Andean soils of variable texture depending on the nature of the parent material. Within the study area, most soils are poorly developed, and, where profile development has occurred, the soils are rocky. The texture of the non- rocky fractions is fine. Most soils are loams and silty loams; clayey soils are rare and localized. Within the Park, the soils are developed on several different parent materials. Those on old morainal material can develop a deep profile, but are quite pebbly or bouldery. Soils on younger morainal material are less developed. Many soils on slopes are formed on a thin layer of weathered material overlying rock; these soils are usually well-developed, with a very shallow profile, and can be extremely drouthy. Soils of the valley bottoms, boggy areas, or depressions are usually deep, rich in organic matter, and saturated. At the higher elevations, the parent materials are little, if at all, modified. In the north of the Park, most of the parent material is a coarse granodiorite, and in the south, a fine-grained colluvial material derived from sedimentary rocks. All the soils in the reserve, especially those of boggy areas, can have relatively high levels of organic matter, which accumulates due to the low rates of decomposition at the high altitudes. Drosdoff et al. (1960) found the soils to be moderately to strongly acid, and with low levels of available nitrogen and phosphorous. The nutrient availability is influenced by the low rates of decomposition, and the low environmental temperatures. At the high elevations, freezing temperatures can be reached any night of the year, and in the presence of adequate moisture, needle ice will form in the soil (Lawler, 1988; Perez, 1984). This phenomenon is widespread throughout the world (Lawler, 1988), and can be encountered at high elevations at low latitudes, and low elevations at high latitudes. It is most commonly seen in high mountain environments. In southern Peru, Hastenrath (1977) encountered the zone of needle ice from 4,400 m upwards. In the Huascarân National Park, it is encountered at similar elevations (D. N. Smith, pers. obs.). Needle ice is an aggregation of slender, needle-like, vertically oriented crystals which form at and below the soil surface. The crystals can penetrate 5-9 cm into the soil at 4,300 m, and perhaps more at higher elevations (Perez, 1984). As the crystals grow, they lift the soil particles vertically; when the crystal is stressed (either by the load in relation to the length of the crystal, or in the process of thawing) it bends down 16

slope, thus transporting soil particles down slope. The particles may be transported further down slope by sliding along the saturated soil surface in the final stages of thawing (Higashi and Corte, 1971). Although the magnitude of the individual events are small, the events are very numerous, and the frequency large; the effects of needle ice contribute a large component to soil creep and mass wasting, thus needle ice is a very important in geomorphic process. Soil texture and soil moisture affect needle ice formation. Adequate moisture must be present or ice cannot form; in the presence of adequate moisture needle ice can form in coarse or fine soils (Lawler, 1988; Perez, 1985, 1987b). But soils which are of very coarse or very fine texture will inhibit needle ice development; soils of medium texture with high levels of organic matter or silt favor needle ice development (Lawler, 1988; Perez 1985, 1987b). In addition to the down slope movement of soil particles, the freezing and thawing sorts the particles by size fraction (Lawler, 1988; Corte, 1968); the sorting action of the ice moves the coarse and fine fractions differently (Lawler, 1988). As the freezing front progresses through the soil, the smaller fractions are pushed before the front and the coarser left behind (Corte, 1968). Sorting can produce soil patterns, including polygons and stripes (Lawler, 1988; Perez, 1984). Although the ice mobilizes mostly smaller particles, even stones of several kilograms in weight can be mobilized (Perez, 1988). The biological significance of needle ice is that it disturbs the soil by keeping it in constant movement. Colonization and seedling establishment are inhibited, and the plants which can colonize needle ice zones must have adaptations that allow them to withstand the movement, or must select microsites which protect them from the soil movement (Lawler, 1988; Perez, 1987a). Within the needle ice zone, the action of the ice can significantly affect the distribution of plants in general and, more specifically, that of certain species (Perez, 1987a). 17

CLIMATE The placement of the Andes is perpendicular to the easterly trade winds which bring the weather systems to tropical South America. As the air streams are forced to rise over the Andes, many different climatic situations are created (Johnson, 1976; Sarmiento, 1986). General Circulation The weather systems of tropical South America are controlled by the tropical easterlies, and oscillations of the ITCZ, the Intertropical Convergence Zone (Gomez- Molina and Little, 1981; Johnson, 1976). The ITCZ shifts northward in June and July and southward in December and January, and with it carries moisture laden air masses to the north and south, causing the tropical rainy seasons (Gomez-Molina and Little, 1981). At irregular intervals, the ENSO, El Niflo-Southern Oscillation, phenomenon occurs (Crane, 1986; Ramage, 1986). This phenomenon is an interruption of the normal atmospheric and oceanic circulation in the Pacific, and is linked to worldwide climatic anomalies and attendant biological consequences (Rasmussen and Wallace, 1983). During an ENSO, the easterly trade winds weaken (Johnson, 1976); studies from the Quelccaya Ice Cap in Peru show a significant decrease in precipitation for the eastern cordillera during an ENSO (Thompson et al., 1984). The situation in the western cordilleras seems to differ from that of the eastern cordilleras. The precipitation and temperature data from the Huascarân National Park are quite variable from year to year; cursory examination of the data show no marked effect in the ENSO years. Evidence from population studies of Andean condors during ENSO years showed no significant effects at the high elevations (Wallace and Temple, 1988) Seasonality Tropical climates are characterized by a constancy of solar radiation, day length, and mean temperature, but have a highly seasonal pattern of cloudiness and precipitation (Sarmiento, 1986; Smith and Young, 1987). While the annual variability of monthly mean temperature in the tropics is low, the variability of the daily temperature cycle can be very large (Johnson, 1976; Sarmiento, 1986). The daily variation increases with altitude, and is affected by cloud cover and humidity (Johnson, 1976). The annual weather patterns in the Huascarân National Park are shown by the Walther diagrams in Figure 1. 18

QUEROCOCHA* (3980)'* 7.3"® 858'' SAFUNA (4275) 5.7® 1398

20.8" 14j' 11.4

12.20 9.8

0.4" -0.8 iiiiiiiiiiiiiiiMiii'iiiniiiiiiiiiiiiiiiniMtiii IllllllUllhlinilllllllllllllllllUlll 9®43'Sl 8® 49'S

LAMPAS ALTO No. 2 (4030) 8® 737

18.8 13.8

13.4

"3.9 -9.0 iiniiiliiiliiiliii 10® ors

® station; ^ elevation (m); ® mean annual temperature ("C); ^ total annual precipitation (mm); ® record maximum temperature; ^ mean maximum temperature; ® temperature range; ^ mean minimum temperature; * record minimum temperature; j latitude. The divisions on the ordinate are lO'C or 20 mm of precipitation. The precipitation curve is barred above the temperature curve, and precipitation in excess of 100 mm per month is graphed at a scale of 1/10 and shaded black. The precipitation curve falls below the temperature curve in months with a water deficit; this is shaded with dots. The abscissa is divided into months, July - June; January is the center line. The black bars below the abscissa indicate months with a mean minimum temperature below O'C, and broken bars months with an absolute minimum temperature below 0°C (Walter, 1973). Figure 1. Huascarân National Park: Climate diagrams 19

There are 10 recording stations nearby or in the Park. The stations at Lampas Alto, Querococha, and Safuna record temperatures, precipitation, hours of day light, humidity, evaporation, and wind velocity; the remaining stations record only précipitation. Currently, ElectroPeru, a state-owned electric company, operates the stations. All data reported here are taken from the unpublished weather records of the Huaraz Office of ElectroPeru, except for the data from Safuna, a station which is no longer operative; those data were published by ONERN (1972). Map 3 shows the location of the recording stations referred to herein. It is important to note that there is no station on the eastern side of the Cordillera Blanca, which is on the windward side of the range. Day Length and Hours of Daylight At the latitudes of the Park, the day length varies from 11.S hours on June 21 to 12.5 hours on December 21. Topography may cause shadowing of some areas up to 2.5 hours after sunrise or before sunset. The daylight data from the Park were collected with Campbell-Stokes heliometers, and are reported as total hours of daylight. This limits the usefulness of the data, as distribution of the cloudy periods throughout the day is important, especially for appraising the magnitude of the topoclimatic effects. Since it does not add significant information, the daylight duration data will not be reported here. Solar Radiation With increasing altitude, the mass of the atmosphere decreases, as do its scattering and absorptive properties (Tranquillini, 1964). The dense, sea-level atmosphere screens out the short wave length spectra, the blues and ultra-violets, but, at high elevation, these spectra are more intense (Tranquillini, 1964). There is controversy regarding the magnitude of the increase in ultraviolet radiation; Caldwell (1968) found the increase small (4-26%), while other authors found increases of up to 120% (Barry and van Wie, 1974). Long-wave emission, infra-red, is little effected by altitude, as the temperature of the emitting surface is cooler and there is little back radiation from the thin atmosphere (Barry, 1981; Geiger, 1969; Tranquillini, 1964). Changes in the radiation balance are due to the increased solar radiation (Geiger, 1969). Radiant flux has not been recorded or studied at the Huascaràn National Park, or any nearby high elevation site. At high elevations, the highest radiant intensity is encountered under partially cloudy conditions, when there are bright, highly reflective clouds at high elevations. In 20

SAFUNA(427Sm) 1400 mm HUASCARAN NATIONAL PARK CUMATE ANNUAL RAINFALL Approx. Scale:

J 25 km

PARON (4188 mj, 830 mm

LLANQANUCO (3850 m) 840 mm

CAHUISH (4550 m) QUEROCOCHA (3980 m) +1 970 mm 960 mm

PACHACOTO (3570 m) 595 mm

RECRETA (3990 m) ^^NACOCHA (4500 m) 470 mm 885 mm

t^^UNTA MOJON (4395 m) 675 mm

LAMPAS ALTO (4030 m) 740 mm

Map 3. Huascaràn National Park: Climate and rainfall 21

these situations, the radiant intensity may exceed the solar constant (Geiger, 1969; Tranquillini, 1964). Temperature The monthly mean temperatures at 4000 m are 6-7.5'C, with the warmer temperatures in the northern part of the park, and the cooler in the southern part. The altitude of the highest recording station is 4275 m; there are no observed data for the higher elevations. Using an estimated lapse rate of 6.S'C/1000 m (Johnson, 1976), the calculated mean monthly temperatures for 4500 and 5000 m are 3-4*C and 1-0'C, respectively. The daily temperature variation at 4000 m averages between 10-13.5''C, at standard observation height; the range at the soil surface is much greater (Hedberg and Hedberg, 1979). The range of variation, and the temperature extremes are greater during the dry season; during the rainy season the cloudiness and positive water balance dampen the daily temperature range and the extremes. Monthly mean, mean minimum, and mean maximum temperatures are shown in Table 1. At elevations of 4000 m and above, the absolute minimum temperature will be below 0*C all months of the year, and frost can occur any night of the year. The frequency and severity of frost increase with increasing elevation. The frost is formed by radiative cooling under clear sky and calm conditions (Geiger, 1964), and is most severe on ridges (Troll, 1968). During the night, cold air will flow down slope and pool in the valley bottoms or behind barriers (Geiger, 1964; Troll, 1968), thus the ridges and the valley bottoms are the coldest topographic features, and tend to have frost before the slopes. The slopes may remain frost free. In the field, I have observed needle ice formed at the soil surface and hoar frost on plant surfaces; both are formed by radiative cooling. Frosts and frozen precipitation seldom last more than a few hours; but, I have seen needle ice on a southern aspect last until 2:00 p.m. by which time the slope was shaded for the rest of the afternoon, and I assume the frost lasted for longer than a 24 hour period. In the equatorial high altitudes, radiative cooling is a relatively short term phenomenon; the cooling period is rarely more than about 14 hours. During the field work, I never saw a deep, hard freeze that suggested formation by cold-air advection. Table 1. Mean monthly, mean minimum, and mean maximum temperatures, and mean percentage of nights with absolute minima below 0°C (Safuna 2 yrs. of observation, Querococha 32 yrs., Lampas Alto 28 yrs.)

MEAN MONTHLY TEMPERATURE Station Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Ave® Abs®

Safuna 5.9 6.0 5.8 6.2 6.0 5.4 4.9 5.5 5.7 6.1 5.7 5.5 5.7 Querococha 7.1 7.0 7.2 7.4 7.5 7.4 7.1 7.5 7.5 7.4 7.5 7.3 7.3 Lampas Alto 6.1 6.0 6.2 6.3 6.0 5.8 5.8 5.9 6.0 6.0 6.0 6.1 6.0

MEAN MONTHLY MINIMUM Safuna 1.2 1.6 1.5 1.9 1.1 0.6 -0.6 -0.1 0.3 1.0 0.8 0.9 0.8 Querococha 2.2 2.5 2.6 2.4 1.6 0.8 0.4 0.6 1.2 1.6 1.5 1.8 1.6 -4.5 Lampas Alto 1.3 1.8 2.0 1.2 -0.7 -3.1 -3.9 -3.2 -1.5 -0.4 -0.4 0.3 -0.6 -9.4

MEAN MONTHLY MAXIMUM Safuna 10.6 10.4 10.3 10.4 10.9 10.3 10.4 11.4 11.2 11.2 10.6 10.2 10.6 Querococha 13.2 13.0 13.0 13.6 14.0 13.9 13.8 14.5 14.4 14.3 14.2 13.7 13.8 20.8 Lampas Alto 12.3 11.9 12.2 12.6 12.9 13.0 13.0 13.6 13.6 13.4 13.0^ 12.8 12.8 18.8

MEAN PERCENTAGE OF NIGHTS WITH ABSOLUTE MINIMUM BELOW 0°C Lampas Alto 20 9 8 20 61 88 92 87 77 49 48 34 Querococha 4 3 3 4 9 21 34 29 13 11 12 9

^Ave - long term annual average; Abs = absolute minimum or maximum temperature. 23

Precipitation The sources of the precipitation falling on the Huascarân National Park are moisture carried from the Amazon basin by the tropical easterlies and clouds formed locally by heating of slopes (Johnson, 1976). The lowland tropical areas within the influence of the ITCZ, that is within about lO^ of the Equator, have two rainfall maxima per year, whereas the tropical highlands influenced by the trade winds have one maximum (Sarmiento, 1986). The rainy season in the Cordillera Blanca lasts 5.5 months, from late November to April. March is the wettest month, and July the driest. Table 2 shows the monthly and annual averages, and Table 3 the monthly distribution of the precipitation expressed as a percent of the annual total. Based on the data from the Safuna station and the recording stations located within the Park boundaries, the precipitation ranges from 640-1400 mm per year, and from 70 to 80% of the annual total falls during the rainy season. The northern part of the Park recieves more precipitation than the southern part. The dry season is stronger in the southern part, that is that a higher percentage of the annual precipitation, closer to 80% or even slightly more, falls during the rainy season. Using the vegetation as an index, it is probable that the eastern side of the Park has the highest precipitation; unfortunately, there are no data to confirm this. The monthly and annual precipitation is quite variable from year to year. Johnson (1976) attributes a large part of this variability to the the important contributions of local convective development and orographic precititation to the annual total at the high elevations. These produce very localized precipitation. In general terms, on mountain ranges rainfall increases with increasing altitude up to the middle elevations, then decreases with altitude (Flohn, 1974; Sarmiento, 1986). The recording stations within the Park, all of which are on the western side of the Cordillera Blanca, show an increase of precipitation with altitude. This situation may not be unusual, Tranquillini (1964) reported an increase in precipitation with altitude for high elevation sites in general, and Johnson (1976) reported the same for the Peruvian Andes. Other local factors may affect this increase. Although there are no recording stations on the eastern side of the Cordillera Blanca, the vegetation suggests that the eastern side receives a higher rainfall than the western side. As the fronts approach from the east, rainfall is produced orographically as the cloud masses rise over the Table 2. Precipitation (mm)

Station^ Jan Feb Mar Apr May Jun Jul Aug Sep Oct NOT Dec AVEB

Safuna 149.6 147.0 207.2 163.5 81.4 40.0 15.1 30.1 53.8 103.3 186.2 218.7 1395.9 Parôn 117.7 141.8 146.5 79.5 34.1 10.6 3.6 10.9 38.7 70.5 80.5 95.3 827.2 Llanganuco 93.7 116.6 137.6 67.2 18.8 1.4 1.4 5.6 20.7 47.0 58.9 72.9 641.8 Querococha 150.3 164.4 161.8 95.0 39.1 8.7 4.5 9.5 32.9 85.4 82.2 123.9 957.8 Cahuish 132.5 156.4 161.0 111.7 56.2 11.1 4.5 11.8 43.9 86.5 84.8 115.6 970.6 Pachacoto 98.3 106.6 134.0 62.5 21.8 1.2 0.5 2.6 13.1 41.6 47.5 65.0 594.5 Recreta 77.7 97.7 117.4 47.9 13.5 0.6 0.5 2.5 8.1 22.6 30.7 50.6 469.8 Yanacocha 135.3 171.4 195.3 92.7 36.9 5.8 1.9 6.5 22.8 56.5 64.9 95.0 885.0 Punta Mojôn 108.7 140.6 154.6 71.1 21.1 3.9 0.8 3.5 14.9 37.9 44.3 75.4 676.8 Lampas Alto 119.6 141.1 161.4 70.8 21.4 4.4 2.8 5.8 18.9 53.6 56.8 82.9 737.4

\ears of observation: 33, except Safuna (2), Lampas Alto (28), Querococha and Llanganuco (32), and Parôn (37). ''long term average annual precipitation. Table 3. Monthly distribution of precipitation (% of annual total)

Station^ Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Rainy^

Safuna 10.7 10.5 14.8 11.7 5.8 2.9 1.1 2.2 3.9 7.4 13.3 15.7 63 Pardn 14.2 17.1 17.7 9.6 4.1 1.3 0.4 1.3 4.7 8.5 9.7 11.5 70 Llanganuco 14.6 18.2 21.4 10.5 2.9 0.2 0.2 0.9 3.2 7.3 9.2 11.4 76 Querococha 15.7 17.2 16.9 9.9 4.1 0.9 0.5 1.0 3.4 8.9 8.6 12.9 73 Cahuish 13.7 16.1 16.6 11.5 5.8 1.1 0.5 1.2 4.5 8.9 8.7 11.9 70 Pachacoto 16.5 17.9 22.5 10.5 3.7 0.2 0.1 0.4 2.2 7.0 8.0 10.9 78 Recreta 16.5 20.8 25.0 10.2 2.9 0.1 0.1 0.5 1.7 4.8 6.5 10.8 83 Yanacocha 15.3 19.4 22.1 10.5 4.2 0.6 0.2 0.7 2.6 6.4 7.3 10.7 78 Punta Mojôn 16.1 20.8 22.8 10.5 3.1 0.6 0.1 0.5 2.2 5.6 6.5 11.1 81 Lampas Alto 16.2 19.1 21.9 9.6 2.9 0.6 0.4 0.8 2.6 7.3 7.7 11.2 78 Average® 15 18 20 10.5 4 0.8 0.4 0.9 3 7 8.5 12 75.5

^ears of observation: 33, except Safuna (2), Lampas Alto (28), Querococha and Llanganuco (32), and Pardn (37). ^percent of precipitation falling in the rainy season. ^monthly precipitation expressed as percent of annual total. 26

eastern side of the Cordillera Blanca. Some of this rain must pass over the crest of the Cordillera and result in higher precipitation at the higher elevations. Although there are frontal storms passing aloft throughout the rainy season, the daily pattern of cloudiness and precipitation is more closely linked to diurnal convective activity than large scale fronts. The mornings are clear, by mid to late-morning local heating begins convective cloud build-up, and rains begin in the afternoon and may continue until evening (Johnson, 1970, 1976). Since the local clouds are of convective origin, there are usually some clouds all year round, but in the dry season the build-up is small and little precipitation is produced. Coile and Culmsee (1953) and Johnson (1970) found the thunderstorms concentrated in the rainy season; there are very few in the dry season, with the minimum being in July. Coile and Culmsee (1953) indicated 3-4 pm as the peak hours for thunderstorm development; my experience is similar. Since orographic precipitation and thunderstorms are common in the Andes, rainfall can be very localized (Johnson, 1976). In the case of strong development of thunder storms, the afternoon storms are often violent, producing heavy rainfall with thunder and lightning, and the evening rains more gentle. Precipitation may fall as rain, snow or graupel. At the lower elevations, rain is the most common form, but above 4600 m it becomes rare (Johnson, 1976). At the higher elevations and in the southern part of the Park, snow and graupel are common. The frozen forms of precipitation are transitory, rarely lasting more than a few hours or overnight. Relative Humidity Absolute humidity (vapor pressure) decreases with altitude, and at the same time, with the altitudinal decrease in temperature, saturation vapor pressure decreases, thus the relative humidity increases and the saturation deficit (evaporative power) decreases (Tranquillini, 1964). Although the increased radiation and wind velocity and lower atmospheric pressure should increase evaporation (Hodge, 1946), studies show evaporative rates at high elevations to be little different from those of the lowlands (Tranquillini, 1964). The mean relative humidity is in the range between the low SO and high 70 percentiles; the lowest relative humidities occur duing the dry season and the highest during the wet (Table 4). Table 4. Average relative humidity and average wind speed

AVERAGE RELATIVE HUMIDITY (%) Station^ Jmn Feb Mar Apr May Jun Jul Aug Sep Oct No? Dec

Safuna 66 70 62 69 60 58 53 52 58 61 63 66 Querococha 69 72 71 67 62 58 57 58 60 64 63 66 Lampas Alto 76 78 77 72 65 59 56 58 61 66 66 71

AVERAGE WIND SPEED (m/sec) Querococha 4.9 4.6 5.0 5.3 5.2 5.7 6.5 6.3 6.1 5.3 4.8 4.7 Lampas Alto 3.4 3.3 3.3 3.2 3.1 3.4 3.6 3.6 3.8 3.6 3.6 3.5

^years of observation: Safuna (2), Querococha (32), and Lampas alto (28). 28

Wind Surface winds in the Andes follow a typical valley wind regime (Johnson, 1976). This is produced by the diurnal heat balance of the slopes. During the day the slopes are differentially heated and up-slope winds created by rise of the buoyant, heated air. The process is reversed in the night when the air cools by long-wave radiation and winds are created by down slope flow of the dense cold air (Flohn, 1969). Mean wind speed data for Lampas Alto and Querococha are included in Table 4. The speeds for Querococha are considerably higher than those of Lampas Alto. The Querococha station is located on a slope in a relatively narrow valley, while the Lampas Alto station is on a more open site. TopocUmates The heat and water balance on slopes is different than those of horizontal surfaces (Geiger, 1969). Slope angle and aspect will affect the amount of radiation received, hence the temperature and moisture regimes of the slope (Geiger, 1964, 1969; Sarmiento, 1986). Topography will affect flow of water and heated or cooled air, and will influence shading of slopes. The varied topography of the Huascarân National Park creates a complex mosaic of topoclimates and microhabitats. The enviromental conditions of the topoclimates and microhabitats are in a state of constant change, and often change rapidly and markedly with respect to the adjoining topoclimates (Sarmiento, 1986). The daily cloud pattern creates a topoclimatic difference between the east- and west-facing aspects. The east-facing aspect receives solar radiation in the morning before the daily cloud build-up, but the west-facing aspects receive direct insolation later in the morning and the insolation is reduced by the afternoon clouds. Thus the east-facing slopes are warmer, drier, and have greater temperature extremes (Sarmiento, 1986; Smith and Young, 1987; Smith, 1980). The higher wind speeds at the Querococha station are the result of topoclimate. These are two examples among many of the effects of topoclimates. 29

VEGETATION OF THE HUASCARAN NATIONAL PARK Life Forms of High Andean Plants Throughout the tropics where high mountains occur, a set of peculiar plant life forms has evolved (Cuatrecasas, 1968; Hedberg and Hedberg, 1979; Rauh, 1978; Smith and Young, 1987) including: giant rosettes (or caulirosettes), sclerophyllous shrubs, cryptofrutices, tussock grasses, acaulescent rosette herbs, and cushion plants. These tropical high elevation life forms are an excellent example of convergent evolution, having evolved independently in different taxonomic groups on widely separated continents (Hedberg and Hedberg, 1979; Smith and Young, 1987). The repeated evolution of these life forms reflects their unique adaptation to the difficulties of the equatorial high mountain climate (Smith and Young, 1987). The nature of the adaptation to the climate has been, and continues to be, the focus of research for many of the students of the Andean type vegetation. A review of the life forms found in the Park follows: Giant rosettes (caulirosettes): These are large, erect herbs with the leaves arranged in a rosette. Although some species do not develop an elongated stem, the stems of many species are elongated and have a thin woody outer layer surrounding a large pith. The rosette of leaves is borne at the apex of the stem, and, following leaf death, the stem becomes sheathed by the marcescent leaves. The inflorescences are terminal or axillary, and well exserted. In the Americas, this life form is typical of the paramo and ceja de selva. Due to its dryness, there are few species in the Park that are clearly caulirosettes. Blechmm loxense, Puya raimondii and Nicotiana thyrsiflora most closely resemble this life form. Sclerophyllous shrubs: The general shape of the plants and the woody stems have a normal shrub-like appearance, however the leaves are highly modified for high stress conditions. Leaf modifications include: small size, coriaceous texture, revolute margins, unusual orientations, thickened epidermis on one or both faces of the blade, thick cuticles, pubescence of various types, various types of glands, and unusually high lipid contents in the leaf tissue. Cuatrecasas (1968) considers the plants of this life form to be short-lived. Many of the shrubs in the Huascaràn flora are of this type, especially species of Baccharis and Senecio. Cryptofrutices: Cuatrecasas (1968) introduced the term cryptofrutex for the dwarf shrubs that are prostrate on, and often immersed in, the soil. Baccharis caespitosa and several species of Astragalus are examples of this life form. 30

Tuisock graues: These are bunch grasses, in which the caespitose habit is taken to extreme development in size and density of shoots. The individual clumps can be very large, often nearly a meter in diameter, the crown widely spreading and hemispherical in shape, much of the foliage dead and marcescent, and the bases of the shoots clothed in a tunic of dead and decomposing leaf bases. The foliage is modified to withstand the stresses of a moisture deficient and frost-prone environment. They are often highly sclerified, with a thick cuticle, and the blades are tightly rolled with a sharp tip. The majority of the grasses in the study area are smaller and of a more typical bunchgrass appearance. They too have a protective tunic of dead leaf bases around the base of the plant. This life form is widespread in the temperate Southern Hemisphere. Acaulescent rosette herbs: Stem development is suppressed in this life form; the stem is usually present as a caudex, with the rosette of leaves arising from the apex. The leaf position varies from erect to prostrate, and the texture and indûment of the blade is also variable. The scapes or peduncles of the inflorescence may be elongate or entirely suppressed, and, if elongate, erect to prostrate. Many of the herbs in the flora are of the rosette type. The life form becomes more common with increasing altitude. Herbs which are neither rosette nor cushion-forming, are often found in dense clones with short-creeping rhizomes and erect leaves. This is especially common at the higher elevations. Cushion plants: Varying in size from a few centimeters to meters in diameter and from a few centimeters to nearly a meter in height, the flattened to hemispherical cushion is composed of densely packed and interwoven stems. Each stem has a rosette of leaves at the apex, and the older leaves remain within the cushion, decaying to form a peaty interior (Heilborn, 1925). The shoots can be so densely packed that the cushion surface is hard and unyielding, even to strong blows. The inflorescences of these species may be exserted or sessile and protected by the rosette of leaves. This life form becomes more common at higher altitude, and can be found in both dry and boggy microsites. Aciachne pulvinata, Azorella spp., Distichia muscoides, Mmdes spp., Nototriche spp., Oreobolus obtusangulatus, Plantago rigida, Plettkea cryptantha, Pycnophyllum spp., Stangea erikae, and Werneria spp. all exhibit this life form. Trees: Several species of trees are present in the Park: Alnus acuminata, Buddleia incana, Gynoxys oleifolia, Polylepis sericea, and P. weberbaueri, and, at times, Myrica pubescens, Weinmannia aff. laxiflora, and Vallea stipularis become tree-sized. These usually have 31

a contorted growth form, are of relatively low stature, and grow in very specific microsites. The epiphytic and lithophytic habit is developed in several species in different families of the Park flora. The features of epiphytism are reviewed in papers by Benzing (1986, 1987) and Madison (1977). Gigantism is also expressed in the Park flora; Krap/ia weberbaueri, Nicotiana thyrsiflora, and Puya raimondii are all exceedingly large representatives of their genera. Environmental Factors and Plant Adaptations Altitudinal ilmlti to niant growth Plants can grow in suitable microsites well above the snowlines in mountain areas; Webster (1961) found 5944 m to be the highest documented altitudinal record of plant growth. Growth of plants in high mountains is limited by lack of soil moisture and/or cold (Halloy, 1981; Perez, 1987a; Webster, 1961). It has been suggested that the only absolute altitudinal limit to plant growth is set by low partial pressures of CO2, however this factor is not limiting until well above the limits set by the water and thermal balances (Gale, 1972; Halloy, 1981). Intense «olar radiation As the mass of the atmosphere decreases, the radiant intensity increases (Tranquillini, 1964). It has been suggested that light colored or appressed pubescence may reflect part of the incident radiation (Johnson, 1975; Smith and Young, 1987), but the results of several studies have given contradictory results. Meinzer and Goldstein (1986) observed that the thermal balance is critical at high elevations, that is, despite high radiant inputs the thermal inputs are low. This suggests that the plants need not adapt to avoiding radiant input, but rather to increase heat capture. Given the high levels of radiant flux, the amount of radiation lost to reflectance may not have any significant consequences. Leaves which are glabrous above and pubescent below may reduce the absorptivity below and increase skyward emissivity (Johnson, 1975). If the solar radiation load is excessive, these adaptations could effectively reduce it. Many species in the Huascarân flora have this pubescence pattern. In addition, there is evidence that high altitude plants have adapted to high radiant intensity and can utilize the intense radiation effectively in photosynthesis (Tranquillini, 1964). At high elevation in the lower latitudes, this increase in radiant intensity is accompanied by a marked increase in the Ultraviolet-B radiation. This is because the 32

global ozone envelope is narrowest at the Equator, the path length the shortest and the optical amplification of the radiation the greatest (Caldwell et al., 1980). UV-B can interfere with shoot elongation, destroy chlorophyll and tissue, and interfere with flowering (Caldwell, 1968; Tranquillini, 1964). Lee and Lowrey (1980) suggest that UV- B could increase rates of speciation by increasing the mutation rate. Robberecht et al. (1980) found species native to regions of high UV-B radiant flux densities better able to attenuate the UV-B spectra in the leaf cuticle and epidermis than species from regions with low UV-B radiant flux densities. The UV-B flux at the leaf mesophyll in equatorial, tropical, and arctic latitudes were very similar, even though the flux density at the leaf surface is far greater in the equatorial latitudes. The UV-B spectra can be reflected at the leaf surface or attenuated within the leaf. In some cases, pubescent or glaucous leaf surfaces were found to reflect the spectra, in others no reflectance could be shown. Alkaloids and pigments (flavonoids and phenolics) were found to absorb UV- B. Lee and Lowrey (1980) suggest that thick cuticles, waxy coatings, and anthocyanin concentrations also attenuate UV-B. Tmnnivn Andean plants are faced with three significant environmental temperature regimes: air, soil surface, and soil. Soil temperature is stable, not characterized by the oscillations of either the air and soil surface, yet is so low that transport or translocation processes occur at suboptimal rates (Meinzer and Goldstein, 1986). The air and soil surface temperatures are highly variable on both a short term and diurnal basis. Nocturnal air temperatures can drop to levels that require physiological adaptation to frost and freezing. The temperature variation at the soil surface is great; the diurnal range can be in excess of 47*C (Hedberg and Hedberg, 1979). The life forms previously discussed are adapted to these temperature stresses. In the giant rosettes, sclerophyllous shrubs, tussock grasses, rosette herbs, and cushion plants, Hedberg and Hedberg (1979) found that the growth points were insulated from the nocturnal ambient air temperature or bare ground temperature, usually in the range of about S*C warmer than the ambient air or soil temperatures. The marcescent foliage sheathing the stem of the giant rosette plants and the tussock grasses is an efficient barrier to heat loss. The sensitive growth points of the tussock grasses are insulated, but the leaf tips are not, and shortly after development are killed and freeze-dried by the diurnal changes. The dead terminal portion of the leaves give the puna its characteristic brown or straw color. Weberbauer (1936) noted a tendency in cushion plants for the 33

protected center to be saturated with water; the water and the peaty, decomposing organic matter in the center can store heat and buffer temperature changes. For years, the role of the thick pubescence found on many species of Andean plants has been debated. It has been suggested that it protects the plant from the intense solar radiation either by reflectance (Baruch and Smith, 1979; Smith and Young, 1987) or by forming a boundary layer. Although the incident solar radiation is very intense in the high Andes, the thermal inputs are low, and ambient and tissue temperatures are usually suboptimal for translocation, growth, and photosynthesis (Meinzer and Goldstein, 1986). In the giant rosette plants in Venezuela (species of Espeletia^ s. 1.), the effect of the indûment in the pubescent species is to create a boundary layer that allows an increase of leaf temperature by decreasing convective and evaporative losses (Meinzer and Goldstein, 1986). A similar rise in leaf temperature for glabrous species was possible only with an increase in leaf width (Smith and Young, 1987). In a comparative study of species of Puya from different elevations. Miller (1986) found that thick pubescence was advantageous with respect to flower and seed development. The pubescence kept the inflorescences warmer than air temperature both during the day and at night. A large, globose inflorescence has a thermal mass that is advantageous for heat conservation. Also, pubescence prevents or retards wetting of the leaf surface, thus further protecting the leaf from freezing (Rada et al., 1985b). Some species, especially the giant rosette plants, can form a night bud by nyctinastic movements of the leaves (Smith, 1974); this insulates the growing tissues at the center of the plant from low night-time temperatures and protects the young leaves from early morning transpiration stress. It has been observed that several species of the giant rosettes have their leaves arrayed in a parabola oriented toward the sun (Smith, 1974); this is an adaptation for increased heat capture. Meinzer et al. (1985) found that the tendency to form a parabola increases with altitude. Some plant species exude a mucilaginous fluid at the the base of the leaves or in the inflorescence (Smith and Young, 1987); this is thought to increase heat storage. Cortaderia sericantha, a species which inhabits bottoms and bogs at 4200-4500 m, exhibits this phenomenon. Given the sharp drops in night-time temperature in the Andes, the unprotected leaves must also have some form of frost hardiness at night, yet maintain physiological 34

activity at low temperatures by day (Larcher, 1975; Rada et al., 1985a). In the Afroalpine region, it has been shown that the local species have evolved freezing tolerance (Bodner and Beck, 1987). Freezing intolerant plants can only be protected by freezing point depression and supercooling (Larcher, 1982). Supercooling, a depression of the freezing point within plant tissues, is a short-lived state, lasting only a few hours, and is most effective in situations where frosts are of short duration and temperatures do not fall much below -10*C (Larcher, 1982; Sakai and Larcher, 1987). Unlike the Afroalpine species, the Andean species are freezing intolerant, and have evolved a tolerance to supercooling (Perez, 1987a; Rada et al., 1985a). Supercooling this can be very effective to very low temperatures, for example, in species of Espeletia^ tolerance of cooling to -16*C has been observed experimentally (Rada et al., 1985a). In comparing the Afroalpine and Andean systems of frost hardiness, the productivity of the Afroalpine, freeze tolerant plants was found to be significantly lower than the Andean, supercooling plants (Rada et al., 1985a). Supercooling is not limited to the giant rosettes. Polylepis sericea has been found to adjust the osmotic potential in the leaves on both a daily and seasonal basis (Rada et al., 1985b). The seasonal adjustment of osmotic potential increases freezing avoidance when the temperatures are lowest. The ability to supercool increases with altitude (Smith and Young, 1987). Sugars are the probable cryoprotectant agents (Rada et al., 1985b). Water relations: Water is frequently a limiting environmental factor, in part due to the rarified atmosphere, the high potential évapotranspiration (Henning and Henning, 1981), the local precipitation patterns, and the low environmental temperatures. High Andean plants are not drought resistant (Heilborn, 1925; Gonzalez, 1985; Perez, 1987a); they have developed adaptations to withstand temporary water shortages, but will be damaged by prolonged drought. Leaves, live or marcescent, can gather and channel water flow from the plant crown during rains, heavy fogs, or melting of snow and graupel (Smith, 1979). The water usually flows down the stem and moistens the soil immediately around the plant. In species with a protective tunic of marcescent leaves, if the tunic is near or in contact with the ground, it will reduce evaporation from the soil beneath it thus improving the water situation in the immediate vicinity of the plant. Excavation of the root systems of Espeletia spp. in Venezuela has shown a concentration of feeder roots in the immediate vicinity of the stem (Smith, 1979). 35

The insulating layers of dead leaves protect the stem of the plant from desiccation and evaporation, as well as temperature changes (Smith, 1979). On the other hand, the insulating effect of the marcescent leaves during the day maintains the stem temperature lower than it would be if exposed to the ambient air, and this maintains the efficiency of water transport in the stem below potential levels (Meinzer and Goldstein, 1986). In caulirosette species, the large pith serves as a moisture reserve and can replace short term transpirational losses (Goldstein et al., 1984; Perez, 1987a; Meinzer and Goldstein, 1986; Smith and Young, 1987). Use of water stored in the pith allows the plant to pass through the early-morning translocational bottleneck when the soil, roots, and lower stem are too cold for efficient translocation. The pith of larger giant rosettes can supply the average transpirational moisture demand for about 2.5 hours (Goldstein et al., 1984). The seriousness of water stress increases with altitude, and the pith volume to leaf area ratio also increases with altitude (Goldstein et al., 1984; Meinzer et al., 1985). Cushion plants can store considerable quantities of water in the protected, peaty center of the cushion; Weberbauer (1936) likened them to sponges. This water is possibly available to the plant in times of moisture stress. The boundary layer created by thick pubescence reduces evaporative losses by reducing air flow at the leaf surface (Meinzer and Goldstein, 1986). The thickness of the pubescence increases with altitude, thus increasing the effectiveness of the boundary layer at altitudes where evaporative demand is increased (Meinzer et al., 1985). The characters associated with sclerophylly (small leaf area, thick cuticle and epidermis, pubescence, glandular exudates, and positioning and number of stomata) are adaptations, at least in part, for water conservation. Flowering depends on an adequate moisture supply; the majority of the species in the Peruvian high Andes flower during the rainy southern summer, especially from January to March (Weberbauer, 1936). Other species flower all year long. Mineral nutrition: High Andean soils have limited development, and tend to be less developed with altitude. The lack of chemical development limits the amounts of mineral nutrients available to plants (Monasterio, 1986), and the limited mineral supply is further complicated by the effect of low soil temperatures on absorption and transport processes. Studies reviewed by Monasterio (1986) indicate nutrient cycling associated with the high altitude giant rosette plants. Arthropods slowly decompose the tunic of marcescent leaves in situ, and the nutrients and organic matter fall to the ground at the 36

base of the plant. Smith (1979) found the live roots concentrated around the base of the plants, especially at the contact between the soil surface and the marcescent leaves. In the same study, invertebrate decomposers were more common in the leaf tunic than in the soil. Members of the the Bromeliaceae and Orchidaceae have roots and trichomes specialized for the absorption of water and nutrients from the atmosphere or precipitation, especially fog and dew. The role of Puya raimondii plants as habitat for several species of birds is well documented (Dorst, 1957; Macedo-Ruiz, 1978; Rees and Roe, 1980; Venero, 1984; Venero and Macedo-Ruiz, 1983). Rees and Roe (1980) speculate that bird feces can be a significant supplement for mineral nutrition, as the nutrients could be absorbed through the leaves or carried by stem flow to the roots. In addition to the droppings, some birds become ensnared by the sharp, recurved, marginal spines on the leaves. The slowly decomposing corpses could also add to the nutrient economy of the plant (Rees and Roe, 1980). Studies of above ground:below ground biomass in the Venezuelan paramos have shown very low below ground biomass ratios as compared to other vegetation types worldwide (Smith and Klinger, 1985). The climate of the paramos is nearly aseasonal, requiring little underground storage of food reserves. The cool soil conditions could inhibit development of large root systems. The puna has a dry season which becomes increasingly severe towards the south (Troll, 1968), and there is a period of drought- induced dormancy or near dormancy (W. F. Franklin, pers. com.). As food reserves are needed during dormancy, it is probable that the above ground:below ground biomass ratios in the puna, especially the dry puna, are lower than those reported by Smith and Klinger (1985). Wind Gentle to gusty winds are common in the high Andean regions. Smith (1972) found evidence of the wind affecting plant growth form and phenology. Shrubs of the genus Hesperomeles had asymmetrical, wind sculptured crowns; the affected sides had suffered breakage of twigs and, probably, increased evaporative stress. In the windy areas, cushions of Aciachne pulvinata were modified into vegetation stripes oriented with the prevailing wind direction. Hypericum laricifolium shrubs showed alteration of flowering pattern; rather than a synchronous flowering, the lee side flowered before the windward. This could be the result of water stress or evaporative cooling. 37

Photoivnthetle nathwav» Little study has been done regarding the photosynthetic pathways employed by plants at high altitudes in the Andes; it is assumed that the majority of the species are C3 plants (Smith and Young, 1987). At the northern end of the Cordillera Blanca, the western slopes have environmental conditions that support a dry grassland with Aristida, Eragrostis, Muhlenbergia, and Sporobolus, all genera which have species. Cyperus hermaphroditus grows in the same grassland, and is a known C4 species (G. C. Tucker, pers. com.). CAM photosynthesis is well known in the Agavaceae, Bromeliaceae, Cactaceae, Crassulaceae, and Orchidaceae (Kluge and Ting, 1978), all of which are represented in the Park. It is probable that some of the Huascaran species are CAM plants, although Mc Williams (1970) found no CAM activity in either Pitcairnia or Paya. CAM has been reported in Isoëtes (Styiites) andicola (Keeley et al., 1984), as well as CO2 absorption through the roots; there are species of Isoëtes in the Park, and it is possible that the same phenomena occur here. Diers (1961) studied the chromosome ploidy levels in plants collected along a transect from the coastal desert to the high Andean zone of central Peru, and found a significant increase in the number of polyploid species in the high Andean region as compared with the coastal desert and the middle elevations. Diers suggests that polyploids have adaptive advantages over diploids under the difficult environmental conditions of the high Sierra. Longevity: The difficulties of seedling establishment and survival at high elevations give an adaptive advantage to perennial plants; once established they can reproduce vegetatively as well as sexually. There are very few annual species in the high Andean flora (Rauh, 1978). Microsite selection Although the differences seem subtle in the field, there are significant differences in the east- and west-facing aspects in the tropical highlands. The daily pattern of cloud development determines that early- and mid-mornings are clear, receiving full radiation, but by late morning, the clouds begin to form, and in the afternoon cloudiness and precipitation are likely, often into the night. As a result, the east-facing slopes are warmer, drier, and have higher minimum and maximum temperatures (J. M. B, Smith, 1977; Smith and Young, 1987). J. M. B. Smith (1977) is of the opinion that plant site preference is controlled by maximum temperatures. The distribution of Polyiepis sericea illustrates this very well; in general it is restricted to the 38

drier and warmer sites at 4000 m or below, but on east-facing aspects it can reach 4200- 4300 m. The distribution of dwarf woods is correlated with old boulder falls. The trees need a warm microsite for establishment and growth. The boulder falls have a current of air flow within them; the dense, cold air drains out by gravity, drawing the warmer air into the boulder fall, and creating a warmer microclimate (Walter and Medina, 1969). In a transplant study, A. P. Smith (1977) found that the protection offered by rocks was significant; part of the site amelioration was an increase in temperature. The Polylepis and Gynoxys woods at higher elevation are clearly restricted to the old boulder falls. Solifluction caused by needle ice continually disturbs the soil on the very high elevation sites (Perez, 1987b). The constant downslope movement and heaving of the soil makes seedling establishment difficult, and it is estimated that solifluction is the main cause of seedling mortality in the needle ice zones (Perez, 1987b). Adaptations for survival in areas of solifluction are: deep, flexible taproots, extensive fibrous root systems, contractile roots, prostrate growth habit, growth in clones or cushions, and aerenchyma-filled receptacles that can withstand daily compression and expansion (Perez, 1987b; Smith and Young, 1987). After initial establishment, vegetative reproduction is a more effective method of propagation; large clones are common in needle ice zones (Perez, 1987b). Plant species lacking specialized adaptations for growth in solifluction zones can only survive in protected microsites, such as in the protection of large stones, cracks in rocks outcrops, pockets of coarse material, and in the clones or cushions of other plants (Perez, 1987b). This site selection can effect the distribution of plant species. Cushion plants are usually found downslope from large stones (Perez, 1987b). The geomorphic effects of needle ice were discussed in the section on geology. Polllnntori Beetles, birds, bumblebees, butterflies (Lamas and Perez, 1983), and flies have been seen visiting flowers in the Park. Of these, only birds and bumblebees were seen clearly pollinating flowers. The low environmental temperatures at high elevations interfere with the action of pollinators, especially regarding energy balance (Heinrich and Raven, 1972). Insects are especially effected, except the bumblebees, which can thermoregulate (Heinrich and Raven, 1972). During cloudy weather, insect pollination virtually ceases (Cruden, 1972; Smith, 1975). Considering the decline in effective insect pollination, bird pollination becomes increasingly important at high elevations (Cruden, 1972). 39

Smith (1975) found that Oritrophium limnophilum, a rosette plant with scapose flowers and a capitulum that is parabolic in shape, to be heliotropic. Smith concludes that the flower shape and heliotropism increase the heating of the capitulum, making it thermally more attractive to visiting insects, and, at the same time, affecting the rate of floral development by warming the flower tissues. The resulting increase in pollination rates should increase the fitness of the plant species (Smith, 1975). I suspect that there is an altitudinal level beyond which the number of pollinating birds decreases. The most commonly observed pollinators in the Park were bumblebees. Prédation: The low environmental temperatures of the high Andean region inhibit, but do not eliminate, insect activity; herbivorous insects can be found at high elevations. Both pubescence and secondary chemical compounds, which inhibit insect prédation, as well as that of other herbivores, are common in many of the species in the Park flora. Both pubescence and secondary chemical compounds have adaptive value for other environmental factors; their entomofugal properties are likely a coincidence. Vegetation Types The vegetation of the Park is a mosaic of different vegetation types, which often intergrade into one another. The principal vegetation types are: dwarf woods, shrublands, grasslands, aquatic and semi-aquatic communities, and high Andean. Each of these types is represented by one or more plant communities. The principal goal of this study was to enumerate and document the flora; there was no time to conduct detailed studies of the plant communities. A general overview of the vegetation types and plant communities follows: Riverine woodland: In the northern valleys at elevations below 3850 m, the microclimate is warmer. A narrow band of mixed woodland dominated by Alnus acuminata is often present, bordering the streams. Myrica pubescens, Vallea stipularis, and Weinmamia aff. laxiflora, which can be either small trees or large shrubs, are the other dominants in the riverine forest; CHharxylum dentatum and Duranta mandonii are the dominant shrubs. These genera are important in the ceja de selva vegetation of the eastern slopes of the Andes. The warmth and humidity in the understory of this forest is an appropriate habitat for bryophytes and larger ferns (Athyrium and Thelypteris). In the western valleys, this forest is distinct, not intergrading into the adjacent vegetation. However, in the eastern valleys, this is continuous with the tall shrubland, and can scarcely be delineated from it. 40

POIVICPII ierlcea woods: These can vary from an open overstory in areas of dry, bouldery shrubland to dense woodlands. Polylepis sericea is found on warm sites, and seems to be quite tolerant of aridity. The situations where P. sericea is an overstory are most common at the mouths of the northern valleys on the western side of the Park, the associated species are determined by the underlying vegetation type, which is usually the dry shrubland. The dense woodlands are at slightly higher elevations, ca. 3800 m, on both sides of the Cordillera. The understory of these woods is species poor when compared with the higher elevation woods, having Micrograma chrysolepis, a few species of Peperomia, and abundant bryophytes. In this woodland, two species of canopy climbers. Clematis sp. and Passi/lora trifoiiata, are well developed. At times Polylepis sericea can be found up to 4200 m high, but then it will be found on east-facing microsites. High elevation woods: At elevations of 4000 to 4500 m, or perhaps higher, the woodlands can be single species stands of Polylepis weberbaueri or Gynoxys oleifolia, or a mixture of the two, and are in stands that vary from open to rather dense. Pure stands of Gynoxys oleifolia are found on the north-facing slopes, Polylepis weberbaueri prefers the higher colder sites, especially the south-facing slopes, and the mixed stands are found in intermediate habitats. These have well developed herbaceous understories with Arenaria lanuginosa. Stellaria ovata, Senecio arachnolomus, S. klattii, all in extensive clones. Several species of tuberiferous Solanum are found growing in soil pockets of the understory. Shrubs, although rare, are represented by Ribes aff. magellanica, Solanum nitidum, and Vaccinium floribundum, and vines are represented by Salpichroa glandulosa and Solanum oxycoccoides, or, less commonly, Passiflora trifoliata. Two species of parasites, Tristerix chodatianus and T. pubescens, are very noticeable elements in these forests. Occasionally, Miconia alpina, a small tree, is found in moist spots in these woods. Buddleia incana is found sporadically in different situations, mostly above 4000 m. It can be encountered as an isolated individual, or in small stands by itself, or near Polylepis-Gynoxys woods. This species can become a very large tree (up to 15 m tall and 50 cm in diameter), and is a typical element of the high Sierran flora (Weberbauer, 1945). Many lithophytic habitats can be found throughout the Park, be they talus slopes, bouldery areas, individual large boulders, rock outcrops, or the massive granodiorite 41

valley walls in the Llanganuco, Parôn, or Santa Cruz valleys. These habitats support most of the populations of Bromeliads, Orchids, and the xeromorphic ferns. Shrublands are very common throughout the Park, and can be divided into the tall-stature shrublands of the moist valleys, and the short shrublands of the drier habitats. Tall shrublands: These are most abundant on the east side of the Cordillera, but can also be found in the Parôn valley, with a tendency to tall shrubland in parts of the Ishinca valley. They inhabit moist sites up to 3800 to 4000 m elevation, and intergrade with the riverine woods. The characteristic shrubs in this type are Baccharis tricuneata, Gynoxys caracensis, and Miconia salicifolia (the most abundant), along with other species which show a strong connection with the ceja de selva: Escallonia resinosa, Fuchsia denticulata, Myrica pubescens, Myrsine dependens, Symplocos aff. sandiae, Vallea stipularis, and Weinmannia aff. laxiflora. Species of Diplostephium may be found in both the tall and dry shrublands. Dry shrublands: In the lower regions of many of the western valleys, there are extensive stands of shrubland dominated by Baccharis tricuneata, Berberis lutea, Calceolaria spp. Gynoxys caracensis, Lupinus aff. tarapacensis, and several species of Senecio. Among the shrubs one finds many species of grasses and herbs. Intermediate between these two shrubland categories are the low shrublands found in the moister portions of the northern valleys on the west side of the Cordillera, which have Coreopsis sherffii, Diplostephium azureum, Cardoquia elliptica and Hesperomeles cuneata as codominants. Three species of shrubs can form small to medium-sized, single-species stands: Buddleia coriacea forms large, dense stands at higher elevations in the southern regions of the Park, especially in the Querococha valley; Chuquiraga spinosa is found in the dry grasslands in the southern portion of the Park; and Loricaria ferruginea can be found from middle to the highest elevations throughout the Park. Other species of shrubs do grow at the highest elevations, but the individuals are widely dispersed. Grasslands: As a group, this vegetation type is the most important in the Park. There are several plant communities included in this category, and perhaps more than 50% of the species known from the Park are grassland plants. A few of the several plant communities can be tentatively identified. On the western slopes of the Cordillera, north of the Santa Cruz valley, the grassland is very arid, and dominated by species from typically C^ genera: Aristida enodis, Eragrostis aff. 42

pastoensis, Muhlenbergia sp., Schizachyrium sanguineunit and Sporobolus lasiophyllus. Near the stands of Puya raimondii in the Rio Pachacoto drainage, there is a tussock grass community dominated by Stipa obtusa, which is virtually the only grass in the community. Stipa obtusa is commonly seen on disturbed sites, especially the spoil bank of the roads in the southern part of the Park, At higher elevations, grasslands dominated by species of Calamogrostis and Stipa hans-meyeri are very common; this community is very tolerant of thin, dry soils. In the gullies and low spots on the same terrain one finds the Caiamagrostis - Stipa hans-meyeri community, there is also a community with low species diversity dominated by Festuca weberbaueri. The grasslands intergrade into the adjoining vegetation types: shrubland, semi- aquatic, and high Andean. At the ecotone with other communities, many species that can subsist in other vegetation types will be found in the grasslands. Aquatic and seml-aquatlc communities: This is a category of convenience, a collection of environmental situations involving impounded water or saturated soils: lakes, springs, vernal pools, streamsides, marshy areas, and bogs. The lakes in the Park are of glacial origin; many are formed by impoundment behind terminal moraines. LOffler (1968) states that most of the high Andean lakes are of glacial origin. LOffler (1968) classifies high elevation lakes into two categories based on water temperature and stratification of temperature, oxygen, and nutrients within the water. Although both categories have high levels of circulation and mixing, lakes at lower elevation are slightly warmer (average temperature higher than 12'C) and have a slight stratification. It is possible that the lower lakes in the Llanganuco and Santa Cruz valleys are of the warmer type. All other lakes in the Park are cold, with frequent circulation, and no stratification of temperature or nutrients. The alternation of daytime heating and strong nocturnal cooling in the high Andes promotes circulation. Studies of high mountain lakes have shown them to be nutrient poor, although a high organic content can be present (LOffler, 1968). Many of the pools and lakes in the Park have very dark water indicative of a high organic content. The lakes fed directly by glaciers have a high level of particulate matter, glacial flour, which gives them a milky to aquamarine color. I found no plants growing in lakes with high particulate loads. Floristically, the lakes can be separated into low elevation, less than 4000 m, and high elevation. The low elevation lakes have a varied flora of emergent, submerged, and 43

floating aquatic plants, including Elodea potamogeton, Isoëtes spp., Jtmcus arcticus var. andicola, Myriophyllum elatinoides, and Schoenoplectus californicus. The high elevation lakes have few plants, usually only Isoetes lechleri. Several springs have been found which have a high volume discharge of clear, cold water, and a distinct flora: Epilobium denticulatum, Lemna minuscula, Mimulus glabratus, Veronica serpylli/olia, and several species of aquatic bryophytes. A large number of ponds and vernal pools are scattered throughout the grasslands. These are often shallow, and are prone to wide temperature variations. Under direct solar radiation, the shallow water can be heated to temperatures warm to the touch, while, at the same time, air temperatures are so cool that a sweater is needed for personal comfort (D. N. Smith, pers. obs.). The pools and ponds may vary in lifespan from very short to permanent in years with normal precipitation. The most ephemeral may have no flora, or a species of Isoitesf while the most persistent have a flora of several species, and may be associated with a marshy area with Jtmcus arcticus and other coarse marsh plants. The pond flora includes several of the following: Alopecurus aequalis, Elatine peruvianus, Callitriche spp., Lilaea scillioides, Potamogeton paramoanus, and Ranunculus iimoselloides. Aquatic animals can be found in the lakes and ponds; fresh water shrimp are sometimes abundant. Lôffler (1968) notes that the non-malacostracan crustaceans show a strong connection with the southern temperate aquatic fauna. Marshy areas will usually have Juncus arcticus and an undescribed species of Carex. Streamsides have a vegetation very similar to the adjacent vegetation, which is often grassland, with an increase in water demanding plants: Calamagrostis spp., Carex spp., Gentiana sedifolia, Gentianella spp., Castilleja spp., Lupinus weberbaueri, and Werneria spp. Boggy spots may be found in depressions or on slopes with a ground water source; all are characterized by very wet to saturated soils and low temperatures; with increasing elevation there is an increase in boggy vegetation. The Andean bogs closely resemble the cushion bogs, which are widespread in the cold parts of the Southern Hemisphere and Pacific Islands (Godley, 1978). The dominant species are cushion plants, Distichia muscoides, Oreoboius obtusangtdatus, and Plantago rigida, and are associated with species in the Cyperaceae (Eleocharis albibracteata, Isolepis inundata, Phylioscirpus deserticola), and the Asteraceae (Senecio and Werneria), as well as species of Aichemilia, Castiiieja, Huperzia, Lysipomia, Ourisia, Plantago, and various grasses. 44

At about 4500 m there is a shift in species dominance in the boggy communities; below that elevation the bogs are of mixed composition, or tend to be dominated by Oreobolus obtusangulatus, and above 4500 m Distichia muscoides becomes increasingly dominant. The Distichia can grow intermixed with other dominant bog species, but at high elevations Distichia becomes completely dominant and forms hummocks to several meters in diameter and nearly a meter tall, standing island-like in vernal pools. In these bogs there are few species other than Distichia: Alchemilla diplophylla, Calamagrostis ovata, Lysipomia montioides, and Ourisia muscosa, among others. The Distichia bogs are always in low spots on relatively level ground, not on slopes. High Andean vegetation: Found at the highest elevations, this vegetation type develops under the most difficult environmental conditions: constant solifluction produced by the action of needle ice and freezing temperatures nearly every night. This is the most severe environment in the the Central Andes. Plants which survive here must be well adapted to withstand the constant substrate movement, extreme temperature fluctuations, and an average temperature so low that it may limit physiological processes. Many have deep, strong taproots or clonal or cushion-plant growth form, and most will be perennial. The important genera in the high Andean zone are: Mnodes, Nototriche, Pycnophyilum, Stangea, Senecio, Valeriana^ and Werneria. Swan (l%8) describes a biotic zone above the limits of plant growth in the high mountains of Mexico and the Himalayas, called the aeolean zone. This zone is inhabited by heterotrophic organisms feeding on wind-borne organic matter from lower life zones. As yet, no aeolean zone has been identified in the Cordillera Blanca. 45

THE FLORA: COMMENTARY AND LIST OF TAXA The flora of the Park includes 104 families, 339 genera, and 799 species of gymnosperms, pteridophytes and flowering plants. The following shows the distribution of the of families, genera and species in each major group and the percent contribution of each taxonomic level to the flora. Families Genera Species Gymnosperms 1 1 1 Pteridophytes 14 - 13% 28 - 9% 67 - 8% Monocots 18 - 17% 75 - 22% 206 - 26% Dicots 71 - 68% 236 - 69% 525 - 66% Although there are a large number of families and genera, there are relatively few which contribute a large number of species to the flora. This was also seen to be the case in the high elevation floras of Colombia (Cleef, 1981), Argentina (Ruthsatz, 1977), East Africa (Hedberg, 1957), and New Guinea (van Royen, 1979b, 1982, 1983). In the Huascar&n flora, only 48% of the families are represented by more than one genus, and 40% are represented by a single species. The 10 largest families (shown below) account for 50% of the genera in the flora and 57% of the species. Together, the Asteraceae and Poaceae contribute 26% of the genera and 34% of the species in the flora. Family Genera: Species 1. Asteraceae 59: 164 2. Poaceae 32: 111 3. Scrophulariaceae 12: 33 4. Orchidaceae 11: 19 5. Brassicaceae 11: 15 6. Caryophyllaceae 10: 25 7. Cyperaceae 10: 20 8. Roscaeae 9: 18 9. Fabaceae (s. s.) 8: 26 10. Solanaceae 8: 26 The Asteraceae are 21 % of the total number of species in the Park, the Poaceae 14%, and the Scrophulariaceae 4%. In the early 1880s, John Ball made a short excursion from Lima to the high mountain pass at Ticlio. Analyzing the results of his collections and observations. Ball (1885) found the same group of principal families in the high elevation zone as those in the Huascaràn study, with the exception of the Cyperaceae, Orchidaceae, and Rosaceae, which he did not encounter. Ball observed that the Asteraceae accounted for about 25% of the flora and the Poaceae another 10%; the Scrophulariaceae was the third most important, although no percentage was estimated. 46

The 10 largest families in the Huascaràn flora are also important in the floras of the Colombian paramos, the Argentinan puna, and the Afroalpine zone of East Africa. However, the contribution and importance of each family is not the same from flora to flora The mean number of species per genus is 2.4, however, 61% of the genera are represented by a single species and 89% by 3 species or fewer. Only 11% of the genera are represented by more than 3 species. The ten largest genera (shown below) account for 22% of the species in the flora, and Senecio, alone, accounts for ca 5% of the species in the flora. Genus Species 1. Senecio (Asteraceae) ca 39 2. Calamagrostis (Poaceae) ca 21 3. Werneria (Asteraceae) ca 18 4. Poa (Poaceae) ca 16 5. Solanum (Solanaceae) ca 16 6. Calceolaria (Scrophulariaceae) ca 15 7. Lupinus (Fabaceae, s. s.) ca 15 8. Baccharis (Asteraceae) ca 13 9. Valeriana (Valerianaceae) ca 13 10. Festuca (Poaceae) ca 12 The following is a list of the families, genera and species included in the flora of the Huascarân National Park:

GYMNOSPERMS

EPHEDRACEAE Ephedra rupestris Benth.

PTERIDOPHYTES

ASPLENIACEAE Asplenium castaneum Schlecht. & Cham. A. fragile PresI A. monanthes L. A. polyphyllum Bertol. A. praemorsum Sw. A. sessilifolium Desv. A. triphyllum Presl

BLECHNACEAE Blechnum cordatum (Desv.) Hieron. B. loxense (Kunth) Hieron. 47

DENNSTAEDTIACEAE Histiopteris incisa (Thunb.) J. Sm. Hypolepis bogotensis Karsten H. obtusata (Presl) Hieron.

DRYOPTERIDACEAE Athyrium dombei Desv. Cystopteris fragilis (L.) Bernh. Eiaphoglossum atrosquamatum Miclcel, ined. E. attenuatum Christ. E. aff. deorsum (Christ.) Vareschi E. engelii (Karsten) Christ. E. lindenii (Borg ex Fée) Moore E. mathewsii (Fée) Moore E. minutum (Pohl) Moore E. aff. paleaceum Miclcel, ined. E. pilosus Miclcel, ined. Polystichum cf. nudicaule Rosenst. P. cf. polyphyllum (Presl) Presl P. cf. rochaleanum Glaz Woodsia montevidensis (Spreng.) Hieron.

EQUISETACEAE Equisetum bogotense Kunth

GLEICHENIACEAE Gleichenia (Sticherus) simplex (Desv.) Hook.

ISOETACEAE Isoétes lechleri Matt. I. triquetra A. Braun Isoétes sp. nov. (-1. "ticlioensis" Fuchs, ined.) Isoétes hybrid (I. lechleri x "ticlioensis")

LYCOPODIACEAE Huperzia crassa (Willd.) Rothm. H. saururus (Lam.) Trevisan H. cf. trencilla (Baker) Holub Lycopodium clavatum L. L. magellanicum (Palisot) Sw. L. thyoides Humb. & Bonpl. ex Willd.

OPHIOGLOSSACEAE Ophioglossum crotalophoroides Walt.

PLAGIOGYRIACEAE Plagiogyria semicordata (Pr.) Christ 48

POLYPODIACEAE Campyloneurum angustifolium (Sw.) Presl ssp. amphostenon Farwell C. angustifolium ssp. solutum (Klotzsch) B. Leôn Grammitis achilleifolia (Kaulf.) R. & A. Tryon G. heteromorpha (Hook. & Grev.) Morton G. magellanica Desv. G. moniliformis (Sw.) Proctor G. pilosissima (Mart. & Gal.) Morton Microgramma chrysolepis (Hook.) Crabbe Polypodium buchteinii Rosenst. P. pycnocarpum C. Chr. P. sessilifolium Desv.

PTERIDACEAE Adiantum poiretii Wikstr. Cheilanthes bonariensis (Willd.) Proctor C. marginata Kunth C. myriophylla Desv. C. pilosa Goldm. C. priunata Kaulf. Eriosorus cheilanthoides (Sw.) A. Tryon E. novogratensis A. Tryon Eriosorus sp. inq. Jamesonia alstonii A. Tryon J. goudotii (Hieron.) C. Chr. J. scammanae A. Tryon Notolaena nivea (Poiret) Desv. Pellaea ternifolia (Cav.) Link

THELYPTERIDACEAE Thelypteris caucaensis (Hieron.) Alston T. rudis (Kunze) Proctor

SALVINIACEAE Azolla filiculoides Lam.

ANGIOSPERMS

AGAVACEAE Furcraea occidentalis Trel.

ALLIACEAE Trichlora peruviana Baker in Hook.

ALSTROEMERIACEAE Alstroemeria pygmaea Herbert Bomarea albimontana D. N. Smith & Gereau, ined. B. dulcis (Hook.) Beauv. 49

Bomarea phyllostachya Masters ex Baker, vel sp. aff. B. zosteraefolia Killip

AMARANTHACEAE Alternanthera macbridei Standley Alternanthera spp. — 2 undetermined species.

AMARYLLIDACEAE Stenomesson luteum (Herbert) Baker in Saund.

ANTHERICACEAE Echeandia eccremorrhiza (Ruiz Lôpez & Pavôn) Cruden, comb. ined.

APIACEAE Azorella biloba (Schlecht.) Wedd., vel sp. aff. A. corymbosa (Ruiz Lôpez & Pavôn) Pers. A. crenata (Ruiz Lôpez A Pavôn) Pers. A. multifida (Ruiz Lôpez & Pavôn) Pers. A. pulvinata Wedd. Bowlesia lobata Ruiz Lôpez & Pavôn B. tenella Meyen Daucus montanus Willd. Eryngium humile Cav. Lilaeopsis macloviana (Gandoger) A. W. Hill Nyphogeton scabra (Wolff) J. F. Macbr. Oreomyrrhis andicola (Kunth) Hook. f.

ARALIACEAE Oreopanax oroyanus Harms

ASCLEPIADACEAE Cynanchum pichinchense K. Schum. C. tarmense Schultr. Cynanchum spp. — 2 undetermined species. Metastelma sp. inq.

ASTERACEAE Achyrocline alata (Kunth) DC. A. ramosissima (Schultz-Bip.) Britton Ageratina azangaroensis (Schultz-Bip. ex Wedd.) R. King & H. Robinson Antennaria linearifolia Wedd. Aphanactis villosa S. F. Blake Aristeguieta discolor (DC.) R. King & H. Robinson Baccharis caespitosa (Ruiz Lôpez & Pavôn) Pers. B. genistelloides Pers. B. latifolia (Ruiz Lôpez & Pavôn) Pers. B. tricuneata (L. f.) Pers. — erect and prostrate forms B. uniflora (Ruiz Lôpez & Pavôn) Pers. Baccharis spp. — 8 undetermined species. so

Barnadesia dombeyana Less. Belloa longifolia (Cuatrec. & Arist.) Sagâst. & Dillon B. piptolepis (Wedd.) Cabrera B. schultzii (Wedd.) Cabrera, vel sp. aff. B. turner: Sagâst. & Dillon Bidens andicola Kunth B. triplinervia Kunth Chaptalia cordata Hieron. Chersodoma diclina (Wedd.) Cabrera C. ovopedata (Cuatrec.) Cabrera Chrysactenium acaule (Kunth) Wedd. Chuquiraga spinosa Less. ssp. huamanpinta C. Ezcurra Conyza artemisiaefolia Meyen & Walp. C. bonariensis (L.) Cronq. C. floribunda Kunth Coreopsis intégra Blake C. senaria S. F. Blake & Sherff C. sherffii S. F. Blake Cosmos peucedenifolius Wedd. Cotula australis (Sieb. ex Spreng.) Hook. f. Cuatrecasasiella isernii (Cuatrec.) H. Robinson Dasyphyllum histerix (Wedd.) Cabrera Diplostephium azureum Cuatrec. D. haenkei (DC.) Wedd. D. sagasteguii Cuatrec. Erigeron rosulatus Wedd. Erigeron sp. inq. Galinsoga sp. inq. Gamochaeta erythractis (Wedd.) Cabrera G. humilis Wedd. G. oreophila Dillon & Sagâst. G. purpurina (L.) Cabrera G. spicata (Lam.) Cabrera Gnaphalium badium Wedd. G. dombeyanum DC. G. polium Wedd. Grosvenoria (Eupatorium) coelocaulis (Robinson) H. King & H. Robinson Gynoxys caracensis Muschler G. oleifolia Muschler Helianthopsis discolor (S. F. Blake) H. Robinson Hieracium mandonii (Schultz-Bip.) Arvet Touvet Hieracium sp. inq. Hypochoeris eriolaena (Schultz-Bip.) Reiche H. cf. graminifolia Hieron. H. taraxacoides (Walp.) Benth. & Hook. Jungia paniculata (DC.) A. Gray J. rugosa Less. J. stuebelii (Hieron.) Crisci Leuceria daucifolia (Don) Crisci 51

Liabum solidagineum (Kunth) Less. Lophopappus peruvianus Cabrera Loricaria ferruginea (Ruiz Lôpez & Pavôn) Wedd. L. iycopodinea Cuatrec. Lucilia kunthiana (DC.) Zardini Mnodes andina A. Gray ex Hook. f. & B. D. Jackson M. aretioides (Schultz-Bip.) Cuatrec. M. pulvinulata Cuatrec. M. sp. nov., sterile Munnozia lyrata (A. Gray) H. Robinson & Brettell Mutisia mathewsii Hook. & Arn. Noticastrum marginatum (Kunth) Cuatrec. forma acaule Cuatrec. Novenia tunariensis (O. Ktze.) Freire Ophryosporus chiica (Kunth) Hieron. Oritrophium hieracioides (Wedd.) Cuatrec. O. limnophilum (Schultz-Bip.) Cuatrec. O. peruvianum (Lam.) Cabrera Paranephelius ferreyrii (Hieron.) H. Robinson P. ovatus Wedd. P. uniflorus Poepp. & Endl. Pentacalia andicola (Turcz.) Cuatrec. Perezia ciliaris Hook. & Arn. P. coerulescens Wedd. P. multiflora (Humb. & Bonpl.) Less. P. pungens (Humb. & Bonpl.) Less. Peryminium sp. inq. Plagiocheilus solivaeformis DC. Polymnia fruticosa Benth. Pseudonoseris szyszylowiczii (Hieron.) H. Robinson & Brettell Senecio adenophylloides Schultz-Bip S. adenophyllus Meyen & Walp. S. arachnolomus Wedd. S. breviscapus DC. S. (Lasiocephalus) campanulatus Schultz-Bip. ex Klatt S. (Culcitium) canescens (Kunth) Cuatrec. S. clavus Cuatrec. S. comosus Schultz-Bip. S. condimentarius Cabrera S. culcitioides Schultz-Bip. S. cf. ferreyrae Cabrera S. hyoseridifolius Wedd. S. klattii Greenm. S. leucophorbius Cuatrec. S. macrorhizus Wedd. S. mathewsii Wedd. S. minesinus Cuatrec. S. modestus Wedd. S. (Culcitium) nivalis Kunth S. (Dorobaea) pimpinellifolius Kunth 52

Senecio praeruptorum Schultz-Bip. ex Klatt S. rauhii Cuatrec. S. repens DC. S. scorzoneraefolius Meyen & Walp. S. scrobicarioides DC. S. (Culcitium) serratifolius (Meyen & Walp.) Cuatrec. S. spinosus DC. S. sulinicus Cabrera S. tephrosioides Turcz. S. tingoensis Cabrera & Zardini S. usgorensis Cuatrec. Senecio spp. — 8 undetermined species Siegesbeckia jorullensis Kunth Sonchus oleraceus L. Stevia sp. inq. Stuckertiella capitata (Wedd.) Cabrera Tagetes multiflora L. Tagetes spp. — 2 undetermined species. Vasquezia oppositifolia (Lag.) Blake Verbesina arborea Kunth Viguiera peruviana Gray Werneria aretioides Wedd. W. caespitosa Wedd. W. dactylophylla Schultz-Bip. W. heteroloba Wedd. W. humilis Kunth W. macbridei Cuatrec. W. nubigena Kunth W. orbygniana Wedd. W. pygmaea Hook. & Arn. W. cf. pygmaea Hook. & Arn. W. sedoides Blake W. strigosissima A. Gray W. villosa A. Gray Werneria spp. — 5 undetermined species.

BASELLACEAE Ullucus aborigineus Bruch.

BEGONIACEAE Begonia pleiopetala A. DC.

BERBERIDACEAE Berberis jelskiana C. Schneider B. lutea Ruiz Lôpez & Pavôn

BETULACEAE Alnus acuminata Kunth ssp. acuminata S3

BIGNONIACEAE Tecoma sambucifolia Kunth

BORAGINACEAE Hackelia mexicana (Schlecht. & Cham.) I. M. Johnston Lithospermum peruvianum A. DC. Plagiobotrys humilis (Ruiz Lôpez & Pavôn) I. M. Johnston

BRASSICACEAE Brassica rapa L. Brayopsis alpaminae Gilg & Muschler subsp. smithii Al- Shehbaz B. calycina (Desv.) Gilg & Muschler Capsella bursa-pastoris (L.) Medicus Cardamine bonariensis Pers. subsp. bonariensis Catadysia rosulans O. £. Schulz Cremolobus chilensis (DC.) DC. Descurainia athrocarpa (Gray) O. E. Schulz D. myriophylla (Willd.) Fries Draba alchemilloides Gilg D. cryptantha Hook. f. D. pickeringii A. Gray Lepidium bipinnatifidum Desv. Mancoa hispida Wedd. Weberbauera spathulaefolia (A. Gray) 0. E. Schulz

BROMELIACEAE Pitcairnia pungens Kunth var. pungens Puya angusta Lyman B. Smith P. herrerae Mez, vel sp. aff. P. membranacea Lyman B. Smith, vel sp. aff. P. raimondii Harms P. rauhii Lyman B. Smith, vel sp. aff. P. reflexiflora Mez Puya sp. inq. Tillandsia cerrateana Lyman B. Smith T. fendleri Griseb. var. reducta (Lyman B. Smith) Lyman B. Smith T. humilis PresI T. ionochroma André ex Mez T. lopezii Lyman B. Smith T. oroyensis Mez T. rubella Baker T. tectorum E. Morren T. waltheri Mez var. herrerae (Harms) Rauh

BUDDLEIACEAE Buddleia coriacea Remy B. incana Ruiz Lôpez & Pavôn 54

CACTACEAE Cereus sp. inq. Cleistocactus sp. inq. Opuntia floccosa Salm-Dyck Opuntia sp. inq. Oreocereus sp. ("Matucana yanganucensis Rauh & Backbg.) Oreocereus sp. ("Oroya borchersii (Boed.) Backbg.)

CAESALPINIACEAE Senna birostris (Vogel) Irwin & Barneby var. helveola (J. F. Macbr.) Irwin & Barneby

CALLITRICHIACEAE Callitriche heteropoda Fassett C. nubigena Fassett

CAMPANULACEAE Hypsella reniformis (Kunth) PresI Lobelia nana Kunth L. tenera Kunth Lysipomia laciniata A. DC. subsp. linearifolia (Wimmer) McVaugh L. montioides Kunth L. sphagnophila Griseb. ex Wedd. subsp. acuta (Wimmer) McVaugh Siphocampylus candolei Wimmer S. tupaeformis Zahlbr. var. stenophyllus Wimmer Wahlenbergia peruviana A. Gray

CAPPARACEAE Cleome glandulosa Ruiz Lôpez & Pavôn

CARYOPHYLLACEAE Arenaria jamesoniana Rohrb. A. lanuginosa (Michaux) Rohrb., sensu lato A. nitida (Bartling) Rohrb. A. parvifolia Benth., vel sp. aff. A. serpens Kunth A. standleyi Baehni & J. F. Macbr., vel sp. aff. Cardionema ramosissimum (Weinm.) Nels. & J. F. Macbr. Cerastium candicans Wedd. C. danguyi J. F. Macbr. C. imbricatum Kunth C. soratense Rohrb., vel sp. aff. C. subspicatum Wedd. Drymaria engleriana (Muschler) Baehni & J. F. Macbr. var. devia (Baehni & J. F. Macbr.) Duke D. stereophylla Mattf. var. exstipulata Mattf. Paronychia andina A. Gray subsp. boliviana Chaudhri P. andina A. Gray subsp. purpurea Chaudhri P. communis Cambess. var. communis forma communis 55

Paronychia mandoniana Rohrb. P. weberbaueri Chaudhri Plettkea cryptantha Mattf. Pycnophyllum aschersonianum Mushier P. molle Remy Silene thysanodes Fenzl in Endl. Spergularia arvensis L. Stellaria cuspidata Willd. ex Schldl. S. ovata Willd. ex Schldl.

CLUSIACEAE Hypericum brevistylum Choisy H. laricifolium Juss.

COLUMELLIACEAE Columellia weberbaueri Schltr.

COMMELINACEAE Commelina fasciculata L.

CONVOLVULACEAE Dichondra microcalyx (H. Hallier) Fabris in Cabrera

CORIARIACEAE Coriaria ruscifolia L. subsp. microphylla (Poiret) L. Skog

CRASSULARIACEAE Echeverria sp. inq. Tillaea connata Ruiz Lôpez & Pav6n T. paludosa Schlecht. Villadia imbricata (Diels) Baeh. & J. F. Macbr.

CUNONIACEAE Weinmannia laxiflora Pamp., vel sp. aff.

CUSCUTACEAE Cuscuta sp. inq.

CYPERACEAE Baeothryon rigida (Boeckeler) D. N. Smith, ined. Bulbostylis juncoides (Vahl) KQk. Carex ancashensis D. N. Smith & Reznicek, ined. C. boliviensis Van Juerck & MttH.-Arg. C. bonplandii Kunth C. collumanthus (Steyerm.) Mora C. crinalis Boott C. ecuadorica Kûk. C. hebetata Boott C. mandoniana Boeckeler 56

Cyperus hermaphroditus (Jacq.) Standley C. sesleroides Kunth Eleocharis albibracteata Nees & Meyen ex Kunth bolepis inundate R. Br. Oreobolopsis tepalifera T. Koyama & Guaglian. Oreobolus obtusangulatus Gaud. Phylloscirpus sp. inq. (-Scirpus aff. atacamensis Boeckeler) P. deserticola (Philippi) D. N. Smith, ined. P. sp. inq. ("Scirpus aff. hieronymi Boeckeler) Schoenoplectus californicus (C. Meyer) Sojak subsp. californicus var. californicus

DIOSCOREACEAE Dioscorea ancashensis Knuth D. larecajensis Uline ex Knuth

ELAEOCARPACEAE Vallea stipularis L. f.

ELATINACEAE Elatine peruviana Baehni & J. F. Macbr.

ERICACEAE Gaultheria brachybotrys DC. G. vaccinioides Wedd. Pernettya prostrata (Cav.) DC. Vaccinium floribundum Kunth

EUPHORBIACEAE Chamaesyce serpens (Kunth) Small Croton ruizianus Mûll. Arg. Euphorbia raphanorhiza (Millsp.) J. F. Macbr. Sebastiania haploclada Briq.

FABACEAE Astragalus brackenridgei A. Gray A. garbancillo Cav. A. peruvianus Vogel A. uniflorus DC. Dalea weberbaueri Ulbr. Lathyrus magellanicus Lam. Lupinus microphyllus Desv. L. cf. mutabilis Sweet L. nubigenus Kunth L. rupestris Kunth L. cf. smithianus Kunth L. cf tarapacensis C. P. Smith L. weberbauerii Ulbr. Lupinus spp. — 8 undetermined species. 57

Otholobium munyensis (J. F. Macbr.) J. Grimes O. pubescens (Poiret) J. Grimes Spartium junceum L. Trifolium amabile Kunth Vicia andicola Kunth Vicia sp. inq.

FLACOURTIACEAE Pineda incana Ruiz Lôpez & Pavôn

GENTIANACEAE Gentiana sedifolia Kunth Gentianella mesembryanthemoides (Gilg) Pringle, ined. G. radicata (Griseb.) Pringle G. roseolilacina (Gilg) Pringle, ined. G. sanctum (Gilg) Pringle G. thyrsoidea (Hook.) Fabris G. tristicha (Glig) Fabris G. weberbauerii (Gilg) Fabris Halenia umbellata (Kunth) Gilg

GERANIACEAE Erodium cicutarium (L.) L'Her. Geranium sessiliflorum Cav. G. cf. peruvianum Geranium sp. inq.

GROSSULARIACEAE Escallonia corymbosa (Ruiz Lôpez & Pavôn) Pers. E. cf. myrtillioides L. f. E. resinosa (Ruiz Lôpez & Pavôn) Pers. Ribes cuneifolium Ruiz Lôpez & Pavôn R. magellanica Poiret, vel sp. aff. Ribes spp. — 2 undetermined species.

HALORAGACEAE Myriophyllum elatinoides Gaud.

HYDROCHARITACEAE Elodea potamogeton (Bertero) Espinosa

HYDROPHYLLACEAE Phacelia secunda J. Gmelin

IRIDACEAE Orthrosanthus occissapungus (Ruiz ex Klatt) Diels Sisyrinchium brevipes Baker S. chilensis Hook., vel sp. aff. S. jamesonii Baker 58

Sisyrinchium junceum E. Meyer ex Presl S. praealtum Krflnzlin S. tinctorium Kunth S. trinerve Baker Tigridia philippiana I. M. Johnston

JUNCACEAE Distichia muscoides Nees & Meyen Juncus arcticus L. var andicola (Hook.) Balslev J. bufonius L. J. cyperoides Laharpe J. ebracteatus; E. Meyer J. imbricatus Laharpe J. pallescens Lam. J. stipulatus Nees & Meyen J. tenuis Willd. var. platycaulos (Kunth) Bucheneau Luzula gigantea Desv. L. racemosa Desv. L. vulcanica Liebm.

JUNCAGINACEAE Lilaea scilloides (Poiret) Hauman

LAMIACEAE Gardoquia elliptica Ruiz Lôpez & Pavôn Gardoquia sp. ("Satureja sericea (Presl) Briq.) Lepechinia meyenii (Walp.) Epiing Salvia oppositifoHa Ruiz Lôpez & Pavôn S. sarmentosa Epiing, vel sp. aff. S. tafallae Benth. Scutellaria gardoquioides (Benth. & Hook.) Benth. in DC. Stachys pusilla (Wedd.) Briq.

LEMNACEAE Lemna minuscula Herter

LINACEAE Linum olygophyllum Willd. ex Schult.

LOASACEAE Cajophora sepiaria (Ruiz Lôpez & Pavôn) J. F. Macbr. Cajophora sp. inq. Loasa grandiflora Desv.

LORANTHACEAE Tripodanthus acutifolius (Ruiz Lôpez & Pavôn) Teighem Tristerix chodatianus (Pacz.) Kuijt T. pubescens Kuijt 59

MALVACEAE Acaulimalva crenata (Hill) Krapov. A. rhizantha (A. Gray) Krapov. A. weberbauerii (Ulbr.) Krapov. Nototriche artemesioides Hill N. coccinea Hill N. hillii Krapov. N. obtusa Hill N. pinnata (Cav.) Hill N. aff. porphyrantha Ulbr. N. (ovarii Krapov. Urocarpidium sp. inq.

MELASTOMATACEAE Brachyotum cf. figueroae J. F. Macbr. B. longisepalum Wurdack B. naudinii Triana B. cf. rosmarinifolium (Ruiz Lôpez & Pavôn) Triana B. rostratum (Naud.) Triana Miconia cf. alpina Cogn. M. chionophila Naud. M. salicifolia (Bonpl.) Naud.

MYRICACEAE Myrica parvifolia Benth. M. pubescens Humb. & Bonpl. ex Willd.

MYRSINACEAE Myrsine dependens (Ruiz Lôpez & Pavôn) Spreng. f.

NYCTAGINACEAE Colignonia weberbaueri Heimerl

ONAGRACEAE Epilobium denticulatum Ruiz Lôpez & Pavôn E. fragile Samuelsson Fuchsia denticulata Ruiz Lôpez & Pavôn Oenothera multicaulis Ruiz Lôpez & Pavôn Oenothera spp. — 2 undetermined species.

ORCHIDACEAE Aa matthewsii (Reichb. f.) Schltr. A. paleacea (Kunth) Reichb. f. Epidendrum cernuum Kunth E. excelsum C. Schweinf. E. inamoenum KrSnzlin E. megagastrium Lindley Gomphichis valida Reichb. f. Malaxis andicola (Ridley) O. Ktze. 60

Masdevallia amabilis Reichb. f. & Warsc. Myrosmodes nubigena Reichb. f. M. paludosa (Reichb. f.) D. N. Smith, ined. Odontoglossum rigidum Lindley Pleurothallis spiralis (Ruiz Lôpez & Pavôn) Lindley P. trilineata Lindley Pterichis triloba (Lindley) Schltr. Stelis cf. cupuligera Reichb. f. & Warsc. S. cf. flexuosa Lindley S. cf. leucopogon Reichb. f. Trichoceros platyceros Reichb. f.

OXALIDACEAE Hypsocharis pimpinelifolius Remy Oxalis sp. inq. (ionoxalis group) Oxalis sp. inq. (scandens complex) Oxalis spp. (tuberosa complex) — 3 undetermined species.

PASSIFLORACEAE Passiflora lobbii Masters P. trifoliata Cav.

PHYTOLACCACEAE Phytolacca bogotensis Kunth

PIPERACEAE Peperomia hartwegiana Miquel P. microphylla Kunth P. rotundata Kunth Peperomia spp. — 2 undetermined species.

PLANTAGINACEAE Plantago australis Lam. subsp. hirtella (Kunth) Rahn P. lamprophylla Pilger P. rigida Kunth P. sericea subsp. sericea var. lanuginosa Griseb. P. tubulosa Decne.

POACEAE Aciachne acicularis Laegaard A. pulvinata Benth. Aegopogon cenchroides Humb. & Bonpl. ex Willd. Agrostis auraucana Philippi, vel sp. aff A. breviculmis Hitchc. A. gelida Trin., vel sp. aff. A. nigritella Pilger, vel sp. aff. A. tolucensis Kunth A. aff. tolucensis Kunth Alopecurus aequalis Sobol 61

Anthochloa lepida Nees & Meyen in Meyen Aristida enodis Hackel Bouteloua simplex Lag. Brachypodium mexicanum Link Bromus berterianus Colla B. catharticus Vahl B. lanatus Kunth B. pitensis Kunth B. villosissimus Hitchc. Calamagrostis antoniana (Griseb.) Steud. ex Hitchc. C. brevifolia (Presl) Steud. C. chrysantha (Presl) Steud. C. curvula (Wedd.) Pilger C. densiflora (Presl) Steud. C. eminens (Presl) Steud. C. fuscata (Presl) Steud. C. glacialis (Wedd.) Hitchc. C. heterophylla (Wedd.) Hitchc. C. jamesonii Steud. C. macbridei Tovar C. nitidula Pilger C. ovata (Presl) Steud. C. recta (Kunth) Trin. C. rigida (Kunth) Trin. C. speciformis Hackel ex Stuckert C. spicigera (Presl) Steud. C. swallenii Tovar C. tarmensis Pilger C. vicunarum Wedd. C. weberbaueri Tovar Cortaderia aristata Pilger C. hapalotricha (Pilger) Conert C. nitida (Kunth) Pilger C. sericantha (Steud.) Hitchc. C. trianae Stapf, vel sp. aff. Critesion muticum (Presl) A. LOve Dielsiochloa floribunda (Pilger) Pilger Dissanthelium calycinum (Presl) Hitchc. D. minimum Pilger Elymus angulatus Presl Elytrigia attenuata (Kunth) Covas Eragrostis pastoensis (Kunth) Trin., vel sp. aff. E. pilgeriana Hitchc. Festuca breviaristata Pilger, vel sp. aff F. casapaltensis J. Bail F. dichoclada Pilger F. dolichophylla Presl F. glyceriantha Pilger, vel sp. aff. F. huamachucensis Infanta, vel sp. aff. 62

Festuca lasiorachis Pilger F. pubigluma Tovar F. guadridentata Kunth F. rigidifolia Tovar F. ulochaeta Steud., vel sp. aff F. weberbauerii Pilger Hierochloê redolens (Sol. ex Vahl) Roem. & Schult. Melica scabra Kunth Muhlenbergia angustata (Presl) Kunth M. fastigata (Presl) Henrard M. flexuosa Hitchc., vel sp. aff. M. ligularis (Hackel) Hitchc. M. peruviana (Beauv.) Steud. Muhlenbergia sp. inq. Nassella brachyphylla (Hitchc.), comb. ined. N. depauperata (Pilger), comb. ined. N. featherstonei (Hitchc.), comb. ined. N. inconspicua (Presl), comb. ined. N. mexicana (Hitchc.), comb. ined. N. mucronata (Kunth), comb. ined. Paspalum bonplandianum FlOgge P. pygmaeum Hackel P. tuberosum Mez Pennisetum clandestinum Hochst. ex Chiov. Piptochaetium indutum Parodi P. panicoides (Lam.) Desv. Poa aff. aeguatoriensis Hackel P. aff aequigluma Tovar P. annua L. P. aff. brevis Hitchc. P. aff. candamoana Pilger P. aff. fibrifera Pilger P. aff. ferreyrae Tovar P. aff. gilgiana Pilger P. aff. glaberrima Tovar P. aff. horridula Pilger P. aff. lilloi Hackel P. aff. pardoana Pilger P. aff. pearsonii Reeder P. aff. perligulata Pilger P. aff. subspicata (Presl) Kunth Poa sp. inq. Polypogon interruptus Kunth Schizachyrium sanguineum (Retz) Alston Sporobolus lasiophyllus Pilger Stipa hans-meyeri Pilger S. ichu (Ruiz Lôpez & Pavôn) Kunth S. obtusa (Nees & Meyen) Hitchc. 63

Trisetum macbridei Hitchc. T. spicatum (L.) Richt. Vutpia bromoides (L.) S. R. Gray v. myuros (L.) K. Gmelin var. hirsuta Hackel

POLEMONIACEAE Gilia laciniata Ruiz Lâpez & Pavôn Microsteris gracilis (Hook.) Greene

POLYGALACEAE Monnina conferta Ruiz Lôpez & Pavôn M. salicifolia Ruiz Lôpez & Pavôn

POLYGONACEAE Muehlenbeckia tamnifolia (Kunth) Meisn. M. volcanica (Benth.) Engl. Rumex acetosella L. R. obtusifoiius L.

PORTULACACEAE Calandrinia acaulis Kunth C. ciliata (Ruiz Lôpez & Pavôn) DC. Portulaca peruviana L M. Johnston

POTAMOGETONACEAE Potamogeton paramoanus Haynes & Holm-Nielsen

PROTEACEAE Oreocallis grandiflora (Lam.) R. Br.

RANUNCULACEAE Clematis sp. inq. Krapfia aff. weberbaueri (Ulbr.) Standley & J. F. Macbr. Ranunculus cymbalaria Pursh R. flagelliformis Smith R. limoselloides Turcz. R. peruvianus Pers. R. praemorsus Kunth ex DC. var praemorsus Thalictrum longistylum DC.

RHAMNACEAE Colletia spinossima Gmelin

ROSACEAE Acaena ovalifolia Ruiz Lôpez & Pavôn Alchemilla diplophylla Diels A. orbiculata Ruiz Lôpez & Pavôn A. paludicola Rothm. A. pinnata Ruiz Lôpez & Pavôn 64

Alchemilla procumbens Rose A. verticillata Fielding & Gardner Hesperomeles cuneata Lindley Kageneckia lanceolate Ruiz Lôpez & Pavôn Margyricarpus pinnatus (Lam.) Kuntze Polylepis incana Kunth P. racemosa Ruiz Lôpez & Pavôn P. sericea Wedd. P. weberbauerli Pilger Potentilla dombeyi Nestler Prunus serrotina Ehrh. subsp. capuli (Cav.) McVaugh Rubus bogotensis Kunth R. sparcifolius J. F. Macbr.

RUBIACEAE Arcythophyllum thymifolium (Ruiz Lôpez & Pavôn) Standley Arcythophyllum sp. inq. Galium sp. inq. Relbunium hypocarpum (L.) Hemsley Relbunium sp. inq.

SANTALACEAE Quinchamalium chilense Lam.

SAPINDACEAE Dodonaea viscosa L.

SAXIFRAGACEAE Saxifraga magellanica Poiret

SCROPHULARIACEAE Agalinis lanceolata (Ruiz Lôpez & Pavôn) D'Arcy Alonsoa linearis (Jacq.) Ruiz Lôpez & Pavôn Bartsia diffusa Benth. B. santolinifolia (Kunth) Benth. Bartsia sp. inq. Calceolaria bicrenata Ruiz Lôpez & Pavôn C. cajabambae KrSnzlin C. deflexa Ruiz Lôpez & Pavôn subsp. deflexa C. glauca Ruiz Lôpez & Pavôn C. inaudita Krflnzlin C. incarum Krftnzlin C. linearis Ruiz Lôpez & Pavôn C. lobata Cav, C. nivalis Kunth C. phaceliaefolia Edwin C. triloba Edwin C. utricularioides Benth. C. virgata Ruiz Lôpez & Pavôn 65

Calceolaria viscosa Ruiz Lôpez & Pavôn C. weberbaueriana Krflnzlin Calceolaria spp. — 3 undetermined species. Castilleja cerroana Edwin C. fissifolia L. f. C. nubigena Kunth C. pumila (Benth.) Wedd. ex Herrera C. virgata (Wedd.) Edwin Limosella subulata Ives Mimulus glabratus Kunth Orthocarpus laciniatus (Hook. & Am.) Keck Ourisia chamaedrifolia Benth. in DC; O. muscosa Benth. in DC. Porodittia triandra (Cav.) D. Don Sibthorpia repens (Mutis ex L.) Kuntze Veronica anagallis-aquatica L. V. persica Poiret V. serpyllifolia L.

SOLANACEAE lochroma umbellate (Ruiz Lôpez & Pavôn) Miers Jaltomata dentata (Ruiz Lôpez & Pavôn) Bénitez Jaltomata sp. inq. Lycianthes lycioides (L.) Hassl. Lycopersicon hirsutum Kunth Nicotiana thyrsiflora Bitter ex Goodsp. Salpichroa glandulosa (Hook.) Miers S. ramosissima Miers Salpichroa sp. inq. Saracha spinosa (Dammer) D'Arcy & D. N. Smith Solanum albicans (Ochoa) Ochoa S. aloysiifolium Dunal, vel sp. aff. S. amblophyllum Hook. S. ambosinum var. orophilum (Correll) Ochoa S. ancophyllum (Correll) Ochoa S. caripense Humb. & Bonpl. ex Dun. S. chomatophilum Bitter S. dolichocremastrum Bitter S. cf. fragile Wedd. S. cf. furcatum Dunal S. glutinosum Dunal S. hastiforme Correll S. nitidum Ruiz Lôpez & Pavôn S. nutans Ruiz Lôpez & Pavôn S. sorgarandinum Ochoa Solanum sp. inq.

SYMPLOCACEAE Symplocos sandiae Brand, vel sp. aff. (ca Weberbauer 7024) 66

URTICACEAE Pilea dauciodora (Ruiz Lôpez & Pavôn) Wedd. Urtica echinata Benth. U. flabellata Kunth U. aff. flabellata Kunth U. magellanica A. L. Juss. ex Poiret

VALERIANACEAE Phyllactis rigida (Ruiz Lôpez & Pavôn) Pen. Phyllactis spp. — 2 undetermined species. Stangea erikae Graebn. S. henricii Graebn. Valeriana condamoana Graebn. v. connata Ruiz Lôpez & Pavôn V. aff. decussata Ruiz Lôpez & Pavôn V. globularis A. Gray V. interrupta Ruiz Lôpez & Pavôn V. nivalis Wedd. V. pilosa Ruiz Lôpez & Pavôn V. pycnantha A. Gray V. aff. sphaerocephala Graebn. V. weberbaueri Graebn. Valeriana spp. — 3 undetermined species.

VERBENACEAE Citharexylum dentatum (Tafalla) D. Don Duranta mandonii Mold., vel sp. aff. Glandularia microphylla (Kunth) Schnack & Covas Lantana reptans Hayek Verbena littoralis Kunth

VIOLACEAE Viola micranthella Wedd. V. pygmaea Juss. ex Poiret

ZANNICHELLIACEAE Zannichellia andina Holm-Nielson & Haynes 67

BIOGEOGRAPHIC RELATIONSHIPS OF THE HUASCARÂN FLORA Based on the genera included in the flora of the Huascardn National Park, an evaluation of their worldwide geographic relationships has been made (data in List 1). Genera have been assigned to the following categories according to the geographic distribution of their component species: American: Genera which are native to the Western Hemisphere alone. This category is broad, and includes several distribution patterns in the New World, including Neotropical, North and South Temperate, Western Cordilleran, Antillean, Andean, or Peruvian Endemic. Given the overview nature of this analysis, those more precise distribution patterns are not useful here. Two subgroups which are useful to this analysis are: Andean: Those genera whose distribution is centered in, or restricted to the Andes. Endemic: Genera whose distribution is restricted to Peru. Austral: Genera which have the bulk of their species in the Southern Hemisphere, i.e., southern South America, South Africa, the Antarctic Islands, New Zealand, Australia and associated islands, and, occasionally, the Pacific Islands and/or Japan. CosmonoHtan: The species of these are broadly distributed throughout the world. Holarctic: The majority of species from these genera inhabit the Northern Hemisphere, with some extensions into tropical highlands. Ruderal: This includes the genera that are introduced and weedy in Peru. Wide Temperate: Included here are the genera whose species are widely distributed in the temperate regions of the World, including the Northern and Southern Hemispheres, and tropical highlands, and are present on two or more extra-American continents. Genera assigned to this category cannot be reliably placed in either the Austral or Holarctic distribution patterns. Wide Tropical: Genera native to the tropical regions of more than one continent, i.e., America-Asia, America-Africa, or pantropical. The distributional relationships of the genera included in the Huascaràn flora are summarized below. Genera of American distribution dominate the flora, with those of Andean distribution making the greatest contribution. Elements of other distinctly temperate areas (Wide Temperate, Holarctic, and Austral) together contribute another 29.8% of the genera. Since the study site is located in South America, and at the highest 68

elevations of the tropical Andes, it is not surprising that the flora would be dominated by both American and temperate floristic elements. Nor is it surprising that tropical genera have contributed 10.6% of the flora. Biogeographical relationships of the genera included in the Huascardn National Park flora are: Recion of concentration No. of genera % of total flora American 162 47.8 Andean 89 26.3 Endemic S 1.5 Wide Temperate 58 17.1 Wide Tropical 36 10.6 Cosmopolitan 27 8.0 Holarctic 27 8.0 Austral 19 5.6 Ruderal 10 2.9

The above categories are distribution categories, and not specifically categories of origin, although in many instances the distribution category implies area of origin. Study of the origin and dispersal of the angiosperms is complex and is discussed in detail elsewhere: Gentry (1982), Raven (1979), and Raven and Axelrod (1974) are recommended. In considering the phytogeographical relationships of the Central and South American floras. Gentry (1982) found the lowland tropical floras dominated by families of Gondwanan origin. As the Isthmus of Panama neared completion, the Gondwanan families from South America diffused throughout the Central American lowlands, displacing the Laurasian families which are now found at middle and higher elevations. There were counter-migrations of the Laurasian families southward. These migrations had little effect on the lowland tropical flora, but added a large component of Laurasian families to the South American upland flora. Following Gentry's classification of the gymnosperm and angiosperm families, in Huascarân National Park: 48.8% were of Gondwanan origin, 29.8% of Laurasian origin, and 21.4% unclassified (List 2). In a further sub-classification of the Gondwanan families. Gentry found that the families fell into three distributional groups: Amazonian-centered, Andean-centered, and dry-area- centered. Following that scheme, the 48.8% of the families found to be of Gondwanan 69

origin included 34.5% Andean-centered, 11.9% Amazonian-centered, and 2.4% dry- area-centered. With respect to the vegetation of Peru, the flora of the Huascardn National Park shares floristic elements with the puna, jalca, paramo, and ceja de selva. Puna is is the characteristic vegetation of the high Andes from central Peru to northern Argentina. It is dry, with aridity increasing south and westward, and the vegetation is either grassland essentially devoid of woody vegetation, or patches of a turf of low growing herbs alternating with patches of bare soil (Weberbauer, 1936). Genera typical of the puna are: Aciachne, Anthochloa, Areneuria, Azoreîla, Brayopsis, Catadysia, Cremolobus, Distichia, Draba, Mancoa, Nototriche, Pycnophyllum, Senecio (Culcitium), Stangea, Weberbauera, and Werneria. In the vegetation of the Park, these are common at the highest elevations. Calamagrostis and Festuca are dominants in the puna, and are also important in the paramo and jalca. Ceja de selva, paramo, and jalca are vegetation types of the humid Andes. Ceja is the transition zone between the montane forest and the high elevation grasslands and shrublands, and is usually located between 3400-3600 m on the eastern slopes (Weberbauer, 1936). Alnus, Begonia, Berberis, Bomarea, Citharexylum, Duranta, Escallonia, Fuchsia, Gynoxys, Hesperomeles, Histiopteris, Miconia, Momina, Myrica, Oreopanax, Passi/lora, Piagiogyria, Symplocos, Vaccinium, Vailea, and Weinmannia are evidence of the influence of the ceja in the Huascarân flora. Two important ceja or paramo elements absent from the Huascaràn flora are Chusquea and a luxuriant array of epiphytic and terrestrial bryophytes. Paramo and jalca are moist grasslands. The former is very humid, and has many elements of the montane forest and more woody species. It is distributed from Venezuela to Ecuador, and then along the eastern edge of the Andes and on the tops of eastward outlying ranges to Bolivia. The latter, which occupies the crest of the Andes in northern Peru, has fewer montane elements and fewer woody species. It is often spoken of as a transitional vegetation type between paramo and puna. In comparing the list of paramo species by Cleef (1981) with the species list of the Park, 86% of the families, 72% of the genera, and 24% of the species in the paramo flora are in common with the Huascaràn flora (List 3 and 4). All the families in the paramo flora not shared with Huascarân are either common in, or typical of the montane forests: Aquifoliaceae, Arecaceae, Clethraceae, Eriocaulaceae, Hymenophyllaceae, 70

Lentibulariaceae, Marsiliaceae, Melanthiaceae, Myrtaceae, Theaceae, and Xyridaceae. The genera Espeletia and Espeletiopsis are characteristic elements of the paramo of the northern Andes that are represented in neither jalca nor puna. The floras of the paramos and of the Park are closely related, and both are distinctly equatorial in nature. Making a similar comparison with the flora of the puna of northern Argentina (Ruthsatz, 1977), 82% of the families, 50% of the genera, and 14% of the species were shared with the Huascarân flora (List S). Most of the families characteristic of the Argentinan puna but not found in the Huascarta flora are those which are more common in arid regions: Acanthaceae, Anacardiaceae (Schinus), Calyceraceae, Chenopodiaceae, Cucurbitaceae, Krameriaceae, Mimosaceae, Oleaceae, and Zygophyllaceae. In addition to these both the Marsileaceae and the Selaginellaceae appear in the puna, but not the Huascarân flora; these families have adapted to both extremely humid and arid habitats. The Andean vegetation is often compared with the high elevation vegetation of both East Africa and New Guinea. Although both the East African and New Guinea sites have base levels similar to that of the Huascarân National Park (ca. 3600 and 3000 meters above sea level, respectively), the upper limits of plant growth do not reach altitudes comparable with Huascarân. The information on the East African high elevation flora (Hedberg, 1957) is complete for all vascular plants, but limited to the "alpine" formation, which is above the montane forest and the ericaceous belt. The study of the New Guinea sites (van Royen, 1979a, 1979b, 1982, 1983), includes all vascular plants except the Pteridophytes, and surveys the Upper Montane, Subalpine, and Alpine Zones. Although the flora of Africa has been impoverished by the effects of continental drift (Raven and Axelrod, 1974), the close relationship of that continent to South America, as well as the facility of introduction of wide temperate elements, at least from Eurasia in the north, would lead one to expect a high degree of similarity between the high Andean and the "Afroalpine" floras. Comparison of the Huascarân species list with the one from East Africa (Hedberg, 1957) shows that 89% of the families and 39.5% of the genera in the "Afroalpine" also occur in the Huascarân flora (List 6). Senecio and Lobelia are prominent genera in the "Afroalpine" flora, which have evolved many species in Africa and life forms convergent with those characteristic of the high Andes, 71

especially of the paramos. Despite the convergence of lifeforms, the "afroalpine" is very dry, and is more appropriately the analogue of the puna than the paramo. When reading the floristic list of the "alpine" flora of New Guinea, the families and genera will be very familiar to botanists who have collected in the Peruvian ceja de selva. The high altitude areas of the island have a high annual precipitation (Barry, 1979), and the lower limit of the study area is 3000 m (van Royen, 1979a). As presented, the "alpine" flora of New Guinea includes upper montane, subalpine, and alpine zones, which are analogous to the high montane forest, ceja de selva and the paramo in the northern and central Andes. The New Guinea "alpine" has 57.8% of its families and 24.8% of the genera in common with the Huascardn flora (List 7). In summary, the Huascarân flora is an equatorial temperate flora principally of American origin, many of the genera being strictly Andean. And, it shows the Gondwanan origin of many of the component families, as well as a strong Laurasian component received with the joining of North and South America. Given the continuity of the Andes, it is closely related to the floras of the paramos to the north and east, and to the puna to the south; of these, the most intimate relationship is found with the paramos. 72

BIOGEOGRAPHIC LISTS^ Page List 1. Geographical affinity of the genera included in the flora of the Huascaràn National Park 73 List 2. Phytogeographic origin of the gymnosperm and angiosperm families included in the Huascardn National Park flora 86 List 3. Families and genera of the Huascardn National Park flora 90 List 4. Families and genera of the Columbian paramos 96 List 5. Families and genera of the Argentinian puna 100 List 6. Families and genera of the Afroalpine flora 105 List 7. Families and genera of the New Guinea alpine flora 108

^ The number following the generic name in List 1 indicates the number of species in the genus, and in Lists 3 through 7, the number of species of the genus that are included in each specific flora. 73

List 1. Geographical affinity of the genera included in the flora of the Huascarân National Park.

TAXQN AtTINKTY SOURCES GYMNOSPERMS EPHEDRACEAE Ephedra (40) Holarctic Cronquist et al., 1972

PTERIDOPHYTES ASPLENIACEAE Asplenium (650) Cosmopolitan Cleef, 1979; Tryon and Tryon, 1982. BLECHNACEAE Blechnum (150) Cosmopolitan Cleef, 1979; Tryon and Tryon, 1982. DENNSTAEDTIACEAE Histiopteris (7) Wide Tropical Tryon and Tryon, 1982; Willis, 1973. Hypolepis (40) Andean Tryon and Tryon, 1982. DRYOPTERIDACEAE Athyrium (100) Holarctic Tryon and Tryon, 1982. Cystopteris (6) Wide Temperate Cleef, 1979; Tryon and Tryon, 1982. Elaphoglossum (500) Wide Tropical Cleef, 1979; Tryon and Tryon, 1982. Polystichum (160) Wide Temperate Cleef, 1979; Tryon and Tryon, 1982. Woodsia (25) Holarctic Tryon and Tryon, 1982. EQUISETACEAE Equisetum (15) Wide Temperate Cleef, 1979; Tryon and Tryon, 1982. GLEICHENIACEAE Gleichenia (110) Wide Tropical Tryon and Tryon, 1982. ISOETACEAE Isofltes (150) Wide Temperate Cleef, 1979; Tryon and Tryon, 1982. LYCOPODIACEAE Huperzia (300) Cosmopolitan 011gaard, 1987. Lycopodium (40) Cosmopolitan Cleef, 1979; Ollgaard, 1987. OPHIOGLOSSACEAE Ophioglossum (30) Cosmopolitan Cleef, 1979; Tryon and Tryon, 1982. PLAGIOGYRIACEAE Plagiogyria (50) Wide Tropical Tryon and Tryon, 1982. POLYPODIACEAE Campyloneurum (25) Andean Tryon and Tryon, 1982. Grammitis (400) Wide Tropical Tryon and Tryon, 1982. Microgramma (13) American Tryon and Tryon, 1982. Polypodium (150) Wide Tropical Tryon and Tryon, 1982. 74

List 1. (continued)

PTERIDACEAE Adiantum (ISO) Wide Tropical Tryon and Tryon, 1982. Cheilanthes (ISO) Wide Tropical Cleef, 1979; Tryon and Tryon, 1982. Eriosorus (2S) Andean Tryon and Tryon, 1982. Jamesonia (19) Andean Tryon and Tryon, 1982. Notolaena (39) American Tryon and Tryon, 1982. Pellaea (3S) Wide Tropical Tryon and Tryon, 1982. THELYPTERIDACEAE Thelypteris (800) Cosmopolitan Tryon and Tryon, 1982. SALVINIACEAE Azolla (6) Cosmopolitan Cleef, 1979; Tryon and Tryon, 1982.

ANGIOSPERMS AGAVACEAE Furcraea (20) American Dahlgren et al., 198S ALLIACEAE Trichlora (1) Endemic Weberbauer, 1945. ALSTROEMERIACEAE Alstroemeria (60) American Dahlgren et al., 198S Bomarea (175) Andean Weberbauer, 1945. AMARANTHACEAE Alternanthera (200) Wide Tropical Eliasson, 1987; van Balgooy, 1971. AMARYLLIDACEAE Stenomesson (40) Andean A. Meerow, pers. com.; Weberbauer, 1945. ANTHERICACEAE Echeandia (100) American R. Cruden, pers com. APIACEAE Azorella (70) Austral Cleef, 1979; Mathias and Constance, 1976; Weberbauer, 1945; Willis, 1973. Bowlesia (14) Andean Mathias and Constance, 1962, 1976; Weberbauer, 1945. Daucus (25) Ruderal Eryngium (200) Cosmopolitan Cleef, 1979; Mathias and Constance, 1962, 1976. Lilaeopsis (13) Austral Affolter, 1985; Cleef, 1979; Mathias and Constance, 1976. Niphogeton (16) Andean Mathias and Constance, 1962, 1976. Oreomyrrhis (23) Austral Cleef, 1979; Mathias and Constance, 1962, 1976; van Balgooy, 1971; Weberbauer, 1945. ARALIACEAE Oreopanax (120) Andean Weberbauer, 1945; Willis, 1973. 75

List 1. (continued)

ASCLEPIADACEAE Cynanchum (150) Cosmopolitan van Balgooy, 1971; Willis, 1973. Metastelma (100) American van Balgooy, 1971; Willis, 1973. ASTERACEAE Achyrocline (20) Wide Tropical Cleef, 1979; Dillon, pers. com. Ageratina (230) American Cleef, 1979; Robinson and King, 1977. Antennaria (50) Holarctic MerxmQller et al, 1977. Aphanactis (4) Andean Cleef, 1979. Aristeguieta (20) Andean Robinson and King, 1977. Baccharis (400) American Cleef, 1979; Grau, 1977. Barnadesia (22) Andean Cabrera, 1977; Weberbauer, 1945. Belloa (18) Andean Cabrera, 1958; Sagâstegui and Dillon, 1985. Bidens (230) Cosmopolitan Cleef, 1979; Stuessy, 1977. Chaptalia (50) American Cabrera, 1977; Cleef, 1979. Chersodoma (9) Andean Cabrera, 1946; Nordenstam, 1977. Chrysactenium (6) Andean Robinson, 1983. Chuquiraga (19) Andean Ezcurra, 1985. Conyza(50) Cosmopolitan Grau, 1977. Coreopsis (114) Wide Tropical Stuessy, 1977. Cosmos (26) American Stuessy, 1977. Cotula (90) Austral Cleef, 1979; Dillon, 1981. Cuatrecasasiella (4) American MerxmQller et al., 1977. Dasyphyllum (36) American Cabrera, 1977. Diplostephium (75) Andean Weberbauer, 1945. Erigeron (200) Wide Temperate Grau, 1977; van Balgooy, 1971. Galinsoga (14) American Stuessy, 1977. Gamochaeta (30) American Willis, 1973. Gnaphalium 0^0) Wide Tropical Cleef, 1979; MerxmQller et al., 1977. Grosvenoria (3) Andean Robinson and King, 1977. Gynoxys (100) Andean Nordenstam, 1977; Weberbauer, 1945. Helianthopsis (35) Andean M. 0. Dillon, pers. com. Hieracium (1000) Wide Temperate Cleef, 1979; Tomb, 1977. Hypochoeris (100) Holarctic Cleef, 1979; Tomb, 1977. Jungia (30) American Cabrera, 1977. Leuceria (79) Andean Weberbauer, 1945. Liabum (37) American Robinson, 1983. Lophopappus(5) Andean Cabrera, 1977; Weberbauer, 1945. Loricaria (17) Andean MerxmQller et al., 1977. Lucilia (20) American MerxmQller et al., 1977. Mnodes (5) Andean M. Dillon, pers com. Munnozia (40) Andean Robinson, 1983. Mutisia (59) American Cabrera, 1977. Noticastrum (19) American Zardini, 1985. Novenia (1) Andean Freire, 1986. Ophryosporus (38) Andean Robinson and King, 1977; Weberbauer, 1945. 76

List 1. (continued)

Oritrophium (15) Andean Grau, 1977. Paranephelius (7) Andean Robinson, 1983. Pentacalia (60) Andean M. O. Dillon, pers. com. Perezia (85) Andean Weberbauer, 1945. Perymenium (26) American Stuessy, 1977. Plagiocheilus (8) Andean Weberbauer, 1945. Polymnia (20) American Stuessy, 1977. Pseudonoseris (3) Endemic Robinson, 1983. Senecio (1600) Wide Temperate Cleef, 1979; Nordenstam, 1977; van (incl. Aequatorium, Aetheolaena, Culcitium, Balgooy, 1971. Dorobea, and Lasiocephalus) Siegesbeckia (9) Wide Tropical Cleef, 1979; Stuessy, 1977. Sonchus (50) Ruderal Stevia (200) American Robinson and King, 1977. Stuckertiella (2) American Merxmûller et al., 1977. Tagetes (50) American Strother, 1977. Vasquezia (10) American Turner and Powell, 1977. Verl^sina (150) American Stuessy, 1977. Viguiera (150) American Stuessy, 1977. Werneria (40) Andean Nordenstam, 1977; Weberbauer, 1945. BASELLACEAE Ullucus (1) Andean Weberbauer, 1945. BEGONIACEAE Begonia (1000) Wide Tropical Smith and Wasshausen, 1986. BERBERIDACEAE Berberis (450) Holarctic Cleef, 1979; Weberbauer, 1945; Willis, 1973. BETULACEAE AInus (35) Holarctic Weberbauer, 1945; Willis, 1973. BIGNONIACEAE Tecoma (12) Andean Gentry, 1977. BORAGINACEAE Hackelia (45) Holarctic Cleef, 1979; Cronquist et al., 1984. Lithospermum (75) Wide Temperate Cronquist et al., 1984. Plagiobotrys (50) American Cronquist et al., 1984; Weberbauer, 1945. BRASSICACEAE Brassica (30) Ruderal Brayopsis (4) Andean Weberbauer, 1945. Capsella (6) Ruderal Cardamine (100) Wide Temperate Cleef, 1979; Standley and Steyermark, 1946a; van Balgooy, 1971; Weberbauer, 1945. Catydysia (1) Endemic Weberbauer, 1945. Cremolobus (7) Andean Willis, 1973. 77

List 1. (continued)

Descurainia (40) Holarctic Standiey and Steyermark, 1946a; Weberbauer, 1945. Draba (250) Holarctic Cleef, 1979; Standiey and Steyermark, 1946a; Weberbauer, 1945. Lepidium (120) Wide Temperate Standiey and Steyermark, 1946a. Mancoa (4) Andean Weberbauer, 1945. Weberbauera (2) Andean Weberbauer, 1945. BROMELIACEAE Pitcairnia (260) American Smith and Downs, 1974. Puya (170) Andean Smith and Downs, 1974. Tillandsia (400) American Smith and Downs, 1977. BUDDLEIACEAE Buddleia (100) Wide Tropical Norman, 1982. CACTACEAE Cleistocactus (?) American lOS Working Party, 1986. Cereus (30) American lOS Working Party, 1986. Opuntia (200) American lOS Working Party, 1986. Oreocereus (IS) Andean lOS Working Party, 1986. CAESALPINIACEAE Senna (240) Wide Tropical Irwin and Barneby, 1981. CALLITRICHIACEAE Callitriche (35) Wide Temperate Cleef, 1979; van Balgooy, 1971. CAMPANULACEAE Hypsela (4) Austral Jeppesen, 1981b; van Balgooy, 1971; Weberbauer, 1945. Lobelia (400) Cosmopolitan Cleef, 1979; Jeppesen, 1981b. Lysipomia (22) Andean Jeppesen, 1981b; McVaugh, 1955; Weberbauer, 1945 Siphocampylus (200) Andean Jeppesen, 1981b. Wahlenbergia (150) Austral Jeppesen, 1981a; van Balgooy, 1971; Weberbauer, 1945. CAPPARACEAE Cleome (75) Wide Tropical Standiey and Steyermark, 1946a. CARYOPHYLLACEAE Arenaria (170) Wide Temperate Cleef, 1979; Standiey and Steyermark, 1946a. Cardionema (6) American Willis, 1973. Cerastium (SO) Holarctic Cleef, 1979; Standiey and Steyermark, 1946a. Drymaria (48) American Cleef, 1979; Duke, 1961. Paronychia (109) Holarctic Chaudhri, 1968; van Balgooy, 1971. Plettkea (4-5) Endemic Weberbauer, 1945. Pycnophyllum (IS) Andean Weberbauer, 1945. 78

List 1. (continued)

Silene (250) Holarctic Standley and Steyermark, 1946a; van Balgooy, 1971. Spergularia (6) Ruderal Stellaria (75) Wide Temperate Cleef, 1979; Standley and Steyermark, 1946a; van Balgooy, 1971; Weberbauer, 1945. CLUSIACEAE Hypericum (400) Wide Temperate Cleef, 1979; van Balgooy, 1971. COLUMELLIACEAE Columellia (5) Andean Brizicky, 1961; Weberbauer, 1945. COMMELINACEAE Commelina (170) Cosmopolitan Dahlgren et al., 1985; van Balgooy, 1971. CONVOLVULACEAE Dichondra (15) Wide Temperate Austin, 1982; van Balgooy, 1971. CORIARIACEAE Coriaria (5) Wide Temperate Skog, 1972, 1987; van Balgooy, 1971. CRASSULARIACEAE Echeverria (90) American Standley and Steyermark, 1946a. Tillaea (20) Wide Temperate Cleef, 1979; Standley and Steyermark, 1946a; van Balgooy, 1971. Villadia (25) American Standley and Steyermark, 1946a. CUNONIACEAE Weinmannia (125) Austral Standley and Steyermark, 1946a; van Balgooy, 1971; Weberbauer, 1945. CUSCUTACEAE Cuscuta (150) Cosmopolitan Austin, 1982; van Balgooy, 1971. CYPERACEAE Baeothryon (10) Holarctic Goetghebeur, 1986. Bulbostylis (200) Wide Tropical Cleef, 1979; Goetghebeur, 1986. Carex (2000) Wide Temperate Cleef, 1979; Goetghebeur, 1986; Weberbauer, 1945. Cyperus (300) Wide Tropical Cleef, 1979; Goetghebeur, 1986. Eleocharis (200) Cosmopolitan Cleef, 1979; Goetghebeur, 1986; van Balgooy, 1971. Isolepis (60) Cosmopolitan Goetghebeur, 1986. Oreobolopsis (1) Andean Koyama and Guaglianone, 1987. Oreobolus (14) Austral Cleef, 1979; Seberg, 1988; van Balgooy, 1971. Phylloscirpus (5) Andean Goetghebeur, 1986. Schoenoplectus (50) Cosmopolitan Goetghebeur, 1986. DIOSCORIACEAE Dioscorea (600) Wide Tropical Dahlgren et al., 1985. 79

List 1. (continued)

ELAEOCARPACEAE Vallea (3) Andean Weberbauer, 1945. ELATINACEAE Elatine (20) Wide Temperate Cleef, 1979; van Balgooy, 1971. ERICACEAE GauUheria (SO) Austral Cleef, 1979. Pernettya (13) Austral Cleef, 1979; van Balgooy, 1971 Vaccinium (140) Holarctic Cleef, 1979; van Balgooy, 1971; Weberbauer, 1945. EUPHORBIACEAE Chamaesyce (250) Cosmopolitan Willis, 1973. Croton (750) Cosmopolitan van Balgooy, 1971; Willis, 1973. Euphorbia (2000) Cosmopolitan Cleef, 1979; van Balgooy, 1971; Willis, 1973. Sebastiania (75) Cosmopolitan van Balgooy, 1971; Willis, 1973. FABACEAE Astragalus (2000) Holarctic Polhill, 1981; Weberbauer, 1945. Dalea (160) American Barneby, 1981; Weberbauer, 1945. Lathyrus (150) Holarctic Cleef, 1979; Kupicha, 1981; Weberbauer, 1945. Lupinus (200) Wide Temperate Bisby, 1981. Otholobium (35) Wide Tropical J. W. Grimes, pers. com. Spartium (1) Ruderal Trifolium (250) Holarctic Cleef, 1979; Heyn, 1981; Weberbauer, 1945. Vicia (140) Holarctic Cleef, 1979; Kupicha, 1981; van Balgooy, 1971; Weberbauer, 1945. FLACOURTIACEAE Pineda (1) Andean Sleumer, 1980; Weberbauer, 1945. GENTIANACEAE Gentiana (200) Wide Temperate Cleef, 1979; Cronquist et al., 1984; Weberbauer, 1945. Gentianella (100) Wide Temperate Cleef, 1979; Cronquist et al., 1984. Halenia (60) Wide Temperate Standley and Williams, 1969. GERANIACEAE Erodium (60) Ruderal Geranium (300) Wide Tropical Cleef, 1979; Standley and Steyermark, 1946b.; Weberbauer, 1945. GROSSULARIACEAE Escallonia (50) Austral Cleef, 1979. Ribes (150) Holarctic Cleef, 1979; Standley and Steyermark, 1946a; Weberbauer, 1945. 80

List 1. (continued)

HALORAGACEAE Myriophyllum (35) Wide Temperate Standley and Williams, 1963. HYDROCHARITACEAE Elodea (5) American Cook and Urmi-KOnig, 1985. HYDROPHYLLACEAE Phacelia (150) American Cronquist et al., 1984; Weberbauer, 1945. IRIDACEAE Orthrosanthus (9) Austral Cleef, 1979; Henrich and Goldblatt, 1987a; Weberbauer, 1945. Sisyrinchium (80) American Henrich and Goldblatt, 1987b. Tigridia (15) American Molseed, 1970. JUNCACEAE Distichia (3) Andean Balslev, 1979a; Weberbauer, 1945. Juncus(220) Wide Temperate Blaslev, 1979a; Cleef, 1979. Luzula (75) Wide Temperate Balslev, 1979a; Cleef, 1979; Weberbauer, 1945. JUNCAGINACEAE Lilaea (1) American Haynes and Holm-Nielsen, 1986. LAMIACEAE Gardoquia (33) Andean Weberbauer, 1945. Lepechinia (55) Andean Weberbauer, 1945. Salvia (500) Cosmopolitan Cronquist et al., 1984. Scutellaria (300) Cosmopolitan Cronquist et al., 1984 Stachys (200) Holarctic Cleef, 1979; Cronquist et al., 1984. LEMNACEAE Lemna (13) Cosmopolitan Landolt, 1986. LINACEAE Linum (80) Wide Temperate Standley and Steyermark, 1952; van Balgooy, 1971. LOASACEAE Cajophora (50) Andean Weberbauer, 1945. Loasa (81) Andean Weberbauer, 1945. LORANTHACEAE Tripodanthus (2) American Kuijt, 1986. Tristerix (8) Andean Kuijt, 1988. MALVACEAE Acaulimalva (19) Andean Krapovickas, 1974. Nototriche (62) Andean Weberbauer, 1945. Urocarpidium (1) Andean Weberbauer, 1945. MELASTOMATACEAE Brachyotum (50) Andean Weberbauer, 1945; Wurdack, 1980. Miconia (1000) American Cleef, 1979; Wurdack, 1980. 81

List 1. (continued)

MYRICACEAE Myrica (40) Wide Temperate Cleef, 1979; Standley and Steyermark, 1952. MYRSINACEAE Myrsine (275) Wide Tropical J. J. Pipoly, pers. com. NYCTAGINACEAE Colignonia (11) Andean Weberbauer, 1945. ONAGRACEAE Epilobium (185) Wide Temperate Cleef, 1979; Solomon, 1982; van Balgooy, 1971; Weberbauer, 1945. Fuchsia (100) Austral Berry, 1982; Cleef, 1979; van Balgooy, 1971. Oenothera (80) Holarctic Cleef, 1979; Willis, 1973. ORCHIDACEAE Aa (30) Andean Garay, 1978; Weberbauer, 1945. Epidendrum (825) American Cleef, 1979; C. Dodson, pers. com. Gomphichis (20) Andean Garay, 1978; C. Dodson, pers. com. Malaxis (300) Wide Tropical Atwood, 1986; van Balgooy, 1971. Masdevallia (350) American Luer, 1986a. Myrosmodes (12) Andean Garay, 1978; C. Dodson, pers. com. Odontoglossum (244) Andean Weberbauer, 1945. Pleurothallis (1500) American Atwood, 1986; Luer, 1986c. Pterichis (15) Andean Garay, 1978; Weberbauer, 1945. Stelis (500) American Luer, 1986a. Trichoceros (5) Andean Dodson, pers. com.; Weberbauer, 1945. OXALIDACEAE Hypsocharis (8) Andean Weberbauer, 1945. Oxalis (800) Cosmopolitan Cleef, 1979; Standley and Steyermark, 1946b. PASSIFLORACEÀE Passiflora (400) American Holm-Nielsen et al., 1988. PHYTOLACCACEAE Phytolacca (25) Wide Tropical Cleef, 1979; Standley and Steyermark, 1946a. PIPERACEAE Peperomia (1000) Wide Tropical Cleef, 1979; Standley and Steyermark, 1952. PLANTAGINACEAE Plantago (250) Wide Temperate Cleef, 1979; Weberbauer, 1945. POACEAE Aciachne (3) Andean Lxgaard, 1987. Aegopogon (3) American Clayton and Renvoize, 1986. 82

List 1. (continued)

Agrostis (220) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer, 1945. Alopecurus (36) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer, 1945. Anthochloa (1) Andean Clayton and Renvoize, 1986; Weberbauer, 1945. Aristide (250) Wide Tropical Clayton and Renvoize, 1986. Bouteloua (24) American Clayton and Renvoize, 1986. Brachypodium (16) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer, 1945. Bromus (ISO) Holarctic Clayton and Renvoize, 1986; Weberbauer, 1945. Calamagrostis (270) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer, 1945. Cortaderia (24) Austral Clayton and Renvoize, 1986; Cleef, 1979. Critesion (36) Wide Temperate LOve, 1984. Dielsiochloa 0) Andean Clayton and Renvoize, 1986. Dissanthelium (16) Andean Clayton and Renvoize, 1986; Weberbauer, 1945. Elymus (140) Wide Temperate LOve, 1984 Elytrigia (20) Wide Temperate LOve, 1984; Nicora and Rùgulo de Agrasar, 1987. Eragrostis (350) Wide Tropical Clayton and Renvoize, 1986. Festuca (450) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979. HierochloB (30) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer, 1945. Melica (80) Wide Temperate Clayton and Renvoize, 1986; Weberbauer, 1945. Muhlenbergia (160) American Clayton and Renvoize, 1986. Nassella (55) American Barkworth and Everett, 1987. Paspalum (330) Wide Tropical Clayton and Renvoize, 1986; Cleef, 1979. Pennisetum (80) Ruderal Piptochaetium (30) American Clayton and Renvoize, 1986. Poa (500) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer* 1945. Polypogon (18) Wide Temperate Clayton and Renvoize, 1986. Schizachyrium (60) Wide Tropical Clayton and Renvoize, 1986. Sporobolus (160) Wide Tropical Clayton and Renvoize, 1986; Cleef, 1979. Stipa (300) Wide Temperate Clayton and Renvoize, 1986. Trisetum (70) Wide Temperate Clayton and Renvoize, 1986; Cleef, 1979; Weberbauer. 1945. Vulpia (22) Wide Temperate Clayton and Renvoize, 1986. POLEMONIACEAE Gilia (60) American Cronquist et al., 1984; Weberbauer, 1945. Microsteris (1) American Cronquist et al., 1984. 83

List 1. (continued)

POLYGALACEAE Monnina (179) Andean Weberbauer, 1945. POLYGONACEAE Muehlenbeckia (20) Austral Cleef, 1979; Standley and Steyermark, 1946a; van Balgooy, 1971; Weberbauer, 1945. Rumex (100) Ruderal PORTULACACEAE Calandrinia (100) American Standley and Steyermark, 1946a. Portulaca (100) Wide Temperate Standley and Steyermark, 1946a. POTAMOGETONACEAE Potamogeton (100) Wide Temperate Cleef, 1979. PROTEACEAE Oreocallis (4) Austral Sleumer, 1954. RANUNCULACEAE Clematis (250) Wide Temperate van Balgooy, 1971; Willis, 1973. Krapfia (8) Andean Willis, 1973. Ranunculus (400) Wide Temperate Cleef, 1979; van Balgooy, 1971; Weberbauer, 1945; Willis, 1973. Thalictrum (ISO) Holarctic Cleef, 1979; Weberbauer, 1945; Willis, 1973. RHAMNACEAE Colletia (45) Andean Weberbauer, 1945. ROSACEAE Acaena (40) Austral Cleef, 1979; Standley and Steyermark, 1946a; van Balgooy, 1971; Weberbauer, 1945. Alchemilla (400) Wide Temperate Standley and Steyermark, 1946a; Weberbauer, 1945. Hesperomeles (20) Andean Weberbauer, 1945; Willis, 1973. Kageneckia (3) Andean Weberbauer, 1945; Willis, 1973. Margyricarpus (10) Andean Weberbauer, 1945; Willis, 1973. Polylepis (15) Andean Simpson, 1979. Potentilla (250) Holarctic Cleef, 1979; Standley and Steyermark, 1946a; van Balgooy, 1971; Weberbauer, 1945. Prunus (200) Ruderal Rubus (400) Wide Temperate Cleef, 1979; Standley and Steyermark, 1946a. RUBIACEAE Arcythophyllum (24) Andean Weberbauer, 1945. 84

List 1. (continued)

Galium (400) Wide Temperate Cleef, 1979; van Balgooy, 1971. Relbunium (30) American Cleef, 1979; Ehrendorfer, 1955; van Balgooy, 1971. SANTALACEAE Quinchamalium (20) Andean Weberbauer, 1945. SAPINDACEAE Dodonaea (54) Austral Standley and Steyermark, 1949. SAXIFRAGACEAE Saxifrage (370) Holarctic Willis, 1973. SCROPHULARIACEAE Agalinis (?) American D. N. Smith, pers. obs. Alonsoa (16) Andean Holmgren and Molau, 1984; Weberbauer, 1945. Bartsia (30) Andean Holmgren and Molau, 1984; Willis, 1973. Calceolaria (366) Andean Holmgren and Molau, 1984; Weberbauer, 1945. Castilleja (200) American Cronquist et al., 1984; Holmgren and Molau, 1984; Weberbauer, 1945. Limosella (12) Wide Temperate Cleef, 1979; Cronquist et al., 1984; Holmgren and Molau, 1984; van Balgooy, 1971. Mimulus (100) Wide Temperate Cleef, 1979; Cronquist et al., 1984; Holmgren and Molau, 1984. Orthocarpus (25) American Cronquist et al., 1984; Weberbauer, 1945. Ourisia (30) Austral Cleef, 1979; Holmgren and Molau, 1984; van Balgooy, 1971; Weberbauer, 1945. Porodittia (1) Endemic Weberbauer, 1945. Sibthorpia (6) Holarctic Cleef, 1979; Holmgren and Molau, 1984. Veronica (200) Wide Temperate Cleef, 1979; Cronquist et al., 1984; Weberbauer, 1945. SOLANACEAE lochroma (27) Andean Hunziker, 1979; Weberbauer, 1945. Jaltomata (24) American Hunziker, 1979. Lycianthes (140) Wide Tropical Gentry and Standley, 1974. Lycopersicon (6) Andean Hunziker, 1979. Nicotiana (67) Wide Tropical Hunziker, 1979. Salpichroa (14) Andean Hunziker, 1979; Weberbauer, 1945. Saracha (4) Andean Hunziker, 1979. Solanum (1000) Cosmopolitan Cleef, 1979; Hunziker, 1979; van Balgooy, 1971. SYMPLOCACEAE Symplocos (300) Wide Tropical Cleef, 1979; Standley and Williams, 1967. 85

List 1. (continued)

URTICACEAE Pilea (200) Wide Tropical Standley and Steyermark, 1952. Urtica (35) Wide Temperate Cleef, 1979; Standley and Steyermark, 1952. VALERIANACEAE Phyllactis (3) Andean Larsen, 1986. Stangea (5) Andean Weberbauer, 1945. Valeriana (200) Wide Temperate Cleef, 1979; Weberbauer, 1945. VERBENACEAE Citharexylum (100) American Nash and Nee, 1984. Duranta (35) American Nash and Nee, 1984. Glandularia (28) American Umber, 1979; Willis, 1973. Lantana (50) American Nash and Nee, 1984; van Balgooy, 1971. Verbena (100) American Nash and Nee, 1984; van Balgooy, 1971. VIOLACEAE Viola (400) Wide Temperate Cleef, 1979; Standley and Williams, 1961; Weberbauer, 1945. ZANNICHELLIACEAE Zannichellia (3) Cosmopolitan Haynes and Holm-Nielsen, 1986c. 86

List 2. Phytogeographic origin of the gymnosperm and angiosperm families included in the Huascaran National Park flora. (Classified following Gentry, 1982).

GYMNOSPERMS EPHEDRACEAE Laurasian ANGIOSPERMS AMARANTHACEAE Unclassified AMARYLLIDACEAE Unclassified (including Agavaceae and Alstroemeriaceae) APIACEAE Laurasian ARALIACEAE Gondwanan - northern Andean ASCLEPIADACEAE Unclassified ASTERACEAE Gondwanan - northern Andean BASELLACEAE Laurasian BEGONIACEAE Gondwanan - northern Andean BERBERIDACEAE Laurasian BETULACEAE Laurasian BIGNONIACEAE Gondwanan - Amazonian BORAGINACEAE Laurasian BRASSICACEAE Laurasian BROMELIACEAE Gondwanan - northern Andean CACTACEAE Gondwanan - dry areas CALLITRICHIACEAE Laurasian CAMPANULACEAE Gondwanan - northern Andean CAPPARACEAE Gondwanan - dry areas CARYOPHYLLACEAE Laurasian CLUSIACEAE Gondwanan - northern Andean COLUMELLIACEAE Gondwanan - northern Andean COMMELINACEAE Unclassified CONVOLVULACEAE Gondwanan - Amazonian (including Cuscutaceae) CORIARIACEAE Gondwanan - southern Andean CRASSULARIACEAE Laurasian CUNONIACEAE Gondwanan - southern Andean 87

List 2. (continued)

CYPERACEAE Unclassified DIOSCORIACEAE Unclassified ELAEOCARPACEAE Gondwanan - Amazonian ELATINACEAE Unclassified ERICACEAE Gondwanan - northern Andean EUPHORBIACEAE Gondwanan - Amazonian FLACOURTIACEAE Gondwanan - Amazonian GENTIANACEAE Laurasian GERANIACEAE Laurasian HALORAGACEAE Gondwanan - northern Andean HYDROCHARITACEAE Unclassified HYDROPHYLLACEAE Laurasian IRIDACEAE Gondwanan southern Andean JUNCACEAE Gondwanan < southern Andean JUNCAGINACEAE Unclassified LAMIACEAE Laurasian ? LEGUMINOSAE, s.l. Gondwanan- Amazonian (including Caesalpinicaeae and Fabaceae) LEMNACEAE Unclassified LILIACEAE Laurasian (including Alllaceae and Anthericaceae) LINACEAE Unclassified LOASACEAE Gondwanan - southern Andean LOGANIACEAE Gondwanan - Amazonian (including Buddleiaceae) LORANTHACEAE Gondwanan - northern Andean MALVACEAE Unclassified MELASTOMATACEAE Gondwanan - northern Andean MYRICACEAE Laurasian MYRSINACEAE Gondwanan northern Andean NYCTAGINACEAE Gondwanan northern Andean ONAGRACEAE Gondwanan southern Andean 88

List 2. (continued)

ORCHIDACEAE Gondwanan - northern Andean OXALIDACEAE Gondwanan - northern Andean PASSIFLORACEAE Gondwanan - northern Andean PHYTOLACCACEAE Unclassified PIPERACEAE Gondwanan - northern Andean PLANTAGINACEAE . Laurasian POACEAE Unclassified POLEMONIACEAE Laurasian POLYGALACEAE Gondwanan - Amazonian POLYGONACEAE Unclassified PORTULACACEAE Gondwanan - southern Andean POTAMOGETONACEAE Unclassified PROTEACEAE Gondwanan - southern Andean RANUNCULACEAE Laurasian RHAMNACEAE Laurasian ROSACEAE Laurasian RUBIACEAE Gondwanan - northern Andean SANTALACEAE Gondwanan - southern Andean SAPINDACEAE Gondwanan - Amazonian SAXIFRAGACEAE Laurasian SCROPHULARIACEAE Laurasian SOLANACEAE Gondwanan - southern Andean SYMPLOCACEAE Laurasian URTICACEAE Gondwanan - northern Andean VALERIANACEAE Laurasian VERBENACEAE Unclassified VIOLACEAE Gondwanan - Amazonian ZANNICHELLIACEAE Unclassified 89

List 2. (continued)

Summary

Gondwanan 48.8% Andean 34.5% Amazonian 11.9% Dry-area 2.4% Laurasian 29.8% Unclassified 21.4% 90

List 3. Families and genera of the Huascaràn National Park flora.

GYMNOSPERMS Jamesonia (3) Notolaena (1) EPHEDRACEAE Peilaea (1) Ephedra (1) THELYPTERIDACEAE PTERIDOPHYTES Thelypteris (2) ASPLENIACEAE SALVINIACEAE Asplenium (7) Azolla (1) BLECHNACEAE Blechnum (2) ANGIOSPERMS DENNSTAEDTIACEAE AGAVACEAE Histiopteris (1) Furcraea (1) Hypolepis (2) ALLIACEAE DRYOPTERIDACEAE Trichlora (1) Athyrium (1) ALSTROEMERIACEAE Cystopteris (1) Alstroemeria (1) Elaphoglossum (9) Bomarea (4) Polystichum (3) Woodsia (1) AMARANTHACEAE Alternanthera (3) EQUISETACEAE Equisetum (1) AMARYLLIDACEAE Stenomesson (1) GLEICHENIACEAE Gleichenia (Sticherus) (1) ANTHERICACEAE Echeandia (1) ISOETACEAE IsoCtes (3) APIACEAE Azorella (5) LYCOPODIACEAE Bowlesia (2) Huperzia (3) Daucus (1) Lycopodium (3) Eryngium (1) OPHIOGLOSSACEAE Lilaeopsis (1) Ophioglossum (1) Niphogeton (1) Oreomyrrhis (I) PLAGIOGYRIACEAE Plagiogyria (I) ARALIACEAE Oreopanax (1) POLYPODIACEAE Campyloneurum (1) ASCLEPIADACEAE Grammitis (5) Cynanchum (2) Metastelma (3) Microgramma (I) Polypodium (3) ASTERACEAE PTERIDACEAE Achyrocline (2) Adiantum (I) Ageratina (I) Cheilanthes (5) Antennaria (I) Eriosorus (3) Aphanactis (1) 91

List 3. (continued)

Aristeguieta (1) Siegesbeckia (1) Baccharis (13) Sonchus (I) Barnadesia (I) Stevia (1) Belloa (4) Stuckertiella (1) Bidens (2) Tagetes (3) Chaptalia (1) Vasquezia (1) Chersodoma (2) Verbesina (1) Chrysactenium (I) Viguiera (1) Chuquiraga (1) Werneria (18) Conyza (3) Coreopsis (3) BASELLACEAE Ullucus (1) Cosmos (1) Cotula (1) BEGONIACEAE Cuatrecasasiella (-Luciliopsis) (1) Begonia (1) Dasyphyllum (1) Diplostephium (3) BERBERIDACEAE Erigeron (2) Berberis (2) Galinsoga (1) BETULACEAE Gamochaeta (5) Alnus (1) Gnaphalium (3) Grosvenoria (Eupatorium) (1) BIGNONIACEAE Gynoxys (2) Tecoma (1) Helianthopsis (1) BORAGINACEAE Hieracium (2) Hackelia (1) Hypochoeris (3) Lithospermum (1) Jungia (3) Plagiobotrys (-Allocarya) (1) Leuceria (1) Liabum (1) BRASSICACEAE Lophopappus (1) Brassica (1) Loricaria (2) Brayopsis (2) Lucilia (1) Capsella (1) Mnodes (4) Cardamine (1) Munnozia (1) Catydysia (1) Mutisia (1) Cremolobus (1) Noticastrum (1) Descurainia (2) Novenia (1) Draba (3) Ophryosporus (1) Lepidium (1) Oritrophium (3) Mancoa (1) Paranephelius (3) Weberbauera (1) Pentacalia (I) BROMELIACEAE Perezia (4) Pitcairnia (I) Perymenium (1) Puya (8) Plagiocheilus (1) Tillandsia (9) Polymnia (1) Pseudonoseris (1) BUDDLEIACEAE Senecio (incl. Culcitium, Dorobea, & Buddleia (2) Lasiocephalus) (39) 92

List 3. (continued)

CACTACEAE Tillaea (2) Cereus (1) Villadia (1) Cleistocactus (1) Opuntia (2) CUNONIACEAE Oreocereus (3) Weinmannia (1) CUSCUTACEAE CAESALPINIACEAE Cuscuta (1) Senna (1) CYPERACEAE CALLITRICHIACEAE Baeothryon (Scirpus, pp.) (I) Callitriche (1) Bulbostylis (1) CAMPANULACEAE Carex (8) Hypsella (1) Cyperus (2) Lobelia (2) Eleocharis (1) Lysipomia (3) Isolepis (1) Siphocampylus (2) Oreobolopsis (1) Wahlenbergia (1) Oreobolus (1) Phylloscirpus (Scirpus, pp.) (3) CAPPARACEAE Cleome (1) Schoenoplectus (Scirpus, pp.) 0) DIOSCORIACEAE CARYOPHYLLACEAE Dioscorea (2) Arenaria (6) Cardionema (1) ELAEOCARPACEAE Cerastium (5) Vallea (1) Drymaria (2) Paronychia (4) ELATINACEAE Plettkea (1) Elatine (1) Pycnophyllum (2) ERICACEAE Silene (1) Gaultheria (2) Spergularia (1) Pernettya (1) Stellaria (2) Vaccinium (1) CLUSIACEAE EUPHORBIACEAE Hypericum (2) Chamaesyce (1) COLUMELLIACEAE Croton (1) Euphorbia (1) Columellia (1) Sebastiania (1) COMMELINACEAE Commelina (1) FABACEAE Astragalus (4) CONVOLVULACEAE Dalea (1) Dichondra (1) Lathyrus (1) CORIARIACEAE Lupinus (IS) Coriaria (1) Otholobium (2) Spartium (1) CRASSULARIACEAE Trifolium (1) Echeverria (1) Vicia (1) 93

List 3. (continued)

FLACOURTIACEAE LOASACEAE Pineda (1) Cajophora (2) Loasa (1) GENTIANACEAE Gentiana (1) LORANTHACEAE Gentianella (7) Tripodanthus (1) Halenia (1) Tristerix (2) GERANIACEAE MALVACEAE Erodium (1) Acaulimalva (3) Geranium (3) Nototriche (7) GROSSULARIACEAE Urocarpidium (1) Escallonia (3) MELASTOMATACEAE Ribes (4) Brachyotum (S) Miconia (3) HALORAGACEAE Myriophyllum (1) MYRICACEAE Myrica (2) HYDROCHARITACEAE Elodea (1) MYRSINACEAE Myrsine (1) HYDROPHYLLACEAE Phacelia (1) NYCTAGINACEAE IRIDACEAE Colignonia (1) Orthrosanthus (1) ONAGRACEAE Sisyrinchium (7) Epilobium (2) Tigridia (1) Fuchsia (1) JUNCACEAE Oenothera (3) Distichia (1) ORCHIDACEAE Juncus (8) Aa (2) Luzula (3) Epidendrum (4) JUNCAGINACEAE Gomphichis (1) Lilaea (1) Malaxis (1) Masdevallia (1) LAMIACEAE Myrosmodes (2) Gardoquia (2) Odontoglossum (2) Lepechinia (I) Pleurothallis (2) Salvia^ Pterichis (1) Scutellaria (1) Stelis (3) Stachys (1) Trichoceros (1) LEMNACEAE OXALIDACEAE Lemna (1) Hypsocharis (1) LINACEAE Oxalis (S) Linum (1) PASSIFLORACEAE Passiflora (2) 94

List 3. (continued)

PHYTOLACCACEAE POLYGONACEAE Phytolacca (1) Muehlenbeckia (2) PIPERACEAE Rumex (2) Peperomia (5) PORTULACACEAE Calandrinia (2) PLANTAGINACEAE Plantago (5) Portulaca (I) POTAMOGETONACEAE POACEAE Potamogeton (1) Aciachne (1) Aegopogon 0) PROTEACEAE Agrostis (6) Oreocallis (1) Alopecurus (1) Anthochloa RANUNCULACEAE 0) Clematis (1) Aristida (1) Bouteloua 0) Krapfia (1) Brachypodium (1) Ranunculus (5) Bromus (S) Thalictrum (1) Calamagrostis (8) RHAMNACEAE Cortaderia (5) Colletia (1) Critesion (1) ROSACEAE Dielsiochloa (1) Acaena (1) Dissanthelium (2) Elymus (1) Alchemilla (6) Hesperomeles (1) Elytrigia (1) Kageneckia (1) Eragrostis (2) Margyricarpus (I) Festuca (12) Polylepis (4) Hierochloê (1) Potentilla (1) Melica (1) Muhlenbergia (6) Prunus (1) Nassella (6) Rubus (2) Paspalum (3) RUBIACEAE Pennisetum (1) Arcythophyllum (2) Piptochaetium (2) Galium (1) Poa (16) Relbunium (2) Polypogon (1) Schizachyrium (1) SANTALACEAE Sporobolus (1) Quinchamalium (1) Stipa (3) SAPINDACEAE Trisetum (2) Dodonaea (1) Vulpia (2) SAXIFRAGACEAE POLEMONIACEAE Saxifraga (1) Gilia (1) Microsteris (1) SCROPHULARIACEAE Agalinis (1) POLYGALACEAE Alonsoa (2) Monnina (2) Bartsia (3) 95

List 3. (continued)

Calceolaria (15) Castilleja (5) Limosella (1) Mimulus (1) Orthocarpus (1) Ourisia (2) Porodittia (1) Sibthorpia (1) Veronica (3) SOLANACEAE lochroma (1) Jaltomata (2) Lycianthes (1) Lycopersicon (1) Nicotiana (1) Salpichroa (3) Saracha (1) Solanum (16) SYMPLOCACEAE Symplocos (1) URTICACEAE Pilea (I) Urtica (4) VALERIANACEAE Phyllactis (3) Stangea (2) Valeriana (13) VERBENACEAE Citharexylum (1) Duranta (1) Glandularia (1) Lantana (1) Verbena (1) VIOLACEAE Viola (2) ZANNICHELLIACEAE Zannichellia (1) 96

List 4. Families and genera of the Colombian paramos (Data adapted from Cleef, 1981)

PTERIDOPHYTES Eryngium (2) Hydrocotyle (4) ASPLENIACEAE Asplenium (2) Lilaeopsis (1) Myrrhidendron (1) BLECHNACEAE Niphogeton (6) Blechnum (1) Oreomyrrhis (1) Ottoa (1) DENNSTAEDTIACEAE Pteridium (1) AQUIFOLIACEAE Ilex (1) DRYOPTERIDACEAE Cystopteris (1) ARECACEAE Dryopteris (1) Geonoma (1) Elaphoglossum (2) ARALIACEAE Polystichum (1) Oreopanax (1) EQUISETACEAE Equisetum (1) ASTERACEAE Achyrocline (1) HYMENOPHYLLACEAE Ageratina (as Eupatorium, p.p.) (?) Hymenophyllum (1) Aphanactis (2) Baccharis (6) ISOETACEAE Isoëtes (11) Bidens (1) Conyza(1) LYCOPODIACEAE Cotula (1) Huperzia (as Lycopodium) (3) Diplostephium (14) Lycopodium (3) Erigeron (3) MARSILEACEAE Espelatia (25) Pilularia (1) Espeletiopsis (7) Eupatorium (s.l.?) (S) OPHIOGLOSSACEAE Floscaldasia 0) Ophioglossum (1) Gnaphalium (2) POLYPODIACEAE Gynoxys (7) Campyloncuron (as Polypodium) (1) Hieracium (I) Grammltis (4) Hinterhubera (1) Hypochoeris (1) PTERIDACEAE Jaegeria (I) Eriosorus (I) Laestadia (1) Jamesonla ((2) Loricaria (I) Nephopteris (1) Lourteigia (1) SALVINIACEAE Lucilia (I) Azolla (1) Noticastrum (1) Oritrophium (4) ANGIOSPERMS Oxylobus (I) Pentacalia (as Senecio) (12) APIACEAE Plagiocheilus (I) Azorella (4) Senecio (IS) 97

List 4. (continued)

Sigesbeckia (I) CUNONIACEAE Stevia (1) Weinmannia (2) Verbesina (1) Werneria (3) CYPERACEAE Bulbostylus (1) BERBERIDACEAE Carex (incl. Vesicarex) (11) Berberis (3) Cyperus (1) BORAGINACEAE Eleocharis (3) Moritzia (1) Isolepis (as Scirpus, p.p.) (1) Oreobolus (1) BRASSICACEAE Rhynchospora (3) Cardamine (2) Schoenoplectus (as Scirpus, p.p.) (1) Draba (4) Lepidium (1) ELAEOCARPACEAE Vallea (1) BROMELIACEAE Greigia (1) ELATINACEAE Puya (6) Elatine (1) BUDDLEIACEAE ERICACEAE Buddleia (1) Befaria (1) Disterigma (1) CALLITRICHIACEAE Gaultheria (3) Callitriche (1) Gaylussacia (1) CAMPANULACEAE Macleania (1) Centropogon (1) Pernettya (2) Hypsela (1) Vaccinium (1) Lobelia (1) ERIOCAULACEAE Lysipomia (2) Eriocaulon (1) Rhizocephalum (1) Paepalanthus (8) Siphocampylus (1) EUPHORBIACEAE CARYOPHYLLACEAE Dysopsis (1) Arenaria (2) Cerastium (2) FABACEAE Colobanthus (2) Lupinus (3) Drymaria (1) Vicia (1) Stellaria(l) GENTIANACEAE CLETHRACEAE Gentiana (1) Clethra (1) Gentianella (3) Halenia (1) CLUSIACEAE Hypericum (23) GERANIACEAE Geranium (4) CORIARIACEAE Coriaria (1) GROSSULARIACEAE Escallonia (1) CRASSULACEAE Ribes(l) Tillaea (incl. Crassula) (2) 98

List 4. (continued)

HALORAGACEAE MYRTACEAE Myriophyllum (2) Myrteola (1) Ugni (1) HYDROCHARITACEAE Elodea (1) ONAGRACEAE Epilobium (2) IRIDACEAE Orthrosanthus (1) Ludweigia 0) Sisyrinchium (3) ORCHIDACEAE JUNCACEAE Aa (as Altensteinia) (1) Distichia (1) Altensteinia (1) Coccineorchis (as Spiranthes) (1) Juncus (5) Luzula (3) Epidendrum (2) Masdevallia (I) JUNCAGINACEAE Myrosmodes (as Altensteinia) (1) Lilaea (1) Odontoglossum (1) Pterichis (1) LAMIACEAE Salvia (1) Stenorrhynchos (as Spiranthes) (1) Satureja (1) OXALIDACEAE Stachys (1) Oxalis (2) LEMNACEAE PIPERACEAE Lemna (I) Peperomia (3) LENTIBULARIACEAE PLANTAGINACEAE Pinguicula (1) Plantago (4) Utricularia (1) POACEAE LORANTHACEAE Aciachne (1) Gaiadendron (1) Agrostis (S) MALVACEAE Aulonemia (1) Acaulimalva (1) Brachypodium (1) Bromus (2) MELANTHIACEAE Calamagrostis (7) Toefeldia (1) Chusquea (incl. Swallenochloa) (2) MELASTOMATACEAE Cinna (1) Brachyotum (1) Cortaderia (2) Bucquetia (1) Danthonia (1) Castratella (2) Festuca (2) Chaetolepis (1) Lorenzochloa (1) Miconia (7) Muhlenbergia (1) Purpurella (1) Neurolepis (1) Paspalum (1) MYRICACEAE Poa (I) Myrica (I) Sporobolus (1) MYRSINACEAE POLYGALACEAE Myrsine (as Rapanea) (1) Monnina (1) 99

List 4. (continued)

POLYGONACEAE URTICACEAE Muehlenbeckia (2) Parietaria (1) Rumex (2) Urtica (1) PORTULACACEAE VALERIANACEAE Calandrinia (1) Phyllactis (1) Montia (2) Valeriana (7) POTAMOGETONACEAE VIOLACEAE Potamogeton (5) Viola (1) RANUNCULACEAE XYRIDACEAE Ranunculus (6) Xyris (2) ROSACEAE Acaena (2) Alchemilla (as Lachemilla) (11) Hesperomeles (2) Polylepis (1) Potentilla (1) Rubus (1) Sericotheca (I) RUBIACEAE Arcythophyllum (3) Galium (2) Nertera 0) Relbunium (1) SCROPHULARIACEAE Aragoa (S) Calceolaria (2) Castilleja (2) Gratiola (2) Limosella (2) Mimulus (1) Ourisia (1) Sibthorpia (1) Veronica (1) SOLANACEAE Acnistis (1) Cestfum (1) Solauum 0) SYMPLOCACEAE Symplocos (1) THEACEAE Ternstroemia (1) 100

List 5. Families and genera of the Argentinan puna (Data adapted from Ruthsatz, 1977)

GYMNOSPERMS APIACEAE EPHEDRACEAE Asteriscium (1) Ephedra (3) Azorella (1) Bowlesia (2) PTERIDOPHYTES Lilaeopsis (1) Mulinum (1) ASPLENIACEAE Oreomyrrhis (1) Asplenium (1) ASCLEPIADACEAE DRYOPTERIDACEAE (genera placed in Aphanostelma (2) diverse families) Philibertia (1) Cystopteris (1) Polystichum (1) ASTERACEAE Woodsia (2) Achyrocline (3) Ageratina (as Eupatorium, p.p.) (1) MARSILEACEAE Aphyllocladus (1) Marsilea (1) Baccharis (8) OPHIOGLOSSACEAE Belloa (S) Ophioglossum (1) Bidens (2) Chaetanthera (3) PTERIDACEAE (as Sinopteridaceae) Chaptalia (1) Cheilanthes (2) Chersodoma (1) Notolaena (2) Chiliotrichiopsis (1) Pellaea (1) Chuquiraga (5) SELAGINELLACEAE Conyza (3) Selaginella (1) Cotula (1) Cuatrecasasiella (as Luciliopsis) (1) ANGIOSPERMS Dinoseris (1) Eupatorium (p.p.) (3) ACANTHACEAE Facelis (2) Justicia (1) Flaveria (1) Stenandrium (1) Flourensia (2) ALLIACEAE Galinsoga (1) Nothoscordum (3) Gamochaeta (6) Gnaphalium (5) AMARYLLIDACEAE Gochnatia (1) Castellanoa (1) Guttierrezia (1) Eustephiopsis (1) Heterospermum (3) AMARANTHACEAE Hieracium (1) Alternanthera (2) Hypochoeris (4) Amaranthus (4) Lophopappus(2) Gomphrena (6) Mutisia (S) Guilleminea (1) Nardophyllum (1) Nassauvia (1) ANACARDIACEAE Parastrephia (4) Schinus (2) Parthenium (1) Pectis (1) Perezia (6) 101

List 5. (continued)

Plazia (1) CACTACEAE Porophyllum (1) Echinopsis (incl. Lobivia and Proustia (1) Trichocereus) (9) Schkuhria (2) Gymnocalycium (1) Senecio (26) Neowerdermannia 0) Sonchus (1) Opuntia (incl. Austrocylindropuntia Stevia (S) and Tephrocactus) (18) Tagetes (1) Oreocereus (2) Taraxacum (1) Parodia (S) Tessaria (I) Rebutia (incl. Mediolobivia) (1) Trichocline (2) Urmentia (1) CAESALPINIACEAE Viguiera (1) Caesalpinia (1) Werneria (6) Cercidium (1) Zinnia (1) Hoffmannseggia (1) Senna (as Cassia) (2) BALANOPHORACEAE CALYCERACEAE Juelia (1) Calycera (1) BIGNONIACEAE Moschopsis (1) Tecoma (1) Nastanthus (1) BORAGINACEAE CAMPANULACEAE Cryptantha (1) Hypsella (1) Heliotropium (2) Wahlenbergia (2) Pectocarya (1) Plagiobotrys (1) CARYOPHYLLACEAE Arenaria (7) BRASSICACEAE Cerastium (2) Aschersoniodoxa (1) Colobanthus (1) Brayopsis (1) Herniaria (1) Capsella (1) Paronychia (3) Cardamine (1) Pentacaena (1) Descurainia (2) Pycnophyllum (4) Eudema (1) Siline (2) Kardamoglyphos (1) Spergularia (2) Lepidium (1) Stellaria (1) Lesquerella (1) Mancoa (1) CHENOPODIACEAE Parodiodoxa (1) Atriplex (5) Pennellia (1) Chenopodium (6) Weberbauera (I) Nitrophila (1) Salicornia (1) BROMELIACEAE Suaeda (1) Abromeitiella (2) Puya (1) COMMELINACEAE Tillandsia (3) Commelina (1) BUDDLEIACEAE Buddleia (3) 102

List 5. (continued)

CONVOLVULACEAE Mastigostyla (2) Dichondra (1) Sisyrinchium (6) Evolvulus (1) Ipomoea (1) JUNCACEAE Distichia (1) CRASSULACEAE Juncus(3) Sedum (1) Luzula (3) Tillaea (as Crassula) (1) Oxychloe (1) Villadia (1) JUNCAGINACEAE CUCURBITACEAE Lilaea (1) Apodanthera (1) Triglochin (2) CYPERACEAE KRAMERIACEAE Baeothryon (as Scirpus, p.p.) (1) Krameria (1) Carex (4) Cyperus (1) LAMIACEAE Eleocharia (5) Salvia (1) Phylloscirpus (as Scirpus, p.p.) (4) Satureja (probably Gardoquia) (1) EUPHORBIACEAE LOASACEAE Euphorbia (3) Cajophora (3) Mentzelia (2) FABACEAE Adesmia(ll) LORANTHACEAE Amicia (1) Tristerix (as PhrygilantI ua) (1) Astragalus (9) MALVACEAE Dalea (1) Abutilon (1) Lupinus (5) Acaulimalva (1) Medicago (1) Herissantia (1) Melilotus (1) Nototriche (6) Neocracca (1) Sida (1) Sesbania (1) Tarasa (1) Trifolium (1) MIMOSACEAE FRANKENIACEAE Acacia (2) Anthobryum (1) Prosopis (3) GENTIANACEAE NYCTAGINACEAE Gentiana (1) Allionia (1) Gentianella (2) Boerhaavia (1) GERANIACEAE Bouganvillea (1) Erodium (1) Mirabilis (I) Geranium (1) Oxybaphus(1) HYDROPHYLLACEAE OLEACEAE Phacelia (4) Menodora (1) IRIDACEAE ONAGRACEAE Cardenanthus (2) Oenothera (1) 103

List S. (continued)

ORCHIDACEAE Sporobolus (2) Myrosmodes (as Aa) (1) Stipa (14) Tragus (2) OXALIDACEAE Hypseocharis (1) Trichloris (1) Oxalis (1) Tridens (1) Tripogon (1) PIPERACEAE Vulpia (1) Peperomia (1) POLEMONIACEAE PLANTAGINACEAE Cilia (2) Bougueria (1) Microsteris (as Gilia) (1) Plantago (3) POLYGALACEAE POACEAE Monnina (2) Agrostis (1) Polygala (3) Anthochloa (1) Aristida (7) PORTULACACEAE Bothriochloa (1) Calandrinia (9) Bouteloua (2) Portulaca (6) Bromus (2) RANUNCULACEAE Calamagrostis (incl. Deyeuxia) (7) Barneoudia (1) Cenchrus (1) Caltha (1) Chloris (1) Clematis (I) Cortaderia (1) Myosurus (1) Cottea (1) Ranunculus (1) Critesion (as Hordeum) (2) Cynodon (1) RHAMNACEAE Dielsiochloa (1) Kentrothamnus (1) Digitaria (1) ROSACEAE Diplachne (1) Alchemilla (1) Distichlis (1) Polylepis (1) Enneopogon (1) Tetraglochin (2) Eragrostis (4) Festuca (9) RUBIACEAE Koeleria(l) Borreria (1) Lycurus (1) Galium (1) Microchloa (1) Mitracarpus (2) Muhlenbergia (3) Relbunium (1) Munroa (2) SCROPHULARIACEAE Nassella (incl Stipa, p.p.) (5) Agalinis (as Gerardia) (1) Oryzopsis (1) Bartsia (1) Panicum (1) Calceolaria (3) Pappophorum (1) Castilleja (I) Pennisetum (1) Euphrasia (1) Piptochaetium (1) Limosella (3) Poa (10) Mimulus (2) Puccinella (1) Setaria (1) 104

List S. (continued)

SOLANACEAE Fabiana (2) Lycium (5) Nicotiana (3) Nierembergia (1) Petunia (1) Salpichroa (1) Sclerophylax (2) Solanum (5) Trechonaetes (1) URTICACEAE Urtica (2) VALERIANACEAE Stangea (1) Valeriana (7) VERBENACEAE Acantholippia (2) Glandularia (as Verbena) (1) Junellia (6) Lampaya (1) Lantana (i) Lippia (3) Verbena 0) VIOLACEAE Viola (1) ZYGOPHYLLACEAE Bulnesia (1) Larrea (1) Tribulus 0) 105

List 6. Families and genera of the Afroalpine flora (Data adapted from Hedberg, 1957)

PTERIDOPHYTES Osteospermum (1) Senecio (inch Dendrosenecio) (26) ASPLENIACEAE Asplenium (6) Stroebe (1) DRYOPTERIDACEAE BALSAMINACEAE Cystopteris (1) Impatiens (1) Dryopteris (1) BORAGINACEAE Elaphoglossum (1) Lithospermum (1) Polystichum (2) Myosotis (3) LYCOPODIACEAE BRASSICACEAE Huperzia (as Lycopodium) (1) Arabidopsis (1) Arabis (1) POLYPODIACEAE Pleopeltis (1) Barbarea (1) Polypodium (1) Cardamine (2) Oreophyton (1) PTERIDACEAE Subularia (1) Aleuritopsis (1) Thlaspi (1)

ANGIOSPERMS CALLITRICHIACEAE Callitriche (1) APIACEAE Anthriscus (1) CAMPANULACEAE Lobelia (11) Cancalis (1) Ferula (1) Monopsis (I) Hapalosciadium (1) Wahlenbergia (2) Heracleum (2) CARYOPHYLLACEAE Pencedanum (2) Cerastium (2) Pimpinella (1) Sagina (2) ASPHODELACEAE (as Liliaceae) Siline (1) Kniphofia (2) Stellaria (1) ASTERACEAE CLUSIACEAE (as Hypericaceae) Anthemis (1) Hypericum (4) Artemesia (1) CRASSULACEAE Carduus (5) Crassula (4) Cineraria (1) Sedum (4) Conyza (3) Umbilicus (1) Coreopsis (1) Cotula (2) CYPERACEAE Crassocephalum (1) Carex (6) Crepis (3) Carpha (1) Dichrocephala (1) Cyperus (1) Euryops (3) Scirpus (2) Gnaphalium (2) DIPSACACEAE Haplocarpa (1) Dipsacus (1) Helichrysum (16) Scabosia (1) Nannoseris (1) 106

List 6. (continued)

ERICACEAE POACEAE Biaeria (2) Agrostis (9) Erica (2) Aira (1) Philippia (3) Andropogon (1) Anthoxanthum (1) ERIOCAULACEAE Calamagrostis (1) Eriocaulon (1) Deschampsia (3) EUPHORBIACEAE Festuca (3) Euphorbia (1) Helictotrichon (2) Keniochloa (2) FABACEAE Koeleria (2) Adenocarpus (1) Pennisetum (1) Astragalus (2) Pentaschistis (3) Lathyrus (1) Poa (4) Trifolium (7) Vulpia (1) GENTIANACEAE PORTULACACEAE Sebaea (1) Swertia (7) Montia (1) GERANIACEAE POTAMOGETONACEAE Geranium (3) Potamogeton (1) Pelargonium (1) PRIMULACEAE Anagallis (1) IRIDACEAE Dierama (1) PROTEACEAE Gladiolus (1) Protea (1) Hesperantha (1) RANUNCULACEAE Romulea (3) Anemone (1) JUNCACEAE Delphinium (1) Juncus (1) Ranunculus (5) Luzula (3) ROSACEAE LAMIACEAE Alchemilla (12) Salvia (2) Satureja (6) RUBIACEAE Anthospermum (1) MALVACEAE Galium (4) Malva (1) Sida (1) SANTALACEAE Thesium (1) ONAGRACEAE Epilobium (1) SCROPHULARIACEAE Bartsia (5) ORCHIDACEAE Celsia (2) Disa (1) Hebenstretia (1) Habenaria (1) Limosella (2) OXALIDACEAE Sibthorpia (1) Oxalis (1) Veronica (3) 107

List 6. (continued)

URTICACEAE Parietaria (1) Urtica (1) VALERIANACEAE Valeriana (2) VIOLACEAE Viola (1) 108

List 7. Families and genera of the New Guinea alpine flora. (Data adapted from van Royen, 1979b, 1982, 1983)

GYMNOSPERMS Lagenophora (2) Myriactis (2) CUPRESSACEAE Papuacedrus (1) Olearia (IS) Piora (1) PODOCARPACEAE Rhamphogyne (1) Dacrycarpus (4) Senecio (13) Dacrydium (1) Sonchus (2) Phyllocladus (1) Tetramolophium (24) Podocarpus (3) BALSAMINACEAE ANGIOSPERMS Impatiens (1) ACTINIDIACEAE (as Saurauiaceae) BORAGINACEAE Saurauia (9) Crucicaryum (1) Cynoglossum (1) APIACEAE (as Araliaceae) Hydrocotyle (1) Myosotis (1) Oreomyrrhis (6) Trigonotis (12) Trachymene (10) BRASSICACEAE Cardamine (4) APOCYNACEAE Alyxia (6) Papuzilla (2) Rorippa (3) AQUIFOLIACEAE Ilex (1) CALLITRICHIACEAE Callitriche (1) ARALIACEAE Harmsiopanax (1) CAMPANULACEAE Schefflera (17) Lobelia (S) Peracarpa (1) ASCLEPIADACEAE Wahlenbergia (3) Taxocarpus (1) CAPRIFOLIACEAE ASTELIACEAE (as Liliaceae) Periomphale (1) Astelia (1) CARYOPHYLLACEAE ASTERACEAE Cerastium (I) Abrotanella (1) Sagina (3) Anaphalis (3) Scleranthus (1) Arrhenechthites (2) Bidens (1) CASUARINACEAE Blumea (2) Casuarina (1) Brachycome (2) CELASTRACEAE Cotula (4) Celastrus (1) Dichrocephala (1) Perrottetia (1) Erigeron (2) Gnaphalium (7) CENTROLEPIDACEAE Ischnea (4) Centrolepis (3) Keysseria (9) Gaimardia (1) Lactuca (2) 109

List 7. (continued)

CHLORANTHACEAE Dimorphanthera (18) Asearina (1) Diplycosia (6) Gaultheria (3) CLUSIACEAE (66) Hypericum (S) Rhododendron Vaccinium (39) CORIARIACEAE ERIOCAULACEAE Coriaria (1) Eriocaulon (10) CUCURBITACEAE Gynostemma (1) EUPHORBIACEAE Claoxylon (1) CUNONIACEAE Glochidion (1) Caldcluvia (2) Macaranga (3) Pullea (1) Mallatus (1) Weinmannia (1) Omalanthus (2) Phyllanthus (1) CYPERACEAE Baeothryon (as Scirpus, p.p.) (1) FABACEAE Bulbostylus (1) Vicia (1) Carex (19) Carpha (1) FAGACEAE Eliogeton (as Scirpus, p.p.) (2) Nothofagus (2) Gahnia (1) GENTIANACEAE Isolepis (as Scirpus, p.p.) (3) Gentiana (23) Machaerina (4) Swertia (1) Oreobolus (2) Schoenoplectus (as Scirpus, p.p.) (1) GERANIACEAE Geranium (11) Schoenus(5) Uncinia (2) GESNERIACEAE Aeschynanthus (2) DAPHNIPHYLLACEAE Daphniphyllum (1) Cyrtandra (6) DIPSACACEAE GOODENIACEAE Triplosegia (1) Scaevola (1) GROSSULARIACEAE (as DROSERACEAE Drosera (1) Escalloniaceae) Carpodetus (1) ELAEOCARPACEAE Polyosma (4) Elaeocarpus (S) Quintinia (6) Sericolea (6) GUNNERACEAE EPACRIDACEAE Gunnera (1) Decatoca (1) Styphelia (2) HALORAGACEAE Trochocarpa (S) Gonocarpus (3) Myriophyllum (2) ERICACEAE Agapetes (8) 110

List 7. (continued)

IRIDACEAE Bulbophyllum (22) Patersonia (1) Calanthe (9) Sisyrinchium (-Libertia?) (1) Ceratostylis (13) Corybas (8) JUNCACEAE Dendrobium (59) Juncus (1) Luzula (1) Epiblastus (6) Glomera (16) LAMIACEAE Glossorhyncha (43) Sature ja (1) Habenaria (1) Kerigomnia (I) LENTIBULARIACEAE Liparis (2) Utricularia (3) Mediocular (11) LOGANIACEAE Microtatorchis (5) Fagraea (1) Oberonia (2) Geniostoma (2) Octarrhena 06) Pedochilus (14) LORANTHACEAE Amyemea (4) Peristylus (I) Phreatia (10) MELASTOMATACEAE Platanthera (2) Astronia (?) Pterostylis (3) Medinilla (?) Sarcanthus (1) MONIMIACEAE Spathoglottis (I) Levieria (2) Spiranthes (1) Palmeria (1) Taeniophyllum (1) Thelymitra (1) MORACÉAE Streblus (1) OXALIDACEAE Oxalis (1) MYRSINACEAE Maesa (2) PIPERACEAE Piper (8) Myrsine (as Rapanea) (14) MYRTACEAE PITTOSPORACEAE Deccaspermum (6) Pittosporum (6) Syzgium (3) PLANTAGINACEAE Uromyrtus (1) Plantago (6) Xanthomyrtus (9) POACEAE OCHNACEAE Agrostis (2) Schuurmansia (1) Anthoxanthum (4) OLEACEAE Brachypodium (2) Jasminum (1) Bromus (1) Calamagrostis (as Deyeuxia) (9) ONAGRACEAE Chionochloa (1) Epilobium (4) Danthonia (7) ORCHIDACEAE Deschampsia (1) Agrostophyllum (1) Dichelachne (2) 111

List 7. (continued)

Festuca (4) SANTALACEAE Imperata (1) Cladomyza (6) bachne (2) Exocarpus (1) Microlaena (2) SAPINDACEAE Nastus (2) Dodonaea (1) Poa (22) Racemobambos (1) SCROPHULARIACEAE Trisetum (1) Detzneria (1) Euphrasia (9) POLYGONACEAE Mazus (1) Muelenbeckia (1) Parahebe (12) Polygonum (S) Veronica (2) PORTULACACEAE SPHENOSTEMONACEAE Montia (1) Sphenostemon (1) PROTEACEAE Helicia (1) STERCULIACEAE Commersonia (1) RANUNCULACEAE SYMPLOCACEAE Clematis (1) Symplocos (1) Ranunculus (30) Thalictrum (1) THEACEAE Eurya (18) RHAMNACEAE %rnstroemia (1) Alphitonia (1) Rhamnus (1) THYMELAEACEAE Drapetes (1) ROSACEAE Acaena (1) URTICACEAE Potentilla (8) Cypholobus (2) Prunus (30 Elatostema (4) Rubus (10) Lecanthus (1) RUBIACEAE Parietaria (1) Amaracarpus (10) Pilea (18) Pïpturus (3) Coprosma (11) Urticà (1) Galium (5) Hedyotis (5) VIOLACEAE Hydnophytum (3) Viola (4) Myrmecodia (2) Nertera (1) WINTERACEAE Psychotria (2) Bubbia (S) Timonius (3) Tasmannia (1) RUTACEAE ZINGIBERACEAE Acronychia (2) Geanthus (1) Kolowratia (2) Evodiella (1) Evodia (3) Riedelia (8) Tetractomia (1) 112

PART II. PRELIMINARY STUDIES OF THE FLORA OF THE HUASCARÂN NATIONAL PARK: MONOCOT FAMILIES (EXCLUDING POACEAE) 113

TO THE READER This work is the first part of a floristic manual of the vascular plants of the Huascardn National Park and International Biosphere Reserve, Ancash, Peru. It includes the introductory material and the taxonomic treatment of the monocots, except the Poaceae. Later parts will contain the taxonomic treatment of the Poaceae, dicots and the ferns. This work has been written primarily for resource managers (park and range managers), scientists studying natural phenomena in the Biosphere Reserve, and interested amateurs. It is meant to be practical and usable with a minimum of equipment (basically a lOX hand lens). As a result there is much botanical information that is excluded. The references cited cover a broad range of topics, including many of the botanical topics not included in these pages. In the taxonomic treatment of each family there is a description of the family, the genera, and the species included in the Park flora. Keys are provided for the families of the monocots, and within each family a key to either the genera in the case of large families, or the genera and species in the case of small families. Within the larger genera a key will be provided for the species. DESCRIPTIONS: With the assumption that this work may have a larger audience than people connected with the Huascaràn National Park, the descriptions of the families and genera are written to give an acurate concept of the taxa as they occur in Peru. The descriptions are specific to Peru, and do not attempt to describe the taxa on a worldwide scale. Those interested in family descriptions of global scope should consult Cronquist (1981) or Dahlgren et al. (1985). The descriptions of species are based on the specimen material collected in the Huascaràn National Park, except when the collections are insufficient, and have been compared with descriptions in the reference works cited under the family and genus. To shorten the work, characters held in common by all members of a family or genus are not repeated in the subsequent descriptions of genera or species. KEYS: All keys in this manual are specific to the Huascaràn National Park. TAXONOMIC CONCEPTS: The family descriptions of the angiosperms are based on and modified from the classification system of Cronquist (1981), with the exception of the group of families constituting his concept of the Liliaceae. Finding Cronquist's Liliaceae too broadly defined, I am following the classification system of Dahlgren et al. (1985) 114

for the the Liliaceae and their allies. In addition, the family descriptions have been compared with the works of Cook et al. (1974), Heywood (1978), and Lawrence (1951). The generic concepts are based on the reference works cited under each family or genus, and on my own experience. Author citation for scientific names is a significant part of the descriptions. For the families, the authors have been cited following Cronquist (1981) and Dahlgren et al. (1985): for the genera, following Index Nominum Genericorum (Plantarum) (Farr et al., 1979); for the species, the abbreviations following the Draft Index of Author Abbreviations (Meikle, 1980). Literature is cited following the abbreviations in Serial Sources for the BIOSIS data base (BIOSIS, 1986), except in the few cases that no abbreviation exists in the Serial Sources. Then either Botanico Periodicum Huntianum (Lawrence et al., 1968) or the standard for citation appropriate to non-botanical disciplines will be used. IDENTIFYING PLANTS: The process of identifying a plant involves the formation of mental concepts, or images, of family and genus. When one has an unknown plant in hand, the first question is: to what family does it belong?; and from the family one goes on to the genus, and from the genus to the species. It is important to begin to form a mental picture of each family and genus. For this reason, the familial and generic descriptions are written to give an accurate concept of the taxa as they occur in Peru. While using this work, it will be useful to look at herbarium specimens. The herbaria of the Museo de Historia Natural (USM), Universidad Nacional de Trujillo (HUT), and the Universidad Nacional de Cajamarca (CPUN) have collections rich in the families and genera represented in the Huascaràn National Park including duplicates of the specimens collected for this study, and the National Park has an on-site herbarium. In addition to the descriptions of the taxa, there are notes on their distribution, habitat, pollination, uses, and miscellaneous comments. The distributional data are taken from the literature, herbarium specimens, and the specimens collected in the Park. The species distribution is listed in three ways: worldwide distribution, distribution within Peru, and within the Park citing major river drainages. Specimens will be cited for the localities within the Park; those collected by Smith and collaborators will be listed by collection number only, and those collected by other collectors by collector and number. For families and genera, only worldwide data are given. The elevational range applies only to the altitudinal distribution of the species within the Park. Habitat data are US specific to the ecological distribution of the families and genera within Peru, and of the species within the Park. Comments on pollination are taken from the literature, label data on herbarium specimens, or personal communications. The uses reported are from the literature, or more often from personal observation or from conversation with co-workers and local residents. Throughout the text the following abbreviations are used: Que. - quebrada (valley), H. N. P. - Huascarân National Park, and diam. - diameter. 116

MATERIALS AND METHODS Study of the Huascarân National Park flora includes two phases: fieldwork and herbarium study. The fieldwork involved 18 months of collecting in the Park from the latter part of 1984 until 1986, during which time 3981 specimen sets were collected. Ideally each set should include 12 replicates, but they vary from a single specimen sheet to 12 or more given abundance of material. Aside from camping equipment, field equipment included an altimeter, binoculars, pruning shears, digging tools, and a field press. Collection trips were made by vehicle, a 4-wheel drive Jeep CJ7, or, more often, on foot with beasts of burden for equipment cartage. One or more assistants usually helped during collecting trips. In the field, collection data, observations, and preliminary determinations were recorded into a fieldbook, and the specimens pressed in newsprint and stored in plastic bags, with alcohol as a preservative, until return to the city of Huaraz, the base of operations. In Huaraz, the specimens were dried in herbarium presses, with aluminum corrugates, over a drier employing 2-250 watt electrical resistance strip heaters. After drying, the specimen sets destined for Peruvian herbaria were segregated and dispersed, and those for non-Peruvian institutions shipped. When possible, the specimens were further studied in Huaraz, at the Museo de Historia Natural in Lima, or the Universidad Nacional at Trujillo. In Fall of 1986, the herbarium study phase began. Initially, specimens from all families were determined in order to bring the list of the flora as close to completion as practical. However, there are a number of genera which pose problems requiring detailed study, and they will be fully determined only after intensive studies have been completed. In the process of studying and writing the treatments for the families included here, standard herbarium techniques have been used: measurement of different plant parts, dissection and observation with a low power microscope, and consultation of published works and comparison with determined herbarium specimens and photographic materials. The microscopic equipment is an Olympus dissecting microscope, 7.5-64 x, zoom, and a fiber optic illuminator with variable intensity control. Given the small size of the plant parts to be dissected, high quality dissection equipment is necessary: watch maker's forceps, fine teasing needles, and micro-scalpels. Prior to dissection, the plant parts are softened by immersing them in water, with a few drops of a softening agent, heated to near boiling. The material for dissection is not boiled, as boiling the specimen can agitate the material excessively and may damage it. 117

While dissecting, the material can be maintained soft by adding drops of a softening agent. Pohl's (Pohl, 1954) solution is very useful, especially for chaffy material such as grasses, sedges, and rushes. The descriptions included in the taxonomic portion of this work are based on observations and measurements taken from the specimens collected in the Huascarân National Park, and were made with metric rulers, or an optical micrometer installed in the dissecting microscope. Data collected in the field and included on the specimen label are often included in the discriptions. Work processing was done on a Zenith microcomputer with WordStar and Microsoft Word software. The following is a list of the monocotyledonous taxa included in the flora: AGAVACEAE Furcraea occidentalis Trel. ALLIACEAE Trichlora peruviana Baker in Hook. ALSTROEMERIACEAE Alstroemeria pygmaea Herbert Bomarea albimontana D. N. Smith & Gereau, ined. B. dulcis (Hook.) Beauv. B. phyllostachya Masters ex Baker, vel sp. aff. B. zosterifolia Killip AMARYLLIDACEAE Stenomesson luteum (Herbert) Baker in Saund. ANTHERICACEAE Echeandia eccremorrhiza (Ruiz Lôpez & Pavôn) Cruden, comb.ined. BROMELIACEAE Pitcairnia pungens Kunth var. pungens Puya angusta Lyman B. Smith P. herrerae Mez, vel sp. aff. P. membranacea Lyman B. Smith, vel sp. aff. P. raimondii Harms P. rauhii Lyman B. Smith, vel sp. aff. P. reflexiflora Mez P. sp. nov. (9196) Tillandsia cerrateana Lyman B. Smith T. fendleri Griseb. var. reducta (Lyman B. Smith) Lyman B.Smith T. humilis Presl T. ionochroma André ex Mez T. lopezii Lyman B. Smith T. oroyensis Mez T. rubella Baker 118

Tillandsia tectorum E. Morren T. waltheri Mez var herrerae (Harms) Rauh COMMELINACEAE Commelina fasciculata L. CYPERACEAE Baeothryon (Scirpus) rigida (Boeckeler) D. N. Smith, ined. Bulbostylis juncoides (Vahl) KQk. Carex ancashensis D. N. Smith & Reznicek, ined. C. boliviensis Van Juerck A MOll.-Arg. C. bonplandii Kunth C. collumanthus (Steyerm.) Mora C. crinalis Boott C. ecuadorica KQk. C. hebetata Boott C. mandoniana Boeckeler Cyperus hermaphroditus (Jacq.) Standley C. sesleroides Kunth Eleocharis albibracteata Nees & Meyen ex Kunth Isolepis inundatus R. Br. Oreobolopsis tepalifera T. Koyama & Guaglian. Oreobolus obtusangulatus Gaud. Phylloscirpus sp. inq. (-Scirpus aff. atacamensis Boeckeler) P. deserticola (Phillipi) D. N. Smith, ined. P. sp. inq. ("Scirpus aff. hieronymi Boeckeler) Schoenoplectus californicus (C. Meyer) Sojak subsp. californicus var. californicus DIOSCORIACEAE Dioscorea ancashensis Knuth D. larecajensis Uline ex Knuth HYDROCHARITACEAE Elodea potamogeton (Bertero) Espinosa IRIDACEAE Orthrosanthus occissapungus (Ruiz ex Klatt) Diels Sisyrinchium brevipes Baker S. chilensis Hook., vel sp. aff. S. jamesonii Baker S. junceum E. Meyer ex Presl S. praealtum Krânzlin S. tinctorium Kunth S. trinerve Baker Tigridia philippiana I. M. Johnston JUNCACEAE Distichia muscoides Nees & Meyen Juncus arcticus L. var. andicola (Hook.) Balslev J. bufonius L. J. cyperoides Laharpe J. ebratceatus E. Meyer J. imbricatus Laharpe 119

Juncus pallescens Lam. J. stipulatus Nees A Meyen J. tenuis Willd. var. platycaulos (Kunth) Bucheneau Luzula gigantea Desv. L. racemosa Desv. L. vulcanica Liebm. JUNCAGINACEAE Lilaea scilloides (Polret) Hauman LEMNACEAE Lemna mlnuscula Herter ORCHIDACEAE Aa mathewsii (Reichb. f.) Schltr. A. paleacea (Kunth) Reichb. f. Epidendrum cernuum Kunth E. excelsum C. Schweinf., vel sp. aff. E. inamoena KrSnzlin, vel sp. aff. E. megagastrium Lindley Gomphichis valida Reichb. f. Malaxis andicola (Ridley) O. Ktze. Masdevallia amabilis Reichb. f. & Warsc. Myrosmodes nubigena Reichb. f. M. paludosa (Reichb. f.) D. N. Smith, comb. ined. Odontoglossum rigidum Lindley Pleurothallis spiralis (Ruiz Lôpez & Pavôn) Lindley P. trilineata Lindley Pterichis triloba (Lindley) Schltr. Stelis cf. cupuligera Reichb. f. & Warsc. S. cf. flexuosa Lindley S. cf. leucopogon Reichb. f. Trichoceros platyceros Reichb. f. POACEAE Aciachne acicularis Lsgaard A. pulvinata Benth. Aegopogon cenchroides Humb. & Bonpl. ex Willd. Agrostis auraucana Philippi, vel sp. aff A. breviculmis Hitchc. A. gelida Trin., vel sp. aff. A. nigritella Pilger, vel sp. aff. A. tolucensis Kunth A. aff. tolucensis Kunth Alopecurus aequalis Sobol Anthochloa lepida Nees & Meyen in Meyen Aristida enodis Hackel Bouteloua simplex Lag. Brachypodium mexicanum Link Bromus berterianus Colla B. catharticus Vahl B. lanatus Kunth 120

Bromus pitensis Kunth B. villosissimus Hitchc. Calamagrostis antoniana (Griseb.) Steud. ex Hitchc. C. brevifolia (Presl) Steud. C. chrysantha (Presl) Steud. C. curvula (Wedd.) Pilger C. densiflora (Presl) Steud. C. eminens (Presl) Steud. C. fuscata (Presl) Steud. C. glacialis (Wedd.) Hitchc. C. heterophylla (Wedd.) Hitchc. C. jamesonii Steud. C. macbridei Tovar C. nitidula Pilger C. ovata (Presl) Steud. C. recta (Kunth) Trin. C. rigida (Kunth) Trin. C. speciformis Hackel ex Stuckert C. spicigera (Presl) Steud. C. swallenii Tovar C. tarmensis Pilger C. vicunarum Wedd. C. weberbaueri Tovar Cortaderia aristata Pilger C. hapalotricha (Pilger) Conert C. nitida (Kunth) Pilger C. sericantha (Steud.) Hitchc. C. trianae Stapf, vel sp. aff. Critesion muticum (Presl) A. LOve Dielsiochloa floribunda (Pilger) Pilger Dissanthelium calycinum (Presl) Hitchc. D. minimum Pilger Elymus angulatus Presl Elytrigia attenuata (Kunth) Covas Eragrostis pastoensis (Kunth) Trin., vel sp. aff. E. pilgeriana Hitchc. Festuca breviaristata Pilger, vel sp. aff F. casapaltensis J. Ball F. dichoclada Pilger F. dolichophylla Presl F. glyceriantha Pilger, vel sp. aff. F. huamachucensis Infanta, vel sp. aff. F. lasiorachis Pilger F. pubigluma Tovar F. guadridentata Kunth F. rigidifolia Tovar F. ulochaeta Steud., vel sp. aff F. weberbaueri Pilger Hierochloë redolens (Sol. ex Vahl) Roem. & Schult. Melica scabra Kunth 121

Muhlenbergia angustata (Presl) Kunth M. fastigata (Presl) Henrard M. flexuosa Hitchc., vel sp. aff. M. ligularis (Hackel) Hitchc. M. peruviana (Beauv.) Steud. Muhlenbergia sp. inq. Nassella brachyphylla (Hitchc.), comb. ined. N. depauperata (Pilger), comb. ined. N. featherstonei (Hitchc.), comb. ined. N. inconspicua (Presl), comb. ined. N. mexicana (Hitchc.), comb. ined. N. mucronata (Kunth), comb. ined. Paspalum bonplandianum FlQgge P. pygmaeum Hackel P. tuberosum Mez Pennisetum clandestinum Hochst. ex Chiov. Piptochaetium indutum Parodi P. panicoides (Lam.) Desvaux Poa aff. aeguatoriensis Hackel P. aff aeguigluma Tovar P. annua L. P. aff. brevis Hitchc. P. aff. candamoana Pilger P. aff. fibrifera Pilger P. aff. ferreyrae Tovar P. aff. gilgiana Pilger P. aff. glaberrima Tovar P. aff. horridula Pilger P. aff. lilloi Hackel P. aff. pardoana Pilger P. aff. pearsonii Reeder P. aff. perligulata Pilger P. aff. subspicata (Presl) Kunth P. sp. inq. Polypogon interruptus Kunth Schizachyrium hirtiflorum Nees Sporobolus lasiophyllus Pilger Stipa hans-meyerii Pilger S. ichu (Ruiz Lôpez & Pavôn) Kunth S. obtusa (Nees & Meyen) Hitchc. Trisetum macbridei Hitchc. T. spicatum (L.) Richt. Vulpia bromoides (L.) S. R. Gray V. myuros (L.) K. Gmelin var. hirsuta Hackel POTAMOGETONACEAE Potamogeton paramoanus Haynes & Holm-Nielsen ZANNICHELLIACEAE Zannichellia andina Holm-Nielsen & Haynes 122

GENERAL KEYS TO THE FAMILIES Key to Flowering Material la. Plants thalloid, leafless, 1-5 mm diameter; free-floating aquatics; flowering parts very small, in lateral or dorsal pouches, seldom seen. Lemnaceae lb. Plants developing leaves and/or stems, not minute; of various habits other than free- floating aquatic. 2. 2a. Petals and sepals wanting or reduced to bristles, scales, or a cupule at base of gynoecium. 3. 2b. Petals and sepals developed and conspicuous. 6. 3a. Gynoecium subtended by a pair of bracts (lemma and palea) and lodicules or by a floral bract and bristles, scales,or a perigynium; inflorescence composed of spikesor spikelets of flowers; mostly terrestrial. 4. 3b. Gynoecium subtended by 1 tepal, or a cupule or perianth wanting; inflorescence composed of individual flowers; aquatics. 5. 4a. Female or perfect flower a gynoecium and/or androecium subtended by a lemma, palea, and lodicules aggregated into I-many-flowered spikelets; plants of various habitats. Poaceae 4b. Female or perfect flower a gynoecium and/or androecium subtended by a floral bract and bristles, scales, a perigynium, or naked; aggregated into 1-many-flowered spikes; plants of mainly marshy or boggy habitats. Cyperaceae Sa. Inflorescences all axillary; perianth reduced to a cupule below the gynoecium; leaves cauline. Zannichelllaceae Sb. Inflorescences scapose-exserted and axillary-sessile; tepal 1, bract-like; leaves basal. Juncaglnaceae 6a. Ovary inferior. 7. 6b. Ovary superior. 13. 7a. Caulirosettes, 1 or more m tall; leaves 1 m long or more,stiff, sword-like; inflorescence scapose, axis several m tall. Agavaceae 7b. Herbs; leaves not as above; inflorescences much shorter. 8. 8a. Flowers strongly zygomorphic, resupinate, the lower petal usually enlarged and showy. Orchldaceae 8b. Flowers actinomorphic to slightly zygomorphic, one tepal not enlarged relative to others. 9. 123

9a. Leaves cauline; terrestrial or aquatic. 10. 9b. Leaves basal; terrestrial, from bulbs, short rhizomes or fleshy roots. 12. 10a. Submerged aquatics; weakly rhizomatous. Hydrocharltaceae 10b. Terrestrial, from tubers or rhizomes with swollen storage roots; vines or diminutive plants. 11. 11a. Flowers unisexual, in racemes, plants dioecious; from tubers Dioscoreaceae lib. Flowers perfect, in umbel-like cymes; from slender rhizomes with swollen storage roots. Alstroemerlaceae 12a. Inflorescence cymose, subtended and partially hidden by a spathe of 1-2 sheathing, membranous bracts; flowers ephemeral; from rhizome or bulb. Irldaceae 12b. Inflorescence umbel-like, subtending bracts small, chaffy, deciduous; flowers persist; from bulbs. Amaryilidaceae 13a. Plants floating-leaved aquatics, rooted on the pond bottom. Potamogetonaceae 13b. Plants terrestrial or semi-aquatic, if aquatic, then large, emergent, self-supporting plants. 14. 14a. Leaves cauline; stem elongate, herbaceous; inflorescence hidden in a folded spathe; flowers ephemeral. Commellnaceae 14b. Leaves basal or rosulate at stem apex; stem short or, if elongate, woody; inflorescence without a spathe. IS. 15a. Perianth chaffy or bract-like, persisting into fruit. Juncaceae 15b. Perianth, at least the corolla, membranous, not persisting. 16. 16a. Short-stemmed epi- or lithophytes, or terrestrial caulirosettes; pubescence of peltate scales; flowers subtended by prominent floral bracts. Bromeliaceae 16b. Terrestrial herbs; if pubescent, not with peltate scales; floral bracts inconspicuous. 17. 17a. Plants from fleshy roots; inflorescence a raceme. Antherlcaceae 17b. Plants from bulbs; inflorescence umbel-like. Alllaceae 124

Key to Fruiting Material la. Fruit fleshy, indehiscent, a berry; plants aquatic. Hydrocharltaceae lb. Fruit dry, dehiscent or indehiscent, a capsule, achene, caryopsis, utricle or druplet; plants terrestrial or aquatic. 2. 2a. Fruit a capsule. 3. 2b. Fruit an achene, caryopsis, drupelet, or utricle. 13. 3a. Derived from an inferior ovary. 4. 3b. Dervied from a superior ovary. 9. 4a. Capsule unilocular, dehiscing centrally by 3-6 longitudinal slits, apex and base remaining entire; seeds very fine, like dust. Orchidaceae 4b. Capsule trilocular (unilocular in Luzula, Juncaceae), dehiscing apically; seeds much larger. S. Sa. Caulirosettes, very large; leaves 1 m or longer, fleshy, sword-like; inflorescence a scapose panicle several m long. Agavaceae 5b. Herbs, smaller in all characters. 6. 6a. Rootstock a tuber; vines or diminutive herbs immersed in soil; capsules trigonous, margins winged. Dloscoreaceae 6b. Rootstock a rhizome, some with fleshy storage roots, bulbs, or corms; capsules round in cross-section. 7. 7a. Leaves basal; from rhizomes or bulbs. 8. 7b. Leaves cauline; from delicate rhizome with fleshy storage roots; vines or dwarf herbs immersed in soil. Alstroemerlaceae 8a. Inflorescence subtended by a spathe of 2 bracts; leaves equitant; from bulbs or rhizomes. Irldaceae 8b. Inflorescence subtended by small, chaffy bracts; leaves several ranked; from bulbs. Amaryllldaceae 9a. Leaves basal; from rhizomes or bulbs. 10. 9b. Leaves cauline or rosulate on stem; rhizomatous. 12. lOa. Plants from bulbs. Alllaceae I Ob. Plants from rhizomes or fibrous roots. 11. I la. Perianth membranous, deciduous. Anthericaceae lib. Perianth chaffy, persistent in fruit. Juncaceae 12a. Leaves cauline; stems herbaceous, elongate, jointed; terrestrial. Commellnaceae 125

12b. Leaves rosulate on short stem, or apical on stout, woody stem; plants epi- or lithophytic, or terrestrial. Bromeliaceae 13a. Fruit an achene or caryopsis. 14. 13b. Fruit a utricle or drupelet. 16. 14a. Fruit a caryopsis surrounded by 2 bracts, the larger 1-several-nerved, the smaller 2-nerved, or wanting. Poaceae 14b. Fruit an achene. 15. 15a. Achenes naked, trigonous; leaves spiralled, hollow septate, complanate, sulcate; stems very short; aquatic, in vernal pools. Juncaglnaceae 15b. Achenes subtended by bristles, scales, or naked, lenticular or trigonous; leaves 3- ranked or reduced; stem trigonous or terete; various habitats. Cyperaceae 16a. Free-floating aquatics; plants leafless, minute; fruit a utricle, seldom seen. Lemnaceae 16b. Rooted aquatics; stems and leaves submerged to floating; fruits drupelets. 17. 17a. Fruits in leaf axils, submerged, endocarp spiny. Zannlchelilaceae 17b. Fruits on erect, emergent spikes; endocarp spomgy, fruit floats. Potamogetonaceae 126

AGAVACEAE Endlicher LILIIDAE: LILIALES CAULIROSETTES, Stem short, or to several meters tall. LEAVES: simple, densely crowded at stem apex, long lasting, sessile, coriaceous to carnose, margins often spinose dentate. INFLORESCENCES: terminal, often large, often scapose, racemes, panicles or heads. FLOWERS: perfect, actinomorphic, epigynous or hypogynous (Yucca); perianth of 6 petaloid segments, equal, often carnose, basally united into a tube in Agave and Furcraea; stamens 6, filaments free, anthers versatile; ovary inferior or superior (Yucca), trilocular, placentation axile, nectaries septal or wanting, style terminal, stigmas 3, wet or dry, ovules several per locule. FRUIT: a loculicidal capsule. This family of ca. 18 genera and ca. 300 species is native to the arid regions of the Americas, introduced throughout the arid regions of the world. Agave, introduced from Mexico, and Furcraea are the only representatives in Peru, unless Yucca has been introduced as an ornamental. HABITAT: Arid regions, especially on the western slopes of the Andes, and natural grasslands. POLLINATION: Animal-mediated: by insects, birds and bats. BREEDING SYSTEM: Many of the species in the family are monocarpic: that is, after a long period of vegetative growth, they flower and fruit once, and then die. USES: Cordage is produced from the fibrous leaves, and the sap of different species of Agave is fermented into pulque, a chicha-like beverage, then distilled to produce tequila or mescal, depending on the species used to produce the pulque (Simpson & Conner- Ogorzaly, 1986). PHYSIOLOGY: CAM photosynthesis is common in the family. TAXONOMIC COMMENTARY: The family needs more, detailed study. It has been most extensively studied in North America, especially Mexico (Gentry, 1972, 1978, 1982). COLLECTING PRACTICES: As the plants are very large and succulent, it is difficult to collect a representative specimen. The following are desirable: very complete field notes, a series of photographs of the habitat, habit, and major parts of the plant, measurements of the leaves and inflorescence, pressed cross-sections of the peduncle, pressed peduncle and inflorescence bracts, and flowers pressed whole and sectioned, with abundant additional material for dissection. If the leaves are too thick to press, at least the apex, sections of the margins, and cross-sections of the blade should be pressed. It 127

is important to note if the plant is producing vegetative propagules, either at the base of the plant or in the inflorescence. Any notes on pollinators are very valuable. I thank Mr. Roy E. Gereau (MO) for review and revision of the manuscript.

Furcraea Ventenat References: Baker, 1888; Standley & Steyermark, 1952. CAULIROSETTES, stems usually short, occasionally developing a columnar trunk. LEAVES: simple, long-lasting, sessile, long and narrow, spinose-dentate to entire at margin, somewhat to strongly succulent, densely crowded at stem apex. INFLORESCENCE: a panicle, large terminal, elongate, branches spreading and lax; peduncle to several meters tall. FLOWERS: epigynous; pedicellate, solitary or grouped along inflorescence branches; perianth of 6 equal, carnose segments, basally united into a short, cylindrical tube, limbs spreading to rotate, whitish; stamens 6 free, inserted at throat of perianth tube, shorter than perianth limbs, filaments broadened and fleshy basally; ovary inferior, trilocular ovary, style terminal, thickened and 3-angled basally, stigmas capitate-trilobate. FRUIT: a loculicidal capsule; seeds flattened. Furcraea is a genus of 20 species, occurring from southern Mexico to Bolivia, the West Indies, and in southern Brazil. BREEDING SYSTEM: The species are reportedly monocarpic. Little seed is produced (Leon, 1968), and reproduction is predominantly vegetative. Unlike Agave, Furcraea does not produce shoots from the base of the plant (Everett, 1981), but rather it produces many bulbils as well as flowers in the inflorescence. The bulbils are the most important propagule. USES: The axis of the inflorescence is used in rustic construction. The fiber of Furcraea is of higher quality than that of Agave (Dahlgren et al., 1985), and is used in bags, cordage, and many other woven fiber artifacts.

Furcraea occidentaHs Trel. Reference: Trelease, 1913. CAULIROSETTE trunkless or at most with a very short trunk. LEAVES: ca. 65 cm long, ca. 10 cm wide, narrowly oblong-lanceolate, margin toothed with spines ca. 1 mm long, deltoid, straight to retrorsely curved, light brown to blackish, leaf apex attenuate, terminating in a brown spine. INFLORESCENCE: a panicle, ca. 6 m tall, open, pyramidal, branches spreading and lax, glabrous; developing abundant bulbils. 128

FLOWERS: perianth ca. 30 mm long, whitish-green; gynoecium: ovary ca. 20 mm long. FRUIT: not described. (This description is adapted from Trelease, 1913). habitat: Arid vegetation types on the western slopes of the Andes. DISTRIBUTION: Peru: Ancash and Lima. H. N. P.: seen at the mouth of Santa Cruz, and at Cerro Cunka on the trail to Auquispuquio. Ca. 3240-3400 m. PHENOLOGY: Flowering reported in November. TAXONOMIC COMMENTARY: The type of this species, Weberbauer 1687, was collected at Matucara, Ancash. There is another species of Peruvian Furcraea, F. andina Trel., published in Bailey's Standard Cyclopedia of Horticulture (Trelease, 1915). It is discussed in passing as part of a group of trunkless species including F. elegans Tod., F. macrophylla Hooker, and F. pubescens Tod. Trelease validly published F. occidentalis in 1913, and it is unclear why F. andina was not published in a similar fashion. I do not consider F. andina to be validly published: it is neither fully described as species in its own right, nor directly compared to any validly published species. The group comparison in Bailey's Cyclopedia (Trelease, 19IS) does not constitute an unequivocal description of the species, and cannot be accepted. Macbride (1936) cites a type without a collection number. Furlong; no type is mentioned in the description (Trelease, 1915). If F. andina were valid, the literature gives us little to adequately distinguish it from F. occidentalis. The comparison in Flora of Peru (Macbride, 1936) is based on leaf shape and marginal spines. The comparison of leaf shapes, narrowly oblong versus oblong-lanceolate, is trivial and in disagreement with the original description of F. occidentalis. This leaves only the spines as the distinguishing character. F, occidentalis is minutely toothed with spines ca. 1 mm long (Trelease, 1913), and F. andina is prominently toothed with spines ca. 12.5 mm long and ca. 89 mm apart (Trelease, 1915). Furcraea occidentalis is from the western slopes of the Andes and F. andina from eastern Peru. This problem can only be resolved by further studies of Peruvian Furcraea. CONSERVATION STATUS: Rare due to scarcity of suitable habitat within the Park. 129

ALLIACEAE J. G. Agardh LILIIFLORAE: LILIALES HERBS, from tunicated bulbs; stems short or elongate. LEAVES: alternate, basal, simple, sessile, linear, entire, sometimes hollow, sometimes with pronounced odor. INFLORESCENCE: terminal, scapose umbels, or solitary, subtended by membranous bracts. FLOWERS: perfect, actinomorphic or zygomorphic, hypogynous; perianth segments 6, alike, or distinctly sepals and petals, free or connate, forming a floral tube; stamens 6, filaments free, or united into a tube, elongate, anthers versatile or basifixed; gynoecium 3-carpelIate, united, superior, 3-locular ovary, placentation axile, style terminal, trilobed or trifid, stigmas dry, papillate, ovules few-many. FRUIT: a loculicidal capsule. The Alliaceae has 30 genera and ca. 720 species, and is nearly cosmopolitan in distribution. Certain subfamilies, tribes, and genera have a much more regionally restricted distribution. The subfamily Gilliesioideae, a group of ca. 9 genera including Trichlora, is restricted to southern South America, with its greatest diversity in Chile.

Trichlora Baker in Hook. Reference: Baker, 1877 This is a monotypic genus endemic to the Andes of Peru.

Trichlora neruviana Baker in Hook. HERB, perennial, to 25 cm tall; bulb 2.S-4.S cm long, 1.4-1.7 cm wide, ovoid, outer tunics chartaceous, deeply buried, with taproot and fibrous roots, leaves and stems separate from the bulb easily. LEAVES: exposed portion 14.5-34 cm long, 3.5-5.5 mm wide, linear, margins minutely crenate, this possibly a resinous exudate, apex attenuate- rounded, glabrous, 4-5 per plant. INFLORESCENCES: scapose, umbel-like, but pedicels unequal, scapes 1-3 per plant, (3) 7-18 cm long, ca. I mm diameter, 1-3- flowered, pedicels subtended by 2, narrowly lanceolate, membranous bracts, 9-22 mm long, ca. 4 mm wide, deciduous. FLOWERS: perfect, regular, hypogynous; pedicels 23- 37 (-85) mm long; flowers nodding; sepals 3, 30-50 mm long, 2.5-5 mm wide at base, basebroadly ovate, apex long attenuate, yellowish green with purplish cast at margins, spreading; petals 3, 3-5 mm long, 2.5-5 mm wide, obtriangular, angles rounded, apex emarginate to 0.8 mm deep, both faces scaly, deep purple; stamens 3, filaments 1.8-2.3 mm long, ca. 0.8-1mm wide, ovate-triangular, probably fleshy when fresh, anthers 1.2- 130

1.5 mm long, ca. 0.3 mm wide, narrowly elliptic; gynoeclum 3-carpellate, united into a 3-locular, 3-lobed, superior ovary, style terminal, 1-1.5 mm long, basally inflated, with 3 horn-like lobes, inner surfaces stigmatic, papillate, purple, placentation axile, ovules 1 2 per locule. FRUIT a loculicidal capsule; mature seeds not seen. TAXOMONIC NOTE: Baker (1877) found the umbels 4-6-flowered, the style cyathiform, and the anthers ovate-globose and versatile. HABITAT: Warm rocky microsites in seasonally dry grassland with intermittant shrubs, steep slopes. 3700-3800 m. DISTRIBUTION: Peru: Ancash, Apurimac, and Lima. H. N. P.: Auquispuquio, (11911) along the trail about 100-200 m lower than the elevation of the ruins. PHENOLOGY: Flowering in April; fruiting just begining in April. CONSERVATION STATUS: Very rare. 131

ALSTROEMERIACEAE Dumortier LILIIFLORAE: ASPARAGALES HERBS, perennial; rhizomatous, some roots fleshy or tuberous; stems very short to elongate, erect to twining. LEAVES: simple, alternate, lanceolate to linear, base attenuate, blades often resupinate. INFLORESCENCE: terminal, umbel-like cymes, rarely solitary, often subtended by leaf-like bracts. FLOWERS: perfect, actinomorphic to slightly zygomorphic, epigynous; pedicels short to elongate; perianth segments 6, free, usually somewhat unequal in size, shape, and often differently colored, spreading or connivent and campanulate, petals decurrent on ovary; nectaries at base of petals; stamens 6, free, anthers basifixed; gynoeclum 3-carpellate, compound, wholly or partially inferior, 3-locular ovary, placentation axile, style terminal, elongate, apex trifid, stigmas wet, ovules several per locule. FRUIT: a loculicidal capsule; seeds sometimes covered with a bright sarcotesta. The family is native to Central and South America, and is most abundant in the Andes. There are 4 genera and ca. 160 species; Alstroemeria and Bomarea are found in Peru. HABITAT: Montane rainforests, shrublands, dry high elevation grasslands, the arid western slopes of the Andes, and the costal lomas formation. POLLINATION: By insects and hummingbirds. USES: Some are used as garden ornamentals. The roots of some species are edible. TAXONOMY: Cronquist (1981) includes this family in the Liliaceae. I thank Mr. Roy E. Gereau (MO) for sharing his knowledge of the family, and for his careful revision of the manuscript.

Key to the species of the Alstroemeriaceae la. Plants acaulescent, nearly submerged in soil; inflorescence a solitary flower; flower yellow with deep red markings. Alstroemeria pygmaea lb. Plants with elongate stems, erect or vining; inflorescence a compound, umbel-like cyme; flowers pink or reddish. 2. 2a. Vines, stems several meters long; inflorescence congested, hemispherical, many-flowered; growing in dense, tall brush-fields. Bomarea aibimontana 2b. Erect herbs, rarely clambering, less than 1 m tall; inflorescence open, few- flowered; growing in grasslands and low brushfields. 3. 132

3a. Leaves flat, narrowly elliptical, 4-13 cm long; sepals and petals subequal. Bomarea phyllostachya 3b. Leaves revolute, linear, less than 7.5 cm long; petals longer than sepals. 4. 4a. Inflorescence branches less than 2 cm long; flowers campanulate. Bomarea dulcis 4b. Inflorescence branches 2.5-3 cm long; flowers tubular. Bomarea zosteraefolia

Alstroemeria Linnaeus HERBS erect, from tuberous roots, acaulescent or caulescent. FLOWERS: slightly zygomorphic; perianth yellowish, orange or reddish, spreading; petals often with longitudinal spots of color; stamen filaments curving, unequal; gynoeclum half-inferior. FRUIT: seeds globose, lacking a sarcotesta. The genus contains 60 species, about half are Andean, found from Peru to Chile, and the remainder in Brazil. There are ca. 6 species in Peru. Alstroemeria aurantiaca is grown as a garden ornamental.

AUtrocmgrlii PYgnmtn Herbert HERB from a cluster of deeply (to 10 cm) buried, tuberous roots; acaulescent, the stems thin and submerged in soil, only leaves and inflorescences above ground surface. LEAVES: 27-48 mm long, 2-4 mm wide, in apical clusters of 4-10 per stem, blue- green, erect, narrowly spatulate, somewhat sheathing at base, thin, papery, blades subsucculent, 5-9-nerved, margins entire, apex acute-revolute. INFLORESCENCE: a solitary, terminal flower. FLOWERS: slightly zygomorphic by unequal placement of perianth segments and stamens; sepals 19-21 mm long, 4-7 mm wide, spatulate, yellow, occasionally with red markings; petals 19-25 mm long, 3-5 mm wide, narrowly spatulate, yellow with dark red dots or longitudinal streaks from base to middle; stamen filaments of slightly unequal lengths, 7.5-8.5 mm long, anthers (2-) 2.5-2.7 mm long, base cordate, apex acute-apiculate, yellow; ovary with a pronounced rim at perianth insertion, style 12 mm long, erect, thin, branches 3 mm long. CAPSULE: ca. 12 mm long, ca. 8mm diam., ellipsoidal, green. HABITAT: Grasslands, or grassland-shrubland, in rich soils, north and northwest aspects. 3900-4400 m. 133

DISTRIBUTION: Peru and Bolivia. It is reported from Patagonia; however the Patagonian material is a separate species, A. patagonica Philippi (Ravenna, 1969). Peru: Ancash, Cuzco, Junin, Lima, Pasco, and Puno. H. N. P.: Auquispuquio (12076), Santa Cruz (9244), and Shallap (9668). PHENOLOGY: Flowering in January and February, and fruiting from January to May. Mature fruit seen in May. CONSERVATION STATUS: Rare.

Bomarea Mirbel in Buffon References: Baker, 1888; Killip, 1936a, 1936b. VINES or erect herbs; from a delicate rhizome with tubers or with roots tuberous- thickened dis tally; stems often elongate. LEAVES: resupinate in subgenera Bomarea and Sphaerine. INFLORESCENCE: umbel-like cymes, simple or compound, often many-flowered, or sometimes flower solitary; subtended by an involucre of leaf-like bracts. FLOWERS: actinomorphic; perianth segments usually connivent and campanulate, slightly spreading in some species, the two whorls differentiated in size, shape and color, yellow to red, rarely green or cream; sepals oblong to oblanceolate, often shorter than petals, thickened; petals spatulate, usually clawed, often with spots of color within; stamen about as long as perianth segments, barely exserted at anthesis; filaments acicular, basally thickened, tip filiform, inserted into a deep basal pit on the anther, the pit unequally rimmed, nearly open on abaxial side, anthers thus sub-versatile; ovary wholly or half inferior. CAPSULE: spherical, often 3-angled; seeds globose, with a bright sarcotesta. A genus of ca. 100, mostly high Andean, species, Bomarea is found in the mountains from Mexico to Chile and Argentina, in the West Indies, and one species grows in Guyana. There are about 60 species in Peru. Some species are grown for ornament. POLLINATION: By hummingbirds. BREEDING SYSTEM: All species examined were protandrous. The pollen is shed, and the anthers nearly ready to fall before the stigmas develop and become receptive.

Bomarea alblmontana D. N. Smith & Gereau, sp. nov. ined. VINE; rootstock not seen; stem several meters long, 2-4.7 mm diameter, stiff, twining, rufous-velutinous, often densely, apex pendant. LEAVES: not resupinate, somewhat 134

dimorphic, those of lower stem (36-) 55-66 mm long, 5-10 (-13.5) mm wide, lanceolate, stiff, arcuate, somewhat appressed to stem, those of apex 44-76 mm long, (8-) 12-19 mm wide, elliptic, flattened, widely spreading horizontally in live material, both types sessile, bases short attenuate, rufous-velutinous, upper face glabrous with many, prominent nerves, lower face white pubescent, nerves pilose, internerves velutinous, the hairs multicellular, flattened, margins revolute, apex acute-revolute, blue-green. INFLORESCENCE: hemispherical; involucral bracts 6-11, leaf-like, (40-) 59-73 mm long, 9-18 mm wide, elliptic, widely spreading horizontally in live material; (6-) 17-29- flowered; branches to 35 mm long, 1-2 (-3) forked, with a bract at each node; peduncles very short, 2-4 (-6.2) mm long; floral bracts 8.5-13 mm long, 2.7-6 mm wide, ovate- attenuate to lanceolate, membranous, many-nerved, papery, persistent rarely much larger and sub-foliaceous. FLOWERS: pedicels (3-) 10-27 mm long, filiform, puberulent; perianth segments appearing of equal length due to higher insertion of petals, connivent and campanulate, decurrent on ovary; sepals 3, 17-25 mm long, 10-13 mm wide, oblanceolate, apex acute-revolute, pink; petals 3, 19-24 mm long, (4.5-) 6.5-7.5 mm wide, spatulate, apex acute, claw papillose at base abaxially, with papillose nectary at base adaxially, yellow, adaxlal face purple spotted, costa plicate along 3 central nerves, pink abaxially; stamen filaments (10-) 17.5-19 mm long, terete, base 1-1.5 mm diameter, puberulent, anthers 5.5-8.2 mm long, 1.5-2.7 mm wide, base cordate, apex acute- apiculate, surface mammilose; ovary half inferior, upper surface papillose, style 12-18.5 mm long, puberulent stigmas papillose. CAPSULE: 11-23 mm long, 11-15 mm wide, sub-spherical, blue-green; seeds ca. I mm diam., globose, surrounded by a fleshy sarcotesta, bright orange. HABITAT: Tall shrublands and Polylepis weberbaueri woodlands. 3700-4600 m. DISTRIBUTION: Peru: Ancash, in the middle and northern Cordillera Bianca. H. N. P.: Alpamayo-Cashapampa trail between Huiscash and Mirador (10,000), Cahuish (11085), Carhuazcancha (12338, 12405), Honda (11662), Ishinca (11245, 11252), Llaca (11148), Llanganuco (8240), Parôn (11468), Rajucolta (12171), Ranincuray (10440), Ruruchinchay (12701), Shallap (10776) and Ulta (11409). PHENOLOGY: Flowering has been observed from March until October, and mature fruit observed in October. CONSERVATION STATUS: The species is seen frequently. It is not endangered. 135

Bommrem dulcls (Hook.) Beauverd HERB, erect, rarely clambering; 12-55 (-100) cm tall; from slender creeping rhizome with spherical tuber to 1 cm diam.; stem stiff, 0.8-3 mm diam., puberulent to glabrous, densely leafy, apex nodding and pendent. LEAVES: 75 mm long or less, 3-10.5 mm wide, linear to narrowly elliptic, stiff, arcuate, strongly nerved above, erect, appressed to stem, not resupinate, sessile, margins revolute, white pubescent below, nerves pilose with multicellular, flattened hairs, internerves with shorter hairs of same nature, apex acute. INFLORESCENCE: compound; (1) 2-5- flowered; involucral bracts 3-7, foliaceous, mostly shorter than leaves, narrowly elliptic, spreading to reflexed in live material; branches 1-2-forked, rarely undivided, peduncle to 19 mm long, terminated by floral bract, spreading laterally in fresh material; floral bracts foliaceous, smaller than involucral bracts, persistent. FLOWERS: pendent; pedicels 8-23 mm long, purplish; perianth segments connivent campanulate; sepals 14-25 mm long, 6.5-12 mm wide, oblanceolate to obovate, carnose, margins slightly revolute, apex cuculate, dark pink with a purple apex; petals 18-30 mm long, 10-19.5 mm wide, spatulate, thinner than sepal, base with adaxial glandular area, ca. 5 mm long, ca. 2 mm wide, papillose, apex acute, yellow, Costa dark pink, at least abaxially, apex green, or green terminated by a purple band, or a band of violet dots, purple dotted within; stamens subequal to perianth at anthesis; filaments 16-23 mm long, ca. 1 mm wide basally, anthers 4-8 mm long, ca. 2 mm wide, base cordate, apex acute-apiculate, purple; ovary half inferior, style 8-18 mm long, filiform, erect, stigmas papillate. CAPSULE: 10-15 mm diam., slightly 3-lobed, upper part papillate; seeds 3-3.5 mm diam., angular-globose, sarcotesta fleshy, bright orange. HABITAT: Grasslands; often in areas of morainal soils, or bouldery areas and talus slopes; often in altitudinal zones with frequent needle ice formation. 4000-4700 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash, Apurimac, Arequipa, Cajamarca, Cuzco, Huancavelica, Junin, La Libertad, Lima, Pasco, and Puno. H. N. P.: Alpamayo (9716, 9769), Alpamayo-Cashapampa trail (9962, 10025), Cahuish (Ldpez et al. 7536), Carhuazcancha (12227), Honda at Vinoyapampa (11599), Ishinca (9511, 11215), Llaca (8982), Llanganuco (8262, 8814; Gentry et al. 37418), Los Cedros (9962), Parôn (Mostacero et al. 542), Quenua Ragra (10671), Rio Pachacoto (Stein et al. 2005), Rurichinchay at Pucaraju (12679) and Ulta (11368). 136

PHENOLOGY: Flowering from December to July, and fruiting from January to May. Mature fruit found in May. CONSERVATION STATUS: Common.

Bommrem Bhvllostachv» Masters ex Baker, vel sp. aff. HERB, erect, to 30-45 cm tall; stem puberulent to glabrous. LEAVES: (2.5-) 4-12.7 cm long, 0.9-1.5 cm wide, narrowly elliptic, mammilose and lustrous above, flat, stiff, erect to slightly spreading, tending to be resupinate, margins (sub-)revolute, nerves prominent on lower face, white hirtellous to pilose, apex acute. INFLORESCENCE: compound, erect; involucral bracts foliaceous; few-flowered; 2-several branches of unequal length 2.5-4 (-10) cm long, once or twice forked, with a foliaceous bract at each node, terminated by floral bract, purplish green; floral bracts to 2-2.6 cm long, to 9-14 mm wide, foliaceous, elliptic, persistent. FLOWERS: pedicels 6-28 mm long; perianth segments subequal, connivent-tubular; sepals 19-22 mm long, ca. 6.5 mm wide, oblong, thickened, pink, apex green terminating in purple band; petals 19-23 mm long, ca. 6.5 mm wide, spatulate, base widened, margins with papillose nectaries, lower half pink, upper half yellowish green with dark violet dots within; stamens not exserted, filaments ca. 11 mm long, slightly unequal, papillate, anthers ca. 4 mm long; ovary half-inferior, obconic, apical half papillose, style about as long as perianth at maturity. CAPSULE: subspherical, mature material not seen. HABITAT: Grassland-shrubland in the vegetation types influenced by the macrothermal coastal climate; steep slopes. 3800-4320 m. DISTRIBUTION: Southern Ecuador to Peru. Peru: Ancash, Cajamarca, and Huanuco. H. N. P.: Auquispuquio (11922) and Llanganuco (8240). Baker (1888) reported the type to be from the Colombian Andes. The Lobb collections, including the type of this species, were reportedly collected in Colombia; they are actually from southern Ecuador and Peru (Killip, 1936). PHENOLOGY: Flowering in April. CONSERVATION STATUS: Rare, as there is limited appropriate habitat within the Park.

Bommrem zosteraefoll» Killip HERB, erect to clamboring, 20-76 cm tall; rootstock not seen; stem stiff, leafy portion ca. 1.5 mm diam., glabrous, densely leafy, apex nodding and pendent. LEAVES: 40-69 137 mm long, 4-5 mm wide, linear, stiff, strongly nerved above, often markedly tesselate, spreading to erect, not resupinate, sessile, margins revolute, blades glabrous to pubescent below, apex acute. INFLORESCENCE: compound; (2-) 13-14-flowered; involucral bracts 3-7, foliaceous, mostly shorter than leaves, narrowly elliptic, spreading to reflexed in live material; branches 25-30 mm long, 1-branched, rarely unbranched, peduncle terminated by floral bract, to 35 mm long, spreading laterally in fresh material; floral bracts foliaceous, smaller than involucral bracts, persistent. FLOWERS: pendant; pedicels 6-20 mm long, glabrous; perianth segments carnose, connivent, cylindrical; sepals 22-25 mm long, 5.5-7 mm wide, oblong, margins revolute, apex cuculate, red with purple apex terminated with green; petals 24-28 mm long, 7-8.5 mm wide, spatulate, claw about 2/3 of petal length, base with adaxial glandular area, papillose, apex acute, red with a green apex terminated by a band of violet dots, violet dots within; stamens subequal in length to perianth at anthesis, at most slightly exserted; filaments 21-25 mm long, anthers 2.7-7 mm long, 2-3 mm wide, base cordate, apex bi- apiculate; ovary half inferior, apical half long papillose, style ca. 20 mm long, filiform, erect, stigmas papillate. CAPSULE: not seen. HABITAT: Grassland and dry shrubland. 3400-4250 m. DISTRIBUTION: Peru; Ancash. H. N. P.: Llanganuco (Sagàstegui et al. 12310), Parôn (8920A) and Rio Pachacoto (8777). PHENOLOGY: Flowering has been observed in December, January and July. Although it is being treated as a distinct species in this flora, this taxon may be little more than an evolutionary tendency within Bomarea dulcis. In comparison with that species, the leaves of B. zosteraefolia are thinner and less pubescent, although pubescence is quite variable, the involucral bracts are little differentiated from the upper leaves, the inflorescence more open, the flowers cylindrical in bud, the sepals and petals have a slightly different color pattern, and the species is found at lower elevations, mostly less than 4200 m (R. E. Gereau, pars. com.). There are several probable hybrids, or intermediates if the species are not distinct: Parôn (11471 A, 11492) and Ranincuray (9123). The relationship of this species and B. dulcis merits further study. CONSERVATION STATUS: Rare. 138

AMARYLLIDACEAE Jaume St. Hilaire LILIIFLORAE: LILIALES HERBS, perennial from bulbs; stem short; plants glabrous. LEAVES: simple, ligulate, somewhat carnose, entire, sessile or petiolate. INFLORESCENCE: scapose, umbel-like or solitary; subtended by scale-like involucral bracts, free or connate; scape solid or hollow. FLOWERS: perfect, regular to slightly zygomorphic, epigynous; pedicels short to elongate; perianth members 6, free or somewhat connate, similar in size and shape, often differently colored, spreading or tubular; a corona develops in some species; stamen (3-) 6, inserted at the base of the perianth or epipetalous, filaments free or basally fused into a tube, anthers versatile; gynoecium 3-carpelIate, compound, inferior, trilocular ovary, placentation axile, septal nectaries, style elongate, stigma punctiform to trifid, dry, papillate, ovules many per locule. FRUIT: a loculicidal capsule; seeds sometimes winged. The Amaryllis family is native to tropical and sub-tropical regions, with centers of diversity in South Africa, the Mediterranean, and the Andes. There are ca. 55 genera and ca. 700 species, 10 of which are Peruvian. HABITAT: mostly in arid areas, especially the coastal lomas formations. POLLINATION: Hummingbirds. USES: Several species are used as ornamentals. TAXONOMY: Cronquist (1981) includes this family in the Liliaceae. CHEMICAL COMPOUNDS: All amaryllids contain alkaloidal compounds, and are used in folk medicine as poltices (A. Meerow, pers. comm.). I thank Dr. Alan Meerow (FLAS) for his advice and revision of the manuscript.

Stenomesson Herbert HERBS, from bulbs. LEAVES: linear-lorate to narrowly elliptic-lanceolate, some petiolate, somewhat carnose, developing during, or following flowering. INFLORESCENCE: scapose, sometimes umbel-like, I-several-flowered, subtended by a pair of membranous, deciduous bracts. FLOWERS: perfect, regular, epigynous; erect; subsessile to pedicellate; perianth united, funnelform to salverform, 6-lobed, red, orange or yellow; stamens 6, inserted at base of perianth lobes, filaments basally united into a membranous staminal cup, toothed between the pairs of filaments, free portion of filaments may arise from the apex or below the rim of the staminal cup, stamens 139 exserted; ovary: style terminal, filiform, bilobed, stigmatic surfaces pappillate. FRUIT: a loculicidal capsule; seeds flattened lenticular. The genus has ca. 40 species, distributed in the coastal and Andean regions of Ecuador, Peru, Bolivia, and Chile. The majority (ca. 85%) of the species are found on the western slopes of the Peruvian Andes, at elevations above 2000 m (A. Meerow, pers. comm.). Species of Stenomesson are found on the lomas formations of the coast, and in the arid regions of the Andes, especially those with a pronounced dry season. There are 30-35 species in Peru. COLLECTING TECHNIQUES: There is little information of taxonomic value in the bulb. As the species are rare and endangered, it is preferable to cut off the leaves and inflorescence at the ground line, leaving the bulb in the soil to regenerate. Considering the rareness of the majority of the species, bulbs should be collected only for study by specialists, and then few should be taken. The fruit characters are relatively similar between species; in this genus flowers are mandatory for a positive determination. Notes should be made regarding: presence or absence of a glaucous bloom on the leaves; and colors of the perianth segments (body and apex), of the staminal cup, and of the pollen. One or more of the flowers should be split open and carefully pressed. If there are sufficient, some flowers should be preserved in liquid. Photographs are desirable.

Stenomesson luteum (Herbert) Baker in Saund. HERB, to 40 cm tall; bulb deeply buried, ca. 5 cm long, ca. 3.5 cm wide, outer tunics dry, chartaceous, dark brown. LEAVES: exposed portion 22-40 cm long, 0.9-1.9 cm wide, broadest above middle, linear to linear-lorate, margins entire, apex rounded, fleshy when fresh, primary nerves prominent in dried material. INFLORESCENCE: 2- flowered scape; scape solid, exposed portion 19-27 cm long, 6-8 mm diam., narrowing to 3-4 mm diameter below inflorescence bracts; bracts 2, 16-18 mm long, ca. 4-5 mm wide at base, lanceolate, membranous, soon deciduous, falling by early fruit stage. FLOWERS: erect; pedicels unequal, lower subsessile-8 mm long, upper 11-18 mm long, both 2-3 mm diam.; perianth constricted basally, salverform, ca. 5 cm long, reported to be yellow, floral tube 3.3-3.4 cm long; lobes 1.4-1.5 cm long, obovate, erect; stamens inserted at perianth throat, exserted 7-8 mm beyond the lobes, basally united into a staminal cup, 12-13 mm long (from base of cup to teeth apices), thin, membranous, terminating in 1, narrowly triangular tooth between each pair of filaments, free portion of filaments 9-11 mm long; anthers 2.8-3 mm long, 0.8-1.5 mm wide, narrowly elliptic; 140 ovary 5-6 mm long, 3-3.5 mm wide, elliptic, style ca. 6.1 cm long, exserted beyond stamens, filiform, apically bilobed, inner surfaces stigmatic, pappilose, placentation axile, ovule: appear to be flattened and vertically stacked like coins. FRUIT: a loculicidal capsule, (in immature specimens) 2 cm long, 1.7-1,9 cm wide, erect, green; seeds (approximating from outlines on immature capsule) ca. 7 mm diam., ca. 1.3 mm thick, flat, obliquely winged. N.B.: the floral details in this description based on observations and measurements taken from H. P. Traub 584 at MO. HABITAT: On steep slopes with many rock outcrops, sometimes growing in cracks in cliffs, the sites in grassland or grassland/ shrubland of the more arid vegetation in the northwest corner of the Park. 3800-ca. 4300 m. DISTRIBUTION: Peru: Ancash. H. N. P.: Auquispuquio (11963) and los Cedros (9911). PHENOLOGY: Flowering dates not known; fruiting March and April. CONSERVATION STATUS: Although it is locally common near the Auquispuquio ruins, it is very rare as there is only a small extension of appropriate habitat in the Park. 141

ANTHERICACEAE J. D. Agardh LILIIFLORAE: ASPARAGALES HERBS, perennial, erect; from rhizomes, small corm-like structures, or roots fibrous, often becoming tuberous thickened. LEAVES: basal, simple, sessile, basally sheathing, linear, entire to ciliate-denticulate. INFLORESCENCES: racemose; terminal, scapose, bracteate, simple or compound. FLOWERS: per feet, regular, hypogynous; each subtended by a delicate floral bract; perianth segments 6, free to connate, equal; stamens 6, filaments free, anthers free or connate, versatile; gynoecium 3-carpellate, compound, superior, 3-locular ovary, placentation axile, nectaries septal or wanting, style terminal, 3-lobed, stigma dry, ovules several-many per locule. FRUIT: a loculicidal capsule; seeds black due to presence of phytomelanin. The Anthericaceae is of subcosmopolitan distribution with ca. 35 genera and ca. 700 species. In Peru, the^family is represented by Diamena, Diora, and Echeandia. Anthericum and Chlorophytum are both cultivated as house and garden plants. TAXONOMY: Cronquist (1981) includes this family in the Liliaceae. The family is in need of a complete revision. Ravenna (1987) recently added the genera Diamena and Diora, both endemic to the northern coast of Peru. Superficially the inflorescence is a raceme, however there are 1-several flowers at the nodes, each flower with a subtending bract. This suggests that the inflorescence is derived from a more complex, possibly paniculate, inflorescence type. I thank Dr. R. W. Cruden (lA) for discussion of the family and review of the manuscript. Echeandia Gômez Ortega Reference: Weatherby, 1910. HERBS, roots arising from a small corm, roots usually thickened, entirely or near the tip. LEAVES: with many nerves, prominent in dried material. FLOWERS: pedicellate, pedicels jointed; perianth segments free, nerves 3, proximate and subparallel, central, spreading, white to yellow; stamen filaments smooth or retrorsely scaly, inserted into a pit or depression at anther base, anthers free or connate; ovary apex 3-lobed, nectaries wanting, style filiform. FRUIT: a loculicidal capsule; seeds irregularly compressed. A New World genus, mostly tropical with slight expansion into temperate regions, Echeandia is found from the southwestern United States to Bolivia. The genus has 90- 100 species, 2-4 of which are Peruvian. POLLINATION: Typically by bees. 142

TAXONOMY: Most of the New World species of Echeandia with free anthers have been placed in Anthericum at one time. Anthericum is an Old World genus with free anthers and smooth filaments; Echeandia is a New World genus with free or connate anthers and smooth or retrorsely scaled filaments, and roots arising from a small corm (Cruden, 1981). In recent discussions of the genera in this family. Marais and Reilly (1978) and Ravenna (1987) limit Echeandia to the species with connate anthers.

Echeandia eccremorrhlia (Ruiz Lôpez & Pavôn) Cruden, comb. ined. HERB, from fibrous roots becoming tuberous at depth. LEAVES (4-) 8-15, basal, 24- 55 cm long, 0.8-1.5 cm wide, lorate, 16-26 prominent nerves, base brown, weathering to a fibrous, persistent indûment, margins entire, or sometimes with trichomes and appearing sub-serrulate, apex long attenuate. INFLORESCENCE: racemose, often compound, producing secondary axes at lower nodes; peduncle 30-88 cm long,.p^(ijincle. bracts 6-19 cm long, membranous, subulate; principal axis ca. 27 cm long, 15-22 (-31) nodes, 1-3 flowers per node; secondary axes 7-20 cm long; inflorescence bracts 4-10 mm long, up to 30 mm long at nodes with secondary axes, membranous with prominent nerves, ovate- attenuate. FLOWERS: pedicels with a swollen joint, lower portion 6-9 mm long, upper 2.5-7 mm long; perianth segments 15-18 mm long, ca. 5 mm wide, elliptic, white; stamen filaments ca. 7 mm long, smooth, inserted in pit at base of anther, white, anthers free, 4 mm long, deeply notched at base, yellow; style terminal, filiform, 7 mm long, persistent in fruit. FRUIT: a loculicidal capsule, 7-9.5 mm long, 3.5-5 mm wide, apex 3-lobed; seeds jet black, surface colliculose. Cruden (pers. com.) has found that the inner perianth segments are usually twice as long as wide; the outer segments are narrower than the inner. HABITAT: Dry areas at middle elevations, on the western slopes of the Andes, often in seasonal vegetation. 3500-3900 m. DISTRIBUTION: Colombia to Bolivia. Peru: Ancash, Arequipa, Cajamarca, Cuzco, Junin, and Lima. H. N. P.: Auquispuquio (12024). PHENOLOGY: Flowering is seen from January to April, and fruiting from May until April. Bernhardt and Montalvo (1977) found that the flowers of E. macrocarpa (-£. formosa), a Central American species, lasted up to a week. CONSERVATION STATUS: Rare. There is a very limited extension of appropriate habitat within the Park. 143

BROMELIACEAE A. L. de Jussieu ZINGIBERIDAE: BROMELIALES REFERENCES: Gilmartin, 1972; Smith and Downs, 1974, 1977. HERBS, mostly short-stemmed epiphytes, some terrestrial, of those some develop a stout stem. LEAVES: alternate, rosulate on short stem, or at stem apex, narrow, simple, spinose serrate or entire, coriaceous or carnose, pubescent with stalked, peltate scales, leaf bases often connivent, forming a reservoir. INFLORESCENCES: simple or compound, bracteate spikes, racemes, heads or sometimes solitary. FLOWERS: perfect or sometimes functionally unisexual, regular or slightly zygomorphic, hypogynous to epigynous; sepals 3, coriaceous to membranous, free or connate below; petals 3, free or connate below, some with scale-like, basal appendages; stamens 6, in 2 whorls of 3, free or connate or adnate to the perianth; gynoeclum 3-carpellate, compound, trilocular, superior or inferior (Bromelioideae) ovary, style terminal, often trifid, stigmas papillate, wet or seldom dry, placentation axile, ovules few to numerous. FRUIT: a berry, capsule, or rarely a fleshy multiple (as in Ananas); seeds in capsular fruits winged or plumose. The Bromeliaceae is Neotropical, with one disjunct species, Pitcairnia feliciana (A. Chev.) Harms & Mildbr. in western Africa. Some species extend into the southern temperate region of central Argentina, and Tillandsia usneoides L. reaches Maryland in the United States. There are three centers of development: Mexico and the West Indies, the Guiana Highlands, and eastern Brazil. The family is relatively rare in the Amazon Basin. The family has ca. 45 genera and 2000 species, of which about 14 genera and 175 species are native to Peru. HABITAT: Bromeliads are adapted to a variety of high stress environments. The Pitcairnioids are terrestrial xerophytes or lithophytes found in the arid areas of the western slopes of the Andes, the high elevation grasslands and into the montane rain forests. The Tillandsioids form two groups: the lithophytes which are densely pubescent and found on the sands of the coastal desert and in lithophytic situations throughout the country, and the epiphytes which are sparsely pubescent and form water reservoirs, and are found in brushfields and forests from the Amazon Basin to the Sierra. The Bromelioids are terrestrial or epiphytic, and are found mostly in the Amazon Basin. POLLINATION: Predominantly by animals: bees (Bombids, Euglossines and others), moths, butterflies, birds (Icterids, hummingbirds and others) and bats; rarely by wind; rarely cleistogamous. In some genera, the petals bear appendages at the base of the 144

interior face, it is thought that these help the nectar to draw up the petal to the height of the appendage. The higher position of the nectar makes it available to pollinators other than those strictly adapted to corolla length (McWilliams, 1974). BREEDING SYSTEM: Most observations are of a general nature, few detailed studies have been made. Outcrossing and inbreeding are documented, as well as self-fertility and self-sterility (McWilliams, 1974). USES: Many species are cultivated for ornament, and Ananas comosus (L.) Merr., the pineapple, for its edible fruit. In the Philippines, fiber extracted from the leaves is woven into a fine-textured cloth. PHYSIOLOGY AND ADAPTATIONS: Bromeliads are adapted to high stress environments, especially regarding water and mineral deficiencies, and temperature regime. In the Pitcairnioideae and Tillandsioideae, the leaves are frequently covered with specialized peltate trichomes, which facilitate rapid water absorption, reduce transpirational losses, insulate, and scatter light (McWilliams, 1974). Many species have CAM photosynthesis, which allows the plant increased control of water balance and tolerance of temperature extremes. Some xerophytes, especially Puya spp., do not have CAM photosynthesis (McWilliams, 1970). In species which lack CAM photosynthesis, the seasonal loss of leaf blades through a deciduous habit helps control water loss (McWilliams, 1974). The formation of reservoirs or tanks by the connivent leaf bases aids in maintaining water balance. In the Pitcairnioideae and Tillandsioideae, the epiphytic species are adapted to wind dispersal by unusually buoyant seeds (McWilliams, 1974). In the Bromelioideae, the seeds, including those of the epiphytic species, are animal dispersed. Birds, ants, and bats are important dissemination vectors (McWilliams, 1974). Benzing (1980) has written a very detailed book on the Bromeliads, treating their taxonomy, anatomy, physiology, reproductive biology, and ecology. The water tanks formed by many epiphytic species are important habitats and breeding sites for many organisms, and are ecologically quite complex. Bacteria, algae, bryophytes, and Utricularia spp. inhabit the tanks, as well as many forms of insects, crustaceans, and frogs (McWilliams, 1974). TAXONOMY: The family is subdivided into three sub-families: Pitcairnioideae, Tillandsioideae, and Bromelioideae. The Pitcairnioideae is the least specialized, especially the terrestrial xerophytes. 145

Despite the work of numerous workers, especially Dr. Lyman B. Smith, the taxonomy of the family is not well understood, and further systematic studies are needed: population dynamics, breeding systems, pollination ecology, gene flow, and ecology in the field, and in the laboratory and greenhouse, crossing and compatibility, chromosomal studies, and more phenetics. In all cases, more high quality specimens are needed. COLLECTING PRACTICES: As they are large plants, it is difficult to make useful Bromeliad specimens. Several things are important a complete and accurate description, including measurements, of the habit, and all parts not included in the specimen; a photograph; several fully grown leaves, folded to show both faces of the leaf and the shapes of the outline, base, and apex; peduncle and inflorescence bracts pressed to show their shapes; and extra flowers and fruits for dissection, preferably preserved in liquid, as important characters of the stigma (Brown and Gilmartin^ 1984), sepals, petals, stamen filaments, and anthers (Gardner, 1986) are not adequately preserved in pressed and dried material. I thank Dr. Harry Luther (SEL) for confirmation of determinations, advice regarding the treatment, and revision of the manuscript.

Key to the genera of the Bromeiiaceae la. Stout caulirosettes; leaf margins spinose dentate; terrestrial or lithophytic. Puya lb. Herbs, stems very short or acaulescent; leaf margins entire; lithophytes or epiphytes. 2. 2a. Herbs from densely scaly rhizomes; leaves dimorphic, lower scale-like, upper foliar and deciduous; flowers large, showy, scarlet. Pitcairnia pungens 2b. Herbs without rhizomes; leaves all alike; flowers purple or coral, usually inconspicuous. Tillandsia

Pltcmlrnlm L'Heritier HERBS from scaly rhizomes; stem short, thick and bulbous, or elongated, often stellate pubescent. LEAVES: often dimorphic, lower leaves scale-like with a broadened, membranous base, blade spinose, upper leaves with a broadened, scale-like, membranous base and a flat, chlorophyllous blade. INFLORESCENCE: terminal, simple or compound, pedunculate racemes and panicles, bracteate. FLOWERS: perfect, regular to slightly zygomorphic, hypogynous; pedicellate, subtended by a bract; calyx coriaceous. 146

petaloid, sometimes winged or keeled; corolla showy, sometimes with basal, adaxial appendages, typically red, rarely white, yellow, violet or blue; stamen filaments free, linear, anther versatile; ovary superior, style filiform, trifid, stigmas papillose; ovules many per locule. FRUIT: an elongate capsule; seeds many, appendaged or unappendaged. This genus of ca. 260 species is distributed from Mexico and the Caribbean to northwest Argentina, northern Paraguay and Uruguay. The genus is common in the Andes, and uncommon in the Amazon Basin. Often terrestrial, lithophytic or epiphytic; common in arid habitats. About 40 species are known from Peru.

Pltcalrnta mumeenm Kunth var. pungens HERB terrestrial, from a branching rhizome with many adventitious roots, scales widely ovate, chartaceous, tightly imbricate, basally dark brown, apically many-nerved, pale brown, mucronate to spine tipped, the longer spine-tips with marginal spines; pubescence of floccose, stalked stellate hairs, white; clones 10-15 cm diam.; stem very condensed, appears acaulescent, base bulbous. LEAVES: in fascicles along rhizome, dimorphic, the lower: scale-like, 1.4-2.5 cm wide, blade spine-like with marginal recurved spines; the upper bases scale-like, 2-5 cm long, 1.3-3.5 cm wide, chartaceous, basally dark brown, distally light brown, many-nerved, apex often curling back on itself, margins with retrorse spines, blades deciduous, the line of abscission well-defined, horizontal, 23-54 cm long, 9-17 mm wide, linear, broadening slightly in distal 1/2, acuminate, margin entire. INFLORESCENCE: a raceme; peduncles 35-39 cm long, bracteate, lower bracts to 9 cm long, foliar, acuminate; axis purple: raceme 15-22 cm long, 12-16-flowered, bracts to 22 mm, acuminate, erect, rachis purple. FLOWERS: slightly zygomorphic; pedicels 4-9 mm long; sepals 23-44 mm long, 4.2-4.8 mm wide, narrowly triangular, keeled, chartaceous, nerves prominent; petals 43-61 mm long, 5.5-6 mm wide, slightly unequal, oblanceolate, within base appendaged with 2 scales, 6 mm long, adnate to petal with scale margin free, arcuate, and joining at the apices; scarlet; stamens included, filaments ca. 40 mm long, anthers ca. 8.2 mm long; ovary superior, style ca. 44 mm long, branches ca. 2.8 mm long. CAPSULE: not seen. HABITAT: Soil pockets in lithophytic situations: cliffs, rocky banks, boulders, and stable talus slopes. 3500-3860 m. 147

DISTRIBUTION: Colombia to Peru. Peru: Ancash, Cajamarca, Cuzco, Huancavelica, Huanuco, La Libertad, Lima, Piura, and Tumbes. H. N. P.: Llanganuco (9405) and Ranincuray (9042, 10474). PHENOLOGY: Flowering from January to April. CONSERVATION STATUS: Although it has a wide distribution, its habitat within the Park is not extensive. It should be considered rare.

Puva Molina CAULIROSETTES, terrestrial to lithophytic; stem short to tall. LEAVES: densely packed at apex of stem; bases broadened and inflated, sheathing, coriaceous, margins entire to slightly spinose distally; blades narrow, elongate, margins serrate with dark spines, one or both faces densely pubescent with peltate scales. INFLORESCENCE: central, terminal, usually erect; simple or compound, pedunculate, bracteate spikes or racemes; peduncles bracteate, usually elongate. FLOWERS: perfect, actinomorphic to zygomorphic, hypogynous; sessile or pedicellate, bracteate; sepals free, equal, membranous, shorter than the corolla, stellate pubescent; corolla showy, petals free, often twisting together after anthesis; stamens free, exserted or included, filaments terete or linear, anthers versatile; ovary superior, 3-angled, style elongate, apex 3-fid, branches spirally twisted, stigmas linear, papillose, ovules many per locule. FRUIT: a septicidal capsule, inflated, triangular; seeds large, lenticular, winged. A predominantly Andean genus of ca. 170 species, Puya is distributed from Venezuela and Colombia to northern Argentina and Chile, with outlying species in the Guiana Highlands and Costa Rica. There are about 60 species in Peru. The terrestrial species have well developed root systems. Many of the species of Puya are localized endemics with very small distributional ranges (H. Luther, pers. com.). This genus needs taxonomic revision based on detailed field studies, especially regarding gene flow and population dynamics. There are several groups of species which, given field studies, may prove to be but one species. More well-collected specimens are needed.

Key to the species of Puva la. Plants solitary, erect; flowers greenish white or pale blue. 2. lb. Plants clonal, decumbent to erect; flowers deep green or greenish white. 5. 148

2a. Plants very large, to 14 m tall; leaves carnose, to 1.25 m long; stem well developed, to 85 cm diam.; inflorescence to several meters long, a compound raceme. P. raimondii 2b. Plants less than 3 m tall; leaves coriaceous, to 6 dm long; stem short, less than 10 cm diam.; inflorescences to 5 dm long, simple racemes. 3. 3a. Floral bracts lanceolate, tips very long attenuate, glabrous, conspicuous; marginal spines antrorse. P, sp. nov. 3b. Floral bracts long attenuate, apices densely woolly, inconspicuous; marginal spines retrorse. 4. 4a. Petals oblanceolate, pale blue; plant solitary. P. aff. membranacea 4b. Petals obovate-acute, whitish green; plant solitary or forming small clones. P. aff. herrerae 5a. Inflorescence open, individual flowers and rachis visible; flower bracts glabrous apically, purple; flowers dark emerald green. 6. 5b. Inflorescence densely flowered, individual flowers and rachis not distinguishable; flower bracts densely woolly, tawny; flowers whitish green. P. aff. herrerae 6a. Raceme short and broad (15-20 cm long, 8.5-14 cm wide). 7. 6b. Raceme elongate and narrow (to 50 cm long, to 10 cm wide); flowers erect to spreading. P. angusta 7a. Flowers 40-44 mm long, erect to spreading. P. aff. rauhii 7b. Flowers 55-72 mm long, broadly spreding to reflexed. P. reflexiflora

Puvm angusta Lyman B. Smith CAULIROSETTE lithophytic to terrestrial, 1.5-2.2 m tall; trunk to 50 cm tall; forming clones to several meters diam.. LEAVES: rosette 50-75 cm tall; bases ca. 5 cm wide, ovate; blades 31-61 cm long, 2.5-3.5 cm wide at juncture with base, linear-triangular, spreading to arcuate, marginal spines uncinate, retrorse to antrorse on same leaf, both faces of blade pubescent, the adaxial glabrate in age. INFLORESCENCE: a spike-like raceme; peduncle 25-150 cm tall, 1.5-2 cm diam., bracts 3.2-6 cm long, 1.7-2.8 cm wide, ovate-acuminate, membranous, margins occasionally irregularly membranous toothed, glabrate abaxially, stellate pubescent adaxially, purple; raceme 31-57.5 cm long, 8-12 cm diam., rachis densely stellate pubescent, flowers erect to spreading. FLOWERS: pedicels ca. 10 mm long; bracts 33-34 mm long, 8.2-11.6 mm wide, lanceolate- acuminate, membranous, stellate pubescent on both faces, densely so abaxially, purple; 149

sepals 21-30 mm long, 8-12 mm wide, elliptic-acute, mucronate, strongly nerved, margins entire, stellate pubescent, strongly so abaxially; petals (36-) 49-51 mm long, 8- 12.5 mm wide, spatulate-acute, strongly nerved, connivent into a tube at anthesis, emerald green; stamens exserted, filaments 30-43 mm long, round distally, drying flat, anthers 10-13 mm long, 1.2-1.4 mm wide, yellow; ovary 8-14 mm long, 3-4 mm diam., style 27.5-36 mm long, branches 2.5-3 mm long. CAPSULE: not seen. HABITAT: Slopes and ledges in arid grasslands, or grassland-shrubland; soils of granitic morainal parent material. 4000-4500 m. DISTRIBUTION: Peru: Ancash and Cajamarca. H. N. P.: Llanganuco (8213), Los Cedros (9878), Parôn (8920), and Ulta (11396). PHENOLOGY: Flowering from January to July. CONSERVATION STATUS: Rare.

Puva aff. herrerae Harms CAULIROSETTE, lithophytic, 95-100 cm tall; forms small clones. LEAVES: rosette 33-40 cm tall; base 2.5-6 cm wide, 5.3-5.5 cm long, ovate, brown; blades 26-28 cm long, 1.5-2.5 cm wide at juncture with base, very narrowly triangular, long attenuate, stiffly arcuate, pubescent on lower face, marginal spines uncinate, retrorse. INFLORESCENCE: a dense, spike-like raceme; peduncle 40-75 cm tall, completely covered by bracts ca. 12 cm long, ca. 5 cm wide, ovate-attenuate, margins entire, both faces densely woolly, tawny; raceme 16-25 cm long, ca. 9 cm diam., flowers densely packed, spreading. FLOWERS: pedicels ça. 4 mm long, stout, conical; bracts 48-62 mm long, 11-26 mm wide, ovate-attenuate to elliptic-attenuate, strongly nerved, membranous, margin entire, adaxial face sparsely pubescent, abaxial face woolly, tawny, longer than the flowers; sepals 35-45 mm long, 11-13.2 mm wide, lanceolate to elliptic-acute, strongly nerved, membranous, woolly abaxially; petals 39-45 mm long, 14-20 mm wide, obovate-acute, strongly nerved, spreading, whitish green; stamens included, filaments 26-34 mm long, flat, anthers 5.5-8.5 mm long, ca. 2 mm wide, mucronate, yellow; ovary 11-16 mm long, ca. 5 mm diam., style 22-31 mm long, branches 3-3.5 mm long. FRUIT: 2.3 cm long, 1.3 cm diam., triangular-inflated, deeply sulcate on sides, seeds many, 2 mm long, reticulate, brown, surrounded by a wing roughly triangular in outline, ca. 4 mm on a side. HABITAT: Growing at edge of cliff. 4200-4970 m. ISO

DISTRIBUTION: Ecuador to Peru. Peru: Ancash, Apurimac, Cajamarca, Cuzco. Pasco, and Puno. H. N. P.: Carhuazcancha (123IS) and Ishinca (11222). PHENOLOGY: Flowering in May and July, and fruiting in July.. CONSERVATION STATUS: Very rare.

Puva mff. membranicea Lyman B. Smith CAULIROSETTE, solitary, terrestrial, ca. 1.3 m tall. LEAVES: rosette SO cm tall; base S-6.5 cm wide, ovate; blade 42-47 cm long, 2.8-3.S cm wide at juncture with base, very narrowly triangular, long attenuate, stiffly arcuate, marginal spines uncinate, retrorse, pubescent on lower face. INFLORESCENCE: a spike-like raceme; peduncle ca. 100 cm tall, completely covered by bracts, 12 cm long, 3.S cm wide, narrowly trullate, margins entire, adaxial face densely pilose, abaxial tawny, woolly; raceme ca. 30 cm long, ca. lO.S cm diam., flowers densely packed, spreading. FLOWERS: pedicels ca. 9 mm long, stout; bracts similar in shape and indûment to peduncle bracts, longer than the flowers, reflexed, not conspicuous; sepals 47-SO mm long, 9-lO.S mm wide, lanceolate-long attenuate, strongly nerved, lanate; petals 4S-S2 mm long, 14-19 mm wide, oblanceolate, strongly nerved, spreading, pale blue; stamens exserted, filaments 36 mm long, flat, anthers 4-7.6 mm long, 0.9-1.S mm wide; ovary 1.8 mm long, style 2.6-2.8 mm long, branches 4 mm long. CAPSULE: 3.1 cm long, 2 cm diam., triangular-inflated, deeply sulcate on sides; seeds many, 2 mm long, reticulate, brown, surrounded by a wing roughly triangular in outline, ca. 4 mm on a side. HABITAT: Rocky spots in high altitude grassland. 4600-4970 m. DISTRIBUTION: Peru: Ancash, Cajamarca, and Cuzco. H. N. P.: Llaca (10813). PHENOLOGY: Observed flowering and fruiting from May to July. CONSERVATION STATUS: Rare.

Puva ralmondii Harms "Queshque" CAULIROSETTE, solitary, terrestrial, to 6 m (-14 m; Macedo-Ruiz, 1978) tall when flowering; trunk to 2 m tall, 40-S0 (-ca. 8S) cm diam.. LEAVES: crown spherical, 2-3 m diam.; bases 12-24.S cm wide; blades ca. I m long, 8-11.S cm wide at juncture with base, S-8 cm wide above, narrowly triangular, long attenuate, carnose-coriaceous, glabrous, stiffly spreading, tips arcuate, marginal spines retrorse. INFLORESCENCE: a compound raceme; columnar, 3-6 m tall, ca. 70 cm diam.; peduncle SO-100 cm long, covered with bracts, 16.S cm long, 11.3 cm wide, broadly elliptic, apex caudate, 12 cm 151

long, secondary racemes 20.5 cm long, spreading, apices sterile, pedunculule 1.5 cm long, subtended by large bracts. FLOWERS: bracts 70-88 mm long, 23-55 mm wide, ovate- acuminate to elliptic attenuate, membranous, pubescent abaxially and at base adaxially; pedicels 4-15 mm long; sepals 33-41 mm long, 7-10.1 mm long, elliptic-acuminate, membranous, strongly nerved, woolly throughout; petals 48-49 mm long, 17-19 mm wide, elliptic-acuminate to sub-spatulate, strongly nerved, spreading, greenish white to cream-colored; stamens exserted at anthesis, filaments 22-29 mm long, flat, anthers 8-17 mm long, 2-2.5 mm wide, apex mucronate, orange yellow; ovary 11-19 mm long, style 31-35 mm long, branches 3-9 mm long. CAPSULE: ca. 2.5 cm long, 2 cm wide, dark brown. HABITAT: In tussock grassland; poor, often stony soils of sedimentary parent material. Macedo-Ruiz (1978) found this species growing on lithosols developed on sandstone. Ca. 4230 m. DISTRIBUTION: Peru and Bolivia. Peru: Ancash, Apurimac, Ayacucho, Cuzco, Huancavilica, Junin, Lima, and Puno. H. N. P.: Queshque and Rio Pachacoto (12751; J. D. Boeke 2848; A. Gentry et al. 37451). The type comes from Aija in Ancash. PHENOLOGY AND DEVELOPMENT: This species is monocarpic; it flowers after a long period of vegetative development, and dies after the fruit is developed. Macedo- Ruiz (1978) estimates that, in the native habitat, there is a period of vegetative development for 120 years before flowering. In the Botanical Garden at the University of California Berkeley Campus, a plant grown from seed flowered after 28 years (UCB Office of Public Information, Memo to the Media, file #10043). The flowering occurs from late October to January. The inflorescence needs 6 months for full development (Macedo-Ruiz, 1978). Long term studies of the breeding system of this species are needed. POLLINATION: Hummingbirds. USES: The inflorescence is very light and has great tensile strength (ex Boeke 2848), and can be used in rustic construction or furniture. Venero (1984) recounts that it can be used to make doors. Nearby the stands of P. raimondii, the local residents use sections of the trunk as stools. Sr. Rolando Gonzales of Huaraz mentioned that during the Spanish colonial occupation, saints were carved from the trunks, and then plastered and painted. They were very light and lasted well. Weberbauer (in Macedo-Ruiz, 1978) cited uses such as posts and columns, storage bins, and that the resin produced at the leaf bases was used for stiffening hats. The leaves can be fed to livestock after burning 152

off the spines, as can the pith when mixed with molasses (Macedo-Ruiz, 1978). Local residents eat the uncooked pith of the inflorescence, or ferment it into chicha (Macedo- Ruiz, 1978). The pith can also be used in coca chewing. The pith is burned, and the ashes mixed with sugar and powered anise (Venero and Macedo-Ruiz, 1983); the product, "toqra," substitutes for quicklime. Many species of birds make their nests in the crown of P. raimondii (Dorst, 1957; Macedo-Ruiz, 1978; Rees and Roe, 1980; Venero, 1984; Venero and Macedo-Ruiz, 1983). In Pampas Galeras, heavy flowering of this species is considered to be a harbinger of good harvests (Venero and Macedo-Ruiz, 1983). CONSERVATION STATUS: Rare. Venero and Macedo-Ruiz (1983) list 27 localities for the species in Peru. Given its localized habitat, Puya raimondii needs protection. Sheep occasionally become entangled in the lower leaves and, if not found and freed, can starve. As a result, shepherds view the plant as a nuisance, and often burn the plants. Fortunately the species has good fire resistance. Few of the larger plants are free of burn scars.

Puva mff rauhli Lyman B. Smith CAULIROSETTE, lithophytic to terrestrial, I m tall; trunk to 50 cm tall; forming clones to several meters diam. LEAVES: rosette 60 cm tall; bases ovate; blades 47-60.5 cm long, 2-2.2 cm wide at juncture with base, linear-triangular, spreading to arcuate, marginal spines uncinate, antrorse, lower faces of blade pubescent with scale trichomes. INFLORESCENCE: a spike-like raceme; peduncle ca. 25 cm tall, ca. 1.3 cm diam., bracts ca. 4.5 cm long, ca. 2.2 cm wide, ovate-acuminate, membranous, margins irregularly toothed, stellate pubescent on both faces, glabrate in age, dark; raceme ca. 15 cm long, rachis densely stellate pubescent, flowers erect to spreading. FLOWERS: 40-44 mm long; pedicels 4-6 mm long; floral bracts 30-33 mm long, ca. 12 mm wide, lanceolate-acuminate, membranous, stellate pubescent on both faces, purple; sepals 22-24 mm long, 9-9.5 mm wide, elliptic-acute, mucronate, strongly nerved, margins entire, stellate pubescent abaxially; petals 36-39 mm long, 8-8.4 mm wide, spatulate-acute, strongly nerved, connivent into a tube at anthesis, emerald green; stamens shorter than petals, exposed by spreading petal tips, filaments ca. 29 mm long, drying flat, anthers 6- 7 mm long, 1.2-1.4 mm wide, yellow; ovary 8-14 mm long, 3-4 mm diam., style 24-25 mm long, branches 7-8mm long. CAPSULE: not seen. HABITAT: Dry ledge in grassland-shrubland. 3900-4070 m. 153

DISTRIBUTION: Peru: Ancash. H. N. P.: Llanganuco (Rauh and Hirsch P2022, type specimen) and Rurichinchay (12724A). PHENOLOGY: Flowering in June. CONSERVATION STATUS: Very rare.

Puva reflexifiora Mez CAULIROSETTE, terrestrial, 1.9 m tall; branched trunk sheathed by marcescent leaf bases; clonal. LEAVES: bases ca. 5 cm wide, ovate; blades 40-44 cm long, 3-3.6 cm wide at junction with base, narrowlj^^iangjalar, stiffly arcuate, marginal spines uncinate, retrorse, lower face pubescent. INFLORESCENCE: a spike-like raceme; peduncle 1.1 m tall, 2 cm diam., reddish brown, bracts 3 cm long, 2 cm wide, ovate-acute, membranous, dark purple, sparcely stellate-floccose; raceme 20 cm long, 13.5 cm diam., cylindric, rachis densely stellate pubescent, flowers spreading. FLOWERS: sub-sessile, 5.5-7.2 cm long; bracts 3.5-3.6 cm long, 1.2-1.5 cm wide, elliptic-attenuate, membranous, margin entire, stellate pubescent on both faces, densely so on the abaxial, purple; sepals 32 mm long, 10-11 mm wide, lanceolate-acute, strongly nerved, stellate pubescent, densely so on abaxial face and at base of adaxial face; petals 50-55 mm long, 12.5-14 mm wide, spatulate-acute, strongly nerved, emerald green, connivent into tube at anthesis; stamens exserted, filaments 40-49 mm long, triangular, drying flat, anthers 12-13 mm long, 2-2.5 mm wide, yellow; ovary 12-13 mm long, 4 mm diam., style 36-38 mm long, branches 4 mm long. CAPSULE: not seen. HABITAT: Steep slopes in arid grassland; soils of granitic morainal parent material. 4000 m. DISTRIBUTION: Peru: Ancash. H. N. P.: Querococha (11049). The type of this species was collected at Hacienda Cochabamba, Ancash. PHENOLOGY: Flowering in early July. CONSERVATION STATUS: Rare.

Puvm SD. nov.. aff. P. pygmaea Lyman B, Smith and P. cryptantha Cuatrec. "Queshque" CAULIROSETTE solitary, terrestrial, 1 m tall. LEAVES: rosette 50 cm tall; base to 7 cm wide, ovate; blades 33-35 cm long, 3-3.5 cm wide at juncture with base, very narrow triangular, nearly linear apically, stiffly arcuate, marginal spines antrorse, ascending, pubescent on lower face. INFLORESCENCE: a spike; peduncle 26 cm long, 5 cm diam. at base, completely covered by bracts, 15 cm long, 3.2-3.5 cm wide. 154

lanceolate-long attenuate, apex linear with a few, irregular antrorse spines, membranous, both faces densely tawny lanate, apex glabrate; spike 24 cm long, 7 cm diam., many, densely packed, spreading flowers. FLOWERS: sessile; bract similar to peduncle bracts in shape and indûment, ca. 7 cm long, margins entire, tips reflexed, much exceeding the flowers; sepals 27-28 mm long, 3.7-4.2 mm wide, lanceolate, strongly nerved, woolly; petals 44-51 mm long, 6.2-8.2 mm wide, narrowly oblanceolate-acute, or occasionally truncate, spreading, pale blue; stamens exserted at anthesis, filaments 32 mm long, terete, anthers 32.-3.5 mm long, 1.1-1.2 mm wide; ovary 8 mm long, 2-3 mm diam., style 15.5 mm long. CAPSULE: not seen. HABITAT: By rivulet in arid grassland; soils of mor^inal parent material. Ca. 4400 m. DISTRIBUTION: Colombia to Peru. Peru: Ancash. H. N. P.: Huaripampa (9196). PHENOLOGY: Flowering in mid-January. CONSERVATION STATUS: Rare.

Tillandsia Linnaeus HERBS; epiphytic or lithophytic, rarely terrestrial; short-stemmed, sometimes caulescent; pubescent with peltate scales or glabrous. LEAVES: usually in a rosette, bases often widened and connivent forming a reservoir, margins entire. INFLORESCENCE: simple or compound, pedunculate racemes or spikes, bracteate. FLOWERS: perfect, actinomorphic or zygomorphic, hypogynous; subtended by bracts; calyx posterior sepals sometimes briefly united basally, keeled or not, usually shorter than the corolla; corolla free, petals of smaller corollas often adhere to each other appearing united; stamens free, filaments flat, slightly flattened, or terete in cross-section, anthers basally attached or versatile; ovary superior, 3-angIed, style short to elongate, apex trifid, stigmas linear, papillose, ovules many. FRUIT: a septicidal capsule; seeds elongate with a basal tuft of many, long, capillary hairs at base. Tillandsia has ca. 400 species distributed from the southern United States to central Chile and Argentina, with few species in the Amazon Basin. The genus is well represented in Peru, with about 120 species. The peltate trichome is an important adaptive feature in the Bromeliaceae, and reaches its greatest development in the Tillandsioideae. The trichome consists of a multicellular stalk and a multicellular wing. The structure of the trichome is such that it can facilitate water adsorption and prevent or slow water loss, in part due to structure and in part to an increase in the leafs boundary layer. It also facilitates absorption of ISS

mineral nutrients in solution, effects gas exchange, and modifies light absorption and heat load through increased leaf surface reflectivity, especially in the species of extremely xeric areas. For a complete discussion of the role of the trichomes see Benzing (1976). The nature and abundance of trichomes is correlated with the specific habitat of each species. Those inhabiting extremely xeric sites have abundant, well developed trichomes, and do not develop water reservoirs. The species of mesic sites have a much lower density of trichomes, the trichomes have smaller wings, and the plants tend to develop reservoirs. In the mesic sites, the properties of water and mineral absorption, light reflectance, and boundary layer effects associated a high density of trichomes are unnecessary or disadvantageous (Benzing et al., 1978). The xeric Tillandsioid species tend to have CAM photosynthesis, while the mesic species do not (Benzing and Renfrow, 1971). Currently, it is thought that the Tillandsioideae is of mesic origin, with the xeric species evolving specialized features in order to revert to xeric habitats. The xeric species have the largest geographic range and are at the periphery of the family distribution (Benzing and Renfrow, 1971). Taxonomically, Tillandsia is considered to be the least evolved genus in the Tillandsioideae (Smith and Downs, 1977).

Key to the species of Tillandsia la. Plants densely pubescent with peltate scales; leaves gray-colored; lithophytes; leaf bases not froming water reservoirs. 2. lb. Plants appearing glabrous; leaves green with purple markings or purple; lithophytes or epiphytes; leaf bases forming water reservoirs. 4. 2a. Scales with lateral wing; leaves appearing hirsute; stem prostrate, creeping; plants clonal. T, tectorum 2b. Scales symetrically peltate, lacking lateral wing; leaves appearing smooth; plants erect or creeping and clonal. 3. 3a. Plants erect, solitary or in small clones; leaves narrowly triangular; secondary racemes complanate. 4. 3b. Plants erect from prostrate, creeping stems, forming extensive clones; leaves ovate- long attenuate, apices filiform; secondary racemes terete. T. humilis 4a. Primary inflorescence bracts less than 1/3 the length of the secordary racemes; leaf apices filiform. T. oroyensis 156

4b. Primary inflorescence bracts 1/2, or more, the length of the secondary racemes; leaf apices long attenuate. T, cerrateana 5a. Floral bracts 3-4 cm long; inflorescence simple or compound, simple raceme or secondary racemes (or spikes) 10-16 cm long. 6. 5b. Floral bracts less than 2 cm long; inflorescence compound, secondary racemes (or spikes) less than 5 cm long. 8. 6a. Inflorescence simple; floral bracts ca. 4 cm long. T. walteri var. herrerae 6b. Inflorescence compound; floral bracts ca. 3 cm long. 7. 7a. Floral bracts dark brown to black. T. lopezii 7b. Floral bracts medium brown. T. fendleri var. reducta 8a. Inflorescence erect; lower primary bracts shorter than secondary racemes; lithophyte or rarely epiphyte. T. rubella 8b. Inflorescence down-curving; lower primary bracts 2-3 times longer than secondary racemes; epiphyte. T. iononchroma

Tlllmndsim cerrateann Lyman B. Smith "Huek'lla" HERB, lithophytic, very short stemmed. LEAVES: basal, numerous, erect, 18-29 cm long, 3.2-5.6 cm wide, not well differentiated into base and blade, narrowly trullate, base light brown, blades long attenuate, distally gray, both faces densely covered with appressed peltate scales. INFLORESCENCE: solitary central, erect; a raceme of spikes; peduncle 16-29.5 cm long, bracts imbricate, narrowly trullate, apices long attenuate to caudate, pubescent with peltate scales; raceme 5-6.5 cm long, 4-5 cm wide when pressed, very short internodes, single spike at each node; primary bracts 15-25 mm long, 14-25 mm wide, widely ellpitic, apex caudate, pubescent with peltate scales, covering 1/2, or more, of spike, red; spikes ca. 25 mm long, ca. 10 mm wide, ellipitc, 7-8- flowered, distichous, laterally flattened, pedunculule 3 mm long. FLOWERS: bracts 11- 13.3 mm long, ca. 10 mm wide, ovate-acute, coriaceous, not prominently keeled, margins chartaceous, nerves prominent, pubescent with brown peltate scales most densely adaxially, equalling the sepals, red; sepals ca. 10 mm long, ca. 7.5 mm wide, ovate- acuminate, coriaceous, margins hyaline, posterior sepals promin ently keeled, all prominently nerved; petals immature, ca. 7.6 mm long, ca. 3.8 mm wide, elliptic-acute; stamen filaments immature, ca. 2 mm long, flat, anthers ca. 5.8 mm long, versatile, yellow; gynoecium immature. CAPSULE: 20-25 mm long, 5-6 mm diam., sulcate 157

laterally, reddish brown; seeds 15-19 mm long, linear, distally dark brown, base with a tuft of many capillary hairs, as long as seeds, white. HABITAT: Cliffs in grassland or shrubland; epiphytic (Smith & Downs, 1977). 3500- ca. 4300 m. DISTRIBUTION: Peru: Ancash and Cajamarca. H. N. P.: Alpamayo trail (10075), Auquispuquio (11982), Llanganuco (fide Smith and Downs, 1977) and Los Cedros (9880). The type is from Ancash. PHENOLOGY: Flowering in March and April, and fruiting in March. CONSERVATION STATUS: This species is found in very xeric habitats, and populations can be easily destroyed. It is probably difficult to reestablish. Although not rare, it should be protected.

THImndsIm fendlerl Griseb. vr. reducfa (Lyman B. Smith) Lyman B. Smith "Machito" HERB, lithophytic, acaulescent, 40-50 cm tall. LEAVES: in basal rosette, 45-58 cm long, ca. 10 cm wide, lanceolate-attenuate, dark purple at the base, bases connivent forming a reservoir, blades spreading, green with abundant purple markings, sparsely pubescent with inconspicuous peltate scales. INFLORESCENCE: solitary, central, erect; a compound raceme; peduncle 24-43 cm long, bracts ca. 7 cm long, ca. 2 cm wide, ovate-acuminate, chartaceous; secondary racemes 2-6 per inflorescence, 13-14.5 cm long, 2.5-3.7 cm wide, IO-18-fIowered, complanate, pedunculules ca. 9 mm long. FLOWERS: bracts 22-32 mm long, 15-18 mm wide, ovate-acuminate, chartaceous, keeled, distichous, rachis hidden, shiny, green with golden cast to reddish; pedicels ca. 1.5 mm; sepals 20-27 mm long, 4-7 mm wide, lanceolate (or elliptic), acute, glabrous without, lepidote within, posterior pair connate for ca. 4 mm basally, keeled, anterior sepal not keeled; petals ca. 35 mm long, ca. 7 mm wide, elliptic-ovate, reducing to a basal claw, carnose, apex cucullate, imbricate, enrolled, violet; stamens included, filaments ca. 24 mm long, anthers ca. 9 mm long, basifixed, yellow, connivent about style; ovary ca. 6 mm long, style ca. 19 mm long, branches 1.5 mm long. CAPSULE: ca. 23 mm long, pale brown; seeds 17-18.5 mm long, dense basal plume of hairs, 15-16 mm long, white. HABITAT: Cliffs. 3600-3840 m. DISTRIBUTION: Colombia and Venezuela to Peru. Peru: Ancash, Junin, and La Libertad. H. N. P.: Honda (11715) and Ranincuray (10477). Actually much more common but not frequently collected. Tillandsia fendleri var. fendleri is wide spread in 158

the Greater Antillies, and in South America from Colombia and Venezuela to Bolivia and Brazil. PHENOLOGY: Flowering seen in October and mature fruit and seed in mid-April. USES: The inflorescences are used in great quantities to decorate crosses, portable altars, and shrines during religious festivals in the region. CONSERVATION STATUS: Abundant. This seems to be the most common species in the Park, but the heavy usage of its inflorescences may endanger it.

Tlllmndmlm humlll» C. B. Presl "Keshque" (fide Cerrate & Tovar #1768) HERB; lithophytic; caulescent with very short internodes, forming extensive creeping clones; densely pubescent with peltate, hyaline scales, plant ashen gray. LEAVES; imbricate, to 25.5 cm long, 2.6-2.8 cm wide, widest just above the base, lower 1/5 ovate, apex long attenuate becoming filiform, margins entire, involute, both faces densely pubescent. INFLORESCENCE: solitary, erect, terminal on each stem; a compound racemose panicle; peduncle 8-12 cm long, bracts foliar; panicle 6-12 cm long, 4-8 nodes, branches loosely overlapping, subtended by bract, elliptic-cuspidate, lower bracts 22.5 mm long, 10 mm wide, apex 5.8 mm long, densely pubescent, the upper bracts progressively smaller, bracts equaling or exceeding the flowers, branches 2-3- flowered, internodes 3-4.3 mm long. FLOWERS: bracts 11.8-12.2 mm long, 7 mm wide, ovate, membranous, densely pubescent; sepals 16 mm long, 6 mm wide, lanceolate, membranous, densely pubescent; petals 12 mm long, 5.3 mm wide, elliptic acute, spreading, pale coral-colored; stamens included, filaments 7.2 mm long, flat, anthers 2.7 mm long, 0.3 mm wide, connivent about the stigmas in dried flowers; ovary superior, 4.5 mm long, triangular in cross-section, style 3.1 mm long, stout. CAPSULE: not seen. HABITAT: On boulders in areas of dry open brushfields. 3600-3800 m. DISTRIBUTION: Peru: Ancash, Cajamarca, Huanuco, Junfn, La Libertad, and Lima. H. N. P.: Honda (11710) and Santa Cruz (Rauh P2072). PHENOLOGY: Flowering in early October. CONSERVATION STATUS: Rare.

Tillandsia ionochrom» André ex Mez "HuekMla" HERB, epiphytic, growing in a clone of several plants; very short stemmed. LEAVES: basal, numerous, spreading, to 64 cm long, ca. 8 cm wide, narrowly trullate, widest ca. 1/5 up from the base, dark purple at widest part, bases connivent, forming a reservoir. 159

margins entire, apex purplish; widely scattered peltate scales on both faces. INFLORESCENCE: solitary, central, down-curving, a raceme of spikes; peduncle ca. 53 cm long, bracts imbricate, foliar, lanceolate, reddish tipped; raceme to 18.5 cm long, 6 cm wide dense, 15-20 short internodes with a single spike at each node; primary bracts to 13 cm long, to 2.8 cm wide, lanceolate at base of raceme, ovate-caudate distally, reddish, longer than the spikes; spikes ca. 3.3 cm long, ca. 2.7 cm wide, ca. 7-flowered, ovate, distichous, laterally flattened; pedunculule 3-4 cm long. FLOWERS: bracts 11- 15 mm long, 9-10 mm wide, broadly ovate, coriaceous, nerves pronounced when dry, obscurely keeled, costa not strongly thickened, convex, shorter than calyx; sepals ca. 12 mm, ca. 8 mm wide, briefly united basally, posterior sepals strongly keeled, costa thickened, anterior sepal not keeled, margins chartaceous; petals ca. 14 mm long, oblanceolate, pale violet; stamens included, filaments ca. 6 mm long, flat, anthers ca. 6 mm long; gynoeclum ca. 11 mm long (immature), style ca. 3mm long. CAPSULE: 2-2.5 cm long, triangular in cross-section; seeds not seen. HABITAT: Epiphyte in Polylepis dwarf woods. 3580-3850 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Amazonas, Ancash, Cuzco, Huanuco, La Libertad, and Piura. H. N. P.: Llanganuco (8849A) and Ranincuray (9075, 10448). PHENOLOGY: Flowering from late December until April; fruiting seen in mid-April. CONSERVATION STATUS: Rare.

Tlllmmdmlm lonezH Lyman B. Smith HERB, lithophytic, stem very short. LEAVES: in basal rosette, ca. 35 cm long, ca. 10 cm wide, trullate-attenuate, base purple, blade green with purple markings, sparsely pubescent with peltate scales. INFLORESCENCE: solitary central, erect, a compound raceme; peduncle erect, bracts leaf-like basally, reduced apically; raceme sub-digitate; primary bracts to 40 mm long, erect, ovate-apiculate; secondary racemes 7-9 cm long, 2 cm wide, 8-9-flowered, flattened. FLOWERS: bracts elliptic-acute, coriaceous, keeled, margins dark brown; pedicels short; sepals to 30 mm long, elliptic, posterior pair keeled, connate for ca. 3 mm at base; petals not seen; stamen not seen; gynoeclum not seen. CAPSULE: not seen. (Description adapted from Smith and Downs, 1977.) HABITAT: Rock outcrops and cliffs. To 3800 m. DISTRIBUTION: Peru: Ancash and La Libertad. H. N. P.: collected near the Park boundary in the vicinity of Cahuish (Lôpez et al. 7563), to be expected within the Park. The type specimen is from La Libertad. 160

CONSERVATION STATUS: Unknown. COMMENT: This name may be a synonym for T. macrodactylon Mez.

THUndsU orovcnslg Mez HERB, lithophytic, very short stemmed. LEAVES: basal, numerous, erect, not well differentiated into base and blade, 19-28.5 cm long, 4-4.8 cm wide, narrowly trullate, base light brown, blades long attenuate, filiform at apex, dis tally gray, both faces densely covered with appressed peltate scales. INFLORESCENCE: solitary, central, erect; an open, sub-paniculate raceme of spikes; peduncle 23-27 cm long, bracts sub- imbricate, narrowly triangular, apices long attenuate to caudate, pubescent with peltate ' scales; raceme ca. 9 cm long, ca. 6 cm wide when pressed, very short internodes, single spike at each node; primary bracts 14-24 mm long, 10-16 mm wide, widely elliptic, apex caudate, 7 mm long, pubescent with peltate scales, less than 1/3 length of the spikes, red; spikes 33-41 mm long 11-18 mm wide, elliptic, 8-10-flowered, distichous, laterally flattened, pedunculule 3 mm long. FLOWERS: floral bracts 12-14 mm long, 8-10 mm wide, ovate-acute, coriaceous, not prominently keeled, margins chartaceous, nerves prominent, pubescent with brown peltate scales, most dense adaxially, shorter than the sepals, red; sepals 12-16 mm long, 7-8 mm wide, ovate-acuminate, coriaceous, margins hyaline, posterior sepals prominently keeled, all prominently nerved; petals ca. IS mm long, ca. 3.6 mm wide, spatulate-acute, pale violet; stamen filaments ca. 9 mm long, terete, anthers 4.5-5 mm long, versatile, yellow; gynoeclum ca. 3.5 mm long, ca. 1.5 mm diam., style ca. 4.5 mm long, branches ca. 1.5 mm long. CAPSULE: 28-30 mm long ca. 5 mm diam., sulcate laterally, reddish brown; seeds 17-19 mm long, linear, distally dark brown, base with a tuft of many capillary hairs, as long as seeds, white. HABITAT: On large boulders in dry shrubland, or shrubland - open Polylepis sericea woodland. 3500-3850 m. DISTRIBUTION: Ecuador and Peru. Peru: Ancash, Arequipa, Cajamarca, Huanuco, Junin, La Libertad, and Lima. H. N. P.: Llanganuco (10531; Gentry et al. 37377) and Parôn (10573). The type is from Junin. PHENOLOGY: Flowering and fruiting have been observed in May. CONSERVATION STATUS: Rare. This species is foun«È in very xeric habitats, and populations can be easily destroyed. It is probably difficult to reestablish. 161

TillaadiU rubella Baker HERB, lithophytlc; very short stem; to 2 m tall. LEAVES: basal, numerous, spreading, to 55 cm long, 8.5-11 cm wide, narrowly trullate, widest at 1/5-1/4 up from base, dark purple at widest point on both faces, bases connivent forming a reservoir, margins entire, blade purple streaked and spotted on the distal 1/4 of both faces, apex purple; widely scattered minute pelate scales on abaxial and, occasionally, the adaxial face. INFLORESCENCE: solitary, central, erect; a compound raceme of spikes, reddish; peduncle 85 cm long, bracts 4.7-10 cm long, ovate-acuminate, overlapping, sheathing the stem; raceme 28-29.5 cm long, 12-14 nodes with 2-3 spikes per node; primary bracts 2.5-3.5 cm long, very widely ovate, apex acute to sub-caudate, shorter than secondary spikes; secondary bracts ca. 1.4 cm long, ca. 1.2 cm wide, broadly ovate, coriaceous; secondary spikes 2.8-3.8 cm long, 1.2-1.5 cm wide, ovate, 7-11-flowered, short distichous, pedunculate. FLOWERS: bracts 11-16 mm long, ca. 10 mm wide, very broadly ovate, coriaceous, cucullate, weakly carinate apically, drying longitudinally rugose; sepals equal, 10-11 mm long, 7 mm wide, ovate-acute, strongly keeled, costa thickened, margin chartaceous, auriculate at base, glabrous, as a unit triangular in cross- section; petals 15 mm long, pale violet; stamens included, filaments 7.2-10 mm long, flat, anthers 4.5 mm long; ovary 4.5-5.5 mm long, style 8.5-9.5 mm long, triangular in cross-section. CAPSULE: not seen. HABITAT: On boulders, or on old rock falls, in openings within dwarf woods. 3600- 3700 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash, Apurimac, Cuzco, Huanuco, and Puno. H. N. P.: Llanganuco (9422) and Parûn (10595). PHENOLOGY: Flowering from January to May. USES: Smith and Downs (1977) report that the fruit of this species is edible. CONSERVATION STATUS: Rare.

Tillandsia tectorum C. J. Morren HERB, lithophytic, rhizomatous, branching at short intervals, branches short; pubescence of peltate scales, the body round with a lateral wing 3-4 times the body diameter, hyaline. LEAVES: basal, numerous, erect to spreading; to 17 cm long, 3.5 cm wide at the very base, base widened chartaceous, straw-colored, blade subulate, densely pubescent on both faces, ashen gray. INFLORESCENCE: solitary, central on each branch, long persistent, blackening with age; a compound raceme of 1-3-flowered 162 spikes; peduncle 13-25 cm long, elongating with age, erect to curving with age, bracts 4- S cm long, lanceolate, apex acute to cuspidate, apex 1-4 cm long, imbricate, sheathing, pubescent, pale magenta; racemes 4-6 cm long, 2-5 cm diam.; primary bracts 14 mm long, 9 mm wide, ovate acuminate, chartaceous, nerves prominent, cucullate, pubescent, magenta; secondary bracts 11-12.5 mm long, 3.7-6.6 mm wide, keeled, margins membranous; spikes 2.5-3.5 cm long, 3-8-flowered, lanceolate, short pedunculate, distichous, rachilla pubescent in axils. FLOWERS: bracts 16 mm long, 4.6 mm wide, keeled, pubescent at base, longer than calyx; sepals 12.5-14 mm long, 5.4-6 mm wide, ovate-acuminate, posterior sepals united basally, strongly keeled, the anterior weakly so, costae and bases chartaceous, margin hyaline, pubescent basally; petals 17-19 mm long, cucullate, bluish purple; stamens included, filaments 3-6 mm long, terete, anthers 3.3- 3.6 mm long, versatile, connivent about style; ovary 7-7.5 mm long, style 7.5-10 mm long, entire, stigma apical, convolute, papillate. CAPSULE: ca. 14 mm long. HABITATS Sheer cliff faces, extremely dry. 3240 m. DISTRIBUTION: Ecuador and northern Peru. Peru: Ancash, Cajamarca, Huanuco, La Libertad, and Lima. H. N. P.: Auquispuquio trail at Cerro Cunka (12127). The type specimen is from Ancash. PHENOLOGY: Flowering in April. USES: In Ecuador, this species is used for Christmas decorations, and is an important horticultural species extensively collected for exportation (H. Luther, pers. com.). CONSERVATION STATUS: Rare. The habitat for this species is very limited within the Park.

Tillandsia walterl Mez var. herrerae (Harms) Rauh HERBS, lithophytic, acaulescent, to 60 cm tall. LEAVES: in basal rosette, 40-52 cm long, 4.5-7 cm wide, lanceolate-attenuate, dark purple near base, bases connivent forming a reservoir, blades spreading, green with purple markings, sparcely pubescent with inconspicuous peltate scales. INFLORESCENCE: solitary, central, erect; a simple raceme; peduncle 27-40 cm long, bracts 40-50 cm long, ca. 3 cm wide, ovate-apiculate, chartaceous; raceme 12-17.5 cm long, 7-12-flowered, laterally compressed. FLOWERS: (material immature) bracts 45-53 mm long, 26-30 mm wide, ovate-apiculate, chartaceous, nerves prominent on drying, margins thin, apex often arcuate-beaked, distichous, pink, concealing rachis; pedicel ca. 6 mm; sepals ca. 42 mm long, 16 mm wide, elliptic-acute, chartaceous, slightly keeled basally, imbricate, tightly enrolled; 163 petals ca. 31 mm long, ca. 10 mm wide, elliptic-ovate, reducing to a basal claw, camose, apex cucullate, imbricate, tightly enrolled, purple; stamens included, filaments ca. 25 mm long, triangular in cross-section, anthers IS mm long, basifixed, yellow, connivent about the style; ovary 8.5 mm long, style 24 mm long. CAPSULE: not seen. HABITAT: Lithophytic sites: talus slopes, boulders, cliffs. 3700-3840 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Amazonas, Ancash, Cajamarca, Cuzco, Huancavilica, La Libertad, and Piura. H. N. P.: Ranincuray (10477A, 10481). PHENOLOGY: Very near flowering in mid-April. CONSERVATION STATUS: Rare. 164

COMMELINACEAE R. Brown COMMELINIDAE: COMMELINALES HERBS, perennial or annual; sometimes succulent, or twining; stems jointed. LEAVES: alternate, simple; sheath closed; blade narrow or expanded, margin entire; sometimes with a psuedopetiole. INFLORESCENCES: thyrses or terminal and/or axillary cymes; sometimes subtended by a folded, spathe-like bract. FLOWERS: perfect, actinomorphic or zygomorphic, hypogynous; nectaries lacking; sepali 3, usually green, sometimes petaloid and colored, rarely connate below; petals 3, ephemeral, free, at times clawed, or sometimes connate into a tube, alike or 1 reduced and/or of a different color; stamens usually 6 in 2 whorls, often dissimilar or some staminodial, filaments usually free, often bearded, anthers basifixed or versatile, connective often expanded; gynoeclum 3- carpellate, compound, superior, bi- or trilocular ovary, placentation axile, style terminal, stigma capitate, penicillate, or trilobed, ovules 1-several. FRUIT: a loculicidal capsule, rarely indehiscent or fleshy. The ca. SO genera and 700 species of the Commelinaceae are principally tropical and subtropical, with some species adapted to temperate regions. There are 11 genera reported for Peru. HABITAT: Mesic sites from the coast and Amazon Basin to middle elevations in the Sierra. POLLINATION: By pollen feeding insects, or self-pollination including cleistogamy (Dahlgren et al., 1985). USES: Some species are cultivated as ornamentals, either in garden or as house plants. I thank Dr. Robert Faden (US) for revision of the manuscript.

Commellmm Plumier ex Linnaeus HERBS, erect to sprawling; stems jointed, somewhat succulent, often brittle. LEAVES: sessile; blades usually constricted at juncture with sheath, narrow. INFLORESCENCE: terminal or leaf-opposed, composed of 1-2 cymes subtended by a folded, spathe-like bract. FLOWERS: sepals unequal, ventral 2 free or connate; petals ephemeral, free, ventral 2 clawed; stameni 6, 3 fertile with anthers normally developed, dorsifixed, 3 staminodial, anthers cruciform, filaments free, glabrous; ovary bi- to trilocular, ovules 1- 2 per locule. FRUIT: a loculicidal capsule, seeds 1-5. A cosmopolitan genus inhabiting the tropical and subtropical regions of the world. Commelina has about 170 species, of which about 10 are reported from Peru. 165

HABITAT: The typical habitats are moist, montane and rainforest vegetation types, however, the single species in the Park is found in seasonally arid vegetation. POLLINATION: By insects, or self-pollination (Dahlgren et al., 1985). BREEDING SYSTEM: Owens (1981) found outbreeding very common in the Commelinaceae, and total inbreeding very rare. In the the Tribe Tradescantieae, outbreeding is enforced by a high incidence of self-incompatibility. In the Tribe Commelineae, self-compatibility is much more common than self-incompatibility, and in Commelina all 13 species examined were self-compatible. Owens makes note that self- compatibility does not imply self-fertilization, and observes that there are many floral modifications in the genera of the Commelineae to promote outcrossing, including modification and positioning of the stamens, separation of the stamens and stigmas, and modifications of the corolla in size and color. COLLECTING PRACTICES: As the flowers are so delicate, and disintegrate rapidly, it is best to make very detailed field notes on flower structure, and preserve several flowers and fruits in fixative (FAA) for laboratory study. Photographs of the plant, especially the flowers, are invaluable.

Commellmm fggclculmtm Ruiz Lôpez & Pavon HERB, erect to 60 cm tall; roots fibrous, stout, tuberous-thickened; stems glabrous, brittle, several stems arising from same base, leaf-bearing nodes 5-6 per stem, internodes to 17.5 cm long, commonly branching. LEAVES: cauline; sheath 18-30 mm long, closed, but sometimes open to 2/3 of length, inflated and loose about stem, green with purple mottling, pubescent along a line opposite the cos ta; base of blade constricted, margins of the constriction ciliate; blades 45-65 mm long, 14-r22 mm wide, long elliptic, base rounded, apex attenuate, abaxial face glabrous, adaxial glabrous or asperous with scattered swollen-based trichomes, margins with prickle-hairs. INFLORESCENCE: terminal on branches; peduncles 3-6 cm long, densely pubescent along a line opposite the costa of the spathe, spathe held perpendicular to peduncle; spathe 26-32 mm long, 18-30 mm wide, folded, foliaceous with prominent cross veins, green with purple nerves, ovate, base cordate, margins free, outside glabrous save a few widely scattered hairs, adaxial with scattered hairs, and swollen trichomes near the margins; cyme peduncle ca. 11 mm long, bearing ca. 3 flowers, pedicels ca. 3mm long, curved. FLOWERS: ventral sepal# connate to about 1/3 length, ca. 10 mm long, ca. 5.5 mm wide, broadly elliptic, dorsal sepal free, ca. 8 mm long, ca. 3 mm wide, obovate, apex cucullate; petals 166

ephemeral, very delicate, free, equal, long clawed, blade broadly oval, ca. 12 mm long, ca. 10 mm wide, spreading, blue; fertile stamens about half as long as petals, staminodes slightly shorter, anthers of both yellow; style terminal, ca. 6 mm long. FRUIT: a loculicidal capsule, ca. 7.5 mm long. HABITAT: In seasonally dry vegetation, growing near boulders. 3240 m. DISTRIBUTION: Peru and northern Argentina. Peru: Ancash, Arequipa, Cajamarca, Cuzco, Huanuco, Huancavelica, Junin, La Libertad, Lima, and Puno. H. N. P.: at Cerro Cunka on trail to Auquispuquio (12120). PHENOLOGY: Flowering and early stages of fructification in April. POLLINATION: Visited and possibly pollinated by a fly. CONSERVATION STATUS: Rare; there is a very limited extension of appropriate habitat within the Park. 167

CYPERACEAE A. L. de Jussieu COMMELINIDAE: CYPERALES References: Geotghebeur, 1986; Tucker, 1987; van Royen 1979b. HERBS, perennial and usually rhizomatous, infrequently annual; stems typically solid, often triangular in cross-section, usually photosynthetic, especially in species with reduced leaves. LEAVES: alternate, typically 3-ranked; sheath closed; blade linear, sometimes terete, reduced, or wanting; a ligule sometimes present between blade and sheath. INFLORESCENCES: solitary, terminal spike or compound and spike-like, racemose, or paniculate; basic inflorescence unit a spike, each subtended by a prophyll and a sterile bract. FLOWERS: small, bisexual or unisexual and plants monoecious, rarely dioecious; sessile in axils of spirally or distichously arranged floral bracts, female flower surrounded by a perigynium in Carex and Undnia; perianth 1-many bristles or scales, or lacking; •tttmeni 3 (1-6), filaments free, anthers basifixed; gynoeclum 2-3-carpellate united into a compound, superior, unilocular ovary, placentation basal, style terminal with as many branches as carpels, base often enlarged and persistent in fruit, stigmas dry, ovules solitary. FRUIT: a trigonous or lenticular achene. The Cyperaceae is a family of cosmopolitan distribution, with about 105 genera and about 4000 species. There are ca. 20 genera known from Peru, found in varible habitats, from tropical rain forests to dry grasslands at high elevation, and frequently in marshy or boggy sites. POLLINATION: principally wind pollinated, at least Dichromena is insect pollinated. USES: As members of this family are common to dominant plants in grasslands and marshlands, the family is important as a forage resource. Many species are used in cottage industries around the globe; in Peru mats and other domestic articles are produced. In the Cyperaceae, the structure of the flower aggregates, often called "spikelets," and inflorescences is complex. The terminology in common usage does not adequately describe many of the cyperaceous inflorescences, and when the terms are combined they are usually unwieldly and confusing. It is assumed that the users of this manual will be more comfortable with the traditional terminology, as a result it will be used in the descriptions. The term spikelet is used to describe the basic inflorescence unit of the Poaceae, and there it has a very precise definition. As the basic inflorescence unit of the Cyperaceae and the Poaceae are of a very different composition, the term spikelet is not used in the descriptions of either the genera or species of the Cyperaceae in this manual. 168

In describing the inflorescences, the basic unit of the inflorescence is an individual spike composed of 1-many flowers with their subtending floral bracts, and the 1 or 2 bracts subtending the individual spike. When the spikes are subtended by two bracts, the lower is simple and the upper is a prophyll, a bract with two, prominent, keeled nerves. Prophylls are also found at many branch-bearing nodes throughout the inflorescence, and are often associated with branch points on the vegetative stem. The term Inflorescence bract refers to the bract which subtends the entire inflorescence. Those interested in more detailed analysis and description of the inflorescences in this family, or in general, should consult the works of Eiten (1976), Kukkonen (1984; 1986), Troll (1964), and Weberling (1965). In addition, the morphology and specialization of the Cyperaceae floral unit is the subject of reinterpretation (Goetghebeur, 1986; Meert and Goetghebeur, 1979; Meeuse, 1975a, 1975b). Key to the genera of the Cyperaceae la. Flowers unisexual; female flower enclosed in a perigynium. Carex lb. Flowers, at least 1 per spike, bisexual; flowers naked on floral bract. 2. 2a. Perianth wanting. 3. 2b. Perianth present, of bristles or scales. 7. 3a. Flowers distichous on spike; inflorescence compound. Cyperus 3b. Flowers spirally arranged; inflorescence simple or compound. 4. 4a. Achene with apical tubercle (the enlarged style base); inflorescence compound; plants of grasslands. - Bulbostylis juncoides 4b. Achene lacking apical tubercle; inflorescence simple or compound. 5. 5a. Inflorescence pseudo-lateral, compound, usually of 3, sub-sessile spikes; plants of boggy habitats. Isolepis imndata 5b. Inflorescence a simple, terminal spike. 6. 6a. Leaves cauline; plants a few cm tall; growing in boggy habitats at high elevation. Phylloscirpus 6b. Leaves basal; plants 10-30 cm tall; of dry grassland habitats. Baeothryon rigida 7a. Perianth of retrorsely barbes bristles; plants less than 10 cm tall; of boggy habitats. 8. 7b. Perianth of scales; size and habitat diverse. 9. 8a.Inflorescence a simple spike; leaves reduced to bladeless sheaths. Eleocharis albibracteata 169

8b. Inflorescence compound, (1-) 2-3 sub-sessile spikes; leaves with developed blades. Phylloscripus 9a. Scales fimbriate; inflorescence large and branched; tall emergent aquatics growing in lakes. Schoenoplectus californicus 9b. Scales not fimbriate; inflorescences small, simple spikes. 10. 10a. Spike with 1 terminal, bisexual flower; achene obconic-truncate; of boggy habitats at high elevation. Oreobolus obtusangulatus 10b. Spike with 3-6 bisexual flowers; achene obovate, trigonous compressed; grassland plants. Oreobolopsis tepalifera

Baeothrvon A. G. Dietrich References: Beetle, 1953; Goetghebeur, 1986; Koyama, 1958; Weber, 1987. HERBS, stiffly erect from caespitose rhizomes; stems terete or nearly so; glabrous. LEAVES: basal, sheaths closed, chartaceous, shiny, with a ligule at juncture of sheath and blade, blades linear, stiff, canaliculate. INFLORESCENCE; a solitary, terminal spike; inflorescence bract widened at base, stem-like above, not conspicuous, often soon caducous; spikes several-flowered, subtending bract elliptic with a stout, elongate costa produced into a blade-like arista. FLOWERS: bisexual; floral bracts spirally arranged; perianth 0-6 filiform bristles; stamens 3, filaments linear, anthers linear; stigmas 3. ACHENES: trigonous, compressed to lenticular, smooth, shiny black, minutely apiculate. The genus is still poorly defined, currently approximately 10 species are included, with only Baeothryon rigida known from Peru. The genus is widely distributed in the grassland areas of the Northern Hemisphere, mountains of tropical Asia, and the Andes.

Baeothrvon rlgfda (Boeckeler) D. N. Smith, comb. nov. ined. ("Scirpus rigidus Boeckeler) HERBS: 10.5-25 cm tall, erect, stiff, rhizomes caespitose, forming bunches (8-) 10-40 cm diam., often interrupted or with hollow centers; stems terete below, laterally compressed above; glabrous. LEAVES: 7-15 cm long, sheaths 4-5.5 cm long, closed, many-nerved, shiny, reddish brown, persistent, becoming fibrous, small ligule at mouth of sheath, blades (3-) 7.5-11.5 cm long, ca. 0.6 mm wide, linear, stiff, canaliculate, margins scabrous, apex pungent, slightly rounded. INFLORESCENCE; a terminal, sessile spike; peduncle 5.5-20 cm long, terete, ribbed; inflorescence bract (4-) 8.3-23 mm long, ca. 4 mm wide, foliaceous, base ovate, widened and sheathing, papery, light brown, blade linear, laterally 170

compressed, keels scabrous; spike 4-9 mm long, 6-7- flowered with 1-3 sterile bracts below flowers, subtending bract 5.8-10 mm long, ca. 4 mm wide, similar to inflorescence bract. FLOWERS: floral bracts 3.8-6 mm long, 2.1-3 mm wide, ovate- elliptic, 1- nerved, chaffy, stiff, light brown with tan margins; stamens 3, filaments 3.4-6 mm long, linear, flattened, long exserted, anthers 1.7-2.6 mm long, base cordate, apiculate, yellow; style 3.8-5.3 mm long, deciduous, stigmas 3, reddish. ACHENES: 1.9-2.4 mm long, 1- 1.5 mm wide, obovate, plano-convex to sub-trigonous, apex with a small apicule, shiny, nearly black. HABITAT: Found growing in the arid grasslands, especially those on deeper soils developed on morainal material. The parent material may be either granitic or sedimentary rock. 3875-4600 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash, Junin, and Puno. H. N. P.: Cahuish (10146), Ishinca (9435), Quenua Ragra (10623), Querococha (11036), Queshque (11852) and Rio Pachacoto (8768). PHENOLOGY: Flowering has been observed in December, March and May, fruiting in the same months, and seed cast in July. USES: This species is impalatable and very resistant to grazing. The flowers are sometimes smutted. CONSERVATION STATUS: Common.

BulboitvHs C. Steven Reference: Krai, 1971. HERBS, perennial or annual, erect from very short creeping rhizomes, stems terete, ribbed, thickened at base. LEAVES: basal, sheath short, closed, pubescent, blades filiform, pubescent. INFLORESCENCES: terminal, compound, umbellate to capitate; inflorescence bracts 1-several, leaf-like, erect; spikes several-many-flowered. FLOWERS: bisexual; floral bracts spirally arranged, glabrous or pubescent; perianth wanting; stamens 1-3, filaments linear, flattened, anthers linear; style base swollen, forming a stylopodium, persistent in fruit, stigmas 3. ACHENE: trigonous, smooth or sculptured. The genus has 80-100 species distributed in warm temperate and tropical regions throughout the World, especially on dry, sandy sites. There are about 7 species in Peru. 171

BulbostvUi luncoldes (Vahl) KOk. (including B. arenaria (Nees) Lindm.) HERBS 6-28 cm tall, erect from short creeping rhizomes, forms bunches 2-5 cm diam., stems thickened and knotty at base. LEAVES: 3-12.5 cm long, sheaths short, papery, brown, with fibrous pubescence at margins and apex, persistent, blades filiform, canaliculate, scabrous to hispid, apex acute. INFLORESCENCE: condensed, umbellate, irregularly (1-) 3-12-branched, branches 1- 7 mm long; peduncle 4.5-27.5 cm long; inflorescence bract 6.5-12 mm long, 1.4-2 mm wide at base leaf-like, base ovate, open and sheathing, papery, with fibrous pubescence above, brown; spikes 4-7 mm long, ca. 2 mm wide, 9-14-flowered, subtending bract 2.5-3.5 mm long, 1.4-2 mm wide, like floral bracts in shape and texture, prophylls ca. 3 mm long, ca. 2 mm wide. FLOWERS: floral bracts 2-3 (-3.5) mm long, 1.4-2.2 mm wide, broadly ovate, costa stout, green, apex often mucronate, blade papery, reddish brown, margins ciliate; stamens 3, filaments ca. 2 mm long, linear, anthers 0.6-1.1 mm long, apiculate, base cordate, yellow; style ca. 2 mm long, stigmas 3. ACHENE: 0.7-1.1 mm long, 0.4-0.6 mm wide, obovate, trigonous, constricted into short neck below the depressed globose stylopodium, body with transverse wavy lines, white. HABITAT: Dry sites in grasslands, and Polylepis sericea woodlands or woodland- shrubland. 3500-3950 m. DISTRIBUTION: Southwestern United States to Honduras, West Indies, Colombia and Venezuela to Argentina, and Brazil and Paraguay. Peru: Ancash, Cuzco, and Junin. H. N. P.: Alpamayo-Cashapampa trail (10074), Llanganuco on Maria Josefa trail (10525), and Ranincuray (9074, 10343). PHENOLOGY: Observed flowering from January to May, and fruiting in January and March. Krai (1971) found this species to be the most variable of the North American Bulbostylis. CONSERVATION STATUS: Uncommon; probably not endangered.

Carex Linnaeus References: Boott, 1968; Hermann, 1974; Kûkenthal, 1909. HERBS, perennial from creeping or caespitose rhizomes, often forming extensive clones; stems solid, or becoming hollow with age, triangular. LEAVES: 3-ranked, linear, sheaths closed, sheath margins hyaline to papery, above decurrent on blade, uniting into a small 172 ligule, blades flattened, variously folded, or terete, costa often prominent and winged, margins usually serrulate and cutting. INFLORESCENCE: simple or compound, appearing to be spikes, racemes, panicles or heads composed of few-many-flowered spikes, branching complex, elongate or condensed; terminal on elongate stem, this either a leafy culm, or leafless peduncle, rarely sessile; inflorescence bracts often leaf-like, base a closed sheath; spikes may be unisexual, or bisexual, then androgynous or gynaecandrous. FLOWERS: unisexual, plants monoecious; floral bracts spirally arranged; perianth wanting; itamlnate flowers composed of 3 free stamens, filaments filiform, exserted, anthers linear; carpellate flowers composed of a bi- or trilocular ovary surrounded by a perigynium, stigmas 2 or 3, papillate. ACHENES: trigonous or lenticular, enclosed in perigynium at maturity. Carex, a genus of sub-cosmopolitan distribution, has ca. 2000 species, of which about 24 are known from Peru. The genus is highly variable. Ecologically the genus is very important, especially in marshy and boggy habitats. Many species of Carex are important sources of forage for wildlife and domestic livestock grazing on rangelands. POLLINATION: By the wind. INFLORESCENCES: These can be simple spikes or complex structures with several orders of branching. See the comments following the family description. Some species of Carex are dioecious; none are reported from Peru. I thank Dr. Anton A. Reznicek (MICH) for his help with this difficult genus.

Key to the species of Carex la. Plants dwarfed, immersed in the soil; inflorescence essentially sessile. C. collumantha lb. Plants erect; inflorescence elevated on leafy culm or peduncle. 2. 2a. Stigmas 3; achenes trigonous. 3. 2b. Stigmas 2; achenes lenticular. 6. 3a. Plants 40-100 cm tall, coarse; spikes androgynous; inflorescences either with more than one branch per node or with compound branches 4. 3b. Plants 40 cm tall or less; spikes gynaecandrous; inflorescence branches single at nodes and not compound. S. 4a. Lower inflorescence bracts sheathing; perigynia obovate, 2-3 x 1-2 mm, apex with a distinct, truncate beak, ciliate at mouth; inflorescence branches filiform, lax, spikes often pendant. C. hebetata 173

4b. Lower inflorescence bracts sheathless; perigynia lanceolate, 2-4 x 0.6-1 mm, apex attenuate, 2-toothed, not beaked; inflorescence branches stiff, ascending. C. ancashensis 5a. Culms stiff, erect; perigynia elliptic, 3-4 x 1-2 mm, apex with a truncate beak. C. crinalis 5b. Culms filiform, flexuous, arcuate curving; perigynia ovate 2-3.2 x 1-1.6 mm, apex attenuate, not beaked. C. boliviensis 6a. Spikes gynaecandrous; plants caespitose or tufted from short creeping rhizomes. 7. 6b. Spikes androgynous; culms arising singly from long-creeping rhizomes. C. ecuadorica 7a. Plants to 60 cm tall, culms erect; perigynia thin margined but wingless. C. bonplandii 7b. Plants less than 25 cm tall, culms arching or spreading; perigynia winged. C. mandoniana

Cmrex ancashensis. D. N. Smith & Reznecek, sp. nov. ined. HERB, coarse, erect, 50-120 cm tall, from a stout creeping rhizome, forming extensive clones; glabrous. LEAVES: (28-) 40-108 cm long, 6-13 mm wide, linear, sheath 6-15 cm long, sub-equitant, thickened, septate, purplish, blade plicate, costa winged, margin scabrous, apex attenuate. INFLORESCENCE: a very complex spicate panicle, 18.5-26 cm long; peduncles 44-113 cm long, triangular, solid, scabrous on keels; inflorescence bracts 15-44.5 cm long, foliar, base closed; bracts subtending inflorescence branches lanceolate, apex attenuate or filiform, triangular, scabrous, dark purple, occasionally the lower 1-2 nodes with bracts like the inflorescence bracts in size and caracter; 7-9 ascending branches, triangular, often scabrous on keels, lowermost branch usually much longer than others, 3-7.5 cm long, middle branches progressively shorter, 43 mm long or less, uppermost sub-sessile, branches and central axis terminating in dense, elongate compound spikes, 8-49 (-90) mm long, these bearing a small simple or compound spike at each node, or some basal nodes with a short, secondary branch of the same nature as the primary branches, spikes of the lower half of the compound spike often compound with spikes at lower nodes and individual flowers above, upper half with simple spikes, each small spikes subtended by a bract and prophyll, simple spikes 4-6 mm long, androgynous, ca. 6 staminate and 4 carpellate flowers per spike. FLOWERS: bracts and flowers very dark 174

purplish brown; floral bracts 2-3 mm long, 1.1-2.2 mm wide, ovate, apex acuminate to aristate; staminate flowers: filaments 2-3 mm long, anthers 1.5-2.4 mm long, base cordate, apex acute to apiculate, yellow; carpellate flowers: perigynia 2.2-3.8 mm long, 0.6-1 mm wide, lanceolate, without a distinct beak, teeth erect, 0.3-0.4 mm long, tip diverging outwards, style ca. 3 mm long, stigmas 2. ACHENE: not seen. HABITAT: Growing in boggy or marshy areas, streamsides, or shallow water in lakes and ponds, those habitats with saturated soil conditions, or standing or slowly flowing water. 3900-4400 m. DISTRIBUTION: Peru: Ancash. H. N. P.: Only on the eastern side of the Cordillera; Cahuish (10118), Carhuazcancha in Laguna Ichicpotrero (12419), and Rurichinchay (12600, 12698). PHENOLOGY: Flowering in May and June. CONSERVATION STATUS: Common on the east side of the Park.

Cmx boUvIenslg Van Juerck & MQll.-Arg. HERBS, foliage to 17 cm tall, from caespitose rhizomes. LEAVES: 7-17 cm long, 1-2.6 mm wide, linear, spreading, sheath 1.1-3 cm long, mostly buried, becomming fibrous in age, blades flat to plicate, often pubescent near juncture with sheath, costa winged, apex attenuate, plicate. INFLORESCENCE: spicate; peduncle 13-48.5 cm long, triangular, filiform, arcuate, flexuous, scabrous on keels; inflorescence bracts 14-50 mm long, erect, base ovate, closed, chartaceous, brown, tip foliaceous, filiform, triangular; a second order peduncle sometimes produced between inflorescence bract and lower spike, to 5.5 cm long; (1-) 2-3 spikes per peduncle, the lower sub-sessile, the upper terminal on an erect, triangular, scabrous axis, 2-14 mm long, the middle, if developed, short pedicellate just below terminal spike, lower spikes 8-13 mm long, middle 7-11 mm long, terminal 10-19 mm long, the terminal gynaecandrous, with 6-10 staminate and 25 or more carpellate flowers, those below carpellate, spikes subtended by bracts similar in size and shape to inflorescence bracts. FLOWERS: floral bracts 2-2.5 mm long, 1.1-2 mm wide, wide ovate, light brown, costa green, narrow, apex acute to rounded, margin erose, whitish; staminate flowers: filaments 3-4.5 mm long, anthers not seen; carpellate flowers: perigynia 2.3-3.2 mm long, 1-1.6 mm wide, ovate, green with prominent, straw-colored nerves, apex attenuate, not beaked, mouth with calloused rim, style ca. 2.2 mm long, stigmas 3. ACHENES: 1.8-2 mm long, 0.9-1.1 mm wide, elliptic to obovate, trigonous, light brown, style base persistent as beak, 0.1-0.2 mm long. 175

HABITAT: Found in moist grasslands in valley bottoms. 3700-4100 m. DISTRIBUTION: Mexico to Bolivia and northern Argentina. Peru: Ancash. H. N. P.: Ranincuray (10396) and Rurichinchay (12558). PHENOLOGY: Flowering observed in April to June, and fruiting in June. USES: This species is palatable to livestock, probably both cattle and sheep. CONSERVATION STATUS: Rare. Perhaps this species has been overlooked during the collecting trips, and may be more common.

Carex bomolmndll Kunth HERB erect, to 60 cm tall, from a densely caespitose rhizome, clones 10-40 cm diam.; glabrous. LEAVES: 8-53 cm long, 2.1-4.2 mm wide, linear, sheath 2.8-11 cm long, brownish, blades flat to folded, margins scabrous, costa prominent, slightly winged, apex attenuate. INFLORESCENCE: condensed, spikelike, 15-30 mm long, rarely paniculate, producing a lateral branch at lowermost node; peduncles 10.5-58 cm long, triangular, central lumen filled with very porous aerenchyma, becoming hollow in age, keels scabrous distally; inflorescence bract (7) 10-47 (65) mm long, base ovate, papery, strongly nerved, brown, margin thin, costa green, apex long, triangular, scabrous on keels; spikes 5-11 per inflorescence, 5-15 mm long, gynaecandrous, 2-7 staminate and 15- 25 carpellate flowers per spike, subtending bract ca. 4 mm long, ca. 2.2 mm wide ovate-acute, chartaceous, 1- nerved, brown, the lowermost like inflorescence bract. FLOWERS: floral bracts 2.6- 3.9 mm long, 1.4-1.8 mm wide, ovate to lanceolate, 1-nerved, margins hyaline, apex attenuate, light brown with green costa; staminate flowers: filaments 2.5-5 mm long, anthers 1.2-1.9 mm long, base cordate, apex with tuft of trichomes, yellow; carpellate flowers: perlgynia 2.7-3.8 mm long, 0.8-2 mm wide, ovate-attenuate, several-nerved, membranous, winged, beak 0.8-1.3 mm long, teeth erect, margin serrate, body green, beak brownish, style 4-5 mm long, stigmas 2. ACHENES: 1.2-1.6 mm long, 0.9-1.1 mm wide, elliptic to obovate; style deciduous. HABITAT: Moist grasslands on slopes and valley bottoms, and streamsides. 3700-4600 m. DISTRIBUTION: Southern United States (Arizona) to Bolivia. Peru: Amazonas, Ancash, Huanuco, Junin, and Puno. H. N. P.: Alpamayo trail at Hatuncocha (9981), Cahulsh (10119), Llanganuco in Que. Demanda (10311), Ranincuray (10355, 10430), Rurichinchay (12561), and Shallap (9688). 176

This species is quite variable throughout its range, and further study may show that there are more than one species included in the name C. bonplandii as currently used. PHENOLOGY: Flowering and fruiting observed from February to June. USES: Apparently palatable to livestock. Several specimens are smutted. CONSERVATION STATUS: Not endangered.

Careic eoilumantha (Steyerm.) Mora HERB submerged in soil, underground stems 10-15 cm long, rhizomes creeping, forming extensive clones; glabrous. LEAVES: 7-12 mm long, ca. 2 mm wide, sheaths 3.8-4 mm long, hyaline with ca. 7 prominent nerves, blades narrowly triangular, carinate, thickened, papillate above, diverging from stem at 45 degree angle, margins and keels scabrous apex blunt. INFLORESCENCE: a condensed panicle, 4-8 mm long; sessile, partially hidden in apical leaves; inflorescence bract not seen; spikes 1 -4 per inflorescence, unisexual, terminal spike staminate, few-(2-3?)-flowered, lower spikes carpellate, 2-3-flowered. FLOWERS: floral bracts ca. 3.5 mm long, ca. 3 mm wide, broadly ovate, papery, 2- nerved, margins hyaline; staminate flowers: anthers ca. 2.5 mm long, apiculate; carpellate flowers: perigynia 3.5-5.1 mm long, 2.5-3 mm wide, elliptic to obovate, thickened, nerves obscure, dusky orange, beak 0.4-0.5 mm long, with 2 small teeth, style ca. 3 mm long, stigmas 2. ACHENES: ca. 1.9 mm long, ca. 1.3 mm wide, obovate, lenticular, dark brown. HABITAT: In boggy or seepy spots on sparsely vegetated, recent terminal moraine with poorly developed soil of granitic parent material; on a steep, SSE-facing slope. 4900 m. DISTRIBUTION: Venezuela, Colombia, and Peru. Peru: Ancash. H. N. P.: Parôn (11507). PHENOLOGY: Achenes mature in late September. CONSERVATION STATUS: Very rare.

Carex crinalls Boott HERB, erect, to 40 cm tall, from caespitose rhizome; leaves and stems strigose to hispid. LEAVES: 4.5-33 cm long, 2.3-4.3 mm wide, linear, sheath 2.1-11 cm long, light brown, dried bases often persistent, blades flat to plicate, costa prominent, marins entire, apex attenuate. INFLORESCENCE: racemose, 12-40 mm long; peduncles 7-36 cm long, triangular, solid, spreading; inflorescence bract 13-50 (-69) mm long, foliar, erect, base closed, margins and costa pubescent; a secondary peduncle sometimes produced between 177

inflorescence bract and lower spike, to 26 mm long; spikes short pedicellate along erect axis 7-16 (- 35) mm long, lowermost sometimes remote, spikes 3-5 per inflorescence, 10- 17 mm long, terminal spike gynaecandrous with 8-12 staminate and 10-15 carpellate flowers, lower spikes carpellate, 10-25-flowered. FLOWERS: floral bracts 2.3-3.6 mm long, 1.6-2.2 mm wide, broadly ovate, membranous, green, apex attenuate to rounded, margin hyaline, erose-ciliate; staminate flowers: filaments 3.5-4.5 mm long, anthers 1.8- 2.2 mm long, base cordate, apex with tuft of trichomes; carpellate flowers: perlgynia 3.0- 3.9 mm long, 1.1-1.7 mm wide, elliptic, green, beak 0.6-1 mm long, truncate, style 2.3- 3.5 mm long, swollen and papillate above base, stigmas 3. ACHENES: 1.7-2 mm long, 1.1-1.5 mm wide, obovate, trigonous, tan; style deciduous. HABITAT: Moist grasslands on slopes and valley bottoms. 3800-4300 m. DISTRIBUTION: Colombia to Peru. Peru: Ancash and Huanuco. H. N. P.: Only on the east side of the Cordillera; Carhuazcancha below Laguna Ichicpotrero (12407), Ranincuray (9127, 10385), and Rurichinchay (12631). PHENOLOGY: Flowering has been observed from January to June, and fruiting from April to June. Perhaps the species flowers nearly all year long. CONSERVATION STATUS: Not endangered.

Carex ecuadorlca Kûk. HERB erect, to 15 cm tall or more, from a deeply buried, long creeping rhizome; glabrous. LEAVES: to 15 cm long or more, 2-3 mm wide, linear, bases subterranean, sheaths 5-6 cm long, blades arcuate-spreading, flat to folded, costa slightly winged, margins and keel scabrous. INFLORESCENCE: spike-like, 7-10 mm long, ovate; peduncles 6-7 cm long, triangular, smooth; inflorescence bract ca. 2.8 mm long, broadly ovate, papery, light brown, margins hyaline; spikes 4-6 per inflorescence, 5-7.5 mm long, andro gynous. FLOWERS: floral bracts 3.1-3.7 mm long, 2.1-2.6 mm wide, broadly ovate, chartaceous, apex acuminate to rounded, brown, margin hyaline, erose; staminate flowers: filaments ca. 3 mm long, anthers 2-2.6 mm long, base cordate, apex with tuft of trichomes, yellow; carpellate flowers: perlgynia 2.5-2.7 mm long, 1-1.1 mm wide, lanceolate, winged, keels scabrous, brown, beak indistinct, 2 obscure teeth, stigmas 2. ACHENES: not seen. HABITAT: Moist grassland in river bottom with sandy soils. 4200 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash. H. N. P.: Rio Pachacoto (9370). PHENOLOGY: Staminate flowers in advanced stage of anthesis in mid-January. 178

USES: Palatable to livestock. CONSERVATION STATUS: Very rare; found only once.

C«rex hebetmtm Boott HERB, erect, 41-131 cm tall, coarse, from stout creeping rhizome, forms extensive clones; glabrous. LEAVES: 39-55 (97) cm long, 3-10 mm wide, linear, sheath 4-20 cm long, purplish, blades flat or plicate, costa prominent, winged abaxially, margins scabrous, apex attenuate. INFLORESCENCES: a panicle; culm 17-86 cm long, triangular, smooth, bearing several leaves, erect; inflorescence bracts 12-57 cm long, foliar, base closed, 5- 16.5 cm long, blade erect; main axis 18-69 cm long, triangular, erect, (1-) 2-3 (-5) branches at each node, 13.5-35.5 cm long at lower node, apical branches much smaller, filiform, flexuous, lax, scabrous; spikes, (4) 10-18 (22) per panicle, 9-46 mm long, densely many-flowered, androgynous, staminate portion usually 40-50 % of length, rarely all pistillate or all staminate. FLOWERS: floral bracts (2.1) 2.4-3.1 (3.9) mm long, 1.3- 1.8 (2.0) mm wide, ovate, 1-nerved, apex acute to attenuate, margins ciliate, dark brown; staminate flowers: filaments 2.5-5 mm long, anthers 1.9-3 mm long, 0.2 mm wide, base cordate, tip apiculate, yellow; carpellate flowers: perlgynla 2-3 mm long, 1-1.8 mm wide, obovate, several-nerved, apex constricted into beak 0.1 mm long, truncate, ciliate at mouth, green often with brown markings, style 1.5-2 mm long, stigmas 3. ACHENE: 1.1-1.5 mm long, 0.6-1.1 mm wide, obovate to elliptic, trigonous, light brown with pale nerves, style base persistent as 0.1-0.4 mm beak. HABITAT: Grows in microsites with moist to saturated soils in grasslands, especially tussock grasslands. Often found in seeps, rivulets, or in standing water. 3700-4800 m. DISTRIBUTION: Peru: Ancash and Lima. H. N. P.: Alpamayo (9871), Huaripampa (9158), Ishinca (11176), Llanganuco in Que. Demanda (10301), Llanganuco on Sendero Maria Josef a (10511), Parôn (11504), Ranincuray (10442), and Rurichinchay (12620). PHENOLOGY: Flowering occurs from January to June, and fruiting from March to September. Spikes can begin to shatter as early as April. CONSERVATION STATUS: Common.

Carex mandoniana Boeckeler HERB, to 22 cm tall, from a caespitose rhizome; glabrous. LEAVES: 5.3-17 cm long, 1.5-3.5 mm wide, linear, erect to arcuate-spreading, sheaths 1.5-3 (-7) cm long, blades folded to plicate, costa winged, margins serrulate, apex attenuate. INFLORESCENCE: 179

dense, spicate, 10-18 mm long, ovate; peduncles 5-26 cm long, arcuate- to sinuous- curving, triangular, central lumen with very porous aerenchyma, becomming hollow with age; inflorescence bract 5-28 mm long, base ovate, apex filiform, triangular, keels scabrous; spikes 3-6 per inflorescence, 5-9 mm long, gynaecandrous, 2-3 staminate and 20-30 carpellate flowers per spike. FLOWERS: floral bracts 3.7-4.4 mm long, 2-2.5 mm wide, ovate- attenuate, brown with green costa, margins hyaline; staminate flowers: filaments 3.5-4 mm long, anthers 1.5-2 mm long, apiculate, bases cordate; carpellate flowers: perlgynla 4.2-4.8 mm long, 1.6-2.3 mm wide, elliptic, winged, margin ciliate, green with brown beak and margins, beak 1.1-1.6 mm long, attenuate, arcuate at maturity, with 2 erect teeth, style 3-3.5 mm long, stigmas 2. ACHENES: 1.7-1.9 mm long, 1.2-1.4 mm wide, elliptic to obovate, lenticular, light brown; style deciduous. HABITAT: Dry grassland or grassland-shrubland. 4000-4400 m. DISTRIBUTION: Mexico to Bolivia. Peru: Ancash and Junin. H. N. P.: Only on the west side of the Cordillera in Llanganuco at Que. Demanda (10256) and Rajucolta (12184). PHENOLOGY: Flowering and fruiting observed in April. USES: Palatable, apparently highly so, as the species is found growing in clones of less palatable plants in the areas which suffer serious overgrazing. CONSERVATION STATUS: Uncommon, endangered by severe overgrazing.

Cvnenut Linnaeus References: KQkenthal, 1936; Tucker, 1983. HERBS, annual or perennial, from caespitose rhizomes or fibrous roots; stems 3-angIed to rounded. LEAVES: linear, mostly basal, most with a well developed sheath and blade, a few species bladeless. INFLORESCENCE: simple or compound, umbellate or capitate, branches unequal, subtended by leaf-like inflorescence bracts, the basic inflorescence unit a distichous, cylindric or flattened spike, each subtended by a prophyll and a sterile bract. FLOWERS: bisexual; floral bracts distichous or, less often, spiralled; perianth wanting; stamens (1-2-) 3; stigmas 2 or 3. ACHENE: trigonous or lenticular. Cyperus, a genus of cosmopolitan distribution, has about 600 species, of which 75 more occur in Peru (G. C. Tucker, pers. com.). USES: Some species of Cyperus are palatable and form part of the forage resource. I thank Dr. Gordon C. Tucker (NYS) for his advice regarding this genus, and for revision of the manuscript. 180

Key to the species of Cvnenis la. Plants very small, mostly immersed in the soil; inflorescence subsessile, capitate; spikes strongly flattened laterally. C. seslen'oides lb. Plants erect, 23 cm or more tall; inflorescence on erect peduncle, umbellate; spikes terete. C. hermaphorditus

Cvnerus hermanhraditus (Jacq.) Standley HERB, erect, 23 cm or more tall, from caespitose rhizomes; stems with knotty, thickened bases; glabrous. LEAVES: 8-19 cm long, 3-5 mm wide, mostly basal, linear, spreading, sheaths ca. 3 cm long, closed, costal region membranous, reddish, marginal region subhyaline, weathering to a fibrous tunic, blades plicate, margins scabrous, tip attenuate. INFLORESCENCE: umbellate; peduncles 5.5-23 cm long, triangular, aerenchyma filled, collapsing on drying; 3-6 rays, unequal, to 2.5 cm long, sometimes branched basally, each terminating in a compound spike, 9-20 mm long, to 10 mm wide, spikes at right angle to rachis; inflorescence bracts to 8 cm long, foliar; spikes many, (2.8-) 5- 6.5 mm long, 2-4- flowered, terete, prophyll 1.2-1.7 mm long, sterile bract 0.9-1.5 mm long, broadly ovate, multi-nerved, papery. FLOWERS: floral bracts 2.6-3.2 mm long, 1.2-2 mm wide, elliptic, membranous, multi-nerved, pale green; stamens 3, filaments 2.4-3.3 mm long, linear, flattened, anthers 0.5-0.7 mm long, elliptic, yellow; style 2.9-3.8 mm long, 3-fid. ACHENES: trigonous, narrowly oblong, mature material not seen. HABITAT: Dry grasslands on the northwestern flanks of the cordillera. 3500-3950 m. DISTRIBUTION: Mexico and Central America; West Indies; Colombia and Venezuela to northern Argentina. KQkenthal (1936) also reports this species from the southern United States and Brazil; these reports are not confirmed in recent surveys. Peru: Ancash, Huanuco, Junin and Lima. H. N. P.: Alpamayo-Cashapampa trail (10071) PHENOLOGY: In March the population collected was in late flowering and early fruiting stage. This species has the anatomy associated with the C4 photosynthetic pathway (G. C. Tucker, pers. com.) CONSERVATION STATUS: Rare, as the habitat of the species is limited within the Park. 181

CvBenia «galerlolde» Kunth (including C. andinus Palla in Buchtein ex KQk.) HERB, short-lived, possibly annual, root crown, stems, and leaf bases buried, forming small clones of 3-4 plants; all parts green, glabrous. LEAVES: 3.5-6.5 cm long, 0.9-1.8 mm wide, linear, arcuate, folded, margins entire. INFLORESCENCE: capitate, the branching structure very condensed, visible only on dissection, subhemispheric, 5-10 mm tall, 7-11 mm wide, sessile to little elevated on a terete, channeled peduncle to 5 mm long; inflorescence bracts 4-9, to 5.5 cm long, foliar; spikes many, tightly crowded, 3-6.7 mm long, 9-11-flowered, strongly flattened laterally, prophyll small, sterile bract 2.9-4.3 mm long, 2-2.4 mm wide. FLOWERS: floral bracts 2-3 (-3.7) mm long, 0.8-2.5 mm wide, ovate, thin-membranous, 3-nerved; stamens 1 (-2), filaments 2-3.2 mm long, linear, flattened, widened distally, anthers 0.5-0.7, yellow; ovary with style 1.2-2.3 mm long, deeply divided into 3 stigmatic lobes. ACHENES: 1-1.5 mm long, 0.5-0.9 mm wide, trigonous, ovoid, pale beige. HABITAT: Disturbed spots in dry grassland or grassland-shrubland, in areas of low vegetative cover, often on poor soils. 3800-4300 m. DISTRIBUTION: Southern United States (Arizona) through Mexico and Guatemala; Colombia and Venezuela to northern Argentina. Peru: Ancash, Arequipa, Cuzco, Huanuco and Lima. H. N. P.: Auquispuquio (11951) and Rio Pachacoto (11803). TAXONOMIC COMMENTARY: At high elevations, the individuals of this species are small with a very short peduncle, more involucral bracts, larger inflorescence diameter, longer spikes, and larger floral bracts. Considering these differences in relation to the overall variation within C. seslerioides. Tucker (pers. com.) did not find it significant, not even justifying of subspecific rank. These small individuals were separated into C. andinus Palla in Buchtein (KQkenthal, 1936); Tucker (pers. com.) considers C. andinus to be a synonym of C. sesierioides. PHENOLOGY: Flowering and fruiting occur in rapid succession in March and April. This species has the anatomy associated with plants of the C3 photosynthetic pathway (G. C. Tucker, pers. com.) CONSERVATION STATUS: Probably common. The species is very small and easily overlooked. It is probably more common than the two collections indicate. 182

Eleocharl» R. Brown Reference: Svensen, 1929. HERBS, perennial, erect from caespitose or creeping rhizomes, stems photosynthetic, stout and septate or filiform, terete or triangular; glabrous. LEAVES: sheaths closed, papery, 1-few per stem, often reddish or purplish, blades wanting. INFLORESCENCES: a simple, terminal, few-many-flowered spike; inflorescence bract wanting; spike subtended by a scale-like bract resembling floral bracts in shape and texture. FLOWERS: bisexual; floral bracti spirally arranged; perianth 3-8 retrorsely barbed bristles, or, rarely wanting; •tamcns 2-3, filaments linear, anthers linear; style base swollen forming a triangular stylopodium persistent in fruit, stigmas 2-3. ACHENES: trigonous or lenticular. This sub-cosmopolitan genus has ca. 200 species, of which about 11 are known from Peru.

Elcoch«rl« «Ibibracteata Nees & Meyen ex Kunth HERB perennial, from a creeping rhizome; stems 2-9 cm long, filiform, erect, some sterile and leaf-like in appearance, then usually shorter than fertile stems. LEAVES: sheaths 0.5-3 cm long, papery, reddish to light brown. INFLORESCENCES: spike 2-3.6 (-4.8) mm long, 0.8-1.5 mm wide, 3-10-flowered, subtending bract 1.2-2.1 mm long, 1- 1.6 mm wide, wide elliptic, apex rounded, like floral bracts in texture and color. FLOWERS: floral bracts 1.8- 2.5 mm long, 1.5-2.2 mm long, elliptic, costa thick, green, blade membranous, dark purple, margins hyaline, apex rounded; perianth 3-6 bristles; stamens 3, filaments ca. 2 mm long, anthers 1.1-1.3, apiculate, base cordate, yellow; stigmas 3. ACHENES: 1-1.5 mm long, 0.6-1 mm wide, obovate, trigonous, stylopodium triangular, body striate, dull white to cream-colored. HABITAT: Boggy spots in grasslands. 3700-4500 m. DISTRIBUTION: Ecuador to southern Argentina and Chile; Islas Juan Fernandez. Peru; Ancash, Cuzco, and Puno. H. N. P.: Alpamayo (9837), Ishinca (9476), Llanganuco on Maria Josef a trail (8861, 10508), Ranincuray (10398), Rurichinchay in Pachachaca (12639, 12642). This taxon may include E. boliviana Palla, E. brehmeriana Boeckeler, and E. nubigena C. B. Clarke ex Urban. Barros (1947) combined E. nubigena as a variety of E. albibracteata. I know of no monographic study of this genera since Svensen's work in the 1920s and '30s. The genus needs critical review employing modern techniques. PHENOLOGY: Found in flower from February until May; mature fruit seen in June. 183

USES: This species is an important forage plant. It is highly selected by young and adult alpacas during the dry season, and during the transition between the dry and wet seasons (Bryant and Farf&n, 1984). CONSERVATION STATUS: Probably very common in boggy habitats, but overlooked due to its small size.

Isslfinii R. Brown References: Beetle, 1944; Clapham et al., 1987; Cook et al., 1974; Haines and Lye, 1971; Koyama, 1958. HERBS, short-lived, from fibrous roots; stems photosynthetic, terete; plants small, often only a few cm tall; glabrous. LEAVES; basal, filiform, often canaliculate. INFLORESCENCE: psuedo lateral, a head of 1-several sessile spikes; inflorescence bract culm-like, erect, usually longer than inflorescence; spikes several-flowered. FLOWERS: bisexual; floral bracts spirally arranged; perianth wanting; stamens 1-2, filaments linear, flat tened, anthers linear; stigmas 2-3. ACHENES: obovate, trigonous, smooth or sculptured. The genus has about 60 species of cosmopolitan distribution; there are 2-3 Peruvian species.

Isolepla inundata R. Br. ("Scirpus inundatus (R. Br.) Poiret) HERB to 6 cm tall, from slender, creeping rhizome, this usually breaking off during collection; stems terete. LEAVES: 17-48 mm long, ca. 0.8 mm wide, sheath 7-10 mm long, closed, hyaline, blades filiform, spreading, apex blunt. INFLORESCENCE: a pseudo-lateral head of 2-3 spikes; peduncles 10-24 (-53) mm long; inflorescence bract 6- 11 mm long 0.6-1.5 mm wide at base, foliar, erect, blade flat or slightly involute, margin dilated and hyaline at base, apex rounded, bract longer than inflorescence; spikes 2.2-4.3 mm long, subtending bracts like inflorescence bracts. FLOWERS: floral bracts 1.4-2 mm long, (0.6-) 1-1.5 mm wide, ovate-acute, costa membranous, green, margins hyaline; perianth wanting; stamens 1, occasionally 3 in lower flowers, filaments 1.3-2 mm long, linear, flattened, anthers 0.4-0.6 mm long, elliptic, base cordate, apex apiculate; style 1- 1.7 mm long, stigmas 3. ACHENES: 0.7-0.9 mm long, 0.4-0.6 mm wide, strongly trigonous. HABITAT: Grows in seepy or boggy spots. 3900-4300 m. I

184

DISTRIBUTION: Costa Rica, Colombia and Venezuela to Chile and Argentina, New Zealand, Australia, and Malaysia. Peru: Ancash and Huanuco. H. N. P.: Cahuish (8743), Ranincuray (10399), and Rajucolta (12151). PHENOLOGY: Flowers from December to April. Sometimes the inflorescence becomes vegetatively proliferous (10399), and can become stolon-like with the flowers remaining attached and functional. CONSERVATION STATUS: Unknown. Given its small size this plant is readily overlooked in the field.

OreoboloDili T. Koyama & E. R. Guaglianone Reference: Koyama and Guaglianone, 1987. HERBS, perennial, erect from very short, creeping rhizomes. LEAVES: basal, sheath short, closed, blades filiform. INFLORESCENCE: a solitary, 3-6-flowered spike, sessile on peduncle apex; inflorescence bract leaf-like, caducous. FLOWERS: perfect, sessile; perianth of 6 scales in 2 whorls, free, persistent, outer scales obovate, 2-lobed, abaxial inner scales obovate, 2-lobed with an awn between lobes, adaxial scale flabellate; stamens 3; ovary obovate, style terminal, 3-fid, erect, narrow, stigmas papillate. ACHENE: obovate, trigonous-compressed, style base persistent as small apicule. Oreobolopsis is monotypic; its characters are those of the type species:

OreoboloBsIs temmllfera T. Koyama & Guaglian. HERB, 3.5-22 (-43.5) cm tall, forming bunches to 23 cm diam.; foliage and inflorescences erect; innovations subtended by hyaline, multinerved, 2-keeled prophyll, ca. 24 mm long, ca. 2.4 mm wide, oblong, apex rounded. LEAVES: 5.5-15 (-25) cm long, 0.7 mm wide or less, basal, linear, thickened, stiff, slightly arcuate, sheath 1-3 (-8) cm long, broadened, with several, prominent nerves, margins joined to form closed, chartaceous sheath, this splitting with age, joined above into a ligule ca. 0.5 mm long, bases somewhat persistent in age, blade 2-12 (-17) mm long, canaliculate basally, laterally compressed distally, margins scabrous, apex rounded. INFLORESCENCE: terminal, a solitary, sessile spike; inflorescence bract 5-22 mm long, erect, surpassing the spike, leaf-like, base widely ovate, papery margined, 1-3.7 mm wide, joined above into small ligule; peduncles 3-22 (-43.5) cm long, stiff, slightly laterally compressed, often little exceeding leaves; spike (2.2-) 5-8.5 mm long, sessile, elliptic, 3-6-flowered, lowermost bract 2.4-6.7 mm long, 1.2-3 mm wide, ovate, costa foliaceous, prolonged into short mucronate tip, margins 185

papery, joined distally into small ligule. FLOWERS: floral bracts 5-8, lowermost 2-3 sterile, uppermost with a flower or a rudiment, 2.9-4.5 mm long, 2-4 mm wide, ovate, papery, light brown, costa short, often half length of bract, thickened, green, margins erose, apex acute to erose; perianth scales chaffy, light brown, outer scales 1.2-2 mm long, 0.6-1.3 mm wide, apex bilobed to 0.5 mm deep, sometimes with a rudimentary arista between the lobes, sometimes entire, margins erose to ciliate, abaxial inner scales 0.5-1.3 mm long, 0.3-1 mm wide, arista 0.2-0.7 mm, between lobes, scale sometimes irregularly shaped, margin erose, adaxial scale 0.6-0.8 mm long, 0.4-0.7 mm wide, irregularly 2-3 lobed, margin erose; stamens 3, opposite outer scales, filaments 3-7.5 mm long, filiform, anthers basifixed, 1.2-2.1 mm long, 0.2-0.5 mm wide, linear, base apiculate, tip apiculate, brownish; style 2.2-5.5 mm long, purple. ACHENE: 1.7-2 mm long, 1.3-1.5 mm wide, brown, style base persistent as small apicule, 0.1 mm. HABITAT: Grasslands, sometimes Calamagrostis-Stipa hans~meyerii dominated, on soils of morainal parent material. 3900-4400 m. DISTRIBUTION: Peru and Bolivia. Peru: Ancash and Cajamarca. H. N. P.: Alpamayo- Cashapampa trail between Huiscash and Mirador (9987), Huaripampa (9182), Ranincuray (9124), and Rurichinchay at Pucaraju (12696). PHENOLOGY: Flowering has been observed from November until July, and fruiting from March to July. ^ CONSERVATION STATUS: Since this plant was confused with Baeothryon rigida in the field, its status and distribution are not well known.

Oreobolus R. Brown Reference: Seberg, 1988. HERBS, perennial, from caespitose rhizomes; forms cushions and rings; stems terete; glabrous. LEAVES: basal, thickened, arcuate, base ovate, wider than blade, open, clasping the stem, persistent, blades linear, canaliculate below, flattened above, margins scabrous, often inrolling below when dried. INFLORESCENCE: a terminal, pedunculate, simple, few-flowered spike, or compound with several pedicellate spikes; inflorescence bract erect, foliar. FLOWERS: terminal flower of spike bisexual, lower flowers staminate; floral bracts spirally arranged; perianth segments 6, in 2 whorls, united basally, stiff, incurving, elliptic to ovate, persistent; stamens 3 in perfect flowers, 1 in staminate flowers, filaments linear, flattened; style terminal, stigmas 3. ACHENE: obconic-truncate or elliptic, apex with a swollen rim. * •»

186

Oreobolus is a genus of austral distribution (van Balgooy, 1971), occurring in the Celebes Islands, Malacca Peninsula, Borneo, New Guinea, Oceania, Hawaii, Australia including Tasmania, New Zealand and the shelf islands, and Central and South America (Cleef, 1978; Seberg, 1988; van Royen, 1979b). In South America, the genus is found principally the Andes, although there are outlying locations in the Guyana Highlands. It has 14 species of which 4 are South American (Seberg, 1988). Oreobolus ecuadorensis, O. obtusangulatus, and O. venezuelensis are known from Peru. HABITAT: In South America the species of Oreobolus are important components of bogs at middle and high elevations in the tropics and the low latitudes towards the southern tip of the continent. POLLINATION: The plants are probably wind pollinated. ANATOMY: Seberg (1986) and Metcalfe (1971) discuss the presence of a pseudopetiole between the leaf sheath and blade. The term pseudopetiole, at least as it is used with respect to the grasses refers to a distinct anatomical structure between the leaf sheath and blade that is like a petiole in external form and internal structure. The lower part of the lamina is thicker than the distal portion, having a greater amount of ground tissue. Some illustrations in Metcalfe (1971:397) show a lumen in the center of the ground tissue. On drying, the ground tissue collapses and the margins roll inward toward the costa, giving the impression of a constriction. If the dried specimen is rehydrated, the leaf flattens out and the constriction disappears. In my view, the proported pseudopetiole is an artifact of drying.

Oreobolus obtusangulatus Gaudich. HERB, from caespitose rhizome, forming clones 15-40 cm diam., often dying in the center; glabrous. LEAVES: 15.5-28 mm long, erect, stiff, arcuate, lustrous, base 5-9 mm long, ovate, papery, brown, eligulate, reddish, blades 9-23 mm long, 3.5-7 mm wide, thickened, dorsiventrally flattened, 3-nerved basally, margins scabrous, lower 1/3-1/2 inrolling when dry, pseudopetiole 4-10 mm long, apex acute; prophylls are interspersed with the leaves. INFLORESCENCE: a terminal spike; peduncle exserted with maturity, 15-36 mm long; inflorescence bract 6.6-7 mm long, 2-2.4 mm wide, erect, stiff, foliar, flattened above, soon caducous. FLOWERS: floral bracts 4-4.8 mm long, 2-2.4 mm wide, ovate to elliptic, stiff, apex acute, margins minutely ciliate, purple; perianth 1.6-1.9 mm long, 0.3-0.6 mm wide, subequal, triangular, stiff, margin ciliate, the inner segments sometimes minutely, aristate, segments long persistent; stamen filaments 3-4 mm long. 187

anthers ca. 2 mm long, yellow; style 3.4-4.2 mm long, stigmas about 1/2 length of style, spreading, purple. ACHENE: 1-1.8 mm long, 0.8- 1.3 mm diam., terete to triquitous, apex rugose-ciliate, purple. habitat: This is a common species, often the dominant, in bogs, seeps, streamsides and stream channels. 3700-4480 m. DISTRIBUTION: Colombia to Tierra del Fuego, Islas Malvinas, Isla de los Estados, and Islas Juan Fernandez. Peru: Ancash. H. N. P.: Alpamayo-Cashapampa trail at Hatuncocha (9979), Huaripampa at Morococha, Ishinca (11171), Parôn (11S3S), Rio Pachacoto (11437), and Rurichinchay in Quebrada Pachachaca (12554). Seberg (1988) recognizes two subspecies, subsp. obtusangulatus from southern South America including the Islas Juan Fernandez and Islas Malvinas, and subsp. unispicatus O. Seberg from Colombia, Ecuador and Peru. The material from the Park is like subsp. unispicatus, although the achene measurements overlap those reported for the two subspecies. PHENOLOGY: Flowering has been observed in July, and fruiting in September and January. The developing flowers and fruits are hidden within the foliage, and are exserted only in the final stages of fructification. Without careful observation, the plants can be readily passed over as sterile. CONSERVATION STATUS: Common.

Phvllosclrnm C. B. Clarke References: Barros, 1947; Clarke, 1908, 1909; Goetghebeur, 1986. HERBS, dwarf perennials, 10 cm tall or less, from creeping rhizomes, often forming extensive cushions. LEAVES; basal and sub-rosetteform or cauline, then usually crowded at base of stem, blades short to elongate, canaliculate. INFLORESCENCE: terminal, pseudolateral in a few species; peduncles short to well exserted; inflorescence bract wanting, or developed in some species; spikes 1-7, crowded into a head. FLOWERS; bisexual; floral bracts imbricate, 6-12 per spike, lowermost 2-3 often sterile, membranous to chaffy; perianth (0) 1-6 long, retrorsely barbed bristles; stamens 3, filaments linear, anthers linear, yellow; style base thickened, stigmas (2-) 3. ACHENE; elliptic to obovate, trigonous, plano-convex, apex often with a small beak. Phyiioscirpus is a small, poorly known genus of ca. 8 species. Most of the species are Andean, distributed from Ecuador to Chile and Argentina. The Atacama Desert appears to be the center of diversity. Phyiioscirpus nevadensis (S. Watson) Goetgh., comb, nov. provis. (Goetghebeur, 1986) has a disjunct distribution in the western United States 188

and Canada and Argentina. Barros (1954) calls attention to the description of Scirpus lenticularis (Spreng.) Kunth in Boeckeler (1870). Barros implies that Scirpus lenticularis is likely related to Scirpus spegazzinianus Barros, a taxon that probably belongs in Phylloscirpus. The distribution reported for Scirpus lenticularis is Cape of Good Hope, Reunion Island and Tasmania. When fully studied, genus may be found to have an austral distribution pattern. TAXONOMY; This group urgently needs systematic study; there are about 16 published names that must be considered in the course of a revision.

Key to the species of Phvifoscirnug la. Perianth of retrorsely barbed bristles; inflorescence compound, 3 sub-sessile spikes. P. deserticola lb. Perianth wanting; inflorescence a simple, solitary spike. 2. 2a. Achenes 1.1-1.7 x 0.8-1.1 mm, circular to obovate; seed coat loose from embryo. P. sp. ('Scirpus aff. atacamensis) 2b. Achenes 0.8-0.9 x ca. 0.5 mm, obovate; seed coat adnate with embryo. P. sp. ("Scirpus aff. hieronymi)

Phvllosclrnus sp. indet. {Scirpus aff. atacamensis Boeckeler) HERB, to 3 cm tall, clones forming extensive cushions; glabrous. LEAVES: 4.1-7.3 mm long, 0.5-0.7 mm wide, cauline, imbricate, sheath closed, ligule ca. 0.6 mm, delicate, blades flattened, ascending, thickened, several-nerved, margins entire, apex blunt, caloused, more or less dorsi-ventrally compressed. INFLORESCENCE; a terminal, simple spike, 1.7-2.5 mm long, 2.5-2.6 mm wide, globose, 6-9-flowered; peduncle 15-20 mm long, 0.2-0.3 mm diam.; inflorescence bract 2.1-2.7 mm long, 2-2.5 mm wide, very broadly ovate, papery, apex rounded and mucronate, light brown with rust colored punctations. FLOWERS: floral bracts 0.9-1.5 mm long, 1.3-2.1 mm wide, very broadly ovate, papery, light brown with rust colored punctations, apex rounded; stamens not seen; style 1.3- 2.4 mm long including stigmas, stigmas (2-) 3. ACHENES; 1.1-1.7 mm long, 0.8-1.1 mm wide, circular becoming obovate, lenticular to sub-trigonous, seed coat collaniculose, loose from the ovule or seed, light brown. 189

The lowermost 2-3 flowers have 2 lateral appendages that are presumed to be stamen filaments. If this is correct, then the lower flowers are perfect, and the upper flowers female. HABITAT: Growing in saturated soil in a Distichia muscoides-Plantago rigida bog. 4500 m. DISTRIBUTION: Not adequately known. Peru: Ancash and Huanuco. H. N. P.: Queshque (11828 pp.). A few individuals of this species are included with a specimen of Distichia muscoides from Yanashallas in Huanuco (MacBride 2484, F), at 4848 m. PHENOLOGY: In late fruiting stage in mid-March. COMMENT: In a vegetative state, it is impossible to distinguish this from S. aff. hieronymi. CONSERVATION STATUS: Rare. Perhaps overlooked during field studies.

PhvllosclrnuB deaertlcol» (Philippi) D. N. Smith, comb. ined. (Scirpus deserticola Philippi) HERBS, 3-5 cm tall, from branched, creeping rhizomes, branches caespitose; forming cushions (3-) 15-40 cm diam., these can form extensive, interconnected clones. LEAVES: cauline, imbricate, base widened, sheathing, ligule wanting, blades (3-) 5-9 mm long, 0.4- 0.5 mm wide, linear-attenuate, folded to rolled, ascending, margins serrulate, apex blunt, calluosed; a few leaves become long-etiolated from the lower nodes; upper leaves often sheath the inflorescence. INFLORESCENCE: terminal, compound, spiciform, subglobose; peduncles 10-40 mm long on exsertion, inflorescence bract 3.5-5.4 mm long, 2-3.3 (-4) mm wide, ovate-mucronate, papyraceous, mid-nerve membranous, maturing brown with red punctations; spikes 4-6 mm long, 1 central spike with 1-3 lateral spikes, the central spikes 10-20-flowered, the laterals 6-10-flowered, subtending bracts 3.2-3.5 mm long, 1.8-3 mm wide, like floral bracts in texture, prophyll ca. 2 mm long, ca. 2 mm wide, bilobed, chaffy. FLOWERS: floral bracts 2-2.6 mm long, 1.5-2.4 mm wide, elliptic, chaffy, costa thickened, apex acute to mucronate or rounded, brown with red punctations; stamen 3, filaments 3.5-6 mm long, filiform, long exserted, anthers 1.1-2.1 mm long, base cordate, apiculate, yellow; gynoeclum subtended by 3-6 unequally developed bristles, ca. twice as long as the achenes, filiform, retrorsely barbed, style ca. 2.3 mm long, deciduous, stigmas 3. ACHENES: 0.9-1.5 mm long, 0.5-0.7 mm wide, trigonous, gray. 190

HABITAT: Grows in seeps and bogs in saturated soil above the water line. It is frequently found in bogs dominated by Oreobolus obtusangulatus, or can occasionally be the dominant species in a bog. 4000-4700 m. DISTRIBUTION: Peru to Chile. Peru: Ancash and Puno. H. N. P.: Alpamayo (9725), Honda at Vinoyapampa (11564), Huaripampa at Morococha (9212), Ishinca (9471), Rajucolta (12154), Santa Cruz at Taullipampa (9322), and Ulta (11363). PHENOLOGY: Flowering occurs from October through July, and fruiting in October through February. CONSERVATION STATUS: Probably common, but overlooked due to its size.

Phvllosclrnus sp. indet. (Scirpus aff. hieronymi Boeckeler) HERB, to 3 cm tall, clones forming extensive cushions; glabrous. LEAVES: 4.1-7.3 mm long, 0.5-0.7 mm wide, cauline, imbricate, sheath closed, ligule ca. 0.6 mm, delicate, blades flattened, ascending, thickened, several-nerved, margins entire, apex blunt, caloused, more or less dorsi-ventrally compressed. INFLORESCENCE: a simple terminal spike, 4.2-6 mm long, 0.7-1 mm wide, 2-3-flowered, elongate; peduncle ca. 26 mm long, thickened at middle, ca. 1 mm diam., tapering toward both ends; inflorescence bract 3.8- 3.9 mm long, 1.6-1.9 mm wide, elliptic, chartaceous, 3-nerved, the laterals obscure, apex acute to mucronate, brown. FLOWERS: floral bracts 2.2-3.6 mm long, 1.2-1.5 mm wide, elliptic, chartaceous, 3-nerved, the laterals obscure, apex rounded, brown; stamens 2, filaments ca. 3.6 mm long, linear, anthers 0.7-0.8 mm long; style 2.8-3.2 mm long, insertion slightly eccentric, 2-3-fid on plants of same population, stigmas about 1/2 length of style. ACHENES: 0.8-0.9 mm long, ca. 0.5 mm wide, obovate, lenticular, slight apical rim about style insertion. HABITAT: Growing in saturated soil in a Distichia muscoides-Plantago rigida bog. 4500 m. A collection from Lima Department (Amstutz 107, MO) was collected in a similar habitat at 4700 m. DISTRIBUTION: Not adequately known. Peru: Ancash and Lima. H. N. P.: Queshque (11828 pp.). PHENOLOGY: Late flowering and fruiting in mid March. The Amstutz collection was flowering and fruiting in July. 191

ADDITIONAL MATERIAL: The Amstutz collection had 3 styles in all examined flowers, and the achenes were 1.6-1.8 mm long, 0.7-0.8 mm wide, wide obovate to elliptic, and lenticular to rounded or sub-trigonous. COMMENT: In a vegetative state, it is impossible to distinguish this from S. aff. atacamensis CONSERVATION STATUS: Rare. Perhaps overlooked during field studies.

SchoenoBlectus (H. G. L. Reichenbach) Palla References: Clapham et ai., 1987; Cook et al., 1974; Koyama, 1963; Lye, 1971; Weber, 1987. HERBS, coarse, tall perennials from stout rhizomes; stems photosynthetic, trigonous or terete; glabrous. LEAVES: sheaths coriaceous, blades much reduced or wanting. INFLORESCENCES: terminal or pseudo-lateral, head-like or branched; inflorescence bract short, foliar, dry, and less conspicuous than the inflorescence or long, well- developed, stem-like, erect, appearing to be a continuation of the stem, then the inflorescence pseudo-lateral; spikes many-flowered, several to many per inflorescence. FLOWERS: bisexual; floral bracts spirally arranged; perianth (0-) 6 fimbriate scales; stamens 3, filaments linear; stigmas (2-) 3. ACHENES: trigonous or lenticular. A genus of about SO species, it has a cosmopolitan distribution.

SchoenoBlectus califoinlcus (C. Meyer) Sojak ('^Scirpus californicus (C. Meyer) Steudel subsp. californicus var. californicus) Reference: Koyama, 1963. HERB, 1.25 m or more tall, emergent aquatic, rhizome system extensive, rhizomes very thick, can become floating; forming extensive clones; stems 3-angled, erect, solid, with pith spongy, flexuous, glaucous; glabrous. LEAVES: sheaths stiff, reddish, bladeless. INFLORESCENCES: a terminal, condensed panicle, 6.5-7 cm long, branches to 45 cm long, ca. 1 mm wide, flattened, scabrous on keels, erect to spreading; inflorescence bracts 40-50 mm long, ca. 1.2 mm wide, narrowly triangular, dry, membranous, erect, sheathing stem tip, round-attenuate; spikes many per inflorescence, many-flowered, 4.5-8.5 mm long, subtending bract 3.8-4.5 mm long, 2.4-3.2 mm wide, ovate, 1-nerved, papyraceous, mucronate, reddish brown. FLOWERS: floral bracts 3.8-4.5 mm long, 2.4-3.2 mm wide, ovate, 1-nerved, papyraceous, mucronate, reddish brown; perianth 2 fimbriate scales, 2.5- 2.7 mm long; stamens 3, filaments 3 mm long, flattened, linear, anthers 2.1-2.3 mm long. 192 base slightly cordate, apex a blunt callus with tuft of trichomes, yellow; style 3.4-4 mm long, stigmas 2. ACHENE: obovate, lenticular with a conical apex. HABITAT: At shores of warmer lakes. 3850 m. DISTRUBUTION: The species is found from the southern United States through Central America, and along the Andes and the coast of southern Brazil and Uruguay to Chile and Argentina, including the Islas Malvinas and Isla de Pasuca. There are two subspecies; this variety is found intermittently from the southern United States through Central America, and along the Andes to northern Chile and Argentina. Peru: Ancash and La Libertad; subspecies totora (Kunth) Koyama is found in central and southern Peru at higher elevations than our subspecies. H. N. P.: Llanganuco (9424) and, possibly, Santa Cruz. PHENOLOGY: Flowering is seen in January. USES: Schoenoplectm cali/ornicus, called totora throughout Peru, is important for construction of "reed" boats, balsas, especially at Lago Titicaca, near Puno, and Huanchaco, near Trujillo. An entire culture, los Uros, depends on totora, as they live on Lago Titicaca on artificial, floating islands made of the "reed," and their livlihood is associated with fishing and the balsas de totora. Heiser (1979) gives detailed information on the uses of this important plant in Ecuador and Peru. CONSERVATION STATUS: Although abundant in other parts of Peru, this species is rare in the Park. 193

DIOSCOREACEAE R. Brown LILIIDAE: LILIALES HERBS, usually twining; from a tuber or tuberous roots. LEAVES: alternate, rarely opposite, petiole commonly twisted or jointed at the base; blade broad, base often cordate, entire to palmately lobed to compound, primary veins arcuate-convergent. INFLORESCENCE: axillary racemes, spikes, panicles, or cymes. FLOWERS: small, unisexual and plants dioecious, regular, epigynous; perianth members 6, subequal, petaloid, usually briefly connate basally, campanulate; nectaries present; stamens 6 in 2 whorls, the inner whorl sometimes reduced to staminodes or wanting, attached to base of perianth, filaments free or basally connate, anthers basifixed, connective often broadened; gynoecium 3-carpelIate, compound, inferior, ovary trilocular, placentation axile, styles free or basally connate, stigmas free, ovules 2 per locule. FRUIT capsular, often triangular and winged; seeds mostly winged. This family of 6 genera and ca.. 630 species is mainly tropical and subtropical. Dioscorea is the only genus of the family represented in South America (Dahlgren et al., 1985). REPRODUCTION: The plants can be reproduced ^egetatively using the tubers, and some species produce bulbils, which can also be used for vegetative propagation. USES: Yams {Dioscorea spp.) are the second most important tuber crop in the world (Simpson and Conner-Ogorzaly, 1986). Different species of Dioscorea have been domesticated in Africa, Asia and South America. Two Central American species are grown commercially to produce the steroid precursors used in the manufacture of oral contraceptives, hormonal regulators, and cortisones and hydrocortisones (Simpson and Conner-Ogorzaly, 1986).

Dioscorea Linnaeus Reference: Knuth, 1924. HERBS: stems usually vining, sometimes erect. LEAVES: alternate, rarely opposite, simple and entire to lobed or compound, base often cordate. INFLORESCENCE: spikes, racemes, panicles, cymes, or solitary. FLOWERS: perianth rotate, campanulate, or tubuliform; stamlnate flowers composed of perianth, stamens, and a vestigial ovary, carpellate flowers of perianth, (0-) 3-6 staminodes, and an ovary; stamens 3-6, filaments free or united into a column, anthers introrse. CAPSULE: triquetrous, 3-winged; seeds flattened, winged. 194

Dioscorea, the largest genus in this family, has about 600 species; most are Neotropical. There are more than SO species in Peru. The species are usually found in light gaps and clearings, riverine sites, and brushfields from the lowlands to high elevations. I thank F. Ayala (AMAZ) for his help with this genus.

Key to the species of Dioscorea la. Plants vining; flowers dark purple; growing in bouldery spots. D. larecajensis lb. Plants prostrate or immersed in soil; flowers green or yellow; growing in grasslands or grassland - shrubland. D. ancachsensis

Dlogcorea ancachsensls Knuth HERB, prostrate or immersed in soil, branches to 9.5 cm long; tuber 1.2-2 cm diam.; glabrous. LEAVES: alternate, petioles 2.7-12 mm long, blades 6.1-10.4 mm long, 5.9- 12.2 mm wide, base cordate, broadly ovate, thickened with glandular dots, nerves obscure, apex broadly acute, often mucronate. INFLORESCENCES: cymes, tending to be subumbellate, open; peduncles 4.5-8 mm long. FLOWERS: pedicels 3.5-7 mm long; floral bracts 1-1.5 mm long, ovate, membranous; perianth members of staminate flowers 1.2-1.3 mm long, 0.6-0.8 mm wide, basally connate, rotate, elliptic, 1-nerved, apex rounded, pale green, yellow-green, or yellow, those of the carpellate flowers not seen; stamens 6, filaments 0.3-0.5 mm long, free, anthers ca.. 0.3 mm long, yellow; gynoeclum not seen. CAPSULE: ca.. 10.5 cm long, ca.. 5 mm wide at maturity. HABITAT: Grasslands and grassland - shrubland. 3800-4250 m. DISTRIBUTION: Peru: Ancash. H. N. P.: Alpamayo (9840), Huaripampa at Que. Paria (9163), Ishinca (9548), Rajucolta (12185), and Shallap (9659). The type of this species is from Mina Huaura in the northeastern corner of Ancash. As currently understood, the species is considered to be endemic to the Department of Ancash, however, considering its small size it is easily overlooked in the field. It probably occurs in the adjoining departments but, as yet, has not been collected there. PHENOLOGY: Flowering specimens have been collected from January to March, and friuting specimens from February to April. The foliage of this species may dry up completely the dry season, and the plant may pass the dry season in dormancy. Plants of this species are often attacked by a rust which affects the foliage and flowering parts. 195

CONSERVATION STATUS: Uncommon? It is likely that this species is more common than herbarium collections indicate, and is neither noticed nor collected due to its small size.

Dioscorea larecaiensis Uline ex Knuth VINE, from tuber, ca.. 7 cm long, ca.. 6 cm wide, shape irregular, taproot-like, woody; stem to 60 cm long or more. LEAVES: petiole 1.6-5.4 cm long, blade 4.4-10.4 cm long, 2.1-6 cm wide, base deeply cordate, ovate to broadly lanceolate, 11-13 prominent, arcuate nerves, primary and secondary nerves with triangular, flattened trichomes, apex attenuate to cuspidate. INFLORESCENCES: racemes, 3.6-10 cm long, many-flowered; peduncles 0.5-1.2 cm long. FLOWERS: pedicels 3-4 mm long; floral bracts 1-2 mm long, ovate, membranous, purple; perianth members of staminate flowers 2.3-2.9 mm long, 1-1.6 mm wide, basally connate, rotate, wide ovate to elliptic, apex rounded to acute, dark purple, those of carpellate flowers not seen; stamens 6, filaments ca.. 0.3 mm long, briefly connate basally, anther ca. 0.2 mm long, yellow; gynoecium not seen. CAPSULE: ca.. 2.2 cm long, ca. 1.2 cm wide at maturity, marcescent corolla remains attached, greenish brown. HABITAT: Growing among boulders in arid shrubland. 3350-3900 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash and Cuzco. H. N. P.: Auquispuquio (11996), Parôn (8942), and Santa Cruz (9352). PHENOLOGY: Flowering in January and April; fruiting in April. CONSERVATION STATUS: Rare. 196

HYDROCHARITACEAE A. L. de Jussieu ALISMATIDAE: HYDROCHARITALES HERBS, perennial, aquatic; rooted on the bottom, rhizomatous or stoloniferous. LEAVES: cauline, alternate, opposite to whorled, base often sheathing or expanded, with intravaginal scales, blade, if differentiated, narrow, sessile, elongate, margins often with prickle trichomes, plants otherwise glabrous. INFLORESCENCES: compact, few- flowered cymes (especially the staminate flowers) or a single flower (the carpellate flowers), subtended by a spathe composed of 2 bracts. FLOWERS: small to large; unisexual and the plants dioecious or monoecious, or perfect, regular, epigynous; sepals 3, free, green; petals 3, free, colored; perianth attached to ovary apex or to an hypanthium; stamens 3-9, variously arranged, some reduced to staminodes; gynoecium 3- carpellate, compound, inferior, unilocular ovary, placentation parietal, styles 3, bilobed or bifid, stigmas dry, ovules many. FRUIT: berry-like, fleshy, splitting irregularly. Cosmopolitan in distribution, this family has ca. IS genera and ca. 100 species. Only Elodea is found in Peru. HABITAT: Growing submerged in ponds and lakes at high elevations. POLLINATION: By a complex water-borne system: the carpellate flowers are often long pedunculate and floating at the surface of the water. The staminate flowers develop submerged, and at maturity either break loose from the plant and rise to the suface where they open, or the pollen grains are shed below the water and rise to the surface (Dahlgren et al., 1985). USES: Aquarium plants; agressive weeds in aquatic habitats.

Ehuka Jaume Saint-Hilaire Reference: Cook & Urmi-KOnig, 1985. HERBS, submerged aquatic perennials; rooted in soil; stems can be quite long, erect to somewhat rhizomatous. LEAVES: cauline, verticellate; sessile. TURIONS (vegetative propagules) are produced in the leaf axils of some species. INFLORESCENCES: cymes subtended by 2 bracts (spathes); axillary. FLOWERS: unisexual, the plants dioecious or monoecious, or, rarely, flowers bisexual when the stamens of a normally female flower are fertile; perianth of both sexes alike; sepals 3, free, green; petals 3, free, delicate, often pale white with purplish cast; staminate flowers with 7-9 stamens, usually lacking vestigial ovary; carpellate flowers with 3 staminodes, ovary with 3, entire or bifid styles. FRUIT: berry-like, splitting irregularly. 197

A genus native to temperate North and South America, it has become an aggressive weed in aquatic habitats throughout the world. There are S species, only E. potamogeton is found in the Andes. POLLINATION: Although the pollen can be transported by the wind, water is the vector, as the pollen grains float to the stigmas on the water surface.

Elodea notamogeton (Bertero) Espin. HERB, stems smooth, glabrous, greater than 20 cm long; branching common; lower internodes elongate, distal ones short, congested. LEAVES: 5-11 mm long, 1.7-3 mm wide, elliptic, blade thin, appears 1-2 cells thick, costa noticeable at base, margins entire or with spinose trichomes, apex acute to rounded, olive green; in verticels of (2-) 3 (-4); leaves and stems often have abundant surficial cristals on dried specimens. TURIONS well developed, 6-10.2 mm long. Only sterile material has been seen from the Huascarân National Park. The inflorescences, flowers and fruits are as described in the generic description. Cook and Urmi-Kdnig (1985) found the sepals and petals of the staminate flowers to measure 5.3- 6.5 mm and 4.5-7.2 mm long, respectively. The carpellate flowers are smaller. HABITAT: Pools in grasslands, probably also occurring in lakes and streams throughout the Park. DISTRIBUTION: Ecuador to Chile. Peru: Ancash, Ayacucho, Cuzco, and Puno. H. N. P.: Honda (11698); probably widespread, but overlooked during the field collections. PHENOLOGY: Reported to flower and fruit all year, (Cook & Urmi-Kônig, 1985). CONSERVATION STATUS: Probably common. 198

IRIDACEAE A. L. de Jussieu LILIIDAE: LILIALES Reference: Innes, 1985. HERBS perennial from rhizomes, corms and bulbs, or annual (JSisyrhinchium spp.). LEAVES: distichous; base open and sheathing, or sheathing, often equitant; blades linear to lanceolate, flat or plicate. INFLORESCENCE: cymose or solitary, simple or compound, usually terminal, composed of rhipidia; pedunculate. FLOWERS: showy, ephemeral, often pedicellate, subtended by a bract; perfect, actinomorphic or zygomorphic, epigynous; perianth segments 6, petaloid, equal or unequal, free, nectaries usually present, except in Sisyrinchiunr, stamens 3, opposite the sepals, filaments free or connate below, anthers basifixed; gynoecium compound, inferior, tricarpellate ovary, placentation axile, style terminal, 3-lobed, branches simple or divided, sometimes expanded and petaloid, stigmas terminal, in Iris on the ventral side of the style branches, dry, papillate, ovules 1-many. FRUIT: a loculicidal capsule. The family is of cosmopolitan distribution, but best developed in the Southern Hemisphere, being most abundant and diverse in Africa; 80 genera and ca. 1500 species. There are 7 genera known from Peru. HABITAT: Coastal deserts, arid regions, and high elevation grasslands. POLLINATION: By insects and birds. USES: Saffron, a culinary seasoning and coloring agent, is the red stigma of Crocus sativus. Many Irises are grown for ornament. RHIPIDIA: The basic unit of the inflorescence is a rhipidium, a condensed scorpoid cyme composed of a sterile subtending bract, an axis, 1-several floral bracts, and flowers. The subtending bract and first floral bract are foliaceous or membranous, and enclose the remaining floral bracts and flowers. Traditionally the foliaceous bracts are called spathes. Careful examination shows that the lower spathe does not subtend a flower; all other bracts within the rhipidium are flower-bearing. The enclosed floral bracts become progressively smaller and prophyll-Iike. In many species the texture of the floral bracts changes from membranous and multi-nerved in the outermost to hyaline and 2-nerved within. In other species, all the enclosed bracts are hyaline and 2-few- nerved. COLLECTING TECHNIQUES: The flowers of this family are very delicate, and last but a few hours. They are very difficult to maintain in a fresh state after picking. As the flowers wilt, the petals crumple or twist about each other, and adhere to each other 199

tightly. Dissection of dried, wilted flowers is very difficult. It is recommendable to press the specimens in the field. Some collectors put sheets of tissue paper on either side of the flower to prevent it from adhering to the coarse pressing paper. If there is adequate material, preserve some flowers in liquid. In all cases, complete field notes should be made, and a photograph taken. I thank Drs. Peter Goldblatt and James Henrich (MO) for discussions of this family and revision of the manuscript.

Key to the genera and species of the Irldaceae la. Plants from bulbs; leaves with numerous longitudinal folds (plicate). Tigridia philippiana lb. Plants from rhizomes, or fibrous or tuberous roots; leaves flat or terete. 2. 2a. Plants growing in dense bunches, 5-20 cm diam.; plants usually 6-10 dm tall; foliage strongly flattened, stiff, erect; inflorescence a racemose panicle or a solitary, pedunculate rhipidium; flowers yellow or white. 3. 2b. Plants solitary or, at most, in small tufts; plants mostly less than 3 dm tall; foliage flat or terete, not stiff, erect; inflorescence a solitary, pedunculate rhipidium or a compound cyme, never a racemose panicle; flowers yellow, blue, purple, or white. 4. 3a. Plants finely puberulent; inflorescence a racemose panicle; flowers white; flowers and fruits sessile, little exserted from the spathes. Orthrosanthus occissapungus 3b. Plants glabrous; inflorescence a solitary, pedunculate rhipidium; flowers white; flowers and fruits pedicellate, exserted from the spathes. Sisyrinchium praealtum 4a. Leaves and stems terete; flowers yellow, purple, or white. S. 4b. Leaves and stems flattened; flowers yellow or blue. 7. Sa. Leaf bases long persistent as a thick, fibrous sheath at base of plant; plants tufted; inflorescence a solitary, pedunculate rhipidium; flowers yellow. 6. 5b. Leaf bases shortly persistent, never developing a thick sheath; plants usually solitary; inflorescence a compound, 2- branched cyme; flowers purple or white. ^ Sisyrinchium junceum 6a. Leaves and peduncles terete; capsules 12-18 mm long. Sisyrinchium trinerve 6b. Leaves and peduncles bifacial, flattened; capsules 3-5 mm long. Sisyrinchium jamesonii 200

7a. Leaves and peduncles stiffly arcuate, down-curving; inflorescence a solitary, pedunculate rhipidium; flowers yellow. Sisyrinchium brevipes 7b. Leaves and peduncles erect; inflorescence a solitary, pedunculate rhipidium, or a compound, 2-branched cyme; flowers yellow or blue. 8. 8a. Leaves strongly equitant, in a fan-shaped array; inflorescence a solitary, pedunculate rhipidium; flowers yellow; plants of boggy or marshy habitats. Sisyrinchium tinctorium 8b. Leaves weakly equitant, not conspicuously fan-arrayed; inflorescence a compound, 2-branched cyme; flowers blue; plants of seasonally arid habitats. Sisyrinchium aff. chilense

OrthrosanthuB Sweet References: Henrich and Goldblatt, 1987a; Standley and Steyermark, 1952. HERBS, coarse perennials, erect from rhizomes. LEAVES: erect to stiffly spreading, mostly basal, equitant, linear, stiff, flattened. INFLORESCENCES: panicles of rhipidia, compound, terminal, pedunclate. FLOWERS: actinomorphic; ephemeral, white or blue; perianth segments subequal, free, spreading; stamen filaments free or united basally, anthers erect; gynoeclum: ovary enclosed by spathes, style branches entire, alternate with stamens, ovules many. CAPSULES: elliptical; seeds globose. Orthrosanthus is of austral origin. It is found in Australia and along the Andes from northern Argentina to Mexico. The genus has 9 species; two are found in Peru.

Orthrosanthus ocdssanungua (Ruiz Lôpez ex Klatt) Diels HERBS, erect, 60-85 cm tall, rhizomatous, forming clones 5-13 cm diam.; puberulent. LEAVES: 46-52 cm long, 3-5 mm wide, many per plant, linear, bases sheathing, equitant, blades flat, strongly ribbed, stiff, apices attenuate; old leaves persist, not forming fibrous tunic at base of plant. INFLORESCENCES: compound; paniculate, superficially racemose, 10-28 cm long, branched at lower nodes; peduncles 42-73 cm long, elliptic in cross-section, not winged, with several leaf-like bracts; inflorescence bracts foliose, 10 cm long or less, margins briefly united basally; second order peduncles 1.4-4.2 cm long, subtended by a prophyll; rhipidia 1-flowered, spathes subequal, 13-25 mm long, 7-8 mm wide, ovate to elliptic, membranous, multi-nerved, apex acute. 201

FLOWERS: sessile; perianth segments ca. 19 mm long, ca. S mm wide, elliptic, carnose, white; stamen filaments ca. 6.5 mm long, united for lower 1/3, anthers ca. 4.5 mm long, yellow; gynoeclum: style ca. 7 mm long, branches free ca. 3.5 mm, ca. 0.5 mm wide, flat, immature ovary enclosed in spathes, puberulent. CAPSULE: 22-25 mm long, 5-6 mm diam., elliptic, exceeding spathes with age, glabrous, green to golden brown. HABITAT: Dry shrubland-grassland, often on morainal soils. 3880-4300 m. DISTRIBUTION: Peru to northern Argentina. Peru: Ancash, Cajamarca, Cuzco, Huancavelica, Huanuco, La Libertad, and Lambayeque. H. N. P.: Ishinca (9600) and Ranincuray (9088). PHENOLOGY: Flowering in January and February, and fruiting in February. CONSERVATION STATUS: Uncommon, but not endangered.

Sisvrlnchlmg Linnaeus References: Henrich and Goldblatt, 1987b; Standley and Steyermark, 1952. HERBS, erect, perennials from fibrous roots, often thickened, rhizomes sometimes developed, a few annual species; glabrous. LEAVES: erect, often strongly equitant, stiff, linear, flattened or terete, mostly basal, bases often persistent as weathered fibrous tunics. INFLORESCENCE: cymes, simple or compound, then second order peduncles subtended by a bract; terminal; peduncles flattened and winged, terete, or elliptic in cross-section; inflorescence bract foHaceous, often stout and erect, appearing to be a continuation of the peduncle, then inflorescence pseudo-lateral. FLOWERS: actinomorphic; ephemeral, yellow, blue, purple, or white; pedicellate; perianth segments subequal, free or briefly united basally, spreading or cupped; stamen filaments partially united into a tube, or free and connivent, anthers deeply notched basally, attachment pseudo-versatile, arranged around style and oriented outwards; androecium and style forming an erect column, typically yellow; gynoeclum: style branches entire, spreading, often apically broadened and flattened, alternate the stamens. CAPSULE: exerted from the subtending bracts at maturity. The genus has 80 species, distributed in North, Central, and South America, including Greenland, Hawaii, New Guinea, and one species in Ireland. The greatest diversity and number of species are found in South America. Some species are cultivated for ornament. Species of Sisyrinchiunt are common in grasslands and marshy or boggy habitats. 202

The inflorescences in this genus are difficult to understand. In all the species included here, the inflorescence is subtended by a foliaceous bract. If the inflorescence is compound, the second order peduncles are subtended by a bract or prophyll. The spathes are always of a rigid texture; the inner floral bracts may become progressively more delicate, or all may be hyaline. The innermost are hyaline, 2-nerved prophylls. Within the rhipidium, the floral bracts become progressively shorter and smaller. Compound inflorescences are composed of a peduncle, inflorescence bract, second order peduncles, one subtended by the inflorescence bract, the other by a foliaceous bract or a prophyll, and the rhipidia, each subtended by a sterile lower spathe. The rhipidia are terminal on the second order peduncles. The simple inflorescences are an inflorescence bract and a sub-sessile rhipidium composed of the sterile lower spathe and the fertile floral bracts. The repetition of inflorescence bract and lower spathe in the simple inflorescence suggests that the simple inflorescence is derived from the compound, and that the simple inflorescence is the advanced character in the genus.

Sisvrinchlum brevlncs Baker HERB, 3-11,5 cm tall, appearing shorter as leaves and peduncles stiffly spreading and downcurving; roots fibrous, thickened. LEAVES: 3-8.8 cm long, about 1 mm wide, 10- 30 per plant, linear, bases sheathing, equitant, old bases persistent, forming a fibrous sheath about base of plant, blades flat strongly ribbed, apex attenuate, blue-green. INFLORESCENCE: simple; 1-4 per plant; peduncles 1-6.7 cm long, elliptic in cross- section; inflorescence bract 10-29 mm long, 1.5-3 mm wide when folded, lanceolate, margins united briefly at base, chartaceous, apex attenuate, bract stiffly erect, pushing flowers to side, inflorescence pseudo-lateral; rhipidia 2-flowered; spathes subequal, 8-11 long, 4-10 mm wide, lanceolate to broadly elliptic, body membranous, margins wide, hyaline. FLOWERS: pedicels 1-2 mm long, only tip of flower exserted at anthesis; perianth segments 6-8 mm long, 2.5-4 mm wide, delicate, obovate, 5-nerved, spreading, yellow; stamen filaments 3.5 mm long, connivent, anthers 1.7 mm long, deeply notched basally, oriented outwards; style 4 mm long, branches 1.3 mm long, entire, broadened distally, spreading. CAPSULE: 7-9.5 mm long, 4-6 mm diam., elliptic; not exserted from floral bracts; reddish. HABITAT: Dry spots in shrublands or grasslands, often on sandy soil. 4100-4600 m. 203

DISTRIBUTION: Peru, Bolivia, and northern Argentina. Peru: Ancash, Cuzco, Junfn, Lima, Pasco, and Puno. H. N. P.: Carhuazcancha (12271, 12332A), Ishinca (9550), Llaca (12431A), Queshque (11825), and Shallap (9656). PHENOLOGY: Flowering in February; fruiting in February and March. CONSERVATION STATUS: Uncommon; not endangered.

SiiYriothlmn ifft chiknsg Hook. HERB, erect, 13-26.5 cm tall; roots fibrous, stout. LEAVES: (5.2) 7.7-15.2 cm long, 1- 3 mm wide, 10-20 per plant, linear, base sheathing, equitant, margins chartaceous to hyaline, blades ca. 7 nerved, keels scabrous, apex attenuate, glaucous. INFLORESCENCE: compound; 1-2 per plant; erect, pseudo-lateral; peduncles 7-16 (19.3) cm long, usually 2 second order peduncles above, (1.1) 3.7-9 cm long, subequal, subtended by foliaceous bract, peduncles winged, scabrous on keels; inflorescence bract 19-31 mm long, 2.2-3.2 mm wide when folded, margins united for lower 1/4-1/3, chartaceous-hyaline, body membranous, several-nerved, scabrous on keel, apex attenuate; rhipidia mostly 4-flowered; spathes 18-29 mm long, membranous at costa, margins hyaline, free, apex acute to obtuse. FLOWERS: pedicels 17-32 mm long, filiform, flowers just barely exserted at anthesis; perianth segments 10-11 mm long, ca. 4 mm wide, obovate, delicate, 5-nerved, apex emarginate-caudate, caudate tip ca. 1 mm long, blue; stamen filaments ca. 4 mm long, connivent, columnar, anthers 1.2 mm long, deeply notched basally, yellow, oriented outward; style deeply trifid, branches entire, ca. 4 mm long. CAPSULE: 5-6 mm diam., globose, green with reddish tint. HABITAT: Grassland-shrubland in vegetation types influenced by the macrothermal coastal climate; on steep slopes, southern aspects. 3800-3900 m. DISTRIBUTION: Mexico to Argentina. Peru: Ancash, Ayacucho, Cajamarca, Cuzco, Huanuco, and Junin. H. N. P.: Auquispuquio (11935). PHENOLOGY: Flowering and fruiting in April. CONSERVATION STATUS: Rare due to limited habitat within the Park.

Slsvrinchlum lamesonii Baker HERB, 20-35 cm tall, erect from fibrous, thickened roots, densely tufted; glabrous. LEAVES: 15-32 cm long, 1 mm wide, often shorter than peduncles, linear, flattened, stiff, bases weathering to loose fibrous tunic, blades bifacial with spongy mesophyll within, margin entire, apex carinate, spreading. INFLORESCENCE: a solitary 204

rhipidium; peduncle 15-26 cm long, narrowly winged on distal half; inflorescence bract 20-28 mm long, erect, foliar, apex carinate, exceeding rhipidium; rhipidium 2-4- flowered; lower spathe 10-11 (-25) mm long, 2.6-3 mm wide, upper subequal. FLOWERS: perianth segments 8-10 mm long, yellow; stamen filaments free. CAPSULE: 3-5 mm long, oblong, included in the spathes. Description adapted from Baker (1888) and Macbride (1936). HABITAT: Polylepis sericea woodlands with small brushfields included; soils of morainal parent material. 3650-3900 m. DISTRIBUTION: Venezuela to northern Argentina. Peru: Ancash, Apurimac, Cuzco, and Puno. H. N. P.: Ranincuray (1035SA). PHENOLOGY: In April, the species was collected in early fruit stage. CONSERVATION STATUS: Very rare, found only once.

SUvrlBchlum lunceum E. Meyer ex Presl HERB, strictly erect, 16-47.5 (73) cm tall; sometimes in small colonies; roots fibrous, thickened. LEAVES: 16-35 (73) cm long, ca. 1 mm wide, 1-3 per plant, linear, terete, bases sheathing, persistent becoming fibrous, scattered papillae on blades. INFLORESCENCE: compound; 1 per plant; peduncles 10-32.5 cm long, foliaceous, not winged; inflorescence bract 4.5-12 (21.5) cm long, 2-5 mm wide when folded, foliaceous, strongly nerved, base ovate, open, margin chartaceous, apex long attenuate, erect, pushed flowers laterally, appearing to be continuation of peduncle; second order peduncles 1 or 2, the first stout, prominent, 10-48 mm long, 1-2 mm diam., flexuous, the second, if produced, short, subsessile, hidden at base of inflorescence bract, subtended by hyaline prophyllum, ca. 10 mm long; rhipidia 4-6-flowered; spathes elliptic-attenuate, membranous, prominently ribbed, margins chartaceous, free, the lower 18-39 mm long, the upper 16-32 mm long; floral bracts 18 mm long or less, all distinctly prophyll-like, hyaline, 2-nerved, elliptic-attenuate. FLOWERS: pedicels 14-48 (68) mm long, filiform, long exserted to irregular lengths from floral bracts; flowers nodding; perianth segments 8-16 mm long, obovate, 5 or more-nerved, delicate, spreading, magenta or white; stamen filaments connate or connivent into column, filaments 5-6.5 mm long, anthers 1.9-2.7 mm long, deeply notched basally, yellow, oriented outwards; style 8-8.5 mm long, branches ca. 1 mm long, spreading, flattened and broadened dis tally, entire. CAPSULE: obovate-globose, immature, 4-5.5 mm long, 3-3.5 mm diameter, with scattered papillae. 205

HABITAT: Grasslands, sometimes in shrublands, on steep slopes, rocky soils, north and south aspects. 3800-4500 m. DISTRIBUTION: Peru to Chile, Argentina, and Paraguay. Peru: Ancash, Cuzco, Junin, Lima, Moquegua, and PUno. H. N. P.: Auquispuquio (11986) and Los Cedros (9925, 9926). PHENOLOGY: Flowering and early stages of fruiting in March and April. Two color forms of this species are present in the Park. The predominant form has magenta perianth segments. The form with white perianth segments (9925) has been seen in Los Cedros, along the trail to Alpamayo. CONSERVATION STATUS: Common in the northwestern part of the Park.

Slivrlnchium araeaitum Kranzlin HERB, forming dense clones, 6-20 cm diam.; plants stiff, erect, 36-112 cm tall; rhizomatous. LEAVES: 13.5-72.5 cm long, 1.5-4 mm wide, many per plant, linear, bases sheathing, strongly equitant, old bases persist as thick, fibrous tunic, blades flattened, strongly nerved, margins scabrous, apex attenuate. INFLORESCENCE: simple; numerous; peduncles 28-99 cm long, winged, stiff; inflorescence bract 7-13.4 cm long, foliaceous, strongly nerved, stiff, margin chartaceous basally, bract erect, prominent, inflorescence slightly pseudo-lateral; rhipidia 6-flowered; spathes foliaceous, stiff, strongly nerved, margins chartaceous, subequal, lower 37-64 mm long, upper 37-63 mm long, green to purplish; floral bracts increasingly prophyll-like towards center of inflorescence, outermost 30 mm long, membranous, several-nerved, others smaller, hyaline, 2-nerved. FLOWERS: pedicels 20-35 mm long, filiform, slightly exserted from spathes at anthesis; perianth segments 13-15.5 mm long, 3.7-4.5 mm wide, elliptic, 8-9- nerved, delicate, apex acute to acuminate, yellow; stamen filaments ca. 4.5 mm long, basally united for about half their length, anthers 3.4-4.8 mm long, deeply notched basally, notch ca. 1.3 mm deep, oriented outwards, yellow; style ca. 6 mm long, branches entire, free at least half their length. CAPSULE: 9-12 mm long, 5-6 mm diam., elliptic, darkening with maturity; seeds ca. 1 mm diam., globose, black. HABITAT: Grasslands, often tussock grassland, on soils of morainal material. 3370- 4300 m. DISTRIBUTION: Peru: Ancash, Apurimac, Cajamarca, and Cuzco. H. N. P.: Auquispuquio at Cerro Pakia (12048), Honda (11610), Los Cedros (9891), and Rurichinchay in Que. Pachachaca (12612). 206

PHENOLOGY: Observed flowering and fruiting in March, April, June, and October. Perhaps this species flowers nearly all year long. CONSERVATION STATUS: Not common, not believed to be endangered.

SlavrlBchium tlactorium Kunth HERB, erect, to 18 cm tall; roots fibrous, thin rhizome sometimes present. LEAVES: 44-96 mm long, 2-4 mm wide, 4-9 per plant, linear, base sheathing, strongly equitant, margins hyaline, blades flattened, S-7-nerved, apex attenuate, blue-green; leaves in fan­ like array. INFLORESCENCE: simple, 1-2 per plant, erect; peduncles 2.2-15.5 cm long, 1.2-2.5 mm wide at apex, winged; inflorescence bract 17-21 mm long, 2.3-3.4 mm wide when folded, margins joined for ca. lower 1/3, chartaceous above, body membranous, several-nerved, apex attenuate; rhipidia 3-4- flowered; spathes 16 mm long, 6 mm wide, or less; floral bracts 15 mm long or shorter, membranous at costa, margins hyaline, free, apex obtuse. FLOWERS: pedicels 18-21 mm long, filiform, well exserted from bracts at anthesis; perianth segments 6.5-9.5 mm long, ca. 2.5-3 mm wide, elliptic, very delicate, several nerved, yellow; stamen filaments ca. 5 mm long, anthers 1.3-2.7 mm long, deeply notched basally, oriented outwards, yellow. CAPSULE: 6-9 mm long, ca. 5mm wide, obovate-pyriform; well exserted from inflorescence bracts, drooping. HABITAT: Grows in saturated soils, in marshy or boggy spots. 3600-4250 m. DISTRIBUTION: Southern Mexico to Bolivia. Peru: Amazonas and Ancash. H. N. P.: Alpamayo (9824) Llanganuco (8862), Paria (9169), and Rurichinchay (12556). PHENOLOGY: Flowering from December to June, with a peak in June; fruiting about the same time. CONSERVATION STATUS: Not often found, but common where collected; probably not endangered.

SIsvrlnchium trinerve Baker HERB, strictly erect, (17) 31-68 cm tall; roots fibrous, thickened. LEAVES: (10.5) 19- 58 cm long, to 1.5 mm diam., (3) 12-22 per plant, linear, terete, bases becoming fibrous in age, persistent. INFLORESCENCE: simple; 1-8 per plant; erect; peduncles (14) 32.5- 63 cm long, terete; inflorescence bract 1.8-6.6 cm long, foliaceous, closed at very base, green to purplish; rhipidia 2-4-flowered; spathes foliaceous, margin sub-hyaline, apex acute, lower 16-27 mm long, upper 24-29 mm long, purplish, more prominent than 207

inflorescence bract; floral bracts similar to spathes, becoming hyaline and prophyll-like, smaller than spathes. FLOWERS: pedicels 16-28.5 mm long, filiform, mostly barely exceeding spathes at exsertion; perianth segments 8-11 mm long, 4-5 mm wide, elliptic, delicate, 5-7-nerved, apex acute to obtuse, yellow; stamen filaments ca. 4 mm long, united into a tube, anthers 3 mm long, deeply notched basally, yellow; style ca. 5.5 mm long, branches spreading, apically broadened and flattened. CAPSULE: 12-18 mm long, 4-6 mm diam., dark green to purplish brown. HABITAT: Grassland and tussock grassland, often on steep slopes, on rocky, shallow soils, or soils of morainal origin, on either igneous or sedimentary rock; found on all aspects. 4000-4500 m. DISTRIBUTION: Costa Rica, Colombia and Venezuela to northern Argentina. Peru: Ancash, Cuzco, Huancavelica, Huanuco, and Lima. H. N. P.: Alpamayo (9777), Auquispuquio at Cerro Pakla (12061), Carhuazcancha (12301), Churup (9636), Huaripampa (9186), Ishinca (11239), Llanganuco (10248A), Manto Mina (10993), Rio Pachacoto (11788), Ranincuray (10429), and Rurichinchay at Que. Pachachaca (12607). PHENOLOGY: Flowering from January to July, and setting fruit in the same months with maturity from April to July. CONSERVATION STATUS: The most common and widespread Sisyrinchium in the Park.

Tlgrldia A. L. Jussieu Reference: Molseed, 1970. HERBS from tunicated bulbs. LEAVES: mostly basal, a few cauline, equitant, base sheathing, linear, plicate. INFLORESCENCE: a panicle of rhipidia, or a solitary rhipidium; terminal; peduncle terete, simple or branched. FLOWERS: actinomorphic; ephemeral; erect or nodding, blue or yellow; perianth bowl-shaped, segments showy, often with complex patterns of spots, sepals much larger than petals, clawed, obovate- elliptic; petals of variable shape, from round to oblanceolate, apex entire to caudate, nectariferous on upper face; stamen filaments connate into a long tube, anthers spreading; gynoeclum: style filiform, branches bifurcate, opposite the stamens, arms of each style branch arching between the subtending anther. Distributed from Mexico to Chile, Tigridia is a genus of 15 species. Tigridia pavonia is widely cultivated. POLLINATION: By wasps or flies. 208

BREEDING SYSTEM: Although both self-compatibility and self-incompatibility are known in the genus, outcrossing is the most common system. Self-pollination and asexual reproduction are relatively rare. In the species studied by Molseed (1970), he found seed to be the dominant form of propagation. No naturally occurring hybrids have been found (Molseed, 1970).

Tlgridl» nhlllpmlmnm I. M. Johnston Reference: Johnston, 1929. HERB erect, to 25-60 cm tall; from ovoid, deeply buried bulb (not seen), 15-20 mm long, 12-15 mm wide, with thick covering of brownish tunics; stem terete; glabrous. LEAVES: 1-2 basal, 30-50 cm long, 3-8 mm wide, base 4 cm long; 2 cauline, reduced, to 14.5 cm long, base 1-4 cm long; blades plicate, 5-6 nerves. INFLORESCENCE: rhipidia 5-flowered; peduncle ca. 17 cm long; plants may produce multiple inflorescences branching from above upper cauline leaf; spathes subequal, lanceolate, 40-50 mm long, 8-12 mm wide, membranous, margins hyaline, apex acute. FLOWERS: ca. 4 cm diam., perianth segments bases white with blue spots, tips blue; sepals 15-18 mm long, 14-16 mm wide, base clawed, apex rounded to acuminate; petals 8-10 mm diam., base lacking claw, apex rounded, entire; stamens ca. 5 mm long, exserted at anthesis; gynoeeium: 0.9-1.1 cm long, 2-3 mm diam., style branches 2-2.5 mm long. CAPSULE: 28 mm long, 7 mm wide, erect, base attenuate, apex truncate, exserted from spathes; seeds ca. 1.2 mm diam., angular, brown. (Description adapted from Johnston, 1929.) HABITAT: In area of open woods dominated by Polylepis sericea and grassy shrubland. Growing on soils of morainal parent material. 3700-3840 m. DISTRIBUTION: Peru and Chile. Peru: Ancash. H. N. P.: Ranincuray (10466). PHENOLOGY: Flowering in April. CONSERVATION STATUS: Very rare, one individual found on one occasion. 209

JUNCACEAE A. L. de Jussieu COMMELINIDAE: JUNCALES Reference: Balslev, 1979a. HERBS, perennial and rhizomatous, rarely annual with fibrous roots; stem photosynthetic: glabrous, except Luzula. LEAVES: alternate, often all basal, sheath open, closed in Luzula, apices of margins often modified into auricles and a ligule, blades flat, sulcate, or terete, sometimes hollow, sometimes septate, sometimes reduced. INFLORESCENCES: terminal, sometimes appearing lateral; paniculate, racemose, cymose, head-like, spike-like, or rarely solitary, simple or compound, subtended by bracts. FLOWERS: perfect, or unisexual and plants dioecious in Distichia and Oxychloi, actinomorphic, hypogynous; subtended by bracts and bracteoles; nectaries wanting; perianth chaffy, 3-merous, in 2 whorls, sub-equal; stamens 3-6, anthers basifixed; gynoeclum 3-carpellate, united, superior, 1-3-locular ovary with axile or parietal placentation, or basal in Luzula, style terminal with 3 branches, ovules 3-many per locule. FRUIT: a loculicidal capsule, indehiscent in Oxychloë. The family has 8 genera and about 300 species. Juncus and Luzula, the two largest genera, are widespread in the temperate and cold regions of the earth, including tropical mountains. The remaining 6 genera are found in the Southern Hemisphere, with the greatest diversity in South America (Dahlgren et al., 1985). In Peru, there are species of Distichia, Juncus, Luzula, and Oxychloë. The genus Rostkovia has been found in Ecuador (Balslev, 1979b), and could be found in Peru. POLLINATION: Principally wind pollinated, some self-pollinating, and some have readapted to insect pollination (Cronquist, 1981). HABITAT: Lacustrine, riparian, marshy and boggy sites, at middle to very high elevations.

Key to the Juncaceae la. Leaves distichous; flowers solitary, unisexual and dioecious; plants cushion-forming, growing in bogs at high altitudes. Distichia muscoides lb. Leaves spirally inserted; flowers in multi-flowered inflorescences and bisexual; plants erect, solitary or bunch-forming, growing in diverse habitats. 2. 2a. Leaf sheaths closed, leaf margins sparsely to densely pubescent; capsules with 3 seeds. Luzula 2b. Leaf sheaths open, leaves glabrous; capsules with many seeds. Juncus 210

DlitIchI» C. G. D. Nees Se F. J. F. Meyen in C. G. D. Nees "Kunkusk" (Ricker, 1977). HERBS forming large, hemispherical cushions, glabrous. LEAVES: distichous, bases open, sheathing, persistent, blades terete. INFLORESCENCE: flowers solitary. FLOWERS: unisexual, plants dioecious; stamlnate flower long pedicellate, stamens 6; carpelUte flower on short gynophore, ovary unilocular, ovules many. FRUIT: a loculicidal capsule, gynophore elongates with fruit ripening. Distichia has 3 species inhabiting the high Andean regions from Colombia to northern Argentina and Chile; only one species occurs in Peru. Distichia muscoides is a dominant element in the high Andean bogs and moors, especially above 4500 m (Cleef, 1978).

Dhtichta mmcolde» Nees & Meyen HERB, forming dense hard cushions, from 60 cm to 2 m or more in diam., to 1.1 m tall; stems erect, tightly packed; rootstock not seen. LEAVES: 6-30 mm long; sheaths 2.5-13 (-18) mm long, broadly elliptic, chartaceous, margins membranous, blades 3.5-11 mm long, 2-3 mm wide, compressed-terete, narrowly triangular, apex calloused or sometimes prolonged into a thick calloused mucro. FLOWERS: stamlnate flower pedicellate, pedicel ca. 10 mm long, exserted at anthesis, perianth segments equal, 5.1-5.3 mm long, 1.1-1.2 mm wide, narrowly triangular, apex rounded, membranous, dark brown; stamen filaments 0.2-0.3 mm long, anthers 2.4-2.8 mm long, yellow; carpellate flowers not seen. CAPSULE: 8-13.5 mm long, ca. 3 mm diam., obovate, elongate, aeremchyma filled, dark brown; seeds 1-1.2 mm long, 0.7-0.8 mm diam., brown, completely covered by aerenchymatous, cream-colored arils. HABITAT: Bogs at high altitudes, in valley bottoms, stream courses, and gentle slopes. 4200-4850 m. DISTRIBUTION: Colombia to Chile and Argentina. Peru: Ancash, Ayacucho, Cuzco, Huanuco, Junin, La Libertad, Lima, Pasco, and Puno. H. N. P.: Cahuish (8282), Quengua Ragra (11750), Rio Pachacoto (11435, 11440, 11444, 11866), and Ulta (11362A). PHENOLOGY: Flowering and fruiting have been observed in September. CONSERVATION STATUS: Uncommon. 211

Juncui Linnaeus HERBS perennial, rarely annual; rhizomatous, fibrous rooted in annual species; glabrous. LEAVES: spirally arranged; sheath open, sometimes with auricles, and a ligule; blades flat, terete, or elliptical, sometimes hollow or septate, sometimes the blade reduced to a mucro at sheath tip. INFLORESCENCE: terminal, sometimes appearing lateral when subtending bract becomes erect and stem-like; paniculate, racemose, head-like, or rarely solitary; subtended by 1-several bracts. FLOWERS: perfect; perianth segments equal or unequal; stamens 3-6; ovary trilocular with axile placentation, or unilocular with parietal placentation, or intermediate, ovules many per locule. FRUIT: a loculicidal capsule. Juncus has 220 species distributed in the cold and temperate regions of the world. South America is the center of diversity for the genus. Species of Juncus are a common, even dominant, element in marshy and boggy habitats. There are ca. IS species in Peru. POLLINATION: Probably by wind. USES: The stems of the "leafless" species are woven into mats and other domestic articles.

Key to the species of Juncus la. Plants with fibrous, or very poorly developed rhizomes; short-lived or perennial. 2. lb. Plants with well-developed rhizomes; perennials. 3. 2a. Plants robust to 40 cm tall; flowers in head-like, many-flowered clusters. J. tenuis var. platycaulos 2b. Plants small and delicate, 10 cm tall or less; flowers solitary at nodes of spreading panicle. J. bufonius 3a. Leaves reduced to a bladeless sheath; plants 1 m or more tall. J. arcticus var. andicola 3b. Leaves with well-developed blades; plants less than SO cm tall. 4. 4a. Rhizome vertical; leaves cauline. S. 4b. Rhizome horizontal; leaves basal. 6. Sa. Blades bifacial, flat and grass-like; inflorescence paniculate. J. cyperoides Sb. Blades terete; inflorescence a head of (1-) 2 flowers. J. stipulons 6a. Plants 30 cm or more tall; rhizomes short-creeping; plants densely caespitose; leaves not septate. 7. 6b. Plants less than IS cm tall; rhizomes long-creeping, shoots dispersed, appearing to arise singly; leaves septate. J. ebracteatus 212

7a. Flowers in clusters of 1-3; leaves many, filiform, strictly erect. J. imbricatus 7b. Flowers in head-like clusters of 5-20; leaves few, linear, somewhat arcuate. /. pallescens

JuBcui arctlcm Willd. vr. mmdicolm (Hook.) Balslev Totora; Ututu" HERB; erect, 1.05-1.82 m tall; rhizome creeping; stems 0.77-1.43 m tall, aeremchyma- filled, terete. LEAVES: reduced to a bladeless sheath, 4-5 per stem, 1-26 cm long, chartaceous, brown, nearly black at base. INFLORESCENCE: congested, psuedolateral, paniculate, branches 3-9.5 cm long, flowers in clusters of 3-5, many small chaffy bracts subtending nodes and flower clusters; inflorescence bract erect, 28-45.5 cm long, appearing to be a continuation of stem. FLOWERS: floral bracts 2, ca. 2 mm long, ca. 2 mm wide, ovate, chaffy; perianth segments narrowly triangular to lanceolate, costa thickened, tan, margins very dark brown; sepals 5-6 mm long 1-1.7 mm wide; petals 4.5-5.3 mm long, 1-1.3 mm wide; stamens 6, filaments 0.9-1.2 mm long, anthers 0.8- 1.5 mm long, yellow; immature ovary ca. 2.5 mm long, style ca. 1 mm long, stigmas ca. 2.5 mm long, contorted. CÀfSULE: not seen. HABITAT: Seeps, marshy areas, streams, and lake shores, often growing in standing water. 3500-4000 m. DISTRIBUTION: Central America; Ecuador to Chile. Peru: Ancash, Cuzco, Huanuco, and La Libertad. H. N. P.: Huaripampa (9154), Llanganuco (8212), Querococha (8275), and Rio Pachacoto (Lôpez et al., 8330). PHENOLOGY: Flowering occurs in August. CONSERVATION STATUS: Uncommon.

JuBCug bufonlus L. HERB, annual or short-lived; to 10 cm tall; roots fibrous. LEAVES: basal and at stem nodes, 2-4.5 cm long, filiform, little differentiated into sheath and blade, little distinction between inflorescence bract and leaf. INFLORESCENCE: a spreading panicle, most of stem and branching structure of plant is floriferous, chaffy bracts at nodes. FLOWERS: solitary at inflorescence nodes; floral bracts 2 per flower, 2-3 mm long, ca. 1.5 mm wide, chaffy; perianth segments lanceolate, becoming subulate and recurved at maturity, costa green margins hyaline; sepals 4.7-8.4 mm long, 0.9-1.4 mm wide; petals 3.3-4.5 mm long, 0.9-1 mm wide; stamens 6, filaments 0.7-1 mm long. 213

anthers ca. 0.8 mm long. CAPSULE: ca. 2.5 mm long, ca. 1.6 mm wide, asymmetrically elliptic, green; teedi 0.4-0.5 mm long, ca. 0.2 mm diam., light brown with dark apices. Some of the perianth segments of several of the flowers appeared to have fallen off as the flowers matured, while others became subulate and recurved. HABITAT: Moist streamside near a trail. 3580-3850 m. DISTRIBUTION: Adventive worldwide. Venezuela and Colombia to Argentina and Chile; Brazil and Uruguay. Peru: Amazonas, Ancash, Cajamarca, Cuzco, Huanuco, and La Libertad. H. N, P.: Llanganuco on Maria Josefa trail (8865). PHENOLOGY: Flowers and fruits in December. CONSERVATION STATUS: Rare; an introduced weed.

Juncua cvneroide» Laharpe HERB; flexuous, to 22 cm tall; rhizome vertical, branching. LEAVES: cauline, more or less two-ranked, 6.5-9 cm long, 2-5.5 mm wide, bifacial, linear, flat at maturity. INFLORESCENCE: panicle, to 8.5 cm long, bracts membranous 2-3 mm long, 0.8-1.3 mm wide. FLOWERS: in clusters of 2-4; floral bracts wanting; perianth segments lanceolate, costa green, margins hyaline; sepals 2.5-3.4 mm long, 0.7-0.9 mm wide; petals 2.7-3.2 mm long, 0.7-0.7 mm wide; stamens 6, filaments 0.8-1.3, anthers 0.6-0.9 mm long, yellow; immature ovary ca. 2.3 mm long, elliptical, style ca. 0.5 mm long, sitgmas 1.5 mm long. CAPSULE: not seen. HABITAT: Growing at streamsides in the understory of Polylepis sericea woods. 3700- 3850 m. DISTRIBUTION: Colombia to Chile and Argentina. Peru: Amazonas, Ancash, Cajamarca, and Huanuco. H. N. P.: Llanganuco on Maria Josefa trail (10510). PHENOLOGY: Flowering in May. The material found in the Park is very small for the normal size of the species. One vegetatively proliferous flower was found on the inflorescence of our specimen. CONSERVATION STATUS: Very rare; collected only once.

Juncm ebr»cte«tus E. Meyer (including J, brmneus Buchenau) HERB, erect, 5.5-10 cm tall; rhizome long-creeping, nodes 2-5 cm apart. LEAVES: sheaths 1.8-2.6 (-5) cm long, chartaceous, auricles at juncture with blade, blades 5.8-9 cm long, septate. INFLORESCENCE: a highly condensed panicle, to 4 cm long, 1-2 214

main branches, flowers in dense, spherical, head-like clusters of 10- 20; inflorescence bract 0.8-5 cm, erect, foliar; scape 2-13.5 cm long. FLOWERS: floral bracts 1 per flower, 1-2.5 mm long, dark purplish brown; perianth members subequal, 3-3.8 mm long, 0.9-1.6 mm wide, lanceolate, dark purplish brown; stamens 6, filaments 0.7-1 mm long, anthers 1.3-1.6 mm long, yellow; immature ovary 1.8 mm long, style elongate, 1.5- 2 mm long, stigmas papillate, magenta. CAPSULE: ca. 2.2 mm long, ca. 1.7 mm diam.; seeds ca. 0.5 mm long, ca. 0.3 mm diam., brown with dark apices. HABITAT: Growing in saturated soils and marshy spots in grasslands. 3600-4100 m. DISTRIBUTION: Mexico and Guatemala; Peru and Bolivia. Peru: Ancash, Cuzco, and Tacna. H. N. P.: Huaripampa (9157), Ishinca (9565), Llanganuco on Maria Josefa trail (8866), Rajucolta (12148), Rio Pachacoto (Lôpez et al. 8331), and Rurichinchay at Pachachaca (12557). PHENOLOGY: Flowering from January to June; fruiting in June. USES: This species, reported as J. brmneus, has value as forage, and was found to be moderately selected by adult alpacas during the wet season (Bryant and Farfdn, 1984). CONSERVATION STATUS; Common.

Juncus imbrlcatus Laharpe HERB, erect, to 32 cm tall; rhizome short-creeping, plants densely caespitose. LEAVES: 2-3 per innovation, lower leaves reduced to sheaths with arista-like blades, sheaths 2-5 cm long, chartaceous with hyaline margins, ligule at juncture with blade, blades 6-18 cm long, filiform, canaliculate below. INFLORESCENCE: an umbellate panicle 8-11 mm long, few branches, bracts on branches membranous, green; inflorescence bract 4-5 cm long, erect leaf-like; scapes 11-30 cm long; flowers in clusters of 1-3. FLOWERS: floral bracts 2, ca. I mm long, chartaceous, ovate, margins hyaline, brown; perianth members subequal, 4.5-5.1 mm long, ca. 1.2 mm wide, lanceolate, costa thickened, green margins brown; stamens 6, filaments ca. 1.2 mm long, anthers 0.6- 0.9 mmlong, yellow. CAPSULE: not seen. HABITAT: Growing in shrubland and Polylepis sericea woods. 3580- 3850 m. DISTRIBUTION: Colombia to Chile and Argentina; Brazil and Uruguay; introduced to South Africa and Australia. Peru: Ancash, Apurimac, Cajamarca, Cuzco, Huanuco, Junin, and Puno. H. N. P.: Llanganuco (8867, 10564). PHENOLOGY: Flowering in December and May. CONSERVATION STATUS: Rare; possibly adventive in the Park. 215

Juncut Mlleacena Lam. HERB, erect, to 43 cm tall; rhizome short-creeping, plants caespitose. LEAVES: sheaths 4-7 cm long, chartaceous, ligule at juncture with blade, ca. 1.5 mm long, blades 15-16 cm long, canaliculate below, compressed terete above. INFLORESCENCE: paniculate, branches to 10 cm long, foliaceous bracts at major branch points, chaffy bracts and prophylls throughout; flowers crowded into head-like clusters of 5-20; scapes 13-34.5 cm long. FLOWERS: floral bracts 1 per flower, 4-4.5 mm long, ca. 1.8 mm wide, ovate-aristate, hyaline, 1-nerved; flower trigonous, on small stipe; perianth segment! lanceolate to narrowly triangular, light brown, margins hyaline; sepals 2.7-4.7 mm long, 1-1.3 mm wide; petals 3-4 mm long, 0.9 mm wide; stamens 6, filaments 0.8- 1.2 mm long, anthers 0.5-0.6 mm long, yellow; immature ovary 1.7- 2.5 mm long, elliptic to obovate, style short, 0.5 mm long. CAPSULE: not seen. HABITAT: Disturbed sites, in a drainage ditch at road side. 3500-3600 m. DISTRIBUTION: Ecuador to Chile and Argentina; Uruguay. Peru: Ancash, Cajamarca, Huanuco, Junin, and La Libertad. H. N. P.: Llanganuco (9396). PHENOLOGY: Flowers in January. CONSERVATION STATUS: Rare, collected only once. This species may be adventive in the Park.

Juncu» stlnuiatua Nees & Meyen HERB, 5-6 cm tall (above ground level); rhizome vertical, branching. LEAVES: cauline, sheaths 6-10 mm long, chartaceous, ligule at juncture with blade, ca. 0.5 mm long, blades 14-30 mm long, terete, apex pungent. INFLORESCENCE: a head of 1-2 flowers; inflorescence bract 4.5-6 mm long, foliar, base ovate, ca. 1.5 mm wide; scapes 11-16 mm long. FLOWERS: perianth segments subequal, 2.4-2.8 mm long, ca. 1.2 mm wide, ovate, purplish brown; stamens 6, filaments ca. 1.5 mm long, anthers ca. 1 mm long, yellow; immature ovary ca. 1.6 mm long, elliptic, style ca. 0.3 mm long, stigmas ca. 0.8 mm long. CAPSULE: not seen. HABITAT: Boggy spots in grassland. 3850 m. DISTRIBUTION: Colombia to the southern tip of South America. Peru: Ancash. H. N. P.: Rurichinchay at Pachachaca (12639 p.p.). PHENOLOGY: Flowering in June. 216

CONSERVATION STATUS: Rare, collected only once. Probably much more abundant, but overlooked due to its small size and superficial resemblance to Eleocharis albibracteata.

Juneu« tenuii Willd. v«r. platicaulos (Kunth) Buchenau HERB, erect, to 42 cm tall; rhizome poorly developed, plants caespitose, bunches ca. 7 cm diameter. LEAVES: basal, sheaths 3-4 cm long, ligule at juncture with blade, ca. 0.5 mm long, blades 19 cm long or less, canaliculate. INFLORESCENCE: paniculate, condensed, 2-4.5 (-7) cm long; flowers in many-flowered capitula; inflorescence bracts to 6 cm long, foliar. FLOWERS: floral bracts 2-2.4 mm long, 1.4-1.7 mm wide, ovate, hyaline, 1-nerved; perianth segments lanceolate, costa thickened, brown, margins hyaline; sepali 4-4.5 mm long, 1.2-1.8 mm wide; petals 3.3-3.5 mm long, 1.2-1.3 mm wide; stamens 6, filaments 1 mm long, anthers 0.6-0.8 mm long, yellow. CAPSULE: 2.7-3 mm long, 1.9-2 mm diam. There are relatively few leaves, and they seem to be of short duration. HABITAT: At trailside in Polylepis sericea woods. 3580-3850 m. DISTRIBUTION: This species is adventive in the Americas, Europe, Australia, and New Zealand. Colombia to Peru. Peru: Ancash, Cajamarca, Cuzco, and Huanuco. H. N. P.: Llanganuco on Maria Josef a trail (8844). PHENOLOGY: Fruiting in December. DISPERSAL: The seed coat becomes sticky on wetting; the seeds stick to passers-by, thus promoting dispersal (Balslev, 1979a). CONSERVATION STATUS: An introduced weed.

Luzulm A. P de Candolle in Lamark & A. P. de Candolle HERBS perennial; often fibrous rooted; pubescent. LEAVES: spirally arranged, sheath closed, blades flat, margins of leaves and bracts pubescent with silky hairs. INFLORESCENCE: panicles, heads, racemes, or spike-like, simple or compound. FLOWERS: perfect; perlanthsegments equal or unequal; stamens 3-6; ovary tricarpellate, unilocular, placentation basal, ovules 3. FRUIT: a loculicidal capsule, trigonous. The genus has 75 species, distributed in temperate and cold regions, and in the Tropics at high elevations. There are 3-4 species in Peru. Members of this genus are easily confused with members of the Cyperaceae. 217

Key to the species of Luzulm la. Inflorescence paniculate, open; leaves flat, 7-14 mm wide, very sparsely pubescent; plants of the understory and margins of dwarf woods, or shrublands. L. gigantea lb. Inflorescence spicate, dense; leaves folded or rolled, or, if flat, 1-2.5 (-S) mm wide, margins pubescent; plants of grasslands. 2. 2a. Bracts subtending the flower clusters conspicuous, longer than the clusters; perianth members narrowly ovate to lanceolate, subcoriaceous, uniformly brown; rootstock never producing tillers. L. racemosa 2b. Bracts subtending the flower clusters inconspicuous, little, if at all, longer than the cluster; perianth members ovate, membranous, margins subhyaline and paler than center; rootstock produces tillers. L. vulcanica

Luzulm gigantea Desv. HERB, erect, coarse, 63-138 cm tall; plants solitary or forming clones 30-50 cm diam., or sometimes to several meters square; glabrous other than the pubescence on leaves. LEAVES: mostly basal, 20-54 cm long, sheaths 6.5-7.5 cm long, pubescent at juncture with blade, blades 14-48 cm long, (7-) 10-14 mm wide, linear, multi-nerved, flat, lax, sparsely pubescent on margins, apex attenuate; culm leaves several, including 1 at each node of the inflorescence, like basal leaves, but smaller. INFLORESCENCE: 19-85 cm long, a central axis with 3-6 nodes and paniculate, open branching structures (7-15 cm long) arising from each node, branches 1-5 cm long, flexuous and lax, chaffy bracts and prophylls, to 1 cm long, at each branch point; peduncle 10-66 cm long. FLOWERS: solitary at branch tips; floral bracts 2, 1-2 mm long, ca. 1 mm wide, ovate-attenuate, chaffy, margin entire to erose; perianth dark brown; sepals 1.8-2.8 mm long, 0.8-1.2 mm wide, lanceolate, membranous, apex mucronate; petals: 1.7-2.8 mm long, 0.6-0.8 mm wide, lanceolate, membranous, apex mucronate; stamens 6. filaments 0.6-0.8 mm long, anthers 0.6-0.7 mm long, yellow. CAPSULE: 1.8-2.3 mm long, 1.2-1.8 mm diam., wide elliptic; seeds 1.2-1.3 mm long, 0.5-0.6 mm wide, elliptic, light brown. HABITAT: Understory and margins of dwarf woods, especially Polylepis weberbaueri - Gynoxys oleifolia woods, and shrublands. 3600-4400 m. DISTRIBUTION: Venezuela and Colombia to Argentina and Chile. Peru: Ancash, Apurimac, Ayacucho, Cajamarca, Cuzco, Huanuco, Junin, La Libertad, Pasco, and San Martin. H. N. P.: Alpamayo trail (10002), Carhuazcancha at Ichicpotrero (12368), 218

Ishinca (9569, 11263), Llaca (10785), Llanganuco (9414), Ranincuray (10402), Rurichlnchay at Pachachaca (12588), and Ulta (11392). PHENOLOGY: Flowering specimens were collected in January and March, and fruiting material from February to July. FUNGAL PATHOGENS: The inflorescences of specimens 10402 and 12368 were infected by a fungus in the Loculoascomycetes, probably in the Pleosporales (L. H. Tiffany, pen. com.). Both of these specimens are from the eastern side of the Cordillera Blanca, and, as yet, no infected specimens have been collected on the western side. CONSERVATION STATUS: Common. ^

Luzulm racemog» Desv. HERB, erect, 13-42 cm tall; caespitose, forming bunches 2-10 cm diam., shoot bases knotty, foliage erect to spreading; pubescent with hairs to 10 mm long. LEAVES: mostly basal, (3-) 7-28 cm long, sheaths (1-) 3-14 cm long, densely pubescent at juncture with blade, blades 2.5-26 cm long, 1-2.5 (-5) mm wide, linear, flat or folded, margin pubescent, sometimes densely. INFLORESCENCE: racemose, 1-6.5 cm long, 5- 12 mm wide, dense and erect to elongate and lax, often nodding; peduncle (3-) 10-38 cm long, 1-2 reduced cauline leaves. FLOWERS: in clusters of 3-4, subtended by a foliar bract, 2-several times longer than cluster, margin ciliate to lacerate; floral bracts 2, ca. 1 mm long, ovate, hyaline; perianth segments ovate-attenuate to lanceolate, membranous, 1-nerved, dark brown, margins of same texture and color, apices often caudate; sepals 2.0-3.8 mm long, 0.9-1.2 mm wide; petals 1.5-3.7 mm long, 0.5-1.1 mm wide, usually shorter than sepals; stamens 3 or 6, filaments 0.4-0.7 mm long, anthers 0.4-0.8 mm long, yellow. CAPSULE: 1.5-1.9 mm long, 1.2-1.5 mm wide, elliptic to obovate, green to brown; seeds 0.9-1.1 mm long, 0.5-0.6 mm diam., brown. HABITAT: Grasslands, and sunny spots in open woodland or shrubland; usually found in dry areas. 3600-4970 m. DISTRIBUTION: Mexico and Guatemala; Colombia and Venezuela to Argentina and Chile. Peru: Ancash, Apurimac, Ayacucho, Cajamarca, Cuzco, Huancavelica, Huanuco, Junin, La Libertad, Lima, Pisco, and Puno. H. N. P.: Alpamayo (9739), Auquispuquio (11961, 12112), Carhuazcancha (12303), Churup (9629), Ishinca (9496, 11191, 11217), Llanganuco (9420, 10285, 11298), Llaca (10834), Los Cedros (9931), Manto Mina (11017), Parôn (11487), Quenua Ragra (10658), Ranincuray (9066, 9128, 10415), Rio Pachacoto (11798), Rurichinchay (12629, 12723), and Ulta (11311, 11338). 219

PHENOLOGY: Flowering was observed from January to May, and in August. Fruiting was seen nearly year round. TAXONOMY: H. Balslev (pers. com.) includes L, chilensis Nees & Mayen ex Kunth and L. peruviana Desv. in this taxon, based oh studies of the variation between material that represents the three concepts. Luzula chilensis is a taller plant with an elongate, flexuous inflorescence and is found in moister microsites; L. peruviana is a short, erect plant with a compact and often erect inflorescence found on dry sites at high elevations. CONSERVATION STATUS: Abundant; found in all parts of the Park, often as a codominant in grassland communities.

Luzula vulcmmlcm Liebm. HERB, erect to 43 cm tall; caespitose, forming bunches to 1.5 cm diam., rootstock producing tillers. LEAVES: mostly basal, 3.8-14 cm long, sheaths 2-5 cm long, densely pubescent with long hairs at juncture with blade, blades 1.8-9 cm long, 3-7 mm wide, linear, tending to be falcate, rolled or folded, margins pubescent, apex attenuate. INFLORESCENCE: spicate, 8-30 mm long, densely many-flowered, nodding; peduncle (4-) 10-38 cm long, 2 cauline leaves, sheaths ca. 3 cm long, blades ca 3 cm long. FLOWERS: in clusters of 2-5, each cluster subtended by membranous bracts, little longer than the cluster, ovate-caudate, margins pubescent; floral bracts 1-2.5 mm long, ovate, membranous to hyaline, margins fimbriate; perianth members ovate to elliptic, membranous center, margins subhyaline, apex acute to mucronate; sepals 1.9-2.5 mm long, 0.9-1.5 mm wide; petals 1.5-2.2 mm long, 0.8-1 mm wide; stamens 3, filaments 0.3-0.8 mm long, anthers 0.3-0.7 mm long, yellow. CAPSULE: 1.7-1.9 mm long, 1.3- 1.5 mm diam., obovate, brown; seeds ca. 1 mm long, 0.5-0.6 mm diam., elliptic, light brown with a dark apex, base with a tuft of crumpled hairs. HABITAT: Grassland, especially humid spots and flood deposits. 3970-4250 m. DISTRIBUTION: Mexico; Colombia to Peru. Peru: Ancash, Huanuco, Lima, and Puno. H. N. P.: Carhuazcancha (12292) and Ishinca (9567). . PHENOLOGY: Encountered in fruit in February and May. TAXONOMY: This species is closely related to L. racemosa, and differs from it in vegetative proliferation by runners, foliage that is shorter, somewhat falcate, and less densely pubescent, the inconspicuous bracts subtending the flower clusters, the perianth segments, which are thinner, more chaffy, and two-toned, and the tuft of basal hairs on 220 the seeds. One capsule was found with 6 seeds, instead of 3, the typical number in the genus. CONSERVATION STATUS: Rare. 221

JUNCAGINACEAE L. C. Richard ALISMATIDAE: NAJADALES Reference: Haynes and Holm-Nielsen, 1986a. HERBS, perennial or annual, semi-aquatic to aquatic, inflorescences emergent or floating; rhizomatous. LEAVES: basal, linear, sheathing below with hyaline margins terminating in a ligule, blade and sheath continuous, undifferentiated; intravaginal scales membranaceous. INFLORESCENCES: terminal, scapose, ebracteate spikes or racemes, in addition Lilaea has sessile 1-flowered female inflorescences at leaf bases. FLOWERS: small, perfect or unisexual; perianth 6, equal, free tepals in 2 whorls in Triglochin, in Lilaea the staminate and perfect flowers with a single tepal, the female flowers naked; stameni in Triglochin 4-6 in 2 whorls, subsessile, in Lilaea solitary, filament sessile, adnate with tepal, anthers elongate; gynoeclum in Triglochin 6-carpellate, 3 fertile, weakly connate, separating at maturity, ovaries 1-locular, in Lilaea the ovary unilocular, trigonous, placentation basal, styles wanting to filiform and elongate in female inflorescences of Lilaea, stigmas plumose or papillose, ovule solitary. FRUIT: a nut (Lilaea), or a schizocarp (Triglochin). The Juncaginaceae is widespread in the temperate and cold regions of the Northern and Southern Hemispheres. There are 4 genera and ca. 20 species; two genera, Lilaea and Triglochin, occur in Peru. HABITAT: Plants of aquatic and sub-aquatic sites, Lilaea is usually found in vernal pools at high elevation, and Triglochin is common in saline and marine littoral sites (Dahlgren et al., 1985) and in fresh to brackish waters (Haynes and Holm-Nielsen, 1986a). POLLINATION: By wind. TAXONOMY: Occasionally, Lilaea has been separated into a monotypic family, but in the recent classification systems the Lilaeaceae is not accepted as valid (Cronquist, 1981; Dahlgren et al., 1985). TOXICITY; .Members of this family comn]ionIy contain the cyanogenic glycoside, triglochinin (Dahlgren et al., 1985). Triglochin can cause livestock poisoning by HON.

Lilaea Humbolt & Bonpland This is a monotypic genus native to the Americas. 222

Lllmem «clllloldg» (Poiret) Hauman HERB, aquatic; roots fibrous, plants reportedly rhizomatous; glabrous. LEAVES: basal, 5-20 per plant, 16-30 cm long, 2-3 mm wide at middle, base pinkish, thin folded, sheathing, margins forming small ligule, dis tally thickened, linear, aerenchymatous, apex obtuse; intravaginal scales thin. INFLORESCENCES of two types: pedunculate spikes bearing staminate and bisexual flowers and solitary, sessile carpellate flowers hidden in leaf bases; peduncles 5.5-12.5 cm long; spikes 5-9 mm long, 10-46-flowered. FLOWERS: uni- or bisexual; perianth a single tepal in the flowers of pedunculate inflorescence, wanting in hidden carpellate flower; stamen a solitary, sessile anther in flowers of pedunculate inflorescence, these either staminate or bisexual; ovary superior, unilocular, style wanting or, at most, to 1 mm in flowers of pedunculate inflorescence, elongate, 5-11.5 cm long, and filiform in the hidden carpellate flowers, stigma oblique, eccentric, cupular. NUT: 6.2 mm long, trigonous in hidden carpellate flower, immature but well developed fruits of pedunculate inflorescence 3.4-4.3 mm long, 1.5-1.8 mm wide. HABITAT: Lilaea grows in vernal pools with muddy bottoms, and is found with Callitriche heteropoda, Elatine peruviana^ Ranunculus limoselloides and Tillaea paludosa. In the vicinity of Carpa in the Rio Pachacoto drainage, this species is found growing in cold water mineral springs. 4200-4540 m. DISTRIBUTION: Western North America from Canada (British Columbia and Alberta) to Mexico, Central America, and South America along the Andes to Chile and Argentina, and Uruguay. Peru: Ancash, Arequipa, and Puno. H. N. P.: Queshque (11863), Rajucolta (12188), Rio Pachacoto (9364), and Shallap (9686). PHENOLOGY: Flowering occurs from January to April, and fruit development has been observed in March and April. Hey wood (1978) observes that the species flowers and fruits as water level drops, and passes the dry season as seed. The populations in the Park appear to be perennial, and die back only in very dry years. CONSERVATION STATUS: Common. 223

LEMNACEAE S. F. Gray ARECIDAE: ARALES Reference: Landolt, 1986. HERBS, annuals, floating to submerged aquatics; plant body a leafless thalloid frond with cavities (budding pouches) at the frond margin, 2 lateral in Lemna and Spirodela, 1 medial in Wolf fia and Wolffiella', 1-several unbranched roots or none; plants solitary or connected in small clones. INFLORESCENCE: borne in one of the 2 budding pouches in Lemna end Spirodela, or in a dorsal cavity in Wolf fia and Wolffiella; the inflorescence composed of 1-2 male flowers and a female flower subtended by a small spathe in Lemna and Spirodela, or of 1 male and 1 female flower without a spathe in other genera. FLOWERS: lacking perianth, unisexual and plants monoecious; stamlnate flower a single stamen; carpellate flower a single, sessile unilocular gynoecium, placentation basal, style terminal, short, ovules 1-7. FRUIT: a small utricle; seeds 1-4. The family, which is cosmopolitan in distribution, has 4 genera and 34 species. Lemna, Spirodela, and Wolffiella are found in Peru; Wolf fia occurs in other parts of South America and may be found in Peru. HABITAT: Growing floating or submerged, and forming extensive masses on the surface of pools of slow or stagnant water. The species pass through winter or drought by forming a resting bud, a frond with dense tissue and a high food reserve. The resting bud sinks to the bottom of the pond during winter and rises to the surface in the Spring, or rests on the soil during drought (den Bar tog and van der Plas, 1970). REPRODUCTION: The most commonly seen form of reproduction is vegetative budding from the budding pouches. The new plant remains attached to the old for a while. USES: Duckweeds are important food resources for water fowl (Martin et al., 1961) TAXONOMY: The species of Wolf fia are the smallest known Angiosperms (Cronquist, 1981).

Lemna Linnaeus HERBS, floating, thallus 1-5 (-7)-nerved, margin entire; budding pouches 2, lateral, daughter thalli remaining attached to mother thallus after budding, forming clones of few-SO thalli; root solitary, subtended by a winged or wingless, tubular sheath, root tip covered by a root cap; glabrous. INFLORESCENCE: sessile, composed of 3 unisexual flowers, 2 staminate, 1 carpellate; developing in 1 of the 2 budding pouches. 224

FLOWERS: ttaniinate: a single stamen; carpellate: a single gynoecium, subtended by a small spathe. FRUIT: a utricle. Lemna is a genus of cosmopolitan distribution. Of the 13 species, four occur in Peru.

Lemna mlnugcula Herter HERBS, floating; thallus 1.5-3.6 mm long, 0.8-2.1 mm wide, elliptic, ovate, or obovate, indistinctly 1-nerved, basal stipe nearly obsolete; daughter thalli remain attached to mother thallus, forming clones of 6-7 thalli; root 6-8.8 mm long, basal sheath 0.3-0.5 mm long, root cap 0.8-1.2 mm long. INFLORESCENCE, FLOWERS, and FRUIT: not seen. HABITAT: Floating in small, slow moving streams, springs, and in ponds. 3600-3800 m. DISTRIBUTION: United States to Guatemala; West Indies; Venezuela to Chile and Argentina; adventive in Europe and eastern Asia. Peru: Ancash, Cajamarca, Cuzco, Huanuco, Junin, La Libertad, Lima, and Moquegua. H. N. P.: Honda (11699), Huaripampa (9153), and Ishinca (11290). Macbride (1936) cites Lemna minima Philippi ex Hegelm. from Peru; this taxon is a synonym for Lemna minuscula. PHENOLOGY: Neither flowering nor fruiting specimens of this taxon were collected during the fieldwork. The size of the thalli seemed to decrease with altitude. CONSERVATION STATUS: Although not commonly encountered, it does not seem to be endangered within its habitat. 225

ORCHIDACEAE A. L. de Jussieu LILIIDAE: ORCHIDALES References: Ames and Correll, 1952, 1953; Dodson et al., 1987; Dressier, 1981; Schweinfurth 1958, 1959, 1961, 1970. HERBS, perennial, terrestrial or epiphytic, or, in Vanilla, vines; some rhizomatous, some with tuberous roots, epiphytic species with aerial roots covered by a layer of specialized cells (velamen), stems erect and cane-like, or modified into a thickened storage organ (pseudobulb), terrestrial species with corms or tubers; extrafloral nectaries present in some species. LEAVES: simple, alternate, distichous or spiralled, cauline or basal or reduced, bases sheathing and closed, or open and petiolate, blades membranaceous to coriaceous, entire, marcescent or deciduous, blade sometimes deciduous from sheath by an abscission layer (articulate). INFLORESCENCE: terminal or axillary, spikes, racemes, panicles, or solitary, scapose or pedunculate. FLOWERS: usually subtended by a bract; perfect, rarely unisexual and plants monoecious or dioecious, strongly zygomorphic, rarely nearly actinomorphic, epigynous; resupinate or not sepals 3, equal or the one opposite the lip (dorsal sepal) different in shape or color, free or 2-all connate; petals 3, the laterals often similar to the sepals, the central petal (lip) strongly differentiated in size, shape and color; lip and sepals often borne on basal outgrowth of column (foot); often having floral nectaries of several types; androeclum, style and stigmas adnate, joined into a stout column terminated by anther and entire to 3-lobed stigma, a portion of the stigma often modified into a tissue (rostellum) that subtends the anther and develops the viscidium and stipe; stamen 1,2 in Phragmipedium, opposite the lip, pollen joined into pollinia, except in Phragmipedium, these often with a caudicle at base, many genera have complex units of pollen dispersal (pollinaria) composed of the pollinia, caudicles, stipe and viscidium; gynoeclum 3-carpellate, ovary compound, inferior, unilocular, placentation parietal, ovules minute and numerous, usually not developing until after pollination. FRUIT: a capsule, rarely winged, dehiscing by . longitudinal ruptures or a fleshy pod; seeds minute, like dust, except in Vanilla. The family has a cosmopolitan distribution, and can be found in all habitats except in extreme arctic and desert environments (Dressier, 1981). There are ca. 800 genera and between 20,000 and 23,000, no more than 25,000, species (Atwood, 1986; Dodson et al., 1987; Dressier, 1981). The Neotropics has the most diverse orchid flora in the world, with ca. 8270 species in ca. 310 genera (Dressier, 1981). Tropical Asia is the region with the next most diverse orchid flora. Schweinfurth (1958, 1959, 1961, 1970) 226 estimated the Peruvian orchids to include 900 species in 120 genera; recent estimates indicate 2000 species (Dressier, 1981) and 1800 species in 165 genera (Dodson et al., 1987). The Asteraceae, Fabaceae, Orchidaceae, Poaceae, and Rubiaceae are the largest families of flowering plants; of these, the Asteraceae and Orchidaceae are the largest, and have approximately the same numbers of species (Atwood, 1986). HABITAT: Terrestrial to epiphytic-Iithophytic in nearly all known habitats in Peru, except the coastal deserts. Most of the epiphytic species are capable of surviving normally on the ground (Dahlgren et al., 1985). Dressier (1981) estimates that 25% of orchid species are primarily terrestrial, another 5% can live equally well on either the ground or epiphytically-lithophytically, the remaining species are primarily epiphytic. In addition to their habitat preferences, most species require a mycorrhizal association, at least during germination and early growth. POLLINATION: Orchid flowers are very highly specialized for insect pollination; pollination by birds and bats is also known (van der PijI and Dodson, 1966). DISPERSAL: The seeds are wind dispersed, and thought to be suitable for long range dispersal. The pseudobulbs are highly effective vegetative diseminules. USES: Thousands of species are highly prized ornamentals, and are cultivated and bred throughout the world. The genus Vanilla is the source of the flavorant vanilla (Simpson and Conner-Ogorzaly, 1986). Since the Orchidaceae is an exceedingly large family, there are many problems to be studied: taxonomic, physiologic, and floristic. Traditionally, the family has been divided into subfamilies, tribes and subtribes. Burns-Balogh and Funk (1986), Garay (Dressier, 1981), and Dressier (1981) have all proposed classification systems for the family, involving 5 to 7 subfamilies and about 20 tribes. I have followed Dressler's classification scheme. In studying a family with many evolutionary specializations, students inevitably develop an equally specialized vocabulary to describe the structures; here are the important terms used: Articulate: leaves having an abscission layer between the sheath and blade, the blades thus deciduous and the sheaths long persisting. Caudicle: a fine, wire-like tail, composed of pollen or viscin of antherial origin, at the base of the pollinia, attached either directly to the viscidium or to the stipe. This is thought to promote detachment of the pollinia from the viscidium. 227

Column: a columnar structure composed of the adnate style, anthers, and stigmas. This is often winged or appendaged. Lip: the morphologically upper petal, which is usually enlarged and showy, and otherwise modified for pollinator attraction. Many orchid flowers are resupinate, and the lip appears to be the lower petal. Pollinia: an aggregation of few-many pollen grains held together by an elastic substance (viscin). These are thought to increase the efficiency of fertilization by delivering a package of many pollen grains to a gynoecium containing many ovules. Pollinaria: a unit of pollen dispersal composed of pollinia, caudicles, stipe and viscidium, at the minimum pollinia and viscidium. Pseudobulb: an aerial stem specialized for storage. These have many shapes and sizes, and 1-several internodes. Resupinate: inversion of a leaf or flower by a 180 degree twist in either a petiole or pedicel. As a result the morphologically lower side is up, and the upper side down. Rostellum: a flange of tissue developed at the apex of the stigma which subtends the anther. Stipe: a small column of tissue which serves as the connection between the viscidium and the pollinia in some orchids. Velamen: a spongy sheath of 1-many layers of dead cells surrounding orchid roots. This can be found on the roots of terrestrial species as well as the roots of epiphytic-lithophytic species. It absorbs water and nutrients, and shelters symbiotic blue-green algae and mycorrhizal fungi. Viscidium (pi. viscidia): a portion of the apex of the rostellum whose cells break down into a sticky substance that glues the pollinaria to the back of the insect pollinator. This tissue is of stigmatic origin. COLLECTING PRACTICES: Since the Orchid flower has a complex structure, it is preferable to preserve some flowers in liquid, either FAA or alcohol. In the small- flowered genera (Aa, Myrosmodes, PleurothalUs, and Stelis) this is indispensable. Photographs are very useful, but not a substitute for spirit collections. I thank Dr. C. H. Dodson (MO) for advice, discussions of the genera, and access to his notes and manuscripts. His help has been invaluable. In Lima, Sr. Ricardo 228

Fernandez, of the Museo de Historia Natural, kindly shared his knowlege of the Orchids with me. Key to the genera of Orchidaceae la. Plants principally terrestrial, from a cluster of tuberous roots; leaves membranous, mostly basal; flowers usually not resupinate. 2. lb. Plants principally epi- or lithophytic, from caespitose or creeping rhizomes; leaves coriaceous (except Malaxis), cauline or basal; flowers, usually resupinate. 6. 2a. Flowers resupinate; lip with a long nectariferous spur at base, often 3-lobed. Habenaria 2b. Flowers not resupinate; lip lacking long spur. 3. 3a. Inflorescence terminal; leaves fully developed during flowering; column bent back 90° to ovary. Comphichis 3b. Inflorescence lateral; leaves absent (Aa) or developed (Myrosmodes and Pterichis) during flowering; column not strongly bent. 4. 4a. Root crown deeply buried, most of plant body and peduncle immersed in soil; inflorescence conical. Myrosmodes 4b. Plants erect, not immersed in soil, peduncle long exserted; inflorescence cylindrical, elongate. S. Sa. Peduncle completely covered by sheath-like, scarious bracts; flowers white. Aa 5b. Peduncle bracts membranous and widely spaced; flowers olive green with a yellowish lip. Pterichis 6a. Leaves membranous, not articulated, sheathing basally thickened stem; growing in organic matter, sometimes terrestrial; flowers green. Malaxis 6b. Leaves coriaceous, articulated; epi- or lithophytes; flowers yellow, purple, red or, rarely, white. 7. 7a. Stems cane-like, erect, not basally swollen; leaves cauline. 8. 7b. Stems modified into swollen pseudobulbs; leaves basal and apical. 11. 8a. Stems elongate, bearing several to many leaves and well developed adventitious roots; lip prominent, united to column to its apex; ovary not articulated with pedicel. Epidendrum 8b. Stems short, bearing 1 terminal leaf with 1-few bracts below; lip inconspicuous, smaller then the sepals, attached to base of the column; ovary articulated with pedicel. 9. 229

9a. Flowers 2-severaI cm long; sepals connate into a tube, lobes caudate; flowers purple, red, or, rarely, white. Masdevallia 9b. Flowers 1 cm long or less; sepals free or connate, not forming a tube, apices not caudate; flowers yellow or deep purple. 10. 10a. Flowers superficially regular, sepals widely spreading, rotate; petals wider than long. Stelis 10b. Flowers distinctly zygomorphic, sepals not widely spreading; petals longer than wide. Pleurothallis 1 la. Petals and column mimicking a fly, bristly and dark purple; lip not large and prominent. Trichoceros lib. Petals and column not modified for insect mimicry; lip enlarged and prominent. 12. 12a. Lip united with the column to its apex; epiphytes. Epidendrum 12b. Lip attached at base of column, the claw free from and paralleling the column, apex reflexed; lithophytes. Odontoglossum NOTE; In addition to the genera treated here, there is an additional genus in the Spiranthinae found in the Parân valley by Mostacero. The specimen has been misplaced. When collected again, the genus can be determined by consulting the revision of the Spiranthinae (Garay, 1980). Should other genera be found that are not in this treatment, one can consult "Los generos de las Orquideas ecuatorianas" (Dodson, 1986).

At H. G. Reichenbach Reference: Garay, 1978 Terrestrial, from a fascicle of fleshy roots. LEAVES: spiraled, usually in a basal rosette, membranous, not articulate, absent at flowering fully developing afterwards. INFLORESCENCE: lateral, a dense spike; peduncle completely sheathed by imbricate, tubular-funnelform, scarious bracts, stem glabrous or pubescent with multicellular hairs. FLOWERS: small, not resupinate, white; floral bracts hyaline, equalling to much longer than flower, sometimes reflexed at middle when mature; sepals equal, free or briefly connate basally, laterals erect and connivent with or variously embracing the lip, dorsal often recurved; lateral petals narrowly oblong, lip deeply galeate, margin lacerate to fimbriate and involute; column short. This is a high Andean genus of ca. 30 species, of which 9 are Peruvian (Dodson, 1988). 230

POLLINATION: The pollination biology of this genus is unknown. The plants are probably largely self-pollinating with occasional outcrossing. TAXONOMY: Most of the species in this genus and Myrosmodes have been included at one time or another in AUensteinia Kunth. Aa, Altensteinia and Myrosmodes are all in need of systematic revision. Dodson (pers. com.) has seen many specimens of the genus, and found the group to be highly variable. There may be many species of very localized distribution, or the variation seen in the genus may be a result of self-pollination and the formation of genetic lines. The species can be divided into two species groups: Aa agyrolepis group - has floral bracts about equal the length of the flowers, erect and appressed to the inflorescence. Aa paleacea group - has floral bracts much longer than the flowers and reflexed just below the flowers. In this treatment, I am accepting two species, Aa matthewsii in the agyrolepis group and Aa paleacea in the paleacea group. As used in this treatment, the name Aa matthewsii probably better represents the agyrolepis species group rather than the individual species. I will not be surprised if more than one valid species has been included under this name; it is a question that only critical study of the genus can resolve. Using the treatment in the Flora of Ecuador (Garay, 1978), some of the cited specimens will key out to species other than those treated herein. Given the superficiality of our understanding of the genus, I prefer to use the names in Orchids of Peru (Schweinfurth, 1958) rather than assume that Ecuadorian species occur in Peru.

Key to species of Aa la. Floral bracts erect, appressed to inflorescence, 1-1.5 times flower length. A. matthewsii lb. Floral bracts reflexed, 2 times flower length or more. A. paleacea

Am matthewsii (Reichb. f.) Schltr. TERRESTRIAL, from a fascicle of tuberous roots, erect, 19.5-53 cm tall. LEAVES: ca. , 14 cm long, ca. 1 cm wide, narrowly elliptic, base scarious, sheathing, blade attenuate and |>etiolate below, a()ex attenuate. INFLORESCENCE: an elongate spike, 4-8.5 (- 11.3) cm long; peduncle 14.5-45 cm long, hidden by imbricate bracts, sometimes 231

partially exposed apically, glabrous or pubescent with multicellular hairs, bracts to 1-6.5 cm long, tubular-funnelform, scarious, inflated and loose on stem, mouth oblique, most with an attenuate lateral limb to 1.5 cm long. FLOWERS: 5-10 mm long, white; floral bract! 7-11.5 mm long, 3.9-6.5 (-7.5) mm wide, erect, ovate, often narrowly so, scarious, nerveless, apex attenuate, margin erose, base reddish; sepals: laterals 3.3-6 mm long, 1.2-1.8 mm wide, oblong to oblanceolate, I-nerved, apex rounded-oblique to sub­ acute, faintly keeled, suberect and diagonally overarching lip to deflected laterally embracing lip and petals, margin entire to minutely toothed, dorsal 2.6-3.2 mm long, 0.9-2 mm wide, elliptic, I-nerved, apex subacute to rounded, reflexed outwards; petals 2.3-3.5 (-4.5) mm long, 0.6- 1.3 mm wide, narrowly oblong, 1-nerved, laterally arcuate, margin erose, apex rounded, reflexed outwards, lip 2.5-4.3 (-5.2) mm long, 2-4 mm wide, hooded and saccate, glabrous within, margin fimbriate, white with green lines within; ovary glabrous or pubescent with multicellular hairs. CAPSULE: not seen. HABITAT: Open woodlands of Polylepis weberbaueri or Gynoxys oleifolia, grassland- shrubland, or grassland; on soils of morainal origin or organic matter on old rockfalls. 4000-4400 m. DISTRIBUTION: Peru and Bolivia. Peru: Ancash, Ayacucho, Cuzco, Huanuco, Junfn, Lima, and Puno. H. N. P.: Cahuish (11095A), Carhuazcancha at Que. Rima Rima (12201), Llaca (10784, 11141A, 12424), Parôn (11526), and Ulta (11395). PHENOLOGY: Flowering from May to September; early fruiting observed in September. CONSERVATION STATUS: Uncommon, perhaps worthy of protection.

Am nmlemcem (Kunth) Reichb. f. TERRESTRIAL, from a fascicle of tuberous roots, erect, 21.5-38 cm tall. LEAVES; base 1.5-4 cm long, tubular, scarious, blade 30-56 mm long, 6-9 mm wide, elliptic to narrowly elliptic, attenuate and petiolate below, principal nerves 3, arcuate apex acute, margin entire. INFLORESCENCE: an elongate spike, 4.2-6 cm long, rachis sparcely pubescent with multicellular hairs; peduncle 17-33.5 cm long, hidden by imbricate bracts, sometimes partially exposed apically, sparcely pubescent with multicellular hairs, bracts to 2-6 cm long, tubular-funnelform, scarious, inflated and loose on stem, mouth oblique, most with attenuate lateral limb to 1.5 cm long. FLOWERS: 7.2-8.3 mm long, white; floral bracts 15-17 mm long, 5-7.5 mm wide, reflexed, lanceolate, scarious, nerve less, apex long attenuate, margin entire, base reddish; sepals: laterals 3-4.2 (-6) mm 232

long, 1.5-2.3 mm wide, asymmetrically obovate, 1-nerved, apex truncate, concave, erect, connivent with and overarching lip, margin minutely erose, dorsal 2.1-2.8 (-4) mm long, 1.6-2 mm wide, elliptic, 1-nerved, apex acute, reflexed outwards; petals 2.1-3 (-4) mm long, 0.7-0.9 mm wide, narrowly oblanceolate, 1-nerved, margin erose, apex acute to cuspidate, reflexed outwards, lip (2.3-) 3.2-3.7 mm long, 2.6-3.3 mm wide, depressed ovate, clawed, base truncate, deeply galeate, dorsi-ventrally compressed basally, keeled above, glabrous within, margin fimbriate, white; ovary glabrous or sparcely pubescent with multicellular hairs. CAPSULE: not seen. HABITAT: Grasslands on the eastern side of the cordillera, often near streams. 4100- 4300 m. DISTRIBUTION: Venezuela and Colombia to Bolivia. Peru: Ancash, Cajamarca, Cuzco and Pasco. H. N. P.: Carhuazcancha at Que. Rima Rima (12202) and Ranincuray (9108, 10434). PHENOLOGY: Flowering observed from January to May, and fruiting from April. CONSERVATION STATUS: Rare.

Enldendrum Linnaeus EPI- and LITHOPHYTIC, rarely terrestrial, with or without rhizome; stems slender, cane-like, simple or branched, erect with cauline leaves, producing adventitious roots, these sometimes tuberous, or pseudobulbs, variable in shape, 1-several-nodes, covered by bladeless sheaths and with leaves at apex. LEAVES: distichous, coriaceous or fleshy, articulate. INFLORESCENCE: terminal, pedunculate, racemose or paniculate, rarely lateral. FLOWERS: resupinate or not, glabrous; floral bracts membranous; sepals equal, spreading; lateral petals narrower than sepals, spreading. Hp adnate to column, often to the apex, entire or 3-lobed; column short to long, winged or not. Epidendrum is distributed from the United States (North Carolina) throughout tropical America to Argentina. This has been considered the largest genus of Neotropical orchids. Ames and Correll (1953) gave an estimate of 800 species; a recent estimate by Atwood (1986) reduces this to 500 species. Dodson (pers. com.) estimates Epidendrum to have 825 species or more. Pleurothallis is the largest genus of Neotropical orchids, and Epidendrum the second largest. There are ca. 195 species of Epidendrum in Andean Peru (Dodson, 1988). POLLINATION: Mediated by moths, butterflies, flies, and hummingbirds. 233

TAXONOMY: Since it is a large genus, Epidendrum can be unwieldly; as yet there are no subgenera. In treating the genus for Ecuador, Dodson et al. (1987) separated the taxa into species groups, which will be very helpful for future work. With further study, some groups will be separated as segregate genera; Dressier (1984) has addressed this problem for the Mexican taxa.

Key to the species of Epidendrum la. Plants epiphytic; stems modified into pseudobulbs. E. excelsum lb. Plants lithophytic or terrestrial; stems cane-like. 2. 2a. Plants terrestrial; leaf sheaths very rugose. E. cernuum 2b. Plants lithophytic; leaf sheaths faintly rugose or smooth. 3. 3a. Inflorescences pendent, 7-11-flowered; leaves narrowly elliptic, erect. E, inamoemm 3b. Inflorescences erect, 1-3-flowered; leaves elliptic, spreading E. megagastrium

Epidendrum cernuum Kunth, vel Sp. aff. TERRESTRIAL, ca. SO cm tall; stems erect, cane-like, frequently branched, with 1-2 sheath-like bracts at base of branch; no adventitious roots at branch points. LEAVES: apically crowded, imbricate, sheaths tubular-funnelform, 1.6-3.3 cm long, chartaceous, long persistent, very rugose, sculpturing dark brown, blades 2.8-8.5 cm long, 0.9-1.9 cm wide, elliptic to lanceolate, spreading, apex asymmetrically acute. INFLORESCENCE: a terminal, pendent, S-16-flowered raceme, 2.5-8 cm long; peduncles very short, hidden in apical leaf, subtended by a sheath-like bract, ca. 11 mm long. FLOWERS: 2.3-2.8 cm long, not resupinate, green with purple markings; floral bracts 3.5-7.2 mm long, ca. 3 mm wide, ovate to narrowly triangular, membranous, purplish; sepals: laterals ca. 9.5 mm long, ca. 5 mm wide, asymmetrically elliptic, 3-4-nerved, dorsal ca. 9.5 mm long, ca. 3 mm wide, oblong, 3-nerved; lateral petals ca. 8J mm long, ca. 3.5 mm wide obovate, lip carnose, ca. 7.3 mm long, ca. 6 mm wide, 3-lobed, center lobe with 3, small lobes, elongate callus along length of lip; column 4-5 mm long; ovary and pedicel continuous, 14-15.5 mm long. CAPSULE: not seen. HABITAT: Grassland, on steep, south-facing slopes. 3600 m. DISTRIBUTION: Venezuela and Colombia to Peru. Peru: Ancash. H. N. P.: Parôn (8952). The distribution of this taxon in Peru is poorly known; it does not appear in Schweinfurth (1959, 1970). 234

PHENOLOGY: Flowers in January. CONSERVATION STATUS: Rare.

Epldcndram excelaum C. Schweinf. EPIPHYTIC, erect from creeping rhizome with adventitious roots; stems subtended by 1-2 chartaceous, sheath-like bracts, 1.8-2.5 cm long, dark brown, pseudobulbs borne on short stem, 1 cm or less, and subtended by a leaf then crowned by 2 apical leaves, these 1 cm apart or less; glabrous. LEAVES: sheaths closed, tubular, sheath of subtending leaf wholly or partly enclosing pseudobulb, 0.8-1.6 cm long, blades 1.2-4.8 cm long, 0.5- 1.1 cm wide, sub-coriaceous, elliptic to lanceolate, sheaths of apical leaves 0.5-1 cm long, blades (2.3-) 4-9 cm long, 0.6-1 cm wide, sub-coriaceous, elliptic to lanceolate to linear. INFLORESCENCE: terminal, 10-32-flowered raceme, typically arcuate. FLOWERS: 11.4-15 mm long, yellowish or greenish, often with a pinkish or purplish tint, resupinate or not; floral bracts 2.8-6.5 mm long, sterile bracts below to 10 mm long, narrowly triangular to ovate, membranous, 1-nerved, erect; sepals: laterals 4.3-5.6 mm long, 2.4-3.5 mm wide, carnose, asymmetrically obovate, acuminate, dorsal 4.3-5.7 mm long, 2.4- 3.3 mm wide, carnose, elliptic, acuminate; lateral petals 4-6 mm long, 1- 1.4 mm wide, membranous, oblanceolate, acute, margin erose, lip 3.5-4.5 mm long, 4.7- 7.9 (-13) mm wide, 3-lobed to sub-entire on same inflorescence, base cordate, center lobe 2- lobed, bi-callose at base of column and longitudinally through center, blade thickened in center, margins finely lacerate; column 4.3-4.7 mm long; ovary and pedicel continuous, 5.3-7.7 mm long. CAPSULE: not seen. HABITAT: Polylepis sericea or Polylepis weberbauerii woodlands. 3700-4000 m. DISTRIBUTION: Peru: Ancash and La Libertad. H. N. P.: Carhuazcancha at Ichicpotrero (12371), Honda (11634) and Ranincuray at Que. Yanapaccha (10467). PHENOLOGY: Found in flower in October, April and May. This species is closely related to Epidendrum rupestre Lindley and E. albomarginatum Reichb. f. (Schweinfurth, 1970). CONSERVATION STATUS: Uncommon.

Enidendrum inamoenum Kraenzlin LITHOPHYTIC to 1.3 m tall; stem erect, cane-like, branched, with 2 closed sheaths subtending branch base, adventitious roots arising from branch points, the roots pendent, tuberous-swollen; glabrous. LEAVES: crowded at apex, sheaths subimbricate, 1.3-3 cm 235

long, closed, very faintly rugose, pinkish, blades 2.1-6.2 cm long, 0.6-1.2 cm wide, narrowly elliptic, coriaceous, erect, apex asymmetrically acute. INFLORESCENCE: a terminal, pendent, 7-11-flowered raceme, 3.5-4.5 cm long; peduncles short, hidden in terminal leaf. FLOWERS: 2.2-3.2 cm long, not resupinate; floral bracts 4.5-9 mm long, ca. 2.4 mm wide, narrowly triangular, membranous, purplish; sepals: laterals 12-12.5 mm long, 4.3-5.2 mm wide, carnose, asymmetrically obovate, apex acuminate, connate for 1/4-1/2 of length, cupping column and lip, dorsal 10.5-13 mm long, 3.5-4.7 mm wide, carnose, elliptic, apex acuminate, all sepals purplish without, within yellow with purple spots; lateral petals 8-11 mm long, 1.1-1.6 mm wide, very narrowly oblanceolate, yellow with purple spots, lip 4-6.6 mm long, 8-10.3 mm wide, carnose, cordate basally, 3-lobed apically with well defined sinuses, 1.7-1.8 mm deep, central lobe somewhat 2- lobed, callus at base, margins erose, yellow with reddish spots; column 7.3-9 mm long; ovary and pedicel continuous, 13-19 mm long. CAPSULE: 1.8-2.5 cm long, 1.2-1.4 cm wide, elliptic, pendent, green. HABITAT: On cliffs, talus slopes, and old boulder falls in Polylepis sericea woods. 3500-3700 m. DISTRIBUTION: Peru: Ancash, Apurimac, Cuzco, Junin and Puno. H. N. P.: Llanganuco (8869) and Ranincuray at Que. Yanapaccha (10479). PHENOLOGY: Flowering has been observed in December and April, and fruiting in December. CONSERVATION STATUS: Rare.

Epidendrum megagastrlum Lindley LITHOPHYTIC, 35 cm tall or more; stems erect, cane-like, branching, 1-few sheath­ like bracts on lower portion of branch; adventitious roots arising from branch points; glabrous. LEAVES: distal on stem, sheaths 1.5-3.5 cm long, closed, tubular-funnelform, chartaceous, smooth, several-nerved, persistent, weathering nearly scarious, blades 3.2-7 cm long, 1.5-2.8 cm wide, elliptic, coriaceous, spreading, apex asymmetrically acute. INFLORESCENCE: a terminal, erect, 1-3-flowered raceme, 1.5-3 cm long; peduncle short, hidden in sheath of apical leaf, subtended by 2 sheath-like bracts, 2.2-2.8 cm long. FLOWERS: 4.3-4.7 cm long, not resupinate, dark yellow; floral bracts 5.8-13 mm long, elliptic-acute, membranous, purple; sepals: sub-equal, 15-16 mm long, 6-8.5 mm wide, carnose, elliptic, apex attenuate; lateral petals ca. 15 mm long, 3-4.5 mm wide, lanceolate, carnose, lip 8-10 mm long, 7-9.2 mm wide, obtusely triangular, carnose; 236

column ca. 8 mm long; ovary and pedicel continuous, 2.8-3.5 cm long. CAPSULE: ca. 3 cm long, elliptic, perianth persistent. HABITAT: Bouldery areas in brushfields. 3700 m. DISTRIBUTION: Ecuador and Peru. Peru: Amazonas, Ancash, Cajamarca, Huanuco and Junin. H. N. P.: Pardn (8943). PHENOLOGY: Collected in flower in January. This taxon is closely related to Epidendrum geminiflorum Kunth of Colombia, Ecuador, and northern Peru (Cajamarca); further study may show that the two are synonymous (Dodson, pers. com.). CONSERVATION STATUS: Rare.

Gomnhlchls Lindley Reference: Garay, 1978. Terrestrial; erect from fascicle of tuberous roots. LEAVES: spiraled, mostly basal and rosulate, membranous, folded, petiolate, not articulated, glabrous; well developed before flowering. INFLORESCENCE: terminal, a dense spike; peduncle leafy below, becomming reduced to leaf-like bracts above. FLOWERS: small, not resupinate, pubescent; floral bracts membranous, equal to or shorter than flowers; perianth and column usually bent back at right angle to ovary; sepals equal, free; lateral petals narrower than sepals, often ciliate, lip free, carnose, clawed, concave, entire to 3-lobed, connivent with column. Except one each in Brazil and Costa Rica, all the species in this genus are found in the Andes. Dodson (1988) estimates that there are 18 Andean species, of which 7 are Peruvian. POLLINATION: Although the pollinators of this group are unknown, the sweet odor of the flowers suggests insect pollination, perhaps flies.

Gomahlchls valida Reichb. f. TERRESTRIAL, 53-115 cm tall, from a cluster of tuberous roots; flowers and inflorescence pubescent with rufous, multicellular, glandular hairs, often densely so. LEAVES: 4-9 per plant, mostly basal, 1-2 cauline at base of peduncle, 18-38 cm long, 1.4-2.5 cm wide, lorate, margin entire. INFLORESCENCE: erect to arcuate, densely many-flowered, 8-14.5 (-17.5) cm long, cylindrical; peduncles 41.5-97.5 cm long, covered by imbricate sheath-like bracts, 1.3-19 cm long, lowermost foliar, uppermost 237

chartaceous, mouth oblique, one side extended and blade-like, margins pubescent. FLOWERS: to 20 mm long; with sweet odor at anthesis; floral bracts 13-17.5 (-19) mm long, (4.5-) 5.6-8.5 mm wide, elliptic to obovate, membranous, apex attenuate, pubescent on margins and back; perianth pubescent on margin and back, except lip which is glabrous save a few haeir at base; sepals: laterals (7.3-) 8-10.2 mm long, (2.7-) 3.1-4.6 mm wide, carnose, asymmetrically obovate, deeply concave, l-(3)-nerved, apex acute, white with green margins, dorsal (7.5-) 8-10.2 mm long, 2.7-4.5 mm wide, carnose, obovate, concave, 1-nerved, apex acute to attenuate, green; lateral petals (7-) 8-9.5 mm long, 2.9-3.5 mm wide, clawed, asymmetrically ovate to rhombic, 1-nerved, apex attenuate, white with green margins. Up 8-9.7 mm long, 5.5-7.5 mm wide, claw short, triangular, 3-lobed, central lobe elongate, often slightly reflexed, with strongly thickened, medial callous, margins and lateral lobes thinner, involute, margins of central I6be nearly obscuring callous, lateral lobes clasping column, apex of central lobe rounded or emarginate, of lateral lobes rounded or with auricles, yellow with green apex and margins; column 6-7.4 mm long; ovary 7.5-9.5 mm long, sessile. CAPSULE: 9-14 mm long, 4.5-5 mm diam., elliptic, pubescent, green. HABITAT: Woodlands (riverine, Polylepis sericea, or Polylepis weberbaueri), dense brushfields, or moist grasslands, often on bouldery sites. 3880-4300 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash. H. N. P.: Carhuazcancha (12247), Ishinca (9591, 11255), Llanganuco (Lôpez et al. 7437), Rurichinchay at Que. Pachachaca (12535), and Santa Cruz (9253). PHENOLOGY: Observed in flower from February to July, and fruiting in January. CONSERVATION STATUS: Uncommon.

Hmbenwlm Willdenow TERRESTRIAL, from tuberous roots and ovoid-fusiform root-stem tuberoids. LEAVES: spiraled, basal and rosulate and/or cauline, membranous, folded, not articulate. INFLORESCENCE: terminal, spikes or racemes. FLOWERS: resupinate, glabrous; floral bracts conspicuous, about as long as ovary, leafy; sepals subequal, free, laterals spreading, dorsal wider, galeate over column; lateral petals free, often connivent with dorsal sepal, entire or 2-lobed, commonly deeply so, lip basally adnate to column, entire or 3-lobed or lacerate, base spurred; column short. 238

Pantropical in distribution, this genus has about 600 species (Atwood, 1986; Garay, 1978), with IS occurring in Andean Peru (Dodson, 1988). Most the species have a preference for acid soils. POLLINATION: by moths and butterflies. As yet this genus has not been found in the Park, but is expected. The only high altitude species reported for Peru is H. pumiloides C. Schweinf.

Mmlmxim Solander ex O. Swartz Terrestrial, or epiphytic; rhizomatous, stem elongate or bulbous-thickened at base. LEAVES: distichous, cauline (Pichinchae group) or 1-several and basal (Fasciola group), membranous, long-sheathing bases, petiolate, not articulated. INFLORESCENCE; terminal, capitate or elongate spike or umbellate raceme; peduncle few-leaved. FLOWERS: small, resupinate or not, glabrous; floral bracts small; sepals subegual, free or the laterals connate basally, spreading; lateral petals free, narrow to filiform, lip sessile, entire to 2-3-lobed, entire to ciliate; column short. This is a large genus, having about 300 species (Atwood, 1986), of pantropical distribution (Dressier, 1981), with its greatest diversity in Asia and Oceania. Dodson (1988) lists 38 species for the Andes; 9 are Peruvian. POLLINATION: The pollination biology is unknown; flies are possible pollinators. TAXONOMY: In the Andes, the taxa fall into two species groups: Fasciola group - plants caespitose, stems developing pseudobulbs. Pichinchae group - plants creeping, stems without pseudobulbs. Our species is in the Fasciola group.

Malaxis mndlcolm (Ridley) Kuntze TERRESTRIAL, 12.5-32 cm tall, erect; pseudobulbs 2.5-4 cm long ovate, with a few scarious sheath-like bracts; glabrous. LEAVES; 1 bladeless sheath, 4-10.5 cm long, foliar, 2 fully developed leaves per shoot, 11-30.5 cm long, bases 6-16 cm long, open sheathing, blades 5-14.5 cm long, 2-7 cm wide, thin, elliptic, several nerved, secondary nerves conspicuously reticulate, apex acute, blade often spreading. INFLORESCENCE; umbellate raceme, 5-30 mm long, many-flowered; peduncles 9.5-30 cm long erect. FLOWERS: spreading, carnose, green; floral bracts 1.5-2.7 mm long, membranous, ovate-attenuate; sepals: laterals 3-3.2 mm long, 1.4-1.8 mm wide, ovate-acute, briefly united basally, pendent, dorsal 3.3-3.7 mm long, 1.3-1.4 mm wide, lanceolate, reflexed 239

onto ovary; lateral petals 3-3.2 mm long, linear, reflexed, lip 3.1-3.4 mm long, 4-4.2 mm wide, deltoid, deeply saccate, margin entire; column very short; ovary and pedicel 12-20 mm long. CAPSULE: 10-14 mm long, 5-7 mm diam. (immature, but well developed), green, perianth persistent on fruit. HABITAT: Growing on soil or in organic matter in understory of Polylepis sericea woods. 3700-3800 m. DISTRIBUTION: Venezuela to Peru. Peru: Ancash, Ayacucho, Cuzco, Huanuco, Junin, and La Liberté,d. H. N. P.: Llanganuco (8819) and Rurichinchay (12667). PHENOLOGY: Flowering has been observed in December and January, and early fruiting in June. CONSERVATION STATUS: Rare.

Mmsdevmlllm Ruiz et Pavôn References: Luer, 1986a, 1986b, 1987. EPI- and LITHOPHYTIC, and TERRESTRIAL; rhizome caespitose to creeping; stems short (termed ramicauls by Luer), base covered by 2-3, scarious, imbricate sheaths, one apical leaf. LEAVES: coriaceous, solitary and terminal on stem, petiolate or narrowed below, articulate. INFLORESCENCE: lateral, a solitary flower or few-flowered raceme; peduncle with 1-3 sheath-like bracts. FLOWERS: resupinate; floral bract# thin, tubular, hooded; sepals connate into a cup or long tube, lobes spreading, caudate, membranous to carnose; lateral petals smaller than sepals, narrow, callous, lip small, callous, attached to column foot, versatile; column short, winged, with short foot; pronounced joint between ovary and pedicel. Native to the high montane riions of the Neotropics from Mexico to southern Brazil, Masdevallia has about 350 species (Luer, 1986a, 1986b), with 85 species known for Andean Peru (C. A. Luer, pers. com.) POLLINATION: Pollination biology is unknown; hummingbirds are the likely pollinators for the red, purple, yellow, and yellow-orange flowered species. TAXONOMY: The genus has been divided into 5 subgenera and 17 sections by Luer (1986b). I thank Dr. C. A. Luer (MO) for advice regarding this genus. 240

MtidtYlllll imiWlll Reichb. f. (including M. purpwrina Schltr. and M. venusta Schltr.) LITHOPHYTIC or TERRESTRIAL, erect from densely caespitose rhizomes, forming clones to IS cm diam.; stems 1.3-4.2 (-5) cm long, bract 1, basal, scarious; glabrous. LEAVES: solitary and terminal on stem, (4.8-) 6.5-14.5 cm long, 0.8-1.7 cm wide, fleshy, oblanceolate, basally petiolate, margins entire. INFLORESCENCE: a solitary flower per stem; peduncle (8-) 15-26.1 cm long, erect, 1 (-2) scarious, sheath-like bracts, ca. 10 mm long; pedicel 2.5-4 cm long. FLOWERS: deep magenta or scarlet; floral bract ca. 10 mm long, scarious, sheath-like; sepals: tube 11-16 (-20) mm long, limb of laterals 26-33 mm long, 13-22 mm wide, basally connate for ca. 1/2 their length above the tube, limb of dorsal 25-34 mm, narrow, all limbs spreading, with 3, arcuate nerves, apices caudate; corolla hidden well within calyx tube, lateral petals ca. 8 mm long, ca. 3 mm wide, oblong, carnose with calloused tip, lip ca. 7 mm long, ca. 3 mm wide, obovate, carnose with calloused tip. CAPSULE: not seen. HABITAT: Growing in soil pockets on old boulder falls and talus slopes. 3580-4420 m. DISTRIBUTION: Northern Peru : Amazonas, Ancash and Cajamarca. H. N. P.: Alpamayo-Cashapampa trail (10034), Auquispuquio at Lago Veronica (12116), Llanganuco (8790; Sagàstegui et al. 12316) and Ranincuray (9146). PHENOLOGY: Flowering observed from late December until June. Luer (1987) states that this species is widespread and variable in size and color. The flowers can be magenta, rose, scarlet or white; to date no white flowered specimens have been found within the Park. Subspecies venusta Schltr. has large bright reddish flowers; the specimens from Llanganuco are of this subspecies. CONSERVATION STATUS: Uncommon; merits protection.

Mvrosmodes H. G. Reichenbach References: Foldats, 1969; Garay, 1978. TERRESTRIAL, from a fascicle of deeply buried tuberous roots; plants dwarfed, mostly immersed in soil. LEAVES: spiraled, in a basal rosette, membranous, sheaths immersed in soil, not articulate, leaves partially to fully developed at anthesis. INFLORESCENCE: lateral, from root crown, dense, often conical, many-flowered spikes; peduncles immersed in soil, covered by imbricate, tubular, sheath-like, scarious bracts, often darkened at apex. FLOWERS: small, not resupinate, glabrous, white to greenish white; floral bracts shorter than flowers, ustulate; sepals subequal, briefly connate basally into 241

short tube, dorsal adnate to column; lateral petals narrowly oblong to oblanceolate, adnate to column. Hp erect, concave, hooded to tubular, margins fimbriate and never involute, base clawed and cordate or sessile and cuneate, blade apex sometimes slightly 3-lobed; column short, winged; apex of ovary either short or elongated into a long, narrow neck. This genus is restricted to the paramos and punas of the Andes, and has ca. 12 species, four of which are Peruvian (Dodson, 1988). POLLINATION: The pollination biology is unknown. It is likely that the populations are self-pollinating, and that any out-crossing is fly mediated. TAXONOMY: This genus has long been confused with-^a and Altensteinia^ and most of its species have been assigned to one or both of them. All three genera are in need of systematic study. Within the genus there are two, unnamed species groups: a. Lip clawed, base cordate; apex of ovary elongated into a long neck; perianth insertion on ovary noticeably oblique, perianth inflated and globose, dorsal sepal and petals reflexed. This group includes M. paludosa. b. Lip sessile, base cuneate; apex of ovary short, not an elongate neck; perianth insertion on ovary oblique, but subtly so, perianth dorsiventrally compressed, dorsal sepal and petals erect. M. nubigena is included here.

Key to the species of Mvrosmodes la'T Apex of ovary elongated into a long, narrow neck; inflorescence conic M. paludosa lb. Apex of ovary not elongated; inflorescence hemispheric, at most slightly elongated. M. nubigena

Mvrosmodes nubigena Reichb. f. TERRESTRIAL from a cluster of deeply buried tuberous roots; plant immersed in soil, only blades and tip of inflorescence exposed; glabrous. LEAVES: 2-5 cm long, exposed portions in a rosette-like array, sheath 1.9-2.5 cm long, closed, tubular-funnelform, scarious, blades 1.3-2.5 cm long, 0.7-1.2 cm wide, membranous, spatulate, petiolate below, apex acute. INFLORESCENCE: 5.5-6.5 total length; peduncle hidden by scarious bracts, tubular, broadly funnelform, mouth oblique; spike 1.5-2 cm long, hemispheric to slightly elongate. FLOWERS: ca. 13 mm long, white; floral bracts ca. 12 mm long, ca. 15 mm wide, broadly flabellate to orbicular, scarious, straw colored with 242

dark apices, clasping the flower; perianth dorsi-ventrally compressed, not conspicuously, inflated, insertion on ovary apex slightly oblique; sepals: laterals 6.5-7.5 mm long, 2.3- 2.9 mm wide, thickened, oblong with a slight constriction below the apex, 1-nerved, carinate, galeate, apex subacute, connivent with lip, dorsal 4.3-4.5 mm long, 2-2.2 mm wide, thickened, oblong with a slight constriction below the tip, 1-nerved, apex sub­ acute, erect; lateral petals 4.2-5 mm long, 0.6 mm wide, narrowly elongate, 1-nerved, laterally arcuate towards center of flower, apex rounded, margin irregularly fimbriate, lip 5-5.5 mm long, 5.9-7.4 mm wide, transversely elliptic, base cuneate, apex shallowly 3-lobed, margin fimbriate apically with moniliform hairs 0.5-1.5 mm long, dorsiventrally compressed with margins folded toward the center of petal and 2 broad, thickened calli at fold axes; ovary apex short, not produced into a neck. CAPSULE: not seen. HABITAT: Moist spots in grasslands. 4200-4400 m. DISTRIBUTION: Ecuador and Peru. Peru: Ancash and Cajamarca. H. N. P.: Honda (11674). PHENOLOGY: Beginning to flower in October. CONSERVATION STATUS: Very rare, collected only once.

MvroimodM ptludosâ (Reichb. f.) D. N. Smith TERRESTRIAL, from a cluster of deeply buried tuberous roots; plant immersed in soil, only blades and tip of inflorescence exposed; glabrous. LEAVES: exposed portions in a rosette-like array, sheaths 1.3-2.5 cm long, scarious, blades 1.7-4 cm long, 0.9-2.1 cm wide, ovate, petiolate below, apex attenuate. INFLORESCENCE: 4.5-7 cm total length, rarely exserted to 15 cm at maturity; peduncle hidden by scarious bracts, tubular- funnelform, mouth oblique; spikes 1.5-3 (-4.3) cm long, conical, rarely elongate and sub-cylindric. FLOWERS: 10.5-14.5 mm long, white to whitish green, with a faint foetid odor; floral bracts 9-13.5 mm long, 4-10 mm wide, broadly obovate to elliptic, scarious; perianth globose, fleshy, insertion on ovary distinctly oblique; sepals: laterals 2.2-3.7 mm long, 1.5-2.2 mm wide, elliptic, I-nerved, erect, concave, margin minutely erose, apex rounded, connivent with lip, dorsal 2.3-3.4 mm long, elliptic, 1-nerved, margin minutely erose, apex rounded, reflexed outwards; lateral petals 2-3.2 mm long, 0.7-1.2 mm wide, oblanceolate, apices reflexed outwards, lip 3-4.5 mm long, 3-5.5 mm wide, claw 0.5-1 mm long, blade cordate, unlobed, galeate, carnose, calli at base of blade, glabrous within, margin densely fimbriate with moniliform hairs to 1 mm long; ovary apex narrowed into an elongate neck 2.5-5 mm long. CAPSULE: not seen. 243

HABITAT: Moist grasslands, on soils of morainal and alluvial origin. 3900-4350 m. DISTRIBUTION: Peru to northern Argentina. Peru: Ancash, Cuzco, Junin, and Lima. H. N. P.: Alpamayo (9768), Ishinca (9549), Rajucolta (12186), Rurichinchay at Pachachaca (12691) and Shallap (9675). PHENOLOGY: Flowering is seen from February to June, and fruit formation from April onwards. CONSERVATION STATUS: Uncommon.

Odontogiosaum Humbolt, Bonpland et Kunth EPI- and LITHOPHYTIC; rhizomes caespitose to short creeping; pseudobulbs I-node, subtended by several pairs of basal leaves, often bladeless, and with 1-3 apical leaves. LEAVES: distichous, coriaceous to carnose, articulate. INFLORESCENCE: lateral and basal, a raceme or panicle or solitary flower; peduncle with a few scarious bracts. FLOWERS: large, showy, resupinate, glabrous; floral bracts small to equalling length of ovary; acpals subequal, free or laterals connate basally, spreading; lateral petals like the sepals, shorter, broader, lip clawed, oriented parallel the column or briefly adnate to it, entire or 3-lobed, callus at base, margins entire to lacerate, apex reflexed; column short to long, winged or not. This Neotropical genus grows in mountainous regions from Mexico and the West Indies to Bolivia, with its center of diversity in Colombia. It has 140 species (Atwood, 1986). Dodson (1988) estimates 176 species of Odontoglossum for the Andes, 40 are found in Peru. The genus is under critical study by various authors. Haibinger (1982, 1983, 1984) has divided the genus so that there is no species of Odontoglossum sensu stricto in Mexico or Central America. Dodson et al. (1987) reduce all Andean members of this genus to a group within Oncidium. POLLINATION: By bees.

Odonfoglossum rigldum Lindley TERRESTRIAL; to 77.5 cm tall; rhizome very short creeping, clones to 40 cm diam.; pseudobulbs 5.5-9 cm long, elliptic, laterally compressed; glabrous. LEAVES: 4-5 basal, sheaths 6-10.5 cm long, chartaceous, ovate-truncate, persistent, blades 5.4-19 cm long, I.3-2 cm wide, oblanceolate-attenuate, 1-2 apical on pseudobulb, sheaths 1.1-2.6 cm long, coriaceous, blades 15.2-25 cm long, 1.2-2.4 cm wide, oblanceolate-attenuate. 244

INFLORESCENCE: (6-) 12-20-nowered panicle, 11.2-25.5 cm long, 1 bract like floral bracts in texture at branch nodes; peduncle 42.5-55.3 cm long, erect, with 3-4 membranous, basally closed bracts, 10-18 mm long, ovate. FLOWERS: yellow; floral bracts 6-12 mm long, membranous, ovate, apex acute to attenuate; sepals: laterals 13.5- 16 mm long, 3.6-4 (-6.5) mm wide, asymmetrically ovate-attenuate, 3-nerved, margins entire, connate into synsepal, joined ca. 1/2 length, apices divergent, notch 6-9 mm deep, dorsal 10.5-15 mm long, 5.4-6.3 mm wide, ovate-acute, 5-nerved, margins entire; lateral petals 10.5-14 mm long, 4.5-5.7 (-7) mm wide, asymmetrically ovate, apex acute to attenuate, 3-nerved, margins entire, lip 15-18 mm long, 13.2-17 mm wide, claw 2.8-4 mm long, blade broadly obovate, base lobed and auriculate, callus well developed, a series of ridges and papillae, margin entire, apex mucronate; column 4.5-5 mm long, wings 1.5-2 mm long, lunate, margin lacerate; ovary and pedicel continuous, 15-28 mm long. CAPSULE: ca. 3.8 cm long, ca. 1.8 cm diam., elliptic, green with reddish tint, perianth persistent. HABITAT: In soil pockets on rocky or bouldery sites in grasslands. 3700-4200 m. DISTRIBUTION: Ecuador to Bolivia. Peru: Ancash and Cuzco. H. N. P.: Llanganuco (11456), Parôn (11539), and Rurichinchay at Pachachaca (12615). PHENOLOGY: Collected in flower in June and September, and in fruit in June. This taxon, O. aureum Lindley and O. mystacimm (Lindley) Lindley, form a closely related species group (C. H. Dodson, pers. com.). Odontoglossum aureum has yellow flowers, a simple inflorescence, and is distributed from Ecuador to Bolivia; O. mystacimm has orange yellow flowers, a branched inflorescence, and its distribution is wholly Peruvian; and O. rigidum has yellow flowers, a branched inflorescence, and is distributed from Ecuador to Peru. CONSERVATION STATUS: Rare.

Pleurothallia R. Brown in W. Aiton et W. T. Aiton References: Luer 1986a, 1986c. EPI- and LITHOPHYTIC, and TERRESTRIAL; rhizomes caespitose or creeping; stems erect, crowded or distant, simple or branched, covered below with tubular, sheath-like bracts, often with flaring hispid mouths. LEAVES: coriaceous, solitary and terminal on stem, sessile to petiolate, articulate. INFLORESCENCE: lateral or terminal, racemose or solitary; peduncles solitary or fasciculate, with tubular bracts. FLOWERS: small to large, resupinate or not, glabrous or pubescent; floral bracts usually inconspicuous. 245

tubular or hooded; sepals: laterals free to entirely connate, concave under column, dorsal free or partly connate with laterals, membranous or fleshy; lateral petals much smaller than sepals, membranous or fleshy, longer than broad, margins often fringed, lip subequal petals, membranous or fleshy, clawed, simple or lobed, margin often fringed; column short to elongate, often winged, sometimes with foot; pronounced joint between ovary and pedicel. This variable genus is found throughout the Neotropics, and is most numerous in mountainous regions. The exact number of species is unknown; estimates range from 600 (Ames and Correll, 1952) to 1500 (Atwood, 1986). Luer (1986c) found over 2000 names published in the genus; he lists ca. 600 as species representing the sections of Pleurothallis. Dodson (1988) lists 135 species of Pleurothallis for Andean Peru; Luer (pers. com.) estimates that the number of species is much higher. This is the largest genus of Neotropical orchids. POLLINATION: Pollination biology unknown; Bradesiid flies are the probable pollinators (Dodson, pers. com.). TAXONOMY: The genus has been divided into 29 subgenera and 22 sections by Luer (1986c). I thank Dr. C. A. Luer (MO) for his advice regarding this genus.

Key to the species of Pleurothallis la. Rhizome creeping, sub-scandent; inflorescences few-flowered. P. spiralis lb. Rhizome caespitose; inflorescences densely, many-flowered. P. triiineata

Pleurothallis snlralls (Ruiz Lôpez et Pavôn) Lindiey LITHOPHYTIC, or terrestrial in organic matter; rhizome elongate creeping, densely covered by membranous bracts, producing abundant adventitious roots; stems short, mostly hidden by rhizome bracts; glabrous. LEAVES: 3.4-9.5 cm long, 5-12 mm wide, obovate, short petiolate, margins coriaceous, apex acute, minutely bilobed, costa projecting as blunt mucro between lobes. INFLORESCENCE: terminal, 3-(few?)- flowered raceme, wiry, flexuous, 4-8 cm long in total appears lateral as stems are hidden. FLOWERS: 12.5-15 mm long, resupinate, yellow with red linear markings; floral bracts 2.5-4.5 mm long, membranous, funnelform, mouths widely flairing, strongly oblique; sepals: membranous, laterals 10,5-11.8 mm long, 2.5-3 mm wide, asymmetrically arcuate, 1-nerved, connate into synsepal, joined for 3/4 of length, notch 246

2.5-3.4 mm deep, bi-keeled, dorsal free, ovate-attenuate, 3-nerved; lateral petals 5-5.5 mm long 1.8-2.6 mm wide, obovate, 1-nerved, margin entire, lip 7-7.7 mm long, 4- 4.8 mm wide, fleshy, hinged to column foot, 3-lobed, central lobe elongate, ligulate, rounded, lateral lobes auriculate, erect, callus central, elongate, margin entire; column 3- 3.7 mm long, arcuate; ovary 3-4 mm long. CAPSULE: immature, ca. 10 mm long, ca. 4 mm diam., green. HABITAT: Growing on bouldery sites in shrublands of the warmer valleys, especially on microsites with adequate moisture. 3580- 3900 m. DISTRIBUTION: Ecuador and Peru. Peru: Ancash, Cajamarca, and Huanuco. H. N. P.: Auquispuquio (12006), and Llanganuco (8820; Mostacero et al. 619; Sagàstegui et al. 1231; Todzia et al. 2607). PHENOLOGY: Flowering and fruiting is seen from December to April. CONSERVATION STATUS: Rare.

Pleurothalll» trlllmemtm Lindley LITHOPHYTIC; rhizome caespitose; forming clones to 25 cm diam.; stems 5-17.5 cm long, with 3 membranous, tubular, sheath-like bracts, to 6.5 cm long, lower bracts overlapping, uppermost separate; stem often branching from same annulus that gives rise to inflorescence; adventitious roots produced at branch points; glabrous. LEAVES: 5.5- 11.9 cm long, 1-1.7 cm wide, oblong, short petiolate, margins coriaceous, apex rounded, minutely bilobed, costa projecting as blunt mucro between lobes. INFLORESCENCE: terminal, erect, densely 26-32-flowered raceme, 8.5-10 cm long, peduncle 1-1.5 cm long, subtended by 1-2 membranous bracts. FLOWERS: 6-8.5 mm long, resupinate, pale yellow with purple lines on petals and lip; floral bracts 2.4-5.2 mm long, membranous, funnelform, mouths widely flaring, strongly oblique; sepals: membranous, laterals 5.3-5.7 mm long, 1.4-1.8 mm wide, asymmetrically arcuate, 3-nerved, connate into synsepal, joined to 1/2 length, bi-keeled, dorsal 4.5-5.9 mm long, 2.6-3.3 mm wide, briefly connate basally with synsepal, ovate-acute, 3-(S)-nerved; lateral petals 2.8-3 mm long, 0.8-1.1 mm wide, obovate, 1-nerved, margin entire, lip 2-2.1 mm long, 1.2-1.7 mm wide, fleshy, 3-nerved, hinged to foot, 3-lobed, central lobe elongate, rounded, lateral lobes at midpoint of lip, auricular, erect, margin entire; column 1.2-1.6 mm long; ovary 2-3 mm long. CAPSULE: not seen. HABITAT: On bouldery sites in Polylepis sericea woodlands and brushfields in the bottom of a relatively arid river valley. 3580-3850 m. 247

DISTRIBUTION: Peru: Ancash, Apurimac, Cerro de Pasco, and Huanuco. H. N. P.: Llanganuco on Maria Josefa trail (8786). PHENOLOGY: Flowering was seen in December and January. CONSERVATION STATUS: Very rare, collected only once.

Pttriclll» Lindley Reference: Garay, 1978. Terrestrial, from a fascicle of tuberous roots; pubescent. LEAVES: in a basal rosette, membranous, not articulated; withered or undeveloped at anthesis. INFLORESCENCE: lateral, a spike or raceme; peduncle erect, unbranched, covered by tubular, sheath-like bracts. FLOWERS: medium sized, not resupinate; floral bracts ovate to lanceolate; sepals subequal, carnose laterals free, dorsal pendulous; lateral petals narrower than dorsal sepal, free, often connivent with dorsal sepal, lip adnate to column, carnose, deeply concave, often 3-lobed, surrounding column at base, margin often has row of glands; column short. Pterichis is an Andean genus with outlying species in Costa Rica and Jamaica. In South America it is found from Venezuela and Colombia to Bolivia, with some representatives in Brazil; Peru is possibly the center of dispersion (Foldats, 1969). It has ca. IS species; 8 of them are Peruvian (Dodson, 1988). POLLINATION: Pollination biology is not known; the glands on the lip suggest fly pollination (C. H. Dodson, pers. com.).

Pterichis triloba (Lindley) Schltr. TERRESTRIAL, 17-39 cm tali, from a cluster of tuberous roots, breaking off readily, often forming clones; peduncles, bracts, ovaries, and sepals pubescent with multicellular, glandular hairs, often densely so. LEAVES: basal, usually not present during flowering, 1 (-2) per plant, 17 cm or more long, 1.4-2 cm wide, spatulate, petiolate portion 5.5-7 cm long, margins entire. INFLORESCENCE: lateral, erect, 7-21-flowered raceme, 5.5- 12.5 cm long; peduncles 10.5-27.2 cm long, 3-5 membranous bracts, the lower sheath­ like, 5-21 mm long, mouth strongly oblique, apex blade-like, 5-17 mm long, the basal sheaths sometimes woolly, uppermost simple, 13-17 mm long. FLOWERS: 14-20 mm long; with a sweet odor at anthesis; floral bracts 11-13 mm long, 5-5.8 mm wide, ovate- attenuate to lanceolate, membranous, 3-nerved; sepals: laterals 9-11 mm long, 5.7-5.9 mm wide, asymmetrically ovate-attenuate, 3-nerved, concave, erect, connivent, margins 248

tending to be involute, olive green, dorsal 11-11.2 mm long, 3.9-4.4 mm wide, elliptic- attenuate, 3-nerved, reflexed downward, olive green; lateral petals 10-11.5 mm long, 2- 2.8 mm wide, asymmetrically lanceolate-attenuate, arcuate, 3- nerved, often reflexed downward and connivent with dorsal sepal, olive green. Hp 5-6.6 mm long, 8.2-11 mm wide, transversely reniform, carnose, deeply saccate, 3-lobed, the laterals erect, auriculate, terminal lobe ovate to oblong, 1.6-2.7 mm long, reflexed, thin at margin, blade densely long-papillose towards margins, calloused below terminal lobe, yellow with dull brownish lines; ovary and short pedicel continuous, 11-13 mm long. CAPSULES: 15 mm or more long (only immature seen), oblong to obovate, green. HABITAT: In soil pockets on talus slopes, or on grassy slopes. 3700-4100 m. DISTRIBUTION: Colombia to Peru. Peru; Ancash, Ayacucho, Cuzco, Huanuco, Lima, and Puno. H. N. P.: Carhuazcancha at Ichicpotrero (12390), Cahuish-Machac road (Ldpez et al. 7560), Ranincuray (10480), and Rurichinchay at Pachachaca (12570). PHENOLOGY: Found in flower and early fruit development in April and June. CONSERVATION STATUS: Uncommon.

StclU Swartz References: Garay, 1979; Luer, 1986a. EPI- and LITHOPHYTIC, from caespitose or creeping rhizomes; stems short, covered by tubular, sheath-like bracts below, branching in some species; glabrous. LEAVES: coriaceous, solitary and terminal on stem, subsessile to petiolate, articulate. INFLORESCENCE: terminal, racemose, 1-several per stem, arising from an annulus slightly below leaf. FLOWERS: small to medium, resupinate or not, rotate or bilabiate; floral bracts often minute; sepals subequal or dorsal larger, all connate into a shallow cup, or dorsal free and laterals connate, usually spreading widely; lateral petals much smaller than sepals, broad with thickened margins, these often with abundant glands, lip small, usually wider than long, subsessile, simple or 3-lobed, carnose; column short, foot present or absent, apex 3-lobed; pronounced joint between ovary and pedicel. Distributed throughout the Neotropics from Mexico to Brazil and Bolivia, Stelis is a genus of ca. 280 species according to Garay (1979). Luer (1986a) estimates 500 species and Dodson (1988) more than 600 species. There are ca. 73 species in Andean Peru (Dodson, 1988). Luer (pers. com.) estimates that the number of species, both for the genus and in Peru, is much higher. 249

POLLINATION: Pollination biology is unknown; flies are the likely pollinators (C. H. Dodson, pers. com.). I thank Dr. C. A. Luer (MO) for his advice regarding this genus.

Key to the species of Stella la. Flowers bilabiate. S. cf. cupuligera lb. Flowers rotate. 2. 2a. Rhizome caespitose; flowers purple. S. cf. leucopogon 2b. Rhizome creeping; flowers yellow with purple petals and lip. S. cf. flexuosa

Stella cf. cupullgera Reichb. f. & Warsc. LITHOPHYTIC, to 30 cm tall; rhizome caespitose; stems 9-13.5 cm tall, 2-3, membranous, tubular, sheath-like bracts, uppermost 4.5-6 cm long. LEAVES: 8.3-10.8 cm long, 0.9-1.5 cm wide, oblanceolate, short petiolate, margin coriaceous, apex acute, minutely bilobed, costa projecting as blunt mucro between lobes. INFLORESCENCE: 1, rarely 2, per stem, 14-25-flowered raceme, 8.5-12 cm long; peduncle 5-7 cm long, subtended by a membranous bract; inflorescence longer than leaves, rachis faintly zig­ zag. FLOWERS: bilabiate; floral bracts 2.5-8 mm long, funnelform, mouths widely flaring, strongly oblique; sepals: yellow, laterals 2.8-4.2 mm long, 2.5-3 mm wide, asymmetrically ovate-acute, 3-nerved, margins entire, connate into synsepal, joined 80- 90% of length, saccate, dorsal 4.5-5.4 mm long, 2.9-3.6 mm wide, ovate-acute, 3-5- nerved, connate with synsepal for 1/3-1/2 of length; corolla hidden within calyx, color not seen; lateral petals 1 mm long, 1.5-1.6 mm wide, tranversely elliptic, 3-nerved, margin entire, apex thickened, lip 0.8-1 mm long, 1.1-1.3 mm wide, broadly ovate, thickened, bicallous, apex acute, margin entire; column ca. I mm long. CAPSULE: immature, 10 mm long, green. HABITAT: Growing on large boulders, in mesic, tall brushfields. 3970 m. DISTRIBUTION: Peru: Ancash. H. N. P.: Carhuazcancha at Ichicpotrero (12410). PHENOLOGY: Found in late flower and early fruit in May. CONSERVATION STATUS: Very rare, collected only once.

Stellg cf. fiexuosa Lindley LITHOPHYTIC,to 12 cm tall; rhizome creeping, producing adventitious roots, clones to 40 cm diam.; stems 3.5-6.2 cm long, 2-3, membranous, tubular, sheath-like bracts; 250

glabrous. LEAVES; 3.9-6.7 cm long, 5.5-8 mm wide, oblanceolate, short petiolate, margins coriaceous, apex acute, minutely bilobed, costa projecting as blunt mucro between lobes. INFLORESCENCE: terminal, 14-22-flowered, erect raceme, 35-61 mm long, rachis zig-zag; peduncle 11-20 mm long, subtended by 1-2, tubular membranous bracts. FLOWERS: fleshy, exserted, nutant; floral bracts 1.6-4.1 mm long, membranous, funnelform, mouths widely Hairing, strongly oblique; sepals: yellow to yellowish green, basally connate, laterals 1.6-2.5 mm long, 1.2-1.9 mm wide, widely ovate, 3-nerved, margins entire, apex acute, dorsal 1.8- 2.2 mm long, 1.2-1.7 mm wide, ovate, 3-nerved, apex acute; corolla purple; lateral petals 0.7-0.8 mm long, 0.8-0.9 mm wide, broadly obovate, apex thickened, lip 0.5-0.8 mm long, 0.6-0.7 mm wide, hinged to column foot, elliptic, bicallous, apex rounded; column 0.7-0.8 mm long; ovary 1.1-1.4 mm long. CAPSULE: not seen. HABITAT: Rocky sites in open Polylepis sericea woodlands, brushfields, and grasslands. 3500-3950 m. DISTRIBUTION: Peru: Amazonas and Ancash. H. N. P.: Alpamayo-Cashapampa trial (10033), Llanganuco (8792; Todzia 2605), and Rurichinchay at Pachachaca (12645). PHENOLOGY: Observed flowering in March. CONSERVATION STATUS: Rare.

Stells cf. leucoBOBon Reichb. f. LITHOPHYTIC; rhizome caespitose, clones large, to 50 x 100 cm; stems 5.5-18 cm long, simple or branched, then the branch arising from same annulus as inflorescence, 2-4, membranous, tubular, sheath-like bracts, to 7,2 cm long; glabrous. LEAVES: 5.2-10.3 cm long, 1-2 cm wide, elliptic, short petiolate, margins coriaceous, apex acute, minutely bilobed, costa projecting as a blunt mucro between lobes. INFLORESCENCE: terminal, erect, 16-26-flowered raceme, 8.5-11 cm long, rachis slightly zig-zag; peduncle 2-4.4 cm long, subtended by I, membranous bract. FLOWERS: dark purple; floral bracts 2.8-8 mm long, membranous, mouths widely flairing, strongly oblique; sepals: basally connate, laterals 3.9-4.8 mm long, 3-3.9 mm wide, 3-(5)-nerved, asymmetrically ovate, apex acute, dorsal 4.6-5.2 mm long, 3.3-4.4 mm wide, ovate, (3)-5-nerved, apex acute; lateral petals 0.8-1.3 mm long, 1.5-2.1 mm wide, transversely lunate, 3-nerved, apex strongly thickened, lip 1-1.6 mm long, (1.3-) 1.7-1.9 mm wide, rhombic, base cuneate, swollen in center, bi-callous, margins entire, apex acute; column subequal lateral petal length. CAPSULE: not seen. 251

HABITAT: Rocky sites, including cliffs and talus slopes, in Polylepis sericea woodlands and shrublands in warmer valleys. 3580-3950 m. DISTRIBUTION: Peru: Ancash. H. N. P.: Alpamayo-Cashapampa trail (10035), Llanganuco (8789; Todzia 2609), and Ranincuray (9041, 10478). PHENOLOGY: Plants have been found flowering from December to March, and in early fruiting stage in April. CONSERVATION STATUS: Rare.

Trichoceros Humbolt, Bonpland et Kunth EPI- and LITHOPHYTES, or TERRESTRIAL; rhizome elongate; pseudobulbs 1-node, ovate flattened. LEAVES: 2 or more, distichous, bases sheathing pseudobulbs, blades small, coriaceous to carnose, articulated. INFLORESCENCE: lateral and basal, racemes; peduncles 1-2 per pseudobulb. FLOWERS: medium, resupinate, glabrous; floral bracts small; sepals subequal, free, spreading; lateral petals free, like the sepals, lip sessile, 3- lobed, lateral lobes spreading to erect, often clasping the column, central lobe prominent; column short, bristly, lacking wings or foot. This genus of 5 species is distributed from Colombia to Bolivia, with 2 species in Peru (Dodson, 1988). POLLINATION: Pseudocopulation by bristly tachinid flies.

Trlchoceroa platvceros Reichb. f. LITHOPHYTIC, rhizome internodes short to elongate; adventitious roots abundant; pseudobulbs 10-16 mm long, 5-9 mm wide, shrivelling markedly on drying. LEAVES: 2 at base of pseudobulb, 1 apical, 23-45 mm long, 5-9.5 mm wide, carnose, elliptic, apex acute. INFLORESCENCE: 2.5-9.5 cm long, 3-7- flowered, erect; peduncles 4-12 cm long, with 1-3 bracts, similar to floral bracts. FLOWERS: to ca. 20 mm long; floral bracts (2.8- ) 3.7-4.7 mm long, membranous, elliptical, ca. 5-nerved, apex acute; sepals: laterals 9-9.8 mm long, (3.4-) 4-4.8 mm wide, dorsal 10.3-11.5 mm long, 5-6.5 mm wide, both elliptic, ca. 5-nerved, apices acute, pale yellow with dark purple, nerve-like markings, dark purple at base; lateral petals 10-11 mm long, 5.8-6.1 mm wide, elliptic, ca. 5-nerved, apex acute, pale yellow with dark purple, nerve-like markings, dark purple at base. Up 10-11.5 mm long, 6.6-8 mm wide, 3-lobed, central lobe prominent, obovate, like sepals and lateral petals in coloration, margin minutely ciliate callous at base, swollen, dark purple, with bristles, lateral lobes 2-2.2 mm long, 1.8-2 mm wide. 252 auriculate, clasping column, with bristles, dark purple; ovary and pedicel continuous, 7- 14 mm long. CAPSULE; not seen. HABITAT: Bouldery areas in dry brushlands or Polylepis sericea woodland. 3870-3700 m. DISTRIBUTION: Colombia to Peru. Peru: Ancash and Cajamarca. H. N. P.: Auquispuquio (12008), Llanganuco (10516), and Santa Cruz (9351). PHENOLOGY: Observed in flower in January and May. In Orchids of Peru (Schweinfurth, 1961), this species is included in Trichoceros muralis Lindiey as a variety. That species is an Ecuadorian endemic with a distribution centered around Cuenca (Dodson, pers. comm.). CONSERVATION STATUS: Rare. 253

POTAMOGETONACEÂE Dumortier ALISMATIDAE: NAJADALES References: Haynes and Holm-Nielsen, 1982, 1986b. HERBS, perennial; aquatic; rhizomatous; stoloniferous to erect; rooted on the bottom, stems and leaves submerged or floating; glabrous. LEAVES: alternate to verticillate, base sheathing to tubular, margins hyaline joining into a ligule, blades linear to broad, venation often arcuate-convergent; leaves dimorphic, the submerged leaves sessile with less developed blades, the floating leaves petiolate with well developed blades. INFLORESCENCES: axillary or terminal, scapose, ebracteate spikes, the upper inflorescences erect and above the water. FLOWERS: small, perfect, regular, hypogynous; perianth 4 tepals, clawed, valvate, carnose; stamens 4, opposite and basally adnate with tepals, anthers sub-sessile; gynoeclum of 4, free carpels, placentation marginal, style short or wanting, stigma dry, ovule solitary. FRUIT: achenes or drupelets, pericarp aerenchymatous and buoyant. The family is of cosmopolitan distribution and has 3 genera and ca. 100 species. Ruppia filifoUa (Philippi) Skottsb. is often found in volcanic crater lakes throughout the Andes (Haynes & Holm-Nielsen, 1986b). HABITAT: In fresh to brackish waters, from sea level to high elevations. POLLINATION: Wind pollinated. USES: Food resource for water fowl (Martin et al., 1961).

Potamogeton Linnaeus HERBS, aquatic, from delicate rhizomes. LEAVES: blades of floating or emergent leaves well developed, membranous, blades of submerged leaves atrophied or partially decomposed, reduced to vascular skeletons, blade shape variable, margins usually entire, petiolate, ligule or stipules present. INFLORESCENCE: axillary, pedunculate spikes. FLOWERS: bisexual; perianth members carnose, clawed; stamens 4; ovaries 4, free, sessile, style short or wanting. FRUIT: druplets. Potamogeton has ca. 100 species widely distributed in the Northern Hemisphere, and with a few species in South America. There are ca. 4 species in Peru.

Potamogeton paramoanus R. Haynes & Holm-Niels. HERB, aquatic, rooted in pond bottom, apical leaves floating; glabrous. LEAVES: slightly dimorphic; submerged leaves distichous, 5.5-7.5 cm long, 2.5-3.4 mm wide. 254 linear, base attenuate, apex long acuminate, margins entire to sinuate, 4-6 lines of lacunae on each side of costa, ligule 3.5-7 mm long, delicate membranaceous, lacerate, floating leaves 3.1-3.8 cm long, 5-8 mm wide, narrowly lanceolate, base attenuate, blade lacking lines of lacunae, margins entire, petiole 1.3-2.2 mm long, ligule 12-15 mm long, delicate membranaceous, ovate, entire, surrounding the stem. INFLORESCENCE: axillary spikes; peduncle 2.7-4 cm long. FLOWERS: perianth members 2 mm long, claw 1 mm long, blade ca. 1 mm dia, widely elliptic, green; stamens ca. 0.8 mm long; carpels ca. 1 mm long, style nearly iobsolete, stigma capitate. FRUIT: not seen, reported to be a laterally winged drupelet. HABITAT: Ponds at high elevation. 4200m. Haynes and Holm-Nielsen (1982) report that this species grows in ponds, lakes and streams in water with pH values of 6.5-7.5. DISTRIBUTION: In the Andes from Venezuela to Bolivia, and with one outlying population in Costa Rica. Peru: Ancash. H. N. P.: Paria, a lateral valley of Huaripampa (9168). PHENOLOGY: Flowering in January. COMMENT: "Potamogeton paramoanus and P. sclerocarpus probably represent relic populations of a once much more widespread taxon which now survives only in the higher mountains of South America," (Haynes & Holm-Nielsen, 1982). Potamogeton sclerocarpus K. Sch. inhabits the Brazilian highlands. CONSERVATION STATUS: Rare. 255

ZANNICHELLIACEAE Dumortier ALISMATIDAE: NAJADALES Reference: Haynes & Holm-Nielsen, 1986c. HERBS, aquatic, plants submerged; stems stoloniferous below, erect, much branched; rooted to the bottom; glabrous. LEAVES: alternate, opposite, and in whorls of 3 on the same plant, blades sessile, linear, entire, 1-3-nerved; intravaginal scales filiform, stipule­ like, a lateral pair at each node. INFLORESCENCE: 2-flowered, one staminate and one carpellate, axillary; separated by 1-3 nodes. FLOWERS: small; unisexual, plants monoecious; perianth wanting; staminate flower a single stamen, anther connective sometimes elongated; carpellate flower a gynoecium of (1-) 4-5 (-8) free carpels subtended by a membranous cupule, placentation pendulous, style short, stigma funnelform, dry, ovule solitary. FRUIT: drupelets, endocarp spiny. This is a cosmopolitan family of 4 genera; only Zamichellia occurs in the Americas. TAXONOMIC NOTES: The membranous cupule subtending the female flower is interpreted as a bract by Haynes and Holm-Nielsen (1986c), and as a perianth by Dahlgren et al. (1985). The fruit is drupaceous. The exocarp and mesocarp are of a softer texture than the hard, spiny endocarp, and often fall away exposing the endocarp.

Zmmnkhelllm Linnaeus This genus of 2-3 species has the same characters as the family. Dahlgren et al. (1985) suggest that the genus may be monotypic, with Z. palustris L. being the only species. As such it would be a highly variable taxon of subcosmopolitan distribution. In the Flora of Peru (Macbride, 1936), there are several reports of Z. palustris for Peru. One of the collections cited is from Callao (Lima Department), which is a sea level site; the rest are from highland sites. From their studies, Haynes and Holm-Nielsen (1986c) have concluded that Z. palustris is a taxon restricted to the Northern Hemisphere, and they have described the high Andean collections as a new species, Z. andina. The species are differentiated, in part, by fruit size, which is larger in Z. andina. I am assuming that the collections cited from Arequipa and Cuzco are Z. andina. If the Callao collection is correct, there are 2 species of Zamichellia for Peru. The collection 256 from Callao, a major seaport since the Spanish conquest of Peru, it is probably a waif. The genus is of subcosmopolitan distribution, and inhabits fresh to brackish waters. USES: Forage resource for waterfowl (Martin et al., 1961).

Zannlchelli» mmdlmm Holm-Niels. & R. Haynes HERB, submerged aquatic; stems elongate, internodes to 51 mm long. LEAVES: alternate and opposite, 26-34 mm long, 0.4-0.5 mm wide, linear, sessile, 1, central nerve, margin entire, slightly thickened, apex acuminate. INFLORESCENCE: axillary. FLOWERS: staminate a solitary, naked stamen, filament ca. 1.5 mm long, anther ca. 1.3 mm long, carpellate pedunclate and subtended by a cup-shaped, membranous bract, carpels 4, short-stipitate, free, ovary ca. 1 mm long, style short, terminating in an eccentric, expanded, funnelform stigma, ovate in outline, ca. 1 x 0.7 mm. FRUIT: drupelets, often 4, joined by common peduncle, 2.5-3 mm long excluding 0.7-0.8 mm long style, ca. 1 mm wide, long elliptic to reniform. HABITAT: Lakes and streams at high elevations. DISTRIBUTION: Peru, Bolivia, and Chile; expected to be found in Ecuador. Peru: Arequipa, Cuzco, and Huanuco. As yet not found in the Park, but to be expected. PHENOLOGY: Flowering reported in April, fruiting in December. 257

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ACKNOWLEDGMENTS Throughout my graduate studies, especially during the field and herbarium work which has lead to the culmination of this dissertation, I have been helped by several institutions and many individuals. I am very grateful for the kindness shown me and acknowledge it here. The Botany Department of the Iowa State University has given me its support during my years of residence in Ames. The following persons deserve special mention; F. G. Smith, R. C. Coolbaugh, and D. L. Glenn-Lewin, the IXpartment Heads during my tenure; the members of my Graduate Studies Committee: R. W. Pohl (Chairman), W. L. Franklin, J. Pleasants, R. Shaw, and W. Wedin; the Faculty of the Botany Department, especially D. Isely; my colleagues in study, past and present: C. Betancourt, L. G. Clark, P. Dâvila, F. Escalona, M. Gabel, B. Gardfalo, T. Killeen, and R. Pinette; special thanks go to K. Klier, who has rescued me and my text files from doom so many, many times; the staff of the Botany Department Office, who have helped me so often over the years; and Mrs. L. Bishop, of the Thesis Office, for sage advice. Funding for the fieldwork was provided by a research grant from the National Geographic Society, and additional costs were paid by the Missouri Botanical Garden, my employer during the course of this research project. In Peru, there are many people who made my years there interesting and productive. M. Romero-Pastor, Director of the Direcciôn General Forestal y de Fauna in Lima, and the staff, especially J. Purisaca^ M Falero, and R. Acero were very helpful. In Huaraz, the Director and Staff of CENEFOR-Huaraz gave their support to this project. The staff of the Huascar&n National Park provided help and support to the best of their ability. Thanks go to L. Champa, G. Mejia, L. Sanchez, and, especially to A. Gonzales and R. Valencia. René Valencia is a lover of the natural world, an enthusiastic collector, and firm friend. M Oropeza, accompanied me on nearly all of my extended trips to the field, as both an employee and friend. K. Goodwin, M. Torres, and H. Vidaurre, visitors to the Park, who took interest in the project and accompanied me to the field. D. Maldonado of Huaraz took similar interest in the work; our field trips and conversations are well remembered. In Lima, the Museo de Historia Natural was my host institution. I express my thanks and best wishes to: H. de Macedo, E. Cerrate de Ferreyra, and R. Ferreyra, past Directors, and G. Lamas, Director, for their help and advice; the members of the staff. 280

for their many courtesies, lessons, and advice; and A. Turpo, without whose willing help my tasks in Lima would have been truly onerous. Other than the Museo de Historia Natural, the Herbarium Truxillensis of the Universidad Nacional de Trujillo was the Peruvian scientific institution which collaborated most in my work. I greatly value both the friendship and professional interchange that I have with A. Sagdstegui, Director of the Herbarium. As an institution the Missouri Botanical Garden fully supported this study. However, the soul of an institution is the group of individuals that form it; I extend my thanks to: P. H. Raven, Director, and E. Forero, Director of Research, for their advice and support; A. H. Gentry, friend and nemesis, teacher and hair shirt, for his advice; R. E. Gereau and R. Liesner for their help and lessons in general plant identification; M. Guzmàn, Coordinator for the Flora of Peru Project, for much hard work and many favors during the years we have worked together; H. Threnn and K. Coates, of the Shipping room, for their courtesy and efficiency in the shipping and receiving of the thousands of specimens related to this project; L. Ganss and N. Winkelmeir, "the secretaries," who have done dozens of small, but significant, requests for me; M. Albright, J. Ricketson, and the "mounters," for mounting with care and efficiency the thousands of specimens used in this study; and my colleagues at the Garden, with whom I have had many interesting discussions. In the fall of 1987, the Field Museum of Chicago provided funding for an extended study visit. I thank J. Fitzpatrick, Chairman of the Awards Committee, and T. Plowman, then Chairman of the Botany Department, for their help prior to and during my visit, and M. Dillon, W. Burger, and R. Stolze for their support and encouragement of my work. Many specialists in different taxonomic groups have given me the benefit of their advice, and have verified the determination of specimens. They include: I. Al- Shehbaz (Brassicaceae), F. Ayala (Diocorea), H. Balslev (Juncaceae), R. Barneby (Astragalus and Senna), K. Barringer (Scrophulariaceae), R. Callejas (Peperomia), M. Chanco (Malvaceae), H. Conert (Cortaderia), R. Cruden (Echeandia), J. Cuatrecasas (Diplostephium), W. D'Arcy (Solanaceae), M. Dillon (Asteraceae), C. Dodson (Orchidaceae), T. Duncan (Ranunculaceae), F. Escalona (Çalamagrostis), R. Faden (Commelina), R. Ferreyra (Asteraceae), A. Gentry (Tecoma), R. Gereau (Monocots, Caryophyllaceae, Urticaceae), P. Goldblatt (Iridaceae), J. Grimes (Otholobium), R. Harley (Lamiaceae), R. Haynes {Lilaea), J. Henrich (Iridaceae), R. Hickey (Isoêtes), M. Huft 281

(Euphorbiaceae), H. litis (Cleome), T. Koyama (Scirpus, s.l.), J. Kuijt (Loranthaceae), B. Leôn-Bocàngel (Pteridophytes), R. Liesner (diverse families), P. Lowrey (Çreopanax), J. Luteyn (Ericaceae), H. Luther (Bromeliaceae), J. MacDougal (Passiflora), A. Meerow (Stenomesson), J. Mickel (Elaphoglossum), U. Molau (Bartsia, Calceolaria), R. Moran (Pteridophytes), M. Nee (Solanaceae), B. 011gaard (Lycopodiaceae), M. Poston (Loasaceae), J. Pringle (Gentianaceae), K. Rahn (Plantage), A. Rezneck (Carex), N. Robson (Hypericum), A. Sagâstegui (Asteraceae), B. Simpson (Polylepis), L. Skog (Coriaria), A. Smith (Pteridophytes), W. Stevens (Asclepiadaceae), B. Stein (Campanulaceae), R. Stoize (Pteridophytes), M. Taylor (Microsteris), O. Tovar (Poaceae), G. Tucker (Cyperus), B. Turner (Baccharis), and J. Wurdack (Melastomataceae). In a more personal vein, I express my gratitude to the following: —To Klaus Heinz Lackschewitz, who introduced me to the Montana alpine vegetation, and nurtured my love of botany. —To my parents, Donald A. Smith and Ann W. Kerchman, for life. —To Marianne, my companion, who accompanied me during the Huascar&n study, and became a botanist along the way.