Biodiversity and Conservation 5, 1169-1190 (1996)

Aquatic plants of Peru: diversity, distribution

and conservation
B L A N C A LEON*
Museo de Historia Natural, Av. Arenales 1256, Apartado 14-0434, Lima 14, Peru and Department of Botany,
Field Museum, Chicago, IL 60504, USA

K E N N E T H R. YOUNG$
Department of Geography, University of Maryland Baltimore County, Baltimore, MD 21228, USA

Received 19 April 1995; revised and accepted 5 September 1995

Currently 177 vascular plant species are known or presumed to be obligately associated with water in
Peru. Their composition and diversity were surveyed in relationship to their distributions among the
natural regions of Peru. Despite considerable aridity, the coastal plains of Peru have important
aquatic ecosystems, including marshes in river deltas and mangroves in the far north, near the border
with Ecuador; 70 species of aquatic plants are found in this coastal region. The Andean highlands
include a great variety of wetlands, plus lakes and rivers; 62 species are found, including eight species
of Isoetes, some of which are potentially threatened by extinction. The Amazon region of Peru
includes both steep montane rivers in the headwaters and wide floodplains formed by meandering
rivers in the lowlands; 102 species are found distributed among these ecosystems, although many
additional species are semi-aquatic. Research and conservation strategies for Peru's aquatic plants
need to take into account these important regional differences.
Keywords: aquatic plants; conservation; natural regions; Peru.

Introduction
Peru's plant resources are found distributed among a great array of environments. Fresh or
brackish water ecosystems, such as lakes, rivers and wetlands, account for perhaps 5% of
the total surface area of Peru. However, despite this relatively small area, these
environments represent one of the main sources of water, food and habitat for a diverse
biota, and for human society.
The array of vascular plants inhabiting these aquatic environments is less diverse than
their terrestrial counterparts (Kahn et al., 1993); however, aquatic vascular plants comprise
a unique assemblage of floristic diversity and ecological adaptations. In addition, aquatic
plants are important because of their ecological roles and their economic importance.
In Peru, most types of continental aquatic environments have been used, managed,
and/or modified over many centuries (Rostworowski de Diez Canseco, 1981; Seibert, 1983;
Smith, 1985; Matos Mar and Matos Lagos, 1990). The impact of human influence is
complex and interacts with changes in the physical environment. For example, evidence
from Lake Titicaca (Richerson, 1993) and some coastal wetlands (Smith, 1985)

*Present address: SystematicBotany and MycologyLaboratory, Agriculture Research Center-USDA, Room

304, Building011A, BARC-West,Beltsville,MD 20705, USA.
STo whom correspondence shouldbe addressed.
0960-3115 © 1996 Chapman & Hall
1170 Le6n and Young
demonstrates that climatic, ecological, and social processes have changed through time, as
have their effects on aquatic ecosystems. Present-day assemblages of aquatic plants may be
the result of these interactions.
Because Peru includes a great variety of ecological zones, the conservation of aquatic
environments and their biota has to take into account regional differences. In this article,
we present regional and overall evaluations of the plants found in continental aquatic
environments and summarize the state of knowledge about these plant resources. We only
discuss the vascular plants; the rich algae flora of Peru is discussed elsewhere (Acleto et al.,
1978). We use this overview to propose conservation approaches that could function at
both species and ecosystem levels.

Aquatic plants of Peru

One hundred and seventy seven species of vascular plants obligately associated with water
are either known from Peru or are believed to be present (Brako and Zarucchi, 1993; Le6n,
1993a, 1993b) (Table 1). This total is tentative because most localities have not yet been
diligently inventoried. Several species included in Table 1 have not been reported in Peru,
but are expected to be found there with further exploration: Echinodorus tenellus,
Eichhornia diversifolia, E. heterosperma, Hydrocleis nymphoides, H. parviflora, Najas
marina, Sagittaria latifolia, Typha angustifolia, T. latifolia, Wolffia brasiliensis, and
Wolffiella lingulata.

Table 1. The obligately aquatic vascular plants of Peru and their global and regional
distributions (nomenclature follows Le6n, 1993a and 1993b, with additional information
from Godfrey and Wooten, 1979,1981; Howard, 1979; Hnatiuk, 1990; Brako and Zarucchi,
1993; Haynes and Holm-Nielsen, 1994). Global distributions are not given for species
whose presence remains undocumented by voucher specimens. Types of global
distribution: E = endemic to Peru; NW = New World (temperate and tropical); N =
Neotropical; SA = South America (temperate and tropical); T = Tropical (tropics of
America and other continents); TSA = Tropics of South America; W = widely distributed
(both temperate and tropical in the Americas and other continents); I = introduced

Presence in Peruvian regions

Type of global
Taxa distribution Coast Highlands Amazon

Pteridophytes
Equisetaceae
Equ&etum giganteum L. NW X X

Isoetaceae
Isoetes andicola (Amstutz)
L.D. G6mez TSA
Isoetes boliviensis U. Weber TSA
Isoetes dispora Hickey E
Isoetes hewitsonii Hickey E
Isoetes lechleri Mett. TSA
Isoetes novo-granadensis
Fuchs TSA
Aquatic plants o f Peru 1171
Table 1. (Continued)
Presence in Peruvian regions
Type of global
Taxa distribution Coast Highlands Amazon

Isoetes panamensis Maxon &

C.V. Morton N x
Isoetes parvula Hickey E x
Isoetes saracochensis Hickey E x
Marsileaceae
Marsilea ancylopoda A. Br. N x
Marsilea deftexa A. Br. N
Marsilea mollis B.L. Rob and
Fernald NW x
Marsilea vestita Hook. and
Grev. NW x
Osmundaceae
Osmunda cinnamomea L. W
Osmunda regalis var.
spectabilis (Willd.) A. Gray NW
Pteridaceae
Acrostichum aureum L. T x
Acrostichum danaeifolium
Langsd. and Fisch. N x
Ceratopteris pteridoides
(Hook.) Hieron. N
Salviniaceae
Azolla caroliniana Svens. NW x
Azolla filiculoides Lam. N x x
Azolla mexicana C. Presl NW x x
Azolla microphylla Kaulf. N x
Salvinia auriculata Aubl. N
Salvinia minima Baker N
Thelypteridaceae
Thelypteris interrupta (Willd.)
K. Iwats. T
Thelypteris scalaris (H. Christ)
Alston N x
Monocotyledons
Alismataceae
Echinodorus berteroi
(Spreng.) Fassett NW x
Echinodorus bolivianus
(Rusby) Holm-Niels. N X
Echinodorus grandiflorus
(Cham. & Schltdl.) Micheli N X
Echinodorus grisebachii Small N X
Echinodorus horizontalis
Rataj TSA X
1172 Le6n and Young
Table 1. (Continued)

Presence in Peruvian regions

Type of global
Taxa distribution Coast Highlands Amazon

Echinodorus paniculatus
Micheli N X
Echinodorus tenellus (C.
Mart.) Buchenau ? ?
Echinodorus tunicatus Small N X
Sagittaria guayanensis Kunth
ssp. guayanensis NW X
Sagittaria Iatifolia Willd. NW x
Sagittaria montevidensis
Cham. and Schltdl. ssp.
montevidensis SA x
Sagittaria rhombifolia Chain. TSA
Sagittaria sprucei Micheli TSA
Araceae
Montrichardia arborescens
(L.) Schott N
Montrichardia linifera
(Arruda) Schott TSA
Pistia stratiotes L. N x
Urospatha sagittifolia (Rudge)
Schott TSA
Arecaceae
Mauritia flexuosa L.f. TSA
Cyperaceae
Cladium ]amaicense Crantz NW x
Cyperus giganteus Vahl NW
Cyperus laevigatus L. W x
Eleocharis elegans (Kunth)
Roem. and Schult. T x
Eleocharis geniculata (L.)
Roem. and Schult. NW x
Eleocharis mutata (L.) Roem.
and Schult. T
Scirpus americanus Pers. W x
Scirpus californicus (C.A.
Mey.) Steud. NW x X
Scirpus cubensis Poepp. and
Kunth T x X

Scirpus inundatus (R. Br.)

Poiret W x X
Scirpus maritimus L. W x
Eriocaulaceae
Paepalanthus pilosus (Kunth)
Kunth TSA
Aquatic plants of Peru 1173

Table 1. (Continued)

Presence in Peruvian regions

Type of global
Taxa distribution Coast Highlands Amazon

Tonina fluviatilis Aubl. N x

Hydrocharitaceae
Elodea granatensbs Humb. and
Bonpl. TSA x
Elodea potamogeton
(Bertero) Espinosa SA x
Limnobium laevigatum
(Willd.) Heine N x
Juncaceae
Distichia muscoides Nees and
Meyen TSA x
Juncus arcticus vat. andicola
(Hook.) Balslev TSA x
Juncus cyperoides Laharpe SA x
Juncus stipulatus Nees and
Meyen TSA x
Luzula vulcanica Liebm. NW x
Oxychloe andina Phil. TSA x
Juncaginaceae
Lilaea scilloides (Poir.)
Hauman T x
Triglochin striata Ruiz and
Pav. W x
Lemnaceae
Lemna aequinoctialis Welw. W x x
Lemna gibba L. W x x x
Lemna minuta Kunth W x x
Lemna valdiviana Phil. NW x x x
Spirodela intermedia W. Koch NW x x x
Spirodela polyrhiza (L.)
Schleid. W x
Wolffia brasiliensis Wedd. ? ?
Wolffia columbiana H. Karst. ? ?
Wolffiella lingulata (Hegelm.)
Hegelm. ? ?
WolffieUa oblonga (Phil.)
Hegelm. NW x x
Wolffiella welwitschii
(Hegelm.) Monod T x
Limnocharitaceae
Hydrocleys nymphoides
(Willd.) Buchenau ? ?
Hydrocleys parviflora Seub. ?
Limnocharis flava (L.)
Buchenau N x
1174 Le6n and Young
Table 1. (Continued)

Presence in Peruvian regions

Type of global
Taxa distribution Coast Highlands Amazon

Limnocharis laforestii Griseb. N x

Marantaceae
Thalia geniculata L. N x
Mayacaceae
Mayaca fluviatilis Aubl. NW X
Mayaca sellowiana Kunth TSA X

Najadaceae
Najas arguta Kunth TSA
Najas guadalupensis (Spreng.)
Magnus NW x
Najas marina L. ?
Orchidaceae
Eulophia alta (L.) Fawc. and
Rendl N
Habenaria sartor Lindl. TSA
Poaceae
Alopecurus aequalis Sobol. W X
Echinochloa polystachya
(Kunth) Hitchc. NW
Hymenachne amplexicaulis
(Rudge) Nees N
lsachne polygonoides (Lain.)
Doell N
Luziola subintegra Swallen N
Oryza grandiglumis (Doell)
Prod~n TSA
Panicum dichotomiflorum
Michx. NW
Panicum elephantipes Nees N
Panicum grande Hitchc. and
Chase TSA
Paspalidium geminatum
(Forssk.) Stapf T x
Paspalum repens Bergius TSA x
Phragmites australis (Cav.)
Trin. ex Steud. W x
Pontederiaceae
Eichhornia azurea (Sw.)
Kunth N x
Eichhornia crassipes (C.
Mart.) Solms W x
Eichhornia diversifolia (Vahl)
Urban
Aquatic plants of Peru 1175
Table 1. (Continued)
Presence in Peruvian regions
Type of global
Taxa distribution Coast Highlands Amazon

Eichhornia heterosperma
Alexander
Heteranthera limosa (Sw.)
Willd. N
Heteranthera reniformis Ruiz
and Pav. NW
Heteranthera rotundifolia
(Kunth) Griseb. NW
Heteranthera spicata C. Presl N
Pontederia rotundifolia Uf. N
Potamogetonaceae
Potamogeton ferrugineus
Hagstr. SA
Potamogeton filiformis Pers. W
Potamogeton illinoensis
Morong NW
Potamogeton paramoanus
R.R. Haynes and
Holm-Niels. TSA X
Potamogeton pectinatus L. W X
Potamogeton punense
Gal~in-Mera TSA X
Potamogeton pusillus L. W X
Potamogeton striatus Ruiz and
Pav. NW X X

Ruppiaceae
Ruppia filifolia (Phil.) Skottsb. SA
Ruppia maritima L. W X

Typhaceae
Typha angustifolia L. ? ?
Typha domingensis Pers. W X
Typha latifolia L.
Zannichelliaceae
Zannichellia andina
Holm-Niels. and R.R.
Haynes SA X
Zannichellia palustris L. W X X

Dicotyledons
Apiaceae
Hydrocotyle bonariensis
Comm. ex Lain. NW X X
Hydrocotyle rununculoides
L.f. NW X X X
Hydrocotyle umbellata L. NW X X X
1176 Le6n and Young
Table 1. (Continued)

Presence in Peruvian regions

Type of global
Taxa distribution Coast Highlands Amazon

Lilaeopsis macloviana
(Gand.) A.W. Hill SA
Lilaeopsis schaffneriana
(Schltdl.) J.M. Coult. &
Rose

Asteraceae
Enydra ftuctuans Lour. N
Enydra sessilifolia (Ruiz and
Pav.) Cabrera N x
Pacourina edulis Aubl. N

Avicenniaceae
Avicennia germinans (L.) L. T x

Brassicaceae
Cardamine bonariensis Pers. NW
Rorippa nasturtium-aquaticum
(L.) Hayek W-I x

Cabombaceae
Cabomba furcata Schult. and
Schult. f. N

Callitrichaceae
Callitriche heteropoda
Engelm. ex Hegelm. SA

Ceratophyllaceae
Ceratophyllum demersum L. W x X
Ceratophyllum submersum L. W X

Combretaceae
Laguncularia racemosa (L.)
C.F. Gaertn. T x

Convolvulaceae
Ipomoea aquatica Forssk. T X

Crassulaceae
Crassula venezuelensis
(Steyerm.) M. Bywater and
Wickens SA

Elatinaceae
Elatine peruviana Baehni and
J.F. Macbr. E
Elatine triandra Schkuhr W
Aquatic plants of Peru 1177
Table 1. (Continued)

Presence in Peruvian regions

Type of global
Taxa distribution Coast Highlands Amazon

Euphorbiaceae
Phyllanthusfluitans Benth. ex
Muell. Arg. TSA
Fabaceae
Neptunia natans (L.f.) Druce T
Haloragidaceae
Myriophyllum aquaticum
(Veil.) Verdc. W x x
Myriophyllum mattogrossensis
Hoehne TSA X
Myriophyllum quitense
Kunth NW x
Lentibulariaceae
Utricularia foliosa L. W X
Utricularia gibba L. W x X
Utricularia trichophylla Oliv. N X

Menyanthaceae
Nymphoides indica (L.)
Kuntze T X

Nymphaeaceae
Nymphaea amazonum C.
Mart. and Zucc. N X
Nymphaea glandulifera
Rodschied N x X
Victoria amazonica (Poepp.)
Sowerby TSA X

Onagraceae
Ludwigia helminthorrhiza (C.
Mart.) Hara N X
Ludwigia peploides (Kunth)
P.H. Raven W x X

Podostemaceae
Apinagia peruviana (Wedd.)
Engl. E X
Marathrum striatifolium R.
Royen E x X

Polygonaceae
Polygonum acuminatum
Kunth W X
Polygonum ferrugineum
Wedd. N X
Polygonum hispidum Kunth N X
1178 Le6n and Young
Table 1.(Continued)
Presence in Peruvian regions
Type of global
Taxa distribution Coast Highlands Amazon

Polygonum punctatum Elliott NW

Portulacaceae
Montia fontana L. W
Ranunculaceae
Ranunculus flagelliformis Sm. NW
Ranunculus limoselloides
Turcz. TSA
Ranunculus trichophyllus
Chaix W
Rhizophoraceae
Rhizophora mangle L. T
Scrophulariaceae
Bacopa monnieri (L.) Pennell W
Limosella aquatica L. W
Limosella subulata Ives NW
Mimulus glabratus Kunth NW
Sphenocleaceae
Sphenoclea zeylanica Gaertn. W-I

Most of these aquatic plants are herbaceous monocotyledons, followed in importance by

dicotyledons and the pteridophytes (Table 1); this is similar to compositions of many other
aquatic plant floras (Cook, 1983).
The best represented families are those of Alismataceae (arrowheads), Poaceae
(grasses), Cyperaceae (sedges), Lemnaceae (duckweeds), Pontederiaceae (pickerel-
weeds), Potamogetonaceae (pondweeds), Juncaceae (rushes), Apiaceae, Isoetaceae
(quillworts), and Salviniaceae. The most speciose genera are Echinodorus (Alismataceae),
Isoetes (Isoetaceae), and Potamogeton (Potamogetonaceae).
The aquatic flora of Peru consists to a great extent of species with a Neotropical
distribution, followed by those with a New World distribution pattern, and those that are
widespread (Table 2). Pteridophytes, when examined separately, have proportionally
fewer species with world-wide distributions, while dicotyledonean species have more.
Peruvian aquatic plant species that are widely distributed or that also occur outside of
tropical America include Acrostichum aureum, Bacopa monnieri, Ceratophyllum
dernersum, C. submersum, Elatine triandra, Lemna gibba, L. minuta, Limosella aquatica,
Myriophyllum aquaticum, Nymphoides indica, Polygonum acuminatum, Ruppia maritima,
Scirpus inundatus, Sphenoclea zeylanica, Thelypteris interrupta, Triglochin striatum, and
Utricularia gibba. Some of these species have doubtless been introduced to Peru, as is the
case for Sphenoclea zeylanica, but for the majority it is unknown if they are native or
introduced (Cook, 1985). Most of these widely distributed species occur in disturbed areas,
often with long histories of human occupancy.
Species restricted to Peru are relatively few (Table 1). These endemic species are
Aquatic plants of Peru 1179
Table 2. Global distribution patterns of Peru's aquatic plants by major taxonomic grouping.
Values given are percentages

Pteridophytes Monocotyledons Dicotyledons Total

Neotropical 37 21 21 24
New World 22 23 18 22
Tropics, South America 15 23 8 17
South America 0 6 6 5
Endemic 15 0 6 4
Tropics 7 7 12 8
Widely distributed 4 20 29 20

Apinagia peruviana, Marathrum striatifolium, Elatine peruviana, Isoetes dispora,

I. hewitsonii, I. parvula, and I. saracochensis. The first two species mentioned belong to the
dicotyledonean family Podostemaceae, whose species are habitat specialists, growing on
walls or rocks in streams (Cook, 1985). The remaining four species are found in ponds and
lakes. There are no aquatic monocotyledonean species endemic to Peru.

Natural regions of Peru

Peru's territory includes a significant portion of the western side of the Amazon basin,
largely covered by humid lowland forest. The Andes mountains, which cross Peru from
north to south, break the Amazonian influence, dividing the country in three natural
regions: coast, Andean highland, and the Amazon (Fig. 1), delimited here using the 2000 m
contour on the western side of the Andes, and the 3500 m contour on the eastern side.
There are three major hydrological basins: Pacific, Atlantic (or Amazon) and Titicaca
(Fig. 2). The coastal region and part of the Andean highlands correspond to the Pacific
basin. The Atlantic basin originates in the eastern highlands in the Andean region and
continues into the lowlands of western Amazonia. The Titicaca basin is the smallest and is
located on a plateau in the southern highlands.
The endemic aquatic species of Peru include four pteridophytes from the genus Isoetes;
all four are found 0nly in the high Andes. The other endemic species are flowering plants:
Marathrum striatifolium occurs on both the coast and in the Amazon region, while
Apinagia peruviana and Elatine peruviana are known only from the Amazon.
Comparatively few species are shared among the natural regions (Table 1). Surprisingly,
the coast and Amazon regions share the largest number of species. Only nine species are
found in all three regions: Azolla filiculoides, A. mexicana, Eleocharis geniculata,
Hydrocotyle ranunculoides, H. umbellata, Lemna gibba, L. valdiviana, Scirpus cubensis,
and Spirodela intermedia.

The coastal lowlands

The coastal region of Peru runs along the Pacific Ocean for nearly 3100 km (Fig. 1), with
variations in width from 170 km at 6°S to as narrow as 70 km at 14°S. This region has an arid
to semi-arid climate, heavily influenced by the subtropical high pressure cell and by the
1180 LeOn and Young

Figure 1. Peru's three natural regions.

cold water currents of the eastern Pacific Ocean (Prohaska, 1973). Two main subregions
can be differentiated based on climatic, geological, and floristic factors. South of 8°S,
precipitation manifests itself seasonally as winter drizzle ('gartia') and mist; mean monthly
temperatures are 15-18°C. To the north, from 3.5-8°S, precipitation falls during the
summer season, and is additionally modified by E1Nifio/Southern Oscillation events; mean
monthly temperatures are 20-23°C.
In the coastal region, aquatic habitats are sparse and have been highly modified by
humans (Fig. 3A and 3B). Wetlands occur at the end of river drainages, near the shore, in a
fashion similar to those described for savanna areas in Africa (Scoones, 1991). There are 44
rivers on the coast of Peru with permanent fow, but only a few of these carry enough water
or have a shallow enough slope gradient to permit the natural formation of wetlands. The
use of some of those coastal rivers for irrigation date back at least 2500 years BP (Moseley,
1978). Archaeological studies indicate that this region was populated and its landscapes
were extensively managed. The construction of irrigation canals and sunken fields (Smith,
1985) made water available in otherwise barren areas close to the ocean; these techniques
Aquatic plants of Peru 1181

Figure 2. Peru's three major drainage basins.

would have created, often unintentionally, conditions adequate for many aquatic plants,
and in some cases might have expanded their distribution.
Seventy species of aquatic plants are expected to grow in this region (Table 1), although
only 62 have been documented with voucher collections deposited in herbaria.
Cyperaceae, Lemnaceae and Pontederiaceae are the largest families. The genera Scirpus,
Heteranthera, Lemna and Wolffiella are the most speciose.
The flora of this region consists principally of widely distributed species (Table 3),
followed in importance by species that occur in temperate and tropical portions of the
Americas. Only one species endemic to Peru (Marathrum striatifolium) is found in the
aquatic environments of the northern coast. Some other plants are rare or of limited
distribution in this region, such as the mangrove-forming species, Avicennia germinans,
Laguncularia racemosa, and Rhizophora mangle.
Mangroves are only found in the northernmost portion of the coast (Weberbauer, 1945;
Ferreyra, 1979). This is the southernmost point of their distribution along the coast of
western South America. The largest area of mangrove is found in the department of
1182 Le6n and Young

Figure 3. Habitats of Peru's three natural regions. (A) coastal wetland in a nature reserve
('Pantanos de Villa') bordered by the houses of metropolitan Lima; (B) coastal marshes
found a short distance from the Pacific ocean; (C) lake found in the highlands of southern
Peru at 3400 m; (D) irrigation channel carrying water to fields and houses in the highlands
of central Peru; (E) the steep montane areas of the eastern Andes of Peru are dominated by
whitewater rivers; (F) the floodplains of the tributaries of the Amazon River include
numerous aquatic and semi-aquatic plant communities.

Tumbes, on the delta of the Puyango-Tumbes and Zarumilla rivers. A small population
of Laguncularia is located further south in the department of Piura. As an ecosystem,
mangroves appear to be threatened by the same factors that affect t h e m in Asia and Africa:
fish and shrimp-pond construction, waste disposal, and land reclamation (Turner, 1991).
Aquatic plants of Peru 1183
Table 3. Global distribution patterns of the aquatic plants of Peru's three natural regions.
Values given as percentages of species within each natural region

Coast Highlands Amazon

Neotropical 17 5 38
New World 32 31 18
Tropics, South America 2 20 17
South America 0 12 1
Endemic 2 7 3
Tropics 1l 5 9
Widely distributed 36 20 14

During an eleven-year period between 1980 and 1991 the surface area of mangrove in
Tumbes diminished by 22% (Cuya Matos and de Prado, 1993).
The aquatic flora of the northern coast includes other species of limited distribution:
Acrostichum aureum, Heteranthera limosa, H. spicata, Isoetes panamensis, and Marsilea
ancylopoda. Another seven species are found both in this area and in the Peruvian
Amazon: Acrostichum danaeifolium, Lemna aequinoctialis, Limnocharis laforestii,
Marathrum striatifolium, Nymphaea glandulifera, Paspalum repens, and Thalia geniculata.
Other than their presence, very little additional information exists about these plants or
their habitats.
The aquatic environments located to the south of 8°S, share a relatively homogeneous
aquatic flora, mostly restricted to wetlands. Common species include Enydra spp.,
Hydrocotyle ranunculoides, Myriophyllum aquaticum, Potamogeton striatus, Scirpus
americanus, and Typha domingensis. These wetlands are constantly affected by humans,
and their plant assemblages are likely to have changed through time. Cattails (Typha
domingensis) and 'totoras' (Scirpus americanus and S. californicus) have been widely
utilized since pre-Columbian times for their fibres, and this tradition persists today. The
habitats maintained or manipulated for these species may have permitted the survival of
other common aquatic plants.
Coastal wetlands vary in size, although few of them are large enough to attract
international attention for their protection and/or management (Scott and Carbonell,
1986). Scattered herbarium collections and archaeological/historical studies document the
existence of at least 100 wetland sites located along the central coast (between 11-13°S);
many of these have completely disappeared.
For a few coastal wetland sites, the aquatic flora is comparatively well known and can be
evaluated for change through time. For example, we (Cano et al., 1993; Le6n et al., 1995)
were able to document local extinctions in one such locality, the 'Pantanos de Villa'
wetland located near Lima (Fig. 3A). Sixty two species have been reported from that
wetland and surroundings over the last 60 years. However, ten of these species are no
longer found there or are now very uncommon. Some of these species are strict aquatics
with a sparse distribution, such as Paspalidium geminatum, known only from three other
localities more than 100 km apart, and Sagittaria montevidensis, known from two other
localities on the coast and another two more from the Amazonian region (Brako and
Zarucchi, 1993; Le6n, 1993a).
Coastal wetland plant species occur in small populations, often much less than 100 ha in
1184 Le6n and Young
size, and usually associated with agriculture. Such sites cannot sustain wild vertebrate
populations and probably are too small or altered for some of the aquatic plant species.
Today, more than half of Peru's 22 million inhabitants live on the coast. Most of these
people live in urban environments, in settlements of more than 10 000 residents. Changes
in the intensity of resource use has affected many water bodies due to overexploitation for
grazing or agriculture, due to the abandonment of irrigation canals, and changes in
groundwater quantity and quality.

The Andean highlands

The Andean highlands comprise about 30% of the surface area of Peru (Fig. 1). This is a
region with rough topography and dramatic elevational changes: deep interandean valleys
and peaks as high as 6500 m. Temperatures vary with altitude, and are often relatively
constant during the year but change dramatically diurnally. Rain falls seasonally: rainy
season usually begins in October and continues until March or April.
The rivers of the coastal (Pacific basin) and Amazonian regions originate in the Andes.
The most common aquatic ecosystems in the highlands are lakes located above 3000 m
elevation (Fig. 3C), and wetlands associated with streams and rivers.
Sixty-two species of aquatic plants probably grow in this natural region (Table 1); 60 of
these have been documented with voucher specimens. Juncaceae, Apiaceae, Lemnaceae,
and Potamogetonaceae are the more common families. The most speciose genera are
Azolla, Isoetes, Juncus, Potamogeton, Ranunculus, and Scirpus.
The aquatic flora of this region is characterized chiefly by species that exhibit temperate
or tropical distributions in the New World (Table 3). Those species that have a tropical
South American distribution pattern are restricted to the Andean region, including such
species as Distichia muscoides, Elodea potamogeton, Isoetes andicola, I. boliviensis,
I. lechleri, and Oxychloe andina. There are very few differences between the highland
portions of the three major drainage basins in terms of floristic composition. However,
Oxychloe andina is found only in the Lake Titicaca basin.
The Andean region is the most diverse in the country in terms of endemic species (Table
1). All these belong to the genus Isoetes and are found in lakes and ponds at high elevations
(Hickey, 1994).
Andean wetlands are often known in Peru as 'occonales' or 'bofedales' (Tapia and
Flores-Ochoa, 1984). In sites above 3500 m, with poor drainage and located around lakes
and near rivers, the cushion plant, Distichia muscoides, can develop large populations,
often accompanied by Plantago rigida, Carex spp., and Juncus spp. Wetland sites below
3500 m (or those without Distichia), contain a wide variety of aquatic herbaceous plants
such as Azolla spp., CaUitriche heteropoda, Elatine triandra, Hydrocotyle ranunculoides,
and Lilaeopsis spp.
On lake edges, Scirpus californicus and Juncus arcticus var. andicola are often dominant
emergents. These two species have rhizomes that can be consumed by people. They are
also widely used as a source of fibre for handicrafts. Some lakes may have a submerged or
floating flora, with Azolla spp., Isoetes spp., Myriophyllum quitense, Potamogeton spp.,
Ranunculus spp., and ZannicheUia andina. Sometimes these species may also be found in
irrigation canals (Fig. 3D). However, there are few detailed studies that document the
aquatic vascular flora of specific localities; exceptions include those done for Lake Titicaca
(Levieil and Orlove, 1991) and Lake Mataracocha (Becker, 1987).
Aquatic plants of Peru 1185
Humans have been modifying the Andes and its vegetation for at least 10000 years.
Highland wetlands were probably crucial for the domestication of South American
camelids (Flannery et al., 1989), because much of their forage comes from aquatic species.
The introduction of cattle by the Spaniards in the sixteenth century increased the use of
wetlands for grazing and for some agricultural activities (Flores-Ochoa, 1968).
Mining has become the most environmentally destructive activity in the highlands,
constituting the main source of water-borne pollutants. Populations of Isoetes andicola
have been destroyed by mining and drainage (Le6n, 1993b; Young and Le6n, 1993). The
introduction of trout to most of the lakes and rivers started in Peru in the 1940s
(Flores-Ochoa, 1968). Faunal extinctions, such as Orestias cuvieri in Lake Titicaca,
resulted from this introduction (Richerson, 1993), but impacts on flora are unknown.

The A m a z o n
This is the largest and the most biologically diverse natural region of Peru. It includes
numerous types of vegetation located both in montane areas, below 3500 m on the eastern
slopes of the Andes, and in lowland areas from 50 to 500 m elevation (Fig. 3E and F). In
montane areas located at 1500-3500 m elevation, mean monthly temperatures range from
9-25°C and annual precipitation from as little as 500 mm to as much as 7000 (Young, 1992).
In the lowlands, mean monthly temperatures are 25-30°C and annual precipitation is
between 1200 and 3200 mm. Precipitation can occur throughout the year, although some
seasonality can be observed with less rain falling between June and September (Kalliola et
al., 1991; Young, 1992).
The montane and lowland belts differ not only in topography and climate, but also in the
number and kind of aquatic environments. Because of steep slopes, habitats suitable for
aquatic plants are scarce in the montane zone (Fig. 3E). Most of these consist of whitewater
habitats, such as waterfalls and rapids; rocks are the main substrate for aquatic plants. This
elevational zone is the least explored for aquatic plants. Habitat specialists, such as those in
the family Podostemaceae, are found here. The few lakes in the lower parts of the montane
zone may present mats of aquatic vegetation, including Elatine peruviana, Thelypteris
scalaris, and Typha domingensis.
The lowlands of the Amazon are very diverse in types of habitats appropriate for aquatic
plants. There are numerous streams, and isolated lakes and ponds. River floodplains are
conspicuous landscape features and include oxbow lakes and swamps (Junk, 1983; Kalliola
et al., 1991). Plant communities in this zone vary with changes in water level and as a
function of nutrients and water clarity. A large number of semi-aquatic plant species may
accompany those communities (Fig. 3F), substantially increasing the total richness of the
flora associated with water. A unique and important aquatic environment is that of
savanna marshes, found in southern Peru (Kalliola et al., 1991) (although it also extends
into Bolivia), which are dominated by sedges, together with Mayaca sellowiana.
One hundred and two species of obligately aquatic plants are expected to grow in the
Amazon region (Table 1), and 94 of these have been documented with voucher
specimens. The best represented families are Alismataceae, Cyperaceae, Poaceae and
Pontederiaceae. The most speciose genera are Echinodorus, Nymphaea, Sagittaria, and
Utricularia.
The aquatic flora of this region shows a remarkable tendency for dominance by species
with a Neotropical distributional pattern, followed in importance by those with New
1186 Le6n and Young
World and tropical South American distributions (Table 3). Those species exhibiting the
last-mentioned pattern are mostly restricted to the Amazonian basin, and include
Echinodorus horizontalis, Oryza grandiglumis, Phyllanthus fluitans, and Sagittaria sprucei.
The occurrence of regional endemism is lower than the Andean region, but higher than
that of the coast (Table 1). Two of Peru's endemic aquatic species are only known from this
region: Elatine peruviana and Marathrum striatifolium.
The most common aquatic plants in the lowlands include AzoUafiliculoides, Eichhornia
crassipes, Pistia stratiotes, and Salvinia auriculata (Le6n and Encarnaci6n, 1993). The
economic importance of aquatic plants has been recognized by people living along rivers
and oxbow lakes, especially when plant growth occasionally closes ports to boat traffic
(Venero, 1987).
Grasses are also important components of these habitats, occurring along the shores of
lakes and rivers (Fig. 3F); nine are recognized here as strictly aquatic (Table 1), but about a
dozen other species could grow in permanently inundated areas, including Hemarthria
altissima, Oryza latifolia, and Reimarochloa brasiliensis (Tovar, 1993). Some of these
grasses are used as forage for cattle or introduced water buffalo.
Similar to other regions in the country, human history in the Amazon predates
European invasions by many centuries (Aramburd, 1984). There are more than 400
indigenous groups inhabiting the Amazon basin (Chririf et al., 1991), of which 100 belong
to different ethnolinguistic groups. These indigenous groups have long used aquatic
environments for transportation and as resource zones. However, human intervention has
not left as conspicuous an effect on natural landscapes in the past as in the other natural
regions of Peru. Few introduced aquatic species are to be found here, and they may have
appeared only recently. Petroleum exploitation, use of water bodies for waste disposal,
and agricultural expansion count as major factors leading to modern human-caused
alterations of these environments.

Conservation implications
The importance of wetlands has been widely discussed in terms of their economic,
agricultural and biological values (e.g. Junk, 1983; Bjork and Digerfeldt, 1991; Scoones,
1991; Turner, 1991). Proposals for their conservation have stressed the need for better
management, assuming that all wetland areas, especially in the tropics, can (and should) be
incorporated within a sustainable economic development scheme (Turner, 1991).
However, a given tropical country is unlikely to be homogeneous in its regional geography
and in the history of wetland use and abuse. Needed are conservation approaches specific
to individual countries and designed for the particular natural regions involved.
In Peru, regional differences in the composition of aquatic vascular plant species reflect
changes in environmental factors related to climate and to specific characteristics of the
aquatic environments. Regional floristic differences still need further documentation.
Additional studies should gather data on the status, size, and variability of the populations
and communities. Little concrete information exists on the physical conditions that govern
these ecosystems.
To date, conservation planning related to aquatic ecosystems in Peru has focused on the
protection of bird habitats. A preliminary inventory (Scott and Carbonell, 1986) gives data
for those wetlands presumed to be 'internationally important', using as criteria such factors
as the number of individual birds that use a particular site (Moser, 1989). Obviously, we
Aquatic plants of Peru 1187
feel that other taxonomic groups should also be considered when making such evaluations.
Specifically, we suggest that conservation planning for aquatic plants be done both at
species and ecosystem levels for each of the three major natural regions.
Given that the Amazon region's aquatic environments are so vast and, as yet, relatively
little modified, it appears that for now a priority is to develop a species-level approach. We
proposed (Young and Le6n, 1993) that the rarest aquatic plants of the region, Apinagia
peruviana and Elatineperuviana, be declared threatened or endangered species. In theory,
at least, this action would encourage researchers to evaluate the population status of these
species. Large tracts of flooded forest and savanna marshes are included within
Pacaya-Samiria National Reserve and the Pampas de Heath National Sanctuary,
respectively (Table 4).
The coast's aquatic ecosystems have been heavily modified by people. However, despite
their uniqueness and the degree of environmental intervention they sustain, these
environments have received little attention from researchers and conservationists. Many
of the aquatic plants found here are of limited distribution in Peru or are shared with
Amazonian sites at least 300 km distant. By international standards these species are not of
immediate concern, because extensive populations exist in neighbouring countries.
However, they should be of special concern within Peru. We thus suggest that their
protection take place at an ecosystem level by including larger areas and complete
watersheds within the national park and nature reserve system. The protected areas on the
coast are few and small (Table 4); none protect complete watersheds. Completely
overlooked so far is the potential for creating new aquatic ecosystems as part of large
irrigation projects.
The Andean highlands include several aquatic plant species from the genus Isoetes,
which we consider in danger of extinction (Young and Le6n, 1993). Special programmes
for these species are therefore required. Contamination and drainage of water bodies in
the highlands are closely tied to one of Peru's most lucrative economic activities: mining.
Impact evaluations are now required by law. We suggest that aquatic plants, and especially
these endangered species, be included and that mitigation practices be implemented. With
few exceptions, populations of highland aquatic plants are not adequately protected within
nature reserves (Table 4). For example, Junin National Reserve itself is severely
contaminated by mine tailings.
Recently, some observers have quarreled over whether the species or the ecosystem
should be the focus of conservation (Franklin, 1993; Orians, 1993; Tracy and Brussard,

Table 4. Protected areas in Peru that include significant freshwater aquatic and/or wetland
ecosystems (Pulido, 1991)

Region Name Size (ha)

Coast Pantanos de Villa Reserved Zone 396

Los Manglares de Tumbes National Sanctuary 2 972
Lagunas de Mejia National Sanctuary 690
Highland Junfn National Reserve 53 000
Titicaca National Reserve 36180
Amazon Pacaya-Samiria National Reserve 2 080 000
Pampas de Heath National Sanctuary 102109
1188 Le6n and Young

1994). We feel that information, policies, and programmes are urgently needed at both
hierarchical levels. In fact, as we have shown in this article, it is also crucial to adapt
research and conservation efforts to regional realities. Aquatic plants are but one group of
organisms that require such evaluations. International programmes need to consider both
national and regional differences in the distribution and representation of biological
diversity.

Acknowledgements
For financial support, we are grateful to the John D. and Catherine T. MacArthur
Foundation, the Jesse Smith-Noyes Foundation, Missouri Botanical Garden,
CONCYTEC (Consejo Nacional de Ciencia y Tecnologia) of Peru, and the Biodiversity
Support Program, a consortium of the World Wildlife Fund, the Nature Conservancy, and
the World Resources Institute with financing from the Agency for International
Development of the USA. We offer a special thanks to Asunci6n Cano and Francis Kahn
for help with this undertaking and to Robert Haynes for comments on the manuscript.
Joseph School prepared the final versions of the graphics.

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