Introduction
to the
Páramo Ecosystem
Checklist
of Páramo
Plants
Lichens
Mosses
Hepatics
Vascular
plants
Gazetteer
of Páramo
Localities
Costa
Rica & Panama
Colombia
Venezuela
Ecuador
Peru
James L. Luteyn
The checklist of vascular plants started
with the lists of Cleef (1981) and Vareschi (1970). Further additions
came after scanning numerous monographs, revisions, and country or regional
floras, such as Flora de Colombia, Flora of Ecuador, Flora Neotropica,
and Flora de Venezuela. Then a very intensive and exhaustive family
by family and genus by genus search for taxa (native and naturalized) represented
in the páramo flora was made at the United States National Herbarium
(US) by myself and Mauricio Gavilanes, followed by lesser general searches
by myself at AAU, NY, and U. Numerous specific herbaria (e.g., COL,
CR, GB, GH, K, MER, MERF, MO, NY, S, VEN, etc.) were also consulted for
particular families (or genera) when a specialist had worked there.
Once the basic family or generic lists were made, copies were sent to individual
specialists for comment, correction, and suggestion. Whenever possible,
I have tried to include in this list only species for which I found specimen
determinations made by recognized authorities. Normally I have not
used species determined with the notations "aff." or "vel aff.", and only
rarely are species included when determined with "cf." Furthermore,
I have used a very conservative approach, including in the list only species
for which I have seen at least one voucher from páramo (as herein
defined), or when a literature citation or personal communication was considered
by me to be extremely reliable.
There is a bit of subjectivity that
went into the compilation of this checklist of páramo vascular plants.
Some may say that the list does not include species "a" or species "b",
etc., for which the label clearly says "Páramo de Colorado" (hypothetical
name herein used for this example). This may be true and points out
how very difficult it is sometimes to decide whether or not to include
a species. In many cases, it is dangerous to include species for
which the label only says "páramo" and nothing else, often because
collector's concepts of páramo may differ. Several things
must be remembered in this regard. First of all, páramo does
not automatically include everything above the general 3000 m elevation
level. As has been stated in the Introduction of this book, montane
forest may reach as high as 4300 m in the northern Andes. Secondly,
just because a collection label says "Páramo de Colorado", does
not automatically mean that the plant was collected in a páramo
habitat as herein defined. Amongst local people, it is very common
for the proper name "Páramo de Colorado" to include the full range
of low to very high elevations and montane forest to superpáramo,
i.e., the term páramo" in itself is not enough of a prerequisite
to include a plant in the checklist. Therefore, I have not included
a species solely because the label says "Páramo de Colorado" unless
the label also tells me something about the habitat (e.g., grass páramo,
etc.), or I know the plant is a typical páramo plant, or I know
the area by personal experience, or I know the collector and can be quite
certain of what concept is used. On the other hand, some may say
that genus "x" or species "c" or "d", etc., that are here included in this
list, ought not to be, because they occur mostly in forest. This
is where my own personal field experience (i.e., knowledge of the area)
or definition of páramo is needed. Here also is where the
difficulty of "what is subpáramo?" comes into play. For the
purposes of this book and the checklist herein, subpáramo includes
elements of small trees and shrubs in transition to grass páramo,
and species of the isolated shrub-tree islands or forest patches within
open grass páramo. It may be debated whether or not the plants
of the shrub-tree islands ought to be included in a páramo checklist,
because they are usually forest species; I have chosen to include
them just as I have species of the Polylepis forest. From
the beginning of this project, my goal has been to present a list of all
plants found in the páramo, whether from the more distinctive open
grass páramo or superpáramo areas to the less distinctive,
more transitional subpáramo zone that includes some forest elements.
I have not, however, included species of plants that are characeristic
of the "southern Ecuadorean scrub" vegetation as defined by Harling (1979),
which some call "subpáramo chaparral" and include in páramo
vegetation, but which I consider a natural vegetation type distinct from
páramo.
The geographical and elevational ranges
given in this checklist are only for species distributions within the páramo
countries here considered (i.e., Costa Rica to northern Peru), not for
total geographical and elevational ranges. This needs to be clarified
with two hypothetical examples. Example 1: the geographical range
given in the list for species "a" is CR, CO, EC, PE; however, it
could also be found in PA and VE but not in páramo there, and it
could also be found in Mexico and Bolivia but these countries are outside
the defined páramo range. Example 2: the elevational range
given in the list for species "a" is 3100-3900 m; however, it may
also occur in PA and VE, but at 2500-2700 m which is not considered to
be páramo, and it may be found in Mexico and Bolivia at 1500-2100
m but these elevations are not included because the countries are outside
the defined páramo range. Because the data were gathered in
this manner, I was not, unfortunately, able to calculate the total number
of endemic páramo species.
The taxa in this checklist are organized
according to the rank of species, and I have not formally listed the ranks
of subspecies or variety, when they occur. However, if a recognized
species consists one or more subspecies and/or varieties, these are formally
recognized at the end of the entry after the word "includes." Furthermore,
certain taxa included in this checklist as synonyms may be recognized at
another rank (perhaps as accepted species) in other publications, such
as Flora Mesoamericana (for example), or vice versa, taxa recognized as
species in Flora Mesoamericana (for example) may be included as synonyms
in this checklist. This is inevitable because species concepts often
vary amongst specialists. I have not tried to give complete synonomy,
but have given only those synonyms in common or regional use, or of historical
importance.
Two genera that are cited as north-temperate
elements reaching their southern limit at the Chirripó páramo
in Costa Rica (Cleef & Rangel Ch., 1984, and Cleef & Chaverri P.,
1992), namely Helianthemum (Cistaceae) and Smilacina (Liliaceae),
are not included in this Checklist, because I have not been able to locate
herbarium specimens that clearly indicate a páramo habitat.
For the many species that are weeds
(ruderals) or that have been cultivated in or near páramo and have
subsequently escaped into páramo habitats, or those that are now
naturalized, I have only included them in this list when I have seen a
herbarium voucher specimen. Many other species that have not yet
become naturalized, or have escaped and are only rarely found in páramo,
or are reported in the literature or by personal communication to me as
occurring in páramo, have not been included because I was unable
to find voucher specimens in herbaria. For discussions of introduced
plants to páramo regions, see Pérez (In press), Sauer (1988),
and Vareschi (1970). The following species may be expected to occur
in páramo following their escape from cultivation, but are not included
in the list.
Apiaceae
Ciclospermum leptophyllum (Pers.) F.Muell.
Brassicaceae
Arabis turrita L. and other spp. (native to Europe)
Brassica campestris L.
Raphanus raphanistrum L. (native to Europe, introduced
and cultivated)
Caryophyllaceae
Silene inflata Sm. (native to Europe)
Fabaceae
Medicago lupulina L. (native to Europe, now cultivated,
naturalized and used as forage)
and M. denticulata Willd.
Melilotus officinalis Willd. (native to Europe
and Asia)
Ulex europaeus L. (used as ornamental)
Vicia faba L. (introduced as crop) and V. sativa
L. (as forage)
Lamiaceae
Mentha (cf. M. viridis L.)
Papaveraceae
Papaver glaucum Boiss. & Hausskn. ex Boiss.
(and perhaps P. somniferum L.) (native
to Middle East)
Phytolaccaceae
Phytolacca octandra L. (native to Asia)
Pinaceae
Pinus radiata D.Don
Poaceae
Avena sativa L.
Hierochloe mexicana Benth. (native to Mexico ?)
Hordeum muticum J.S.Presl (cultivated as forage)
Lolium multiflorum Lam. and L. perenne L.
(cultivated as forage)
Melinis minutiflora P.Beauv. (native to Africa,
introduced via Brazil, cultivated as forage)
Pennisetum clandestinum Hochst. ex Chiov. (introduced
from Africa, now escaped)
Rhynchelytrum repens (Willd.) Hubb. [syn.: Tricholaena
repens (Willd.) Hitchc.] (native
to South Africa)
Setaria parviflora (Poir.) Kerguélen [syn.:
S. geniculata (Lam.) P.Beauv.] (cultivated as
forage)
Triticum aestivum L. (cultivated as crop)
Polygonaceae
Rumex crispus L. (native to Europe)
The families given in the checklist are
divided alphabetically within the larger artificial groups: ferns
and fern allies, gymnosperms, and angiosperms (including both monocots
and dicots). The taxonomic system used is basically that of Cronquist
(1981), with modifications. Certain families such as Asteraceae and
Poaceae, because they are so large, will always need additional work and
updating. As mentioned above, I feel most unsure about the Orchidaceae,
because, in my opinion, they still need much basic taxonomic study, the
nomenclature is confusing, the herbarium specimens have labels with poor
or little habitat data, are few in number, and were generally unavailable
for my study, and they are very much undercollected at páramo elevations.
I am indebted to Cal Dodson for sharing with me unpublished information
from his database of Ecuadorean orchids.
The list ends with literature citations
of a strictly taxonomic nature that have been cited in the checklist.
These references are not meant to be exhaustive, but are restricted to
those which have emphasis on páramo taxa, in which many páramo
taxa are treated, or are of historical and/or nomenclatural importance.
Finally, when questions arose concerning
proper author citation or spellings of some species names, I generally
followed the TROPICOS database of the Missouri Botanical Garden, and for
this help I am very much indebted to Peter M. Jørgensen.
As mentioned above, I have tried to be consistent in the way I recognize taxa in this checklist and have usually followed the taxonomy of one author or one institutional system (US National Herbarium for Asteraceae, for example); but new revisions, monographs, and floras appear every day.