Manual de Plantas de Costa Rica 127 
The Costa Rican paramos cover a total area of some 64,000 ha (Cleef & Chaverri, 
1992). The largest paramo is found on the massif of Cerro Chirrip6 and covers about 
7250 ha, accounting for 14.5% of the total area of Chirrip6 National Park (Kappelle, 
1993). In terms of climate (seasonal, with a marked dry season and under the influence 
of trade winds from the northeast), the Costa Rican paramos are related to those of the 
northern Andes of Venezuela and the Sierra Nevada de Santa Marta in northern Colom- 
bia (Luteyn, 1999). Luteyn (1999) indicated that the paramo habitat attains its west- 
ernmost geographical distribution on Cerro de La Muerte, 1.e., that it is at its extreme 
Central American limit there. Various recent studies (e.g., Cleef & Chaverri, 1992; 
Kappelle et al., 1992; Luteyn, 1999) have corroborated the details of the phytogeo- 
graphic relationship between the Cordillera de Talamanca (particularly its highest parts) 
and Andean South America, previously suggested by Standley (1937b). An analysis of 
the genera in the Talamancas (Kappelle et al., 1992) concluded that they have two prin- 
cipal origins, tropical (75%) and temperate (17%), plus 8% cosmopolitan. Some of 
the temperate genera are northern (holarctic) in origin, including Alnus, Berberis, Chi- 
maphila, Cirsium, Comarostaphylis, Cornus, Holodiscus, Morella, Prunus, Quercus, 
Rhamnus, Romanschulzia, Vaccinium, and Viburnum; others are of southern (australo- 
antarctic) origin, e.g., Acaena, Desfontainia, Drimys, Escallonia, Fuchsia, Gaiaden- 
dron, Muehlenbeckia, Nertera, Pernettya, Podocarpus, Prumnopitys, Ugni, and Wein- 
mannia (Cleef & Chaverri, 1992; Kappelle et al., 1992). The genera of tropical 
(essentially Andean) origin include Arcytophyllum, Greigia, Jamesonia, and Puya. See 
also the sections on Geology in the chapter on The physical environment, and Origin 
of the flora, in the present chapter. 
This phytogeographical relationship was also supported by a comparison (Islebe & 
Kappelle, 1994) of the generic floras of the subalpine forests of Guatemala (3150— 
3800 m) and Costa Rica (3100-3500 m). This study indicated that nearly 30% of the 
total of 178 genera is shared between these two floras. Guatemala has more temperate 
elements (60%), mostly widely distributed herbs, while Costa Rica has more tropical 
elements (487%), a relatively high percentage (26%) of which are trees (only Buddleja 
in Guatemala), with fewer herbs of temperate origin. 
Toward the south, along the Continental Divide, the Cordillera de Talamanca grad- 
ually diminishes in elevation approaching the border with Panama. Noteworthy are 
some characteristic elements of the 2400—3000 m elevational band, such as Anthurium 
chiriquense*, Begonia brevicyma*, B. parviflora*, Blechnum buchtienti, Cobaea minor*, 
Desfontainia splendens, Greigia columbiana*, G. sylvicola*, Gunnera insignis, G. tala- 
mancana*, Gonocalyx megabracteolatus*, Maytenus woodsonii, Quercus spp., and Wim- 
meria sternii*. The species marked with an asterisk (*) are restricted, in Costa Rica, to 
this southern region, thus distinguishing its flora. Standing out above this gradient, 
Cerro Kamuk, on the Caribbean slope and at 3549 m (higher than the nearby divide), 
harbors a small paramo with species such as Agrostis bacillata, A. subpatens, Arcyto- 
phyllum lavarum, Castilleja irasuensis, C. talamancensis, Chionolaena costaricensis, 
