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Urban vegetation change after a hundred years in a tropical city (San Jose de Costa Rica).

Urban vegetation has mostly been studied in central Europe, where about 50% of species are alien, half of them introduced before the 15th century. Despite the heavy traffic among European cities, five centuries have not been enough to homogenize their flora: the communities of species introduced after the year 1500 are characteristic of each city (Frank et al. 2008).

For animals, which particular species occur in cities is predicted by the "environmental filtering model" that in turn is based on plants. The model states that (1) there is natural selection of species living in urban ecosystems, (2) plants define key habitat characteristics and (3) habitats define which animals can live in the city (Williams et al. 2009). Generally, moderate urbanization produces some increase in plant biodiversity but is deleterious for invertebrates and mammals. A high level of urbanization is correlated with fewer species of plants, invertebrates, amphibians, reptiles, birds and mammals, possibly because humans willingly introduce plant species, but not animals, to their gardens (McKinney 2008).

Outside central Europe, urban vegetation is understudied but there are some recent data from Plymouth, England, where alien species increase with urbanization (Kent et al. 2001). Also in England, gardens in Sheffield have a total of 1 166 plant species (70% alien) and twice the garden size means 25% more species. In these gardens there are 63% biennials/perennials, 18% shrubs, 10% annuals and 8% trees (Smith et al. 2006).

In Anglosaxon and French North America, there is a surprising scarcity of recent studies on urban biotas, but some work has been done. In the city of Halifax (Nova Scotia, Canada), soil moisture and light determine which species are present. Taxa adapted naturally to rock, grassland and flooded habitat find an analog habitat in the city and thrive (Lundholm & Marlin 2006).

The New York metropolitan region has 556 woody species and non-native invasive species are becoming more common (Clem ants & Moore 2004). In the Pelham Bay Park,

New York City, native species went from 72% to 60% and 26% of natives disappeared in 50 years (especially herbaceous and meadow-type plants, DeCandido 2004).

Even though recent studies are scarce, a meta-analysis found 79 studies of species richness with geographic data for New York City; of these, 17 studies found a decrease in species richness, six an increase and three found no change. However, all studies reported an increasing number of exotic species (Puth & Burns 2008).

Tropical cities are in areas of high biodiversity but little is known about Asian and especially African cities regarding urban flora. In Latin America the situation is better but worldwide no floral lists exist for the 50 most populated cities (Clemants 2002). In Jinan City, China, a methodological comparison found that gradient analysis from the urban center to the fringe gives better estimates of the urban flora than the traditional block-area analysis (Kong & Nakagoshi 2005). In Taipei, green areas have 164 tree species (few shared among sites) and large evergreen native species dominate. Larger parks have higher richness, more landscape fidelity to the original vegetation, and more rare and endemic species (Jim & Chen 2007). In Africa, the urban areas of the Nile Delta (Egypt) have vegetation that is mainly correlated with moisture, pH, fertility and texture gradients, but plants always occupy sites similar to their natural habitats (Shaltout & El-Sheikh 2002).

Latin America has a long history of scientific study of urban biota, particularly the plants and there are several recent studies from Mexico, Peru, Brazil, Chile and Argentina.

In the city of Ensenada, Mexico, there are 161 species, 61% non-native (Garcillan et al. 2009). In Mexico City, trees are stressed from dry wind and unfavorable water flow caused by the pavement (Barradas 2000).

Brazil has the largest urban forest in world (Tijuca: 3 300 hectares) but it is being stressed by roads because roads are surrounded by invasive species that burn easily. The fires in turn open adjacent areas to more invasive vegetation and the damage spreads (Matos et al. 2002). There are very few studies of plants that grow on walls but in Jundiai, Brazilian, walls have a biodiversity of 28 species (dos Reis et al. 2006).

Normally, satellites are not used to study urban vegetation but in Arequipa, Peru, satellite images show that desert vegetation is being lost because of urban expansion (Polk et al. 2005). However, only ground work can reliably identify species and this kind of work has shown that temperate South America is not different from North America and Europe: at least half of the plant species in the Argentinean cities of Mendoza and Rosario are introduced. In Lujan de Cuyo, Mendoza, 61 species were identified: 69% introduced (Mendez 2005). The vacant lots of Rosario each have one dominant species, a few abundant species and many rare species. Therophytes predominate and the proportions of indigenous and introduced species are similar (Franceschi 1996).

Chile is the Latin American country with the largest number of recent studies. Synanthropic communities in an urban footpath of Valdivia represents six associations and two communities (Finot & Ramirez 1998). In Concepcion, green areas are dominated by nonnative ornamental species (Paucharda et al. 2006). The distribution of urban vegetation reflects social inequalities. In Santiago, poor areas can have ten times less plant cover than rich neighbourhoods (Hernandez 2008), similar to other countries (Pedlowski et al. 2002). However, workshops in poor areas of cities can result in an improvement of their vegetation (Garzon et al. 2004).

In Costa Rica, there is a long history of study of urban plants that began with the National Museum's collection efforts in the late 19th century, but little has been published. Mendez & Fournier (1980) and Monge-Najera et al. (2002a,b) studied the lichens and their relationship with air pollution. The use of European lichens proved succesful when they were transplanted to this Tropical city (Gruninger & Monge-Najera 1988). Francisco Fallas made checklists and abundance estimates of urban herbs in the late 1970's but to our knowledge he did not publish them. The program "Costa Rica: Jardin Botanico de America Tropical" produces manuals and labels for urban vegetation ( and there is a program to provide urban parks with butterflies and their host plants ( plas/index.html).

The biodiversity in patches of urban vegetation can be surprinsingly high, at least in Costa Rica. For example, after 50 years, in only one hectare of urban vegetation in San Jose, there are 432 plant species (Di Stefano et al. 1995, Nishida et al. 2009), a full new lichen family with a novel symbiotic lifestyle (Eremithallus costaricensis; Lucking et al. 2008), 200 butterfly species (Nishida et al. 2009) and a new species of Hymenoptera, Meteorus oviedo (Shaw & Nishida 2005).

We present an estimate of how the vegetation has changed in the city of San Jose, Costa Rica, after about one century, with a technique called repeat photography. We do not know of repeat photography studies on urban Costa Rican vegetation of San Jose, but the technique was used in Costa Rica by Horn (1989) to assess changes in the paramo habitat.


We used a collection of photographs taken in the late 19th and early 20th centuries from the Jose Fidel Tristan Fernandez Collection in the Archivo Nacional de Costa Rica and others reproduced by Leiva (2004). The sites were re-photographed on December 11, 2008 with a Nikon Coolpix 8800 camera (8 megapixels; Fig. 1). The repeat photography technique, developed in 1880 (Webb et al. 2010) is good for detailed analysis (Hendrick & Copenheaver 2009) and is cost- effective (Robert et al. 2010). We used digital repeat photography, which is fast, detailed and reliable; can be stored for future corroboration and comparison; includes rich data that may become useful in the future; and can classify and measure information automatically (Crimmins & Crimmins 2008).

We re-photographed nine sites (exact year of original photograph included when known): Site 1 Catedral Metropolitana Street: 0 Avenues: 2-4 (1896); Site 2 Catedral Metropolitana St. 0 Av. 0-2 (c. 1910); Site 3 Kiosco Parque Morazan St. 5-7 Av. 3 (1914); Site 4 Colegio de Senoritas St. 3-5 Av. 4-6 (1914); Site 5 Paseo Colon St. 36 Av. 0, looking east (1899); Site 6 Paseo Colon St. 36 Av. 0 looking west; Site 7 Antigua Casa Presidencial St. 15 Av. 7; Site 8 Estacion del Ferrocarril al Atlantico St. 21-23 Av. 3; Site 9 Hospital de Ninos St. 20 Av. 0.

We calculated % cover by clipping and weighing sections from photographs printed on standard bond paper; for example, if the clippings from buildings represented 20 % of the total weight of the photograph, we recorded that buildings represented 20% of the image. This technique is compared with others by Monge Najera et al. (2002b).

We also present an analysis of plants collected in canton de San Jose from 1885 through 1945 by the Museo Nacional staff (Base de Datos, Herbario Nacional, updated to April 13, 2009).


After about a century, the main change in San Jose city photographs is the much larger number of people. The reduction in vegetation affects grasses, shrubs and trees, but is small; the increase in buildings, streets, vehicles, sidewalks, lamps and signs is also small (Fig. 2).

A total of 95 families and 458 species were collected in the late 19th and early 20th century.



The families with most species were Asteraceae, Fabaceae, Poaceae, Lamiaceae, Euphorbiaceae, Solanaceae, Cyperaceae, Acanthaceae, Malvaceae, Piperaceae and Verbenaceae (Appendix 1).


Repeat photography was used in Costa Rica by Horn (1989) to assess changes in the paramo habitat (she found very little change) but we were unable to find a similar study of urban vegetation. Nevertheless, studies about plant biodiversity after long periods suggest that our results are not unusual. For example, after 50 years, Brussels has the same total number of plant species that it had in 1940, albeit the individual species change and the same applies to other areas in much longer time spans (Chocholouskovaa & Pysek 2003). We cannot make a comparison of species from circa 1900 to circa 2000 in San Jose because urban vegetation has rarely been collected in recent decades.

The increase in human presence in the city landscape is explained by the population growth of downtown San Jose or Canton Central (from about 30 000 when the first photographs used in this study were taken circa 1900 to 356 000 when the sites were rephotographed in 2009; see: Centro Centroamericano de Poblacion and Instituto Nacional de Estadistica y Censos 2002). The small increase in the number of vehicles is an underestimation: the 2008 photographs were purposefully taken in low traffic periods to obtain a better view of the scenes.

Central European cities have a mean of 646 plant species/city and larger cities have more species (Pysek 1998), thus, considering that San Jose was and is a small city, the total of 458 species recorded is within the expected range.

Successful urban plants tend to belong to species adapted to natural habitats with strong sunlight, abundant nitrogen and low water levels (Shaltout & El-Sheikh 2002, Chocholouskovaa & Pysek 2003, Lundholm & Marlin 2006), so the presence of many Asteraceae, Fabaceae, Poaceae, Lamiaceae, Euphorbiaceae, Solanaceae, Cyperaceae, Acanthaceae, Malvaceae, Piperaceae and Verbenaceae is not surprising. Many of the plant species in Costa Rica are introduced (Chacon & Saborio 2005) and here again the situation is similar to that in other countries.

Urban trees mitigate global warming (Abdollahi et al. 2000, McPherson et al. 2008), significantly reduce urban heat islands (Huang et al. 2009) and can sequester about 100kg of air pollutants per hectare of urban forest (Vilela-Lozano 2004). Furthermore, urban vegetation protects many species in the five kingdoms (Dana et al. 2002, Smith et al. 2006), including valuable rare species (Maurer et al. 2000, Williams et al. 2009). For these and other reasons, the study and management of urban vegetation is of great importance.

Genetic diversity is low in urban plants and they are less prepared to cope with environmental change (Knapp et al. 2009), so periodic monitoring is needed to conserve original species as well as any others in need of protection (Godefroid 2001). Citizens can learn to effectively take advantage of urban vegetation (Garzon et al. 2004) and to recognize historical changes in the city scene (e.g. the repeat photography groups in, not only for cultural reasons, but also to influence the administration of the urban flora by local governments.

Future studies of urban vegetation in San Jose could investigate these hypotheses:

* Floristic composition results from the interaction of human density, water availability, temperature, altitude and soil (Dana et al. 2002, Chocholouskovaa & Pysek 2003, Fanelli & Tescarollo 2006, Altobelli et al. 2007).

* Ecological succession starts with ruderal annual plants, followed by perennials (Prach et al. 2001).

* Alien species benefit more from human activity (Niggermann 2009).

* With global warming, species adapted to colder climate will become less common and vice versa.

Species collected in the late 19th and early 20th century

Species                          Date               Collector

Acalypha macularis            1906-10-28  Oton Jimenez 46
Acalypha amentacea            1906-12-03  Oton Jimenez 95
Acanthus mollis               1931-03-25  Juvenal Valerio 73
Achimenes grandiflora         1888-07-31  Paul Biolley
Achimenes longiflora          1888-07-31  Paul Biolley
Acmella oppositifolia         1889-06-20  H. Pittier 1091
Acmella radicans              1892-11-30  A. Tonduz 7185
Acnistus arborescens          1893-06-30  A. Tonduz
Adiantum colpodes             1896-07-31  A. Tonduz 8064
Adiantum patens               1888-01-15  H. Pittier 48
Adiantum concinnum            1945-08-06  Richard W. Holm 991
Aeschynomene villosa          1890-12-31  Paul Biolley
Agave wercklei                1907-02-13  Anastasio Alfaro
Ageratum conyzoides           1931-01-03  Fernando Solis 2
Ageratum microcarpum          1893-01-31  A. Tonduz 7281
Albizia adinocephala          1890-07-31  Paul Biolley
Aleurites moluccanus          1906-10-30  Oton Jimenez 127
Alternanthera laguroides      1896-06-30  A. Tonduz 10135
Amaranthus hybridus           1888-07-31  H. Pittier 398
Amphilophium paniculatum      1888-01-31  H. Pittier 975
Anagallis pumila              1893-02-28  A. Tonduz
Anemia hirsuta                1888-07-31  Paul Biolley 912
Anoda cristata                1931-01-03  Fernando Solis 9
Anredera cordifolia           1937-10-21  Jose Antonio Echeverria 4007
Anthurium acutangulum         1896-09-30  A. Tonduz 10360
Aphelandra deppeana           1896-12-31  A. Tonduz 10401
Archibaccharis torquis        1892-11-30  A. Tonduz 1535
Archibaccharis schiedeana     1889-12-12  A. Tonduz 1496
Arenaria lanuginosa           1889-06-20  A. Tonduz
Arracacia xanthorrhiza        1904-11-29  A. Tonduz 17454
Arthrostemma ciliatum         1888-01-15  H. Pittier
Asclepias curassavica         1892-07-31  A. Tonduz 436
Asplenium pumilum             1888-07-31  Paul Biolley 906
Asplenium aethiopicum         1889-06-20  A. Tonduz 1095
Asplenium myriophyllum        1888-07-31  Paul Biolley 516
Asplenium formosum            1888-07-31  Paul Biolley 914
Aster spinosus                1888-07-31  Paul Biolley 990
Athyrium palmense             1888-07-31  H. Pittier 905
Axonopus compressus           1890-08-31  A. Tonduz 2828
Baccharis braunii             1896-09-30  A. Tonduz 10882
Begonia cuculata              1896-12-31  A. Tonduz 10399
Bidens pilosa                 1888-07-31  Paul Biolley 980
Blechnum occidentale          1888-07-31  Paul Biolley 921
Blechum brownei               1890-12-07  A. Tonduz 3204
Blechum pyramidatum           1931-03-25  Juvenal Valerio 79
Bletia purpurea               1903-05-31  H. Pittier 16726
Bouvardia glabra              1896-09-30  A. Tonduz
Brassica campestris           1889-10-31  A. Tonduz 3535
Browallia americana           1892-11-30  A. Tonduz 7183
Brugmansia candida            1892-07-31  A. Tonduz 442
Byttttneria aculeata          1888-07-31  Paul Biolley
Byttneria carthaginensis      1888-09-30  Paul Biolley
Calea axillaris               1889-11-23  A. Tonduz 1430
Calea urticifolia             1931-01-03  Fernando Solis
Calliandra calothyrsus        1888-08-31  H. Pittier
Calyptocarpus wendlandii      1890-05-15  A. Tonduz 2334
Calyptocarpus wendlandii      1931-01-03  Fernando Solis
Calyptranthes pallens         1929-05-21  Manuel Valerio
Campyloneurum irregulare      1888-07-31  Paul Biolley 922
Campyloneurum xalapense       1902-03-31  Anastasio Alfaro
Canavalia villosa             1889-11-23  A. Tonduz
Canna indica                  1906-11-30  101
Capsicum annuum               1895-09-30  Paul Biolley
Cassia tonduzii               1896-06-30  A. Tonduz
Cassia patellaria             1888-10-31  H. Pittier
Cassia pubescens              1889-09-30  A. Tonduz
Casuarina cunninghamiana      1906-09-29  Oton Jimenez 84
Cedrela tonduzii              1903-06-30  H. Pittier
Centaurium quitense           1890-12-07  A. Tonduz
Centradenia inaequilateralis  1888-07-31  Paul Biolley
Cestrum aurantiacum           1888-07-31  Paul Biolley
Cestrum lanatum               1893-01-31  A. Tonduz
Cestrum macrophyllum          1901-02-28  A. Tonduz
Cestrum warscewiczii          1888-01-15  H. Pittier 47
Cestrum glanduliferum         1929-05-05  A. M. Brenes 37
Cestrum tomentosum            1941-12-09  Jose Antonio Echeverria 230
Cestrum nocturnum             1936-04-29  Juvenal Valerio
Chamaedorea costaricana       1905-03-30  Paul Biolley
Chamaesyce lasiocarpa         1892-02-28  A. Tonduz
Chenopodium ambrosioides      1888-01-31  H. Pittier
Chiococca alba                1890-07-31  Paul Biolley
Chloris virgata               1890-09-15  A. Tonduz 2937
Chromolaena odorata           1896-08-31  A. Tonduz 10894
Chrysophyllum cainito         1902-09-15  H. Pittier
Cinchona pubescens            1937-09-07  A. M. Brenes
Cissampelos pareira           1896-06-22  A. Tonduz
Citharexylum donnell-smithii  1895-01-31  A. Tonduz 9623
Cleome costaricensis          1889-05-20  A. Tonduz 1086
Cleome pilosa                 1889-11-28  A. Tonduz 1450
Clerodendrum thomsonae        1928-04-29  Manuel Valerio 64
Clibadium surinamense         1885-07-31  Paul Biolley 976
Cnidoscolus urens             1903-05-31  H. Pittier
Coelogyne speciosa            1932-11-11  A. M. Brenes 366
Commelina diffusa             1890-10-31  A. Tonduz 3049
Commelina leiocarpa           1931-01-03  Fernando Solis 45
Consolida ajacis              1940-11-20  Maria del Carmen Roviralta 10
Conyza bonariensis            1892-10-31  A. Tonduz 857
Conyza schiedeana             1892-10-31  A. Tonduz 856
Conyza canadensis             1931-01-03  Fernando Solis
Cordia spinescens             1896-06-30  A. Tonduz 8879
Cordia eriostigma             1933-04-15  Fernando Solis 527
Crinum erubescens             1893-08-31  A. Tonduz 8214
Crotalaria rotundifolia       1892-06-30  A. Tonduz
Crotalaria vitellina          1892-09-30  A. Tonduz
Crotalaria cajanifolia        1888-03-31  Paul Biolley
Crotalaria acapulcensis       1942-09-30  Jose Antonio Echeverria
Croton hoffmannii             1896-06-30  A. Tonduz
Croton draco                  1892-12-31  A. Tonduz 7261
Cuphea carthagenensis         1891-06-30  A. Tonduz
Cuphea wrightii               1889-07-27  A. Tonduz
Cuphea appendiculata          1888-07-31  Paul Biolley
Cyclanthera tonduzii          1889-11-28  A. Tonduz 1449
Cyclospermum leptophyllum     1889-08-12  A. Tonduz 1287
Cynoglossum amabile           1940-11-13  Maria del Carmen Roviralta 8
Cyperus papyrus               1906-09-29  Oton Jimenez 76
Cyperus flavescens            1892-10-31  A. Tonduz 1531
Cyperus niger                 1890-12-07  A. Tonduz
Cyperus tenuis                1892-07-31  A. Tonduz 433
Cyperus involucratus          1896-09-30  A. Tonduz 10888
Cyperus mutisii               1906-11-11  Oton Jimenez 69
Cyperus hermaphroditus        1906-11-11  Oton Jimenez 68
Cystopteris fragilis          1888-07-31  Paul Biolley 414
Dahlia rosea                  1888-01-15  H. Pittier 58
Dalea cliffortiana            1890-07-31  A. Tonduz
Delilia biflora               1896-12-31  A. Tonduz 10893
Delilia biflora               1931-01-03  Fernando Solis
Dennstaedtia distenta         1922-08-07  Manuel Valerio
Desmodium affine              1888-07-31  Paul Biolley
Desmodium distortum           1902-11-30  H. Pittier
Desmodium intortum            1892-11-26  A. Tonduz
Desmodium tortuosum           1902-12-31  H. Pittier
Desmodium maxonii             1931-01-03  Fernando Solis
Dicliptera unguiculata        1931-01-03  Fernando Solis 10
Dieffenbachia oerstedii       1890-05-15  A. Tonduz 2558
Dietes grandiflora            1931-04-11  A. M. Brenes 89
Dorstenia contrajerva         1896-07-31  A. Tonduz
Dracaena fragrans             1934-02-27  Juvenal Valerio 1078
Drymaria cordata              1889-10-31  A. Tonduz
Drymaria villosa              1892-06-30  A. Tonduz
Drymonia serrulata            1896-07-31  A. Tonduz
Dryopteris opposita           1888-07-31  Paul Biolley 413
Dryopteris patula             1888-07-31  Paul Biolley 923
Dryopteris litigiosa          1896-09-30  A. Tonduz 132088
Duranta erecta                1934-12-30  A. M. Brenes 22868
Echinochloa crus-galli        1888-12-31  H. Pittier 229
Echinochloa crus-pavonis      1890-09-08  3016
Echinocystis coulteri         1889-11-30  A. Tonduz
Eclipta prostrata             1892-10-31  A. Tonduz 9570
Eleocharis elegans            1906-11-11  Oton Jimenez 74
Elephantopus mollis           18-11-1931  Manuel Quiros 371
Elephantopus scaber           1889-11-30  A. Tonduz
Elephantopus spicatus         1889-12-12  A. Tonduz 1494
Encyclia livida               1890-03-10  A. Tonduz 2176
Eragrostis ciliaris           1890-09-15  A. Tonduz 2935
Eragrostis hypnoides          1890-05-15  2477
Eragrostis mexicana           1890-09-15  A. Tonduz 2932
Erechtites hieraciifolius     1888-08-26  H. Pittier 462
Erigeron cuneifolium          1888-11-30  H. Pittier 649
Eryngium carlinae             1889-06-30  A. Tonduz 1124
Erythrina berteroana          1899-12-31  A. Tonduz
Erythrina berteroana          1934-03-14  Alvaro Sanchez
Erythrina crista-galli        1931-04-01  Fernando Solis
Eucalyptus globulus           1889-11-30  A. Tonduz
Eucalyptus globulus           1941-08-20  Jose Antonio Echeverria 25
Euphorbia heterophylla        1889-10-31  A. Tonduz
Euphorbia heterophylla        1931-01-03  Fernando Solis
Euphorbia pulcherrima         1931-03-25  A. M. Brenes
Ficus jimenezii               1906-11-29  A. Tonduz 17536
Ficus velutina                1901-11-30  H. Pittier
Ficus pertusa                 1901-11-30  H. Pittier
Ficus goldmanii               1897-10-31  A. Tonduz
Ficus costaricana             1901-11-30  H. Pittier
Fimbristylis dichotoma        1893-12-31  A. Tonduz 1812
Galeana pratensis             1889-07-27  A. Tonduz 1237
Galeana pratensis             1931-01-03  Fernando Solis
Galinsoga quadriradiata       1888-07-31  Paul Biolley 994
Galinsoga quadriradiata       1889-08-12  A. Tonduz 253
Gonolobus edulis              1888-07-31  Paul Biolley 938
Gossypium barbadense          1893-05-31  A. Tonduz
Graptophyllum pictum          1931-03-25  A. M. Brenes 24416
Grevillea banksii             1938-01-11  Jorge Leon 1003
Habenaria clypeata            1903-08-31  H. Pittier 16722
Hamelia patens                1889-07-31  A. Tonduz
Hauya elegans                 1930-07-08  Manuel Valerio 51
Heliotropium indicum          1889-10-31  A. Tonduz 1146
Heliotropium procumbens       1890-05-31  A. Tonduz 2430
Heteranthera reniformis       1888-10-31  H. Pittier 553
Heterocentron glandulosum     1892-11-30  A. Tonduz 1541
Hibiscus rosa-sinensis        1896-12-31  A. Tonduz
Hibiscus rosa-sinensis        1931-04-23  Vitalia Saenz 96
Holmskioldia sanguinea        1941-11-10  Jose Antonio Echeverria 92
Homalocladium platycladum     1940-12-07  Jose Antonio Echeverria 4015
Hydrocotyle ranunculoides     1889-12-31  A. Tonduz 1688
Hymenocallis littoralis       1905-06-29  A. Tonduz
Hypericum uliginosum          1888-07-31  Paul Biolley
Hypoxis decumbens             1893-06-30  A. Tonduz 8028
Hyptis suaveolens             1889-10-31  A. Tonduz
Hyptis mutabilis              1893-01-02  A. Tonduz
Hyptis capitata               1931-01-03  Fernando Solis
Hyptis pectinata              1931-01-03  Fernando Solis
Hyptis urticoides             1931-01-03  Fernando Solis
Indigofera costaricensis      1892-10-31  A. Tonduz
Indigofera tephrosioides      1888-07-31  Paul Biolley
Inga densiflora               1934-05-19  Collector data missing
Inga vera                     1888-07-31  H. Pittier
Inga leptoloba                1888-05-31  H. Pittier
Inga oerstediana              1888-05-31  H. Pittier
Ipomoea alba                  1932-01-06  Fernando Solis 405
Ipomoea aristolochiifolia     1933-01-01  Manuel Quiros 608
Ipomoea batatas               1932-01-06  Fernando Solis 407
Ipomoea carnea                1931-03-25  Juvenal Valerio 81
Ipomoea parasitica            1931-01-04  Fernando Solis 91
Iresine diffusa               1893-01-01  A. Tonduz 7241
Jaegeria hirta                1888-0-731  Paul Biolley 940
Jaltomata repandidentata      1928-10-05  A. M. Brenes
Jasminum revolutum            1896-09-30  A. Tonduz
Jatropha gossypiifolia        1906-12-03  Oton Jimenez 94
Justicia crenata              1887-05-31  Paul Biolley 1013
Justicia aurea                1940-11-13  Maria del Carmen Roviralta 9
Koellikeria erinoides         1888-07-31  Paul Biolley
Kyllinga odorata              1892-07-31  A. Tonduz 434
Lagerstroemia indica          1891-03-31  H. Pittier
Lantana urticifolia           1889-08-31  A. Tonduz
Lantana hirta                 1888-07-31  A. Tonduz
Laportea aestuans             1936-11-14  Juvenal Valerio 1388
Lasiacis sorghoidea           1893-01-01  A. Tonduz 7234
Lasiacis swartziana           1888-02-20  H. Pittier 81
Lasiacis sorghoidea           1931-01-03  Fernando Solis
Lastreopsis effusa            1889-07-27  A. Tonduz 1128
Leersia hexandra              1889-10-31  A. Tonduz 1383
Leonurus japonicus            1890-09-30  A. Tonduz 1399
Leonurus japonicus            1931-01-03  Fernando Solis 25
Lepidium bipinnatifidum       1892-07-31  A. Tonduz 437
Ligustrum vulgare             1931-04-14  Fernando Solis 192
Limonium sinuatum             1931-04-11  A. M. Brenes 87
Lindernia diffusa             1893-02-28  A. Tonduz
Lipocarpha micrantha          1941-11-06  Jose Antonio Echeverria 133
Lippia myriocephala           1892-11-26  Paul Biolley
Lippia alba                   1888-07-31  Paul Biolley
Lippia cardiostegia           1888-12-31  H. Pittier
Lobelia xalapensis            1892-10-31  A. Tonduz
Loeselia glandulosa           1931-01-04  Fernando Solis 80
Lophospermum erubescens       1930-06-10  Vitalia Saenz
Ludwigia octovalvis           1889-10-31  A. Tonduz
Ludwigia peruviana            1889-10-31  A. Tonduz
Ludwigia peruviana            1888-01-15  H. Pittier 41
Ludwigia octovalvis           1931-01-03  Fernando Solis 19
Malachra radiata              1889-11-30  A. Tonduz
Malachra alceifolia           1928-10-05  A. M. Brenes 19
Malpighia glabra              1890-04-30  A. Tonduz
Malpighia mexicana            1891-08-31  H. Pittier
Malva parviflora              1891-08-31  H. Pittier
Malvaviscus arboreus          1892-12-31  A. Tonduz
Mangifera indica              1888-05-31  H. Pittier 388
Manihot glaziovii             1890-11-30  A. Tonduz
Marsypianthes chamaedrys      1892-07-31  A. Tonduz
Marsypianthes chamaedrys      1931-01-03  Fernando Solis
Masdevallia ecaudata          1890-11-30  Paul Biolley 3127
Maurandia barclayana          1928-04-29  Manuel Valerio
Mauria heterophylla           1894-01-31  Paul Biolley 8475
Mazus pumilus                 1937-04-11  Jorge Leon
Mecardonia procumbens         1892-07-31  A. Tonduz 7113
Melampodium divaricatum       1892-07-31  A. Tonduz 6960
Melampodium gracile           1931-01-03  Fernando Solis
Melanthera nivea              1893-11-30  A. Tonduz 1558
Melanthera nivea              1931-01-03  Fernando Solis
Mikania micrantha             1931-01-03  Fernando Solis
Mildella intramarginalis      1892-11-26  A. Tonduz 7220
Mimosa diplotricha            1893-02-28  A. Tonduz
Mimosa pudica                 1888-07-31  Paul Biolley
Mimosa sensitiva              1889-09-28  A. Tonduz 1279
Mirabilis jalapa              1931-01-03  Fernando Solis 27
Mitracarpus hirtus            1931-01-03  Fernando Solis 36
Mollugo verticillata          1941-03-06  Jose Antonio Echeverria 94
Monstera adansonii            1890-07-31  Paul Biolley 2846
Montanoa guatemalensis        1893-02-28  A. Tonduz 7331
Moringa oleifera              1896-07-31  Paul Biolley
Moritzia lindenii             1896-09-30  A. Tonduz 10886
Muhlenbergia tenella          1890-10-09  A. Tonduz 3015
Muhlenbergia tenella          1906-10-30  Oton Jimenez 50
Myriophyllum aquaticum        1941-10-18  Vitalia Saenz 52
Myrtus communis               1928-04-29  Manuel Valerio
Nasturtium mexicanum          1891-08-31  H. Pittier 6906
Nectandra martinicensis       1896-06-22  A. Tonduz 10104
Nectandra turbacensis         1897-06-24  H. Pittier
Nephrolepis occidentalis      1908-10-06  Humberto Bertolini
Nephrolepis undulata          1922-08-07  Manuel Valerio
Nicotiana tabacum             1905-06-29  A. Tonduz
Ochroma pyramidale            1941-08-20  Jose Antonio Echeverria 27
Oenothera biennis             1905-06-29  A. Tonduz 17490
Oenothera rosea               1936-04-29  Juvenal Valerio 1119
Oleandra costaricensis        1908-09-04  Humberto Bertolini
Ophioglossum reticulatum      1888-07-31  Paul Biolley 925
Oplismenus burmannii          1890-12-07  A. Tonduz 3124
Oplismenus burmannii          1906-11-27  Oton Jimenez 9
Ornithocephalus bicornis      1904-12-30  Paul Biolley 17512
Oxalis latifolia              1896-07-31  A. Tonduz
Pachira aquatica              1933-04-15  Fernando Solis 526
Pachyrhizus erosus            1905-05-30  A. Tonduz
Paspalum costaricense         1890-10-31  A. Tonduz 3028
Paspalum saccharoides         1891-08-31  H. Pittier 6907
Paspalum orbiculatum          1889-06-30  H. Pittier 1183
Paspalum disttichum           1888-07-20  H. Pittier 306
Paspalum notatum              1906-11-29  Oton Jimenez 29
Paspalum conjugatum           1892-09-30  A. Tonduz 758
Passiflora adenopoda          1929-08-30  Jose Maria Orozco 42
Passiflora apetala            1896-02-28  John Donnell Smith
Passiflora edulis             1905-04-29  A. Tonduz
Paullinia barbadensis         1893-01-31  A. Tonduz
Pecluma plumula               1888-07-31  Paul Biolley 920
Pedilanthus tithymaloides     1933-04-06  Manuel Quiros
Pellaea ovata                 1908-10-06  Humberto Bertolini
Peperomia tetraphylla         1891-11-30  A. Tonduz 3198
Peperomia lanceolatopeltata   1892-10-05  A. Tonduz 7262
Peperomia cooperi             1888-07-31  Paul Biolley 524
Peperomia lignescens          1898-09-30  A. Tonduz
Peperomia galioides           1892-11-26  A. Tonduz
Peperomia angularis           1893-01-01  A. Tonduz
Peperomia deppeana            1890-07-31  A. Tonduz
Phaseolus lunatus             1893-01-02  A. Tonduz
Phaseolus lunatus             1888-01-15  H. Pittier 52
Phaseolus lunatus             1932-12-25  Fernando Solis
Philadelphus myrtoides        1889-12-31  A. Tonduz 1492
Phlebodium pseudoaureum       1888-07-31  Paul Biolley 900
Phthirusa pyrifolia           1892-08-31  A. Tonduz
Phthirusa pyrifolia           1904-10-30  A. Tonduz
Phyla scaberrima              1889-07-27  A. Tonduz
Phyllanthus niruri            1888-07-20  H. Pittier
Piper bredemeyeri             1889-06-20  H. Pittier 1088
Piper hispidum                1896-07-31  A. Tonduz 10154
Piper aduncum                 1896-07-31  A. Tonduz
Piper umbellatum              1892-06-30  A. Tonduz 693
Pittiera longipedunculata     1890-07-12  A. Tonduz 3200
Plantago australis            1894-07-31  A. Tonduz
Plantago major                1894-07-31  A. Tonduz
Plectranthus amboinicus       1942-04-29  Manuel Quiros
Pleopeltis macrocarpa         1889-07-27  A. Tonduz 1229
Pleopeltis astrolepis         1892-07-31  A. Tonduz 7121
Pleurothallis listerophora    1890-07-31  Paul Biolley 2986
Poa annua                     1888-05-31  H. Pittier 230
Polyclathra cucumerina        1890-07-12  A. Tonduz 3200
Polygala platycarpa           1888-05-31  Paul Biolley
Polygala costaricensis        1889-11-28  A. Tonduz 1450
Polygala violacea             1888-07-26  Paul Biolley 348
Polygonum punctatum           1892-07-31  A. Tonduz
Polypodium triseriale         1888-07-31  Paul Biolley 899
Polypodium polypodioides      1905-10-01  Oton Jimenez 291
Polypodium triseriale         1888-07-31  Paul Biolley 417
Ponthieva racemosa            1890-12-31  Paul Biolley 3240
Portulaca oleracea            1890-03-31  H. Pittier
Prockia crucis                1891-05-31  A. Tonduz
Pseuderanthemum cuspidatum    1888-07-31  Paul Biolley 942
Psidium solisii               1932-12-25  Fernando Solis 509
Psidium cattleianum           1909-09-21  A. Tonduz
Psidium cattleianum           1931-06-07  Manuel Quiros
Psychotria pubescens          1896-07-31  A. Tonduz
Pterolepis pumila             1888-12-12  H. Pittier 10
Punica granatum               1905-04-29  A. Tonduz 17469
Pycreus rivularis             1906-10-11  Oton Jimenez 75
Rhynchelytrum repens          1906-11-19  Oton Jimenez 58
Rhynchosia longeracemosa      1892-11-26  A. Tonduz
Rhynchospora nervosa          1906-11-29  Oton Jimenez 124
Richardia scabra              1896-07-31  A. Tonduz
Ricinus communis              1889-11-30  A. Tonduz
Rivina humilis                1890-10-31  A. Tonduz
Robinsonella divergens        1889-11-28  A. Tonduz
Robinsonella lindeniana       1931-01-12  Fernando Solis 110
Rorippa mexicana              1887-12-20  H. Pittier 29
Rosa multiflora               1892-06-30  A. Tonduz
Rubus urticifolius            1888-07-31  H. Pittier
Rumex obtusifolius            1896-07-31  A. Tonduz 10123
Russelia sarmentosa           1889-07-27  A. Tonduz
Rytidostylis carthaginensis   1888-07-31  Paul Biolley
Sabazia urticifolia           1896-07-31  A. Tonduz 10143
Salvia wagneriana             1889-11-30  A. Tonduz 1476
Salvia occidentalis           1893-01-31  A. Tonduz
Salvia costaricensis          1895-10-31  A. Tonduz
Salvia lasiocephala           1893-01-02  A. Tonduz
Salvia polystachya            1888-02-20  H. Pittier
Salvia alvajaca               1892-07-31  A. Tonduz 701
Salvia occidentalis           1931-01-03  Fernando Solis
Salvia officinalis            1938-03-30  Manuel Quiros
Salvia polystachya            1931-01-03  Fernando Solis
Salvia tiliifolia             1931-01-03  Fernando Solis
Salvia wagneriana             1940-10-19  Maria del Carmen Roviralta
Sambucus mexicana             1931-07-07  Fernando Solis 288
Sambucus mexicana             1931-07-07  Fernando Solis 288
Sansevieria hyacinthoides     1941-02-18  Jose Antonio Echeverria 4057
Sapium macrocarpum            1893-07-31  A. Tonduz 8209
Scleria hirtella              1888-11-30  H. Pittier 648
Scoparia dulcis               1889-07-31  A. Tonduz
Scoparia dulcis               1931-09-04  Fernando Solis
Scutellaria purpurascens      1891-06-30  A. Tonduz
Sechium tacaco                1902-10-31  K. Werkle 16674
Selaginella serpens           1888-07-31  Paul Biolley 927
Selaginella cuspidata         1888-07-31  Paul Biolley 929
Senecio hoffmannii            1888-01-15  H. Pittier 39
Senna papillosa               1896-07-31  A. Tonduz
Serjania acuta                1897-03-31  A. Tonduz
Setaria geniculata            1890-10-04  A. Tonduz 3008
Sicyos sertuliferus           1932-12-25  Fernando Solis 1100
Sida haenkeana                1891-05-31  H. Pittier
Sigesbeckia jorullensis       1931-01-03  Fernando Solis
Simarouba glauca              1944-06-28  Jose Antonio Echeverria 478
Smallanthus maculatus         1892-07-31  A. Tonduz 699
Smilax spinosa                1889-11-28  A. Tonduz 1466
Solanum lanceolatum           1892-09-30  A. Tonduz
Solanum americanum            1890-03-25  A. Tonduz
Solanum americanum            1931-01-03  Fernando Solis 33
Solanum umbellatum            1934-03-17  Estrella U. de Pacheco
Solanum wendlandii            1932-03-23  Collector data missing
Sonchus oleraceus             1890-11-30  A. Tonduz 3069
Sorghum bicolor               1888-07-31  H. Pittier 383
Spananthe paniculata          1896-09-30  A. Tonduz 10883
Spananthe paniculata          1931-01-03  Fernando Solis 64
Spathodea campanulata         1901-11-30  H. Pittier 16212
Spermacoce assurgens          1896-07-31  A. Tonduz
Spigelia splendens            1888-07-31  Paul Biolley
Sporobolus indicus            1890-05-15  A. Tonduz 2335
Stachys costaricensis         1887-12-20  H. Pittier 34
Stachys costaricensis         1896-07-31  A. Tonduz
Stelis tricuspis              1888-07-31  Paul Biolley 949
Stellaria prostrata           1890-10-31  A. Tonduz
Stellaria ovata               1931-01-03  Fernando Solis 34
Stemodia verticillata         1895-01-31  A. Tonduz 9621
Struthanthus orbicularis      1893-01-31  A. Tonduz
Stylosanthes guyanensis       1896-11-30  A. Tonduz
Synadenium grantii            1941-11-10  Jose Antonio Echeverria 103
Syzygium jambos               1889-10-31  A. Tonduz
Tabebuia rosea                1890-02-28  Paul Biolley 2215
Tagetes microglossa           1889-11-28  A. Tonduz 1451
Talinum paniculatum           1889-07-27  A. Tonduz
Tecoma stans                  1889-12-12  A. Tonduz 1495
Tetrapterys schiedeana        1898-07-31  Paul Biolley
Thalictrum lankesteri         1888-07-31  Paul Biolley
Thelypteris resinifera        1886-01-31  Paul Biolley 1067
Thunbergia alata              1936-04-29  Juvenal Valerio 1136
Thunbergia erecta             1940-11-13  Maria del Carmen Roviralta 5
Tibouchina longifolia         1893-12-31  A. Tonduz
Tinantia erecta               1888-07-31  H. Pittier 391
Tithonia longiradiata         1890-12-03  A. Tonduz 3136
Tournefortia glabra           1889-06-20  A. Tonduz 1089
Tragia volubilis              1892-07-31  A. Tonduz
Trichilia havanensis          1941-01-01  Jose Antonio Echeverria 4023
Trichilia martiana            1896-06-22  A. Tonduz
Trifolium amabile             1889-08-12  A. Tonduz
Tripogandra purpurascens      1889-07-27  A. Tonduz 1252
Tripogandra serrulata         1889-06-20  A. Tonduz 1186
Tripsacum andersonii          1901-08-31  Brade 16174
Verbena litoralis             1936-08-19  Rafael Roig 9
Vigna adenantha               1931-01-03  Fernando Solis
Vismia baccifera              1896-06-30  A. Tonduz
Xylosma flexuosa              1895-01-31  A. Tonduz
Xylosma velutina              1902-03-31  Paul Biolley
Zephyranthes carinata         1893-06-30  A. Tonduz 8045
Zexmenia longipes             1890-12-03  A. Tonduz 3135
Zexmenia costaricensis        1892-07-31  A. Tonduz 7122
Zexmenia frutescens           1909-12-30  A. Tonduz 17977


We thank Hubert Blanco (Archivos Nacionales), Colegio Superior de Senoritas, Maria Jose Guerra Araus (Herbario Nacional), Sergio Aguilar, Sergio Quesada, Karla Vega, Andrea Sanchez and Maria Acuna for their assistance. We specially thank Sally P. Horn (University of Tennessee, Knoxville) for suggestions to improve the manuscript and for advice on repeat photography.


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Julian Monge-Najera & Gabriela Perez-Gomez

Laboratorio de Ecologia Urbana, Vicerrectoria de Investigacion, Universidad Estatal a Distancia, 474-2050 San Pedro de Montes de Oca, San Jose, Costa Rica;,

Received 01-VI-2010. Corrected 28-VII-2010. Accepted 06-VIII-2010.
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Author:Monge-Najera, Julian; Perez-Gomez, Gabriela
Publication:Revista de Biologia Tropical
Article Type:Report
Date:Dec 1, 2010
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