Flora of Barro Colorado Island

Phenological Characteristics

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Leaf fall and leaf flushing. Phenological observations of the three classical sorts-flowering, fruiting, and leaf fall-have been made for most species. Observations of leaf fall were based on field notes made on BCI and in adjacent areas of the Canal Zone during the years 1967-74. Observations of flowering and fruiting were made at the same time, and were supplemented by studies of herbarium specimens (Croat, 1975d).

Many plants, particularly the trees and lianas, lose their leaves in the dry season. In many species, the leaves fall at or near the onset of the dry season, but in some, leaf shedding is continuous throughout the dry season. Studies conducted by the U.S. Army Tropic Test Center (1966) at the Albrook Air Force Base test site on the Pacific slope in Panama show that litter fall declines sharply in February and March. Litter accumulation increases until May, then drops sharply. Similar studies by the Smithsonian Environmental Monitoring Program (Rubinoff, 1974) show maximum leaf-litter accumulations in December and January, followed by a rapid decrease in February and further diminishment in April and May. By February, the forest canopy begins to look bare, at least relative to its appearance in the rainy season, and the atmospheric humidity is much lower. Winds, which increase markedly during the dry season, may be felt even at ground level in the depths of the forest.

Leaf litter, which includes falling flowers and fruits and other debris, accumulates to a depth of several inches in some places by the end of the dry season (personal observation). Measurements by Woods and Gallegos on BCI (1970) show that more than 10 metric tons per hectare of litter accumulate during the months June through August.

The beginning of the rainy season brings a rapid increase in the decay of the leaf litter, for the increased soil moisture and atmospheric humidity greatly increase the number of decomposing organisms. The largest part of the leaf litter decomposes within a few weeks of the first rains (I. Healey, pers. comm.). At least in the early stages of the rainy season, some leaf litter may be washed away, for the water currents in the streams can become quite strong-during heavy rainstorms, debris is carried by water currents along trails even in the flat areas of the forest. The rains, however, serve mostly to compact the litter.

Williams (1941) reported that there is a renewal of litter organisms in May, with the beginning of the rains, and that by the early part of July there is a marked increase in the number of forms present. Fungal organisms as well, which are not common during the dry season, are abundant during the rainy season.

Since many nutrients become available shortly after the onset of the rainy season, it can be assumed that plants are absorbing them at a greater rate during the early weeks of the rainy season, though it is not known how long it takes the plants to assimilate these nutrients. If the assimilation were sufficiently rapid, this influx of nutrients might have some effect on seed germination, leaf maturation, or flower and fruit production. Indeed, emerging seedlings appear to be the most abundant at the end of May and the beginning of June (N. Garwood, pers. comm.), but this may reflect only the increase in soil moisture-that is, it may be unrelated to nutrient availability. Frankie, Baker, and Opler (1974) have shown that for lowland wet forest in Costa Rica, the peak of leaf flushing occurs during the major dry season, especially in February, and a second peak occurs in September, just after the minor dry season. On BCI my general impression is that most flushing of new leaves occurs early in the rainy season. However, random observations on 103 shrubs and trees show that there is no marked difference between the number of species that put on new leaves early in the rainy season and those that put them on in the dry season. Six species show leaf flushing both early in the rainy season and in the dry season, whereas 45 show leaf flushing in the dry season and 42 early in the rainy season. An additional 10 species show leaf flushing both late in the dry season and early in the rainy season, and should be considered as rainy-season leaf flushers. But even if these are included with the rainy-season species, the difference between 45 in the dry and 52 in the rainy is not significant.

Although some species lose and replace their leaves more or less regularly throughout the year, and are never completely leafless, those species that probably contribute most to the accumulation of leaf litter in the dry season are the deciduous species that lose all or nearly all of their leaves for all or part of the dry season; they are the following:

Annona spraguei

Dalbergia retusa

Topobaea praecox
Bauhinia guianensis Enterolobium cyclocarpum Trichilia hirta
Bombacopsis quinata Erythrina fusca Xylophragma
B. sessilis Jacaranda copaia Xylosma chloranthum
Bursera simaruba Pseudobombax septenatum Zanthoxylum belizense
Cavanillesia platanifolia Pterocarpus officinalis Z. panamense
Cedrela odorata Sapium caudatum Z. setulosum
Ceiba pentandra seemannianum Tabebuia guayacan Zuelania guidonia
Cochlospermum vitifolium    

A few species are leafless during the rainy season. Among these are:

Cordia alliodora Ochroma pyramidale
Erythrina costaricensis Triplaris cumingiana

Many species are leafless for only a short time, usually just prior to flowering; often, the leaves are replaced while the plant is in flower. Among the species in this group are the following:

Anacardium excelsum Lindackeria laurina Randia armata
Antirrhoea trichantha Lonchocarpus velutinus Schizolobium parahybum
Apeiba membranacea Luehea seemannii Sloanea terniflora
A. tibourbou L. speciosa Spachea membranacea
Casearia corymbosa Machaerium arboreum Omphalea diandra
C. guianensis Malouetia guatemalensis Spondias mombin
Cassia fruticosa Margaritaria nobilis S. radlkoferi
Castilla elastica Eugenia nesiotica Sterculia apetala
Coccoloba acapulcensis Ormosia coccinea var. subsimplex Strychnos panamensis
C. manzanillensis Peltogyne purpurea Tachigalia versicolor
Combretum decandrum Pisonia aculeata Terminalia amazonica
Dendropanax arboreus Pithecellobium macradenium Tetrathylacium johansenii
Dipteryx panamensis Platymiscium pinnatum Trattinnickia aspera
E. oerstedeana Platypodium elegans Trichospermum mexicanum
Genipa americana Poulsenia armata Trophis racemosa
Hura crepitans Psidium anglohondurense Virola surinamensis
Inga fagifolia    

Some species lose their leaves more than once per year. These include Tabebuia rosea and Quararibea asterolepis, which lose their leaves twice a year, and Ficus spp., which lose all leaves several times a year. Other species, such as Beilschmiedia pendula, Byrsonima crassifolia, Jatropha curcas, and Guazuma ulmifolia, replace their leaves gradually, but may at times be almost completely leafless, as well.

Frankie, Baker, and Opler (1974) studied leaf production in a number of species from lowland wet forest in Costa Rica. Many of these same species occur on BCI and may react similarly on BCI, though the seasons are not exactly comparable.

Flowering and fruiting. The data presented in this section are the result of field observations and herbarium studies made between 1967 and 1974, and include observations made during more than three years in Panama and a survey of more than 50,000 herbarium specimens from BCI and adjacent areas.

In these data no attempt has been made to outline the phenology of individual plants, though numerous individuals were repeatedly observed. Instead, the data represent what is thought to be the normal phenological variation for each species-its historical pattern of flowering. No attempt has been made to include the "broad outlier," especially when the phenology of the species involved is well known. In the better-known species, 95% or more of the flowering or fruiting probably falls within the timespan indicated.

The flowering or fruiting period given for most species is broader than that for any single year; because plants have probably evolved a phenology that is compatible with a particular climatic condition (as opposed, say, to calendar month), I have chosen to look at overall phenological patterns rather than what might happen in any one year. The year-to-year variation in phenological pattern for any given species is considerable, and variation can also be great between individuals in a given year, in terms of both timing and duration.

Although other phenological studies have been made (Rovirosa,1892; Fournier & Salas,1966; Janzen,1967b; Smythe,1970; Foster, 1974; Frankie, Baker & Opler, 1974), this is the only known attempt to define an entire flora in a phenological manner.

Excluded from most aspects of this study were the 53 cultivated species; excluded altogether were the 104 species of vascular cryptogams. Earlier studies (Croat, 1969a) have shown that different habit types exhibit different phenological behavior. In this section and in an earlier version of it (Croat, 1975d), different habitats are also shown to produce different phenological behavior. Graphs of flowering and fruiting activity have been prepared for all major habit-and-habitat classes for the flora:

In the graphs, the number of species in flower or fruit in any month is recorded, though months for evident deviates were not tallied in cases where the phenology of a species is well known.

Herbaceous plants, as a single class of organisms, are quite diverse in terms of both habit and habitat, and as a result are more finely subdivided here than the woody plants are. In all, there are 560 herbaceous plants in the BCI flora, accounting for 42.6% of the native flora. Of these, there are 94 vines, 135 epiphytes, 330 terrestrial herbs (including 18 suffruticose herbs and four saprophytic herbs), and one herb parasitic on trees. Because they are inconsequential, saprophytes and parasites are included in the class "forest herbs."

Graphs 2 and 3 show flowering and fruiting curves for all types of herbs studied. I believe that phenological patterns are at least in part determined by fluctuations in climatic conditions. Aquatic herbs and suffruticose herbs are aseasonal, perhaps because they are less subject to the effects of a severe dry season. Aquatics would not be expected to be seasonally cued by availability of precipitate water, but suffruticose herbs, with their well developed woody root system and underground stems, apparently are also little affected by seasonal changes.

The remaining subclasses of herbs are seasonal. The onset of the dry season, with its reduction of soil moisture and atmospheric humidity, as well as its high insolation, appears to act as a cue to flowering. Flowering times for the different habit-and-habitat classes seem to correlate well with their capacity for withstanding conditions of drought.

Graph 2. Numbers of herb species in flower, by month and habit-and-habitat class.
  
Graph 2. Numbers of herb species in flower, by month and habit-and-habitat class.

 

Graph 2. Numbers of herb species in flower, by month and habit-and-habitat class.

Clearing herbs, for example, being the class most exposed to changes in the environment, reach their peak of flowering activity early in the dry season, in December. Flowering then drops off to a relatively steady rate throughout the remainder of the year, except for a slight dip in May and a deeper decline at the end of the rainy season, in November. Because the fruits of most species are small and develop quickly, the fruiting curve closely resembles the flowering curve. Flowering activity in the clearing herbs wanes most at the beginning and end of the rainy season.

Forest herbs, by contrast, reach the peak of their flowering activity early in the rainy season, and their peak of fruiting midway to late in the rainy season. As suggested by Foster (1974), these groups are probably triggered to flower by intense rains following a period of drought.

He has shown that typically rainy-season trees will flower in the dry season if a dry period is followed by heavy rains.

For the herbs of forest light-gaps and forest edges, the amount of light received is relatively more stable, and they are protected from excessive insolation. Their flowering and fruiting activity therefore peaks during the rainy season.

Epiphytic herbs do most of their flowering midway to late in the dry season, with small peaks in February and April, which are perhaps a response to the advancing aridity caused by the increasingly leafless canopy of the forest. Most epiphyte fruiting also occurs in the dry season, and the small airborne fruits are dispersed during the same dry season. A smaller peak of fruiting, early in the rainy season in July, consists principally of the animal-dispersed fruits. Of all epiphyte species whose fruiting is restricted to the dry season, 97% produce principally wind-dispersed seeds, the remainder principally animal-dispersed seeds. Rainy-season epiphytes, by contrast, produce wind-dispersed seeds in only 23% of their species, animal-dispersed fruits in 77%. These figures correlate well with the markedly stronger winds of the dry season-to which wind-dispersed seeds or fruits are particularly well adapted. According to Foster (1974), the leaflessness of the canopy may be more important than winds in the dispersal of airborne diaspores, since many are dispersed after the rains are renewed but before the trees have put on new leaves.

Perhaps because herbaceous vines in the forest usually occur in well-lighted areas and are often restricted to exposed surfaces of the canopy, they do not react appreciably differently from those that occur in clearings. I have therefore treated all herbaceous vines as a group. The flowering peak for herbaceous vines is in December and January, with a second much smaller peak in June. The curve for fruiting in vines, though lacking strong peaks, shows major activity in the dry season. The June peak represents species that appear to be triggered by wet rather than dry conditions.
Of all habit types, the herbs are the least phenologically variable. As many as 224 species (40%) flower and fruit most or all of the year.

The graph for all species of herbs (Graph 2) shows a decrease in flowering activity late in the rainy season, but from the low of 267 species flowering in November to the peak of 307 flowering in July is only a 15% increase. Certain categories of herbs do, however, show significant increases in flowering activity. For example, there is an increase of 58% for vines, 65% for epiphytes, and 79% for all forest herbs (excluding vines).

When not restricted to open areas, such as in clearings or along the lakeshore, woody plants tend to be more seasonal than herbaceous plants (Graphs 4 and 5). As a class, the trees and shrubs of open areas are not very seasonal. Three of the forest habit classes-lianas, large and medium-sized trees, and small trees-reach their principal peak of activity in the dry season. Flowering in the lianas is most active from January to March, especially in February, substantially ahead of the flowering peak for large and medium-sized trees. This disparity probably reflects the fact that the bulk of the leaf biomass of lianas is restricted to the surface of the canopy and is thus quickly affected by conditions of drought. The fruiting peak for lianas occurs late in the dry season, in March and April, and is stronger than the flowering peak -a circumstance perhaps due to the fact that many species of lianas produce wind-dispersed fruits, even though they may flower in the rainy season. For example, there are six species of bignoniaceous lianas that flower in the rainy season and fruit in the dry season. Of liana species that fruit only in the dry season, as many as 80% have wind-dispersed seeds, whereas just 22% of the liana species that fruit only in the rainy season have wind dispersed seeds.

Graph 3. Numbers of herb species in fruid, by month and habit-and-habitat class.
  
Graph 3. Numbers of herb species in fruid, by month and habit-and-habitat class.

 

Graph 3. Numbers of herb species in fruid, by month and habit-and-habitat class.

Large and medium-sized trees reach their peak of flowering activity from February to June, especially in March and April, at the end of the dry season. Though triggered by conditions of drought, they do not react so quickly to changes in the environment as the herbs, vines, and lianas do, perhaps owing to the fact that they are much less exposed to the environment than these other classes are. It may be that a great many of the trees flowering in the dry-wet transition period are triggered to flower by the first heavy rains, as suggested by Foster (1974). Fruiting activity in the large and medium-sized trees shows two peaks, one in April and a second, smaller, one in August. The earlier peak is made up for the most part by wind-dispersed species, whereas the second, rainy-season, peak is made up of species that are mostly animal-dispersed.

The flowering peak of March and April for large and medium-sized trees on BCI contrasts rather sharply with the May and July peaks in the wet forest of Costa Rica (Frankie, Baker & Opler,1974). However, the April and August fruiting peaks on BCI compare well with the May and September peaks in Costa Rica. The flowering and fruiting curves for small trees and shrubs would be somewhat flatter if they also included the class "arborescent in open areas."

Graph 4. Numbers of woody plant species in flower, by month and habit-and-habitat class.
 
Graph 4. Numbers of woody plant species in flower, by month and habit-and-habitat class.

 

Graph 4. Numbers of woody plant species in flower, by month and habit-and-habitat class.

Graph 5. Numbers of woody plant species in fruit, by month and habit-and-habitat class.
  
Graph 5. Numbers of woody plant species in fruit, by month and habit-and-habitat class.

Graph 5. Numbers of woody plant species in fruit, by month and habit-and-habitat class.

Lianas and herbaceous vines are very similar phenologically. If all climbing plants are compared with all arborescent plants (Graph 6), some interesting differences become apparent. As a group, the climbing-plant species reach their peaks of flowering and fruiting earlier than the arboreal plants do. Climbers share the ability to position themselves where they are exposed to light. By the same token, of course, they are subjected to a high degree of exposure when climatic conditions become harsh, as at the beginning of the dry season. It is, I believe, the onset of the dry season that precipitates flowering in many species, including a great many of the lianas, herbaceous vines, and epiphytes, as well as many trees-though most of the trees flower in the dry-wet transitional period and may be induced to flower prematurely by unseasonally wet conditions. Whether this dry-season phenomenon is in general the result of the drying conditions, photoperiodicity, or otherwise is unknown, but the fact that the flowering period of many species coincides with the dry season is no mere coincidence. The flowering curve for all BCI species considered jointly shows a pronounced peak of activity in the dry season (Graph 7). Fruiting shows two peaks, one in the dry-wet transition period and one in the middle of the rainy season. There is a dearth of activity in both flowering and fruiting during October and November, but acute reactivation in December, with the onset of the dry season.

These conclusions differ significantly from those of Foster (1974), who held that the peak month for overall flowering occurs from one to two months after the start of the rains, i.e., in May or June. My studies for overall flowering show significantly fewer species in flower during May and June than in March and April. Foster's conclusion may derive in part from the fact that he was dealing not with the entire flora but with an area of mostly mature forest containing few of the species that are common in open areas or forest edges. Moreover, his detailed sampling procedures deal principally with fruiting, rather less with flowering.

Notwithstanding the overall flowering peak in the dry season, a number of habit-and-habitat classes and a substantial proportion of the species as a whole are apparently cued to flower sometime after the rainy season begins. Foster (1974) has documented this finding for the BCI Psychotria species. Shrubs, forest herbs, and herbs of light-gaps and forest edges all show increased activity in the rainy season. The same can be said for many individual families, especially monocotyledonous herbs such as Marantaceae, Musaceae, Zingiberaceae, and Amaryllidaceae.

It is enlightening to compare the seasonal behavior of these habit-and-habitat classes by examining the number of species in flower in a given month as a percentage of the total number of species in the class (Graphs 8 and 9). Thus, although 96 species of trees more than 10 m tall flower in April, this figure represents only 43% of all such trees, whereas the 43 aquatic herbs that flower in April constitute 75% of all aquatic-herb species. The percent-flowering curves follow the same contours as the absolute flowering curves, but the heights of the curves are substantially different. Even at their peak of flowering activity, the percentage of trees and lianas in flower is smaller than the flowering percentage of any other category. A significantly larger percentage of small trees are flowering or fruiting in every season, peak or low. Several habitat classifications have been combined here for simplification: all small trees and shrubs, including those restricted to open areas, are combined; the curve for all species includes both woody and herbaceous plants, the herbs showing a greater percentage of activity than the woody plants do; and the forest herbs and herbs of forest light gaps and edges are combined, as well as clearing, epiphytic, and suffruticose herbs.

Graph 6. Numbers of species of trees and climbers in flower and fruit, by month.

Graph 6. Numbers of species of trees and climbers in flower and fruit, by month.
Graph 7. Numbers of vascular plant species in flower and fruit, by month.

Graph 7. Numbers of vascular plant species in flower and fruit, by month.

Graph 8. Percentage of species flowering, by month, four groups.

Graph 8. Percentage of species flowering, by month, four groups.

Comparing the fruiting percentages yields a similar pattern, i.e., the same contour as the absolute fruiting curves and at levels corresponding to the percentages for flowering. Of all large and medium-sized trees (excluding bimodal species) whose fruiting occurs strictly in the rainy season, 85% are animal-dispersed and only 12% are wind-dispersed. Of the comparable species that fruit strictly in the dry season, only 36% are animal-dispersed and 57% are wind-dispersed. Comparable figures for small trees and shrubs are 35% animal-dispersed fruits and 21% wind-dispersed fruits in the dry season (the others are mechanically dispersed or are not clearly adapted for either animal or wind dispersal) and nearly 100% animal-dispersed fruits in the wet season.

Small trees are those less than 10 m tall, excluding plants that are always shrubs (i.e., plants usually 1-2 (3) m tall) and excluding, as well, all small trees and shrubs that are restricted to clearings. Small trees as a class have a strong peak of activity late in the dry season, in April; being understory trees they produce, as one would expect, mostly animal-dispersed seeds, for winds are usually not strong enough in the lower levels of the forest, even during the dry season, to disperse successfully most windborne seeds or fruits. The fruiting of the small trees, although not as conspicuously peaked as their flowering, occurs mostly in the rainy season, with a small peak in July and another in September.



The flowering activity of shrubs is not very seasonal, but more flowering occurs early in the wet season than at any other time. Shrub fruiting, by contrast, shows a fairly definite peak in September. Finally, trees and shrubs restricted to open areas show no marked seasonal variation as a class.

In contrast to the 15% difference between high and low points for flowering activity in herbs, there is a difference of 67% for all arborescent plants. This finding includes a difference of 134% for large and medium-sized trees, 123% for small trees, 83% for shrubs, and 68% for lianas.

Another useful means of comparing plants of different habits and habitats is to compare the average lengths of the flowering or fruiting seasons. Since these are taken from what I have termed the usual pattern of flowering, they do not represent the length of flowering or fruiting for any individual or even for any species in a given year, but rather the length of time over which a species has been known to be in flower or fruit. Table 5 shows these figures for most of the classes already discussed. The aquatic herbs, which are markedly aseasonal, are shown to flower and fruit more than 9 months per year on the average, whereas such classes as epiphytic herbs, lianas, and trees, which are all seasonal in their flowering behavior, flower and fruit on the average for substantially shorter periods of time.

Despite the fact that many species are distinctly "dryseason species" or "wet-season species," by no means all of the species that are seasonal are restricted to one of these two periods. Table 6 shows a categorization of seasonality types. It can be seen that at least 346 species are transitional between the two seasons, in either flowering or fruiting behavior. Some of these conclusions can clearly be disregarded, because of year-to-year fluctuations in the onset of the rainy season, but field observations have confirmed that many typically dry-season species, such as Cochlospermum vitifolium, may begin flowering in November, the wettest month of the year, or extend into a period that is definitely rainy season. The same is of course true of wet-season species that begin before or end after the rainy season.

A total of 126 species were excluded from consideration here because too little is known about their phenological patterns; a number of these are cultivated species. But all cultivated species for which the phenology is known are included here. Because the bimodal species are so difficult to classify into seasonality types, no attempt has been made to do so, for many of these are also transitional in their flowering or fruiting behavior.
There are 294 species that flower and fruit all year, 164 of which exhibit no discernible peak. For others, there is a peak of activity in the dry or wet season, and in a few cases flowering is bimodal (e.g., with a peak of activity at the beginning of the dry season and another at the beginning of the wet season). In still other cases, the peak of activity is transitional, straddling both the dry and wet seasons.

The most heavily represented seasonality types flower and fruit either in the wet season (176 species) or in the dry season (133 species).

Graph 7. Numbers of vascular plant species in flower and fruit, by month.
Graph 9. Percentage of species flowering, by month, three more groups.

Flowering and fruiting extension, by habit-and-habitat class (in months)

Habit-and-habitat class Average flower extension Average fruit extension
Large trees (to 30 m or more)
4.3
4.4
Medium-sized trees (10-30 m)
3.5
3.3
Small trees or shrubs (less than 10 m)
6.3
6.1
Lianas
4.0
3.8
Vines
6.9
6.5
Epiphytic herbs
4.8
4.0
Clearing herbs
8.6
8.5
Aquatic herbs
9.4
9.1
Forest herbs
7.4
7.2
     
ALL CLASSES
6.4
6.1

TABLE 6

Flowering and fruiting seasonality types


Seasonality types		 Number of species
In flower less
than 9 months		 In flower more
than 9 months
(peak-period type)		 total
1. Flowers and fruits dry season
133
 
133
2. Flowers and fruits dry-wet
46
 
46
3. Flowers and fruits wet season
177
 
177
4. Flowers and fruits wet-dry
24
 
24
5. Flowers dry; fruits dry-wet
41
1
42
6. Flowers dry; fruits wet
57
7
64
7. Flowers dry; fruits wet-dry
4
 
4
8. Flowers dry-wet; fruits wet
90
4
94
9. Flowers dry-wet; fruits wet-dry
12
 
12
10. Flowers dry-wet; fruits dry
14
1
15
11. Flowers dry; fruits dry 1 year later
7
 
7
12. Flowers dry-wet; fruits dry-wet 1 year later
7 
 
 7
13. Flowers wet; fruits wet-dry
29
1
30
14. Flowers wet; fruits wet 1 year later
3
 
3
15. Flowers wet; fruits dry
71
7
78
16. Flowers wet; fruits dry-wet
10
2
12
17. Flowers wet-dry; fruits dry
27
4
31
18. Flowers wet-dry; fruits dry-wet
17
2
19
19. Flowers wet-dry; fruits wet
9
 
9
20. Flowers and fruits bimodally
 
 
36
21. Flowers and fruits all year, no peak
 
 164
164 
22. Flowers and fruits all year, Especially wet
 
 81
 81
23. Flowers and fruits all year, Especially dry
 
24 
24 
24. Flowers and fruits all year, wet-dry peak
 
 9
9 
25. Flowers and fruits all year, dry-wet peak
 
 11
 11
26. Flowers and fruits all year, bimodal peaks
 
 5
 5
27. Seasonality incompletely known
 
 
126
28. Flowers dry; fruits all year
 
1
1
29. Flowers dry-wet; fruits all year
 
1
1
    TOTAL
778
325
1,265

Some species that flower for more than 9 months, and thus overlap the established seasonal-type categories, nevertheless exhibit a peak of activity corresponding to a particular seasonal type; these appear in a separate column in Table 6.

From all of these data we can determine whether more species take advantage of the dry season, for flowering or fruiting, than prefer the rainy season. A total of 509 seasonal species (excluding bimodal species) do all or part of their flowering during the dry season; of these, 242 (47%) restrict their flowering to the dry season. A total of 559 seasonal species (excluding bimodal species) do all or part of their flowering in the rainy season; of these, 290 (52%) restrict their flowering to the rainy season.

Comparable figures for the seasonally fruiting species are 462 that set all or part of their fruits in the dry season, 253 (55%) that restrict their fruiting to the dry season, and 542 that set all or part of their fruits in the rainy season (341(63%) restrict their fruiting to the rainy season).

On the strength of general observations in the field, one might conclude that more species take advantage of the dry season to flower, since it is during this period when many of the more conspicuous species come into bloom. Even the flowering curves indicate that there is a decided preference for flowering in the dry season, since most habit-and-habitat classes reach their peak of activity at that time. However, many trees that attain their peak of flowering late in the dry season and into the dry-wet transitional period may in fact be triggered to flower by the onset of the rainy season. Two other groups, the understory shrubs and forest herbs, definitely reach their peak of flowering activity in the rainy season (Foster, 1974); and both groups are relatively inconspicuous by contrast with the showier dry-season species.

Since the rainy season is substantially longer than the dry season, more species of plants flower and fruit during the rainy season than during the dry season, notwithstanding the fact that there is an overall peak of flowering and fruiting activity in the dry season.
The following lists, by family, all of the species assignable to each of the seasonality types* given in Table 6:


*The superscript numbers following some of the species names indicate the following: 1rarely flowers late wet; 2 rarely fruits early dry; 3rarely fruits early wet; 4 rarely flowers and fruits early wet; 5rarely flowers late dry; 6 rarely flowers and fruits early dry; 7rarely fruits late dry; 8rarely flowers early wet.

1. Flowers and fruits in the dry season
       
Typhaceae Typha dominguensis Rhamnaceae Colubrina glandulosa
Gramineae Bothriochloa pertusa Vitaceae Cissus pseudosicyoides
  Chloris virgata Elaeocarpaceae Sloanea terniflora
  Ischaemum indicum Tiliaceae Heliocarpus popayanensis3
  I. rugusum1   Luehea seemannii3,1
  Oplismenus burmani   L. speciosa
  Orthoclada laxa   Triumfetta lappula
  Streptogyne americana Malvaceae Pavonia dasypetala
Cyperaceae Scleria macrophylla   P. paniculata3
Bromeliaceae Pitcairnia heterophylla Bombacaceae Bombacopsis quinata3
Dioscoreaceae Dioscorea trifida   B. sessilis3
Orchidaceae Aspasia principissa   Cavanillesia platanifolia3
  Bulbophyllum pachyrrhachis   Ceiba pentandra
  Caularthron bilamellatum   Pseudobombax septenatum
  Chysis aurea Sterculiaceae Melochia lupulina
  Cochleanthes lipscombiae1   M. melissifolia
  Dichaea panamensis   Waltheria glomerata
  Epidendrum imatophyllum4 Marcgraviaceae Marcgravia nepenthoides1
  E. lockhartioides   Souroubea sympetala3
  E. schlechterianum Cochlospermaceae Cochlospermum vitifolium3
  Gongora quinquenervis Violaceae Hybanthus prunifolius3
  G. tricolor Combretaceae Terminalia chiriquensis3
  Leochilus scriptus1 Myrtaceae Eugenia galalonensis
  Lockhartia pittieri   E. oerstedeana
  Maxillaria variabilis Gentianaceae Schultesia lisianthoides
  Mormodes powellii Convolvulaceae Ipomoea batatas
  Notylia barkeri   I. squamosa
  N. pentachne Labiatae Hyptis capitata
  Oncidium ampliatum   H. mutabilis
  O. stipitatum4   Salvia occidentalis
  Ornithocephalus bicornis4 Bignoniaceae Tabebuia guayacan4
  Polystachya masayensis1   T. ochracea
  Scaphyglottis graminifolia Acanthaceae Aphelandra sinclairiana
  Sobralia panamensis  

Blechum brownei
  Trigonidium egertonianum4   B. costaricense
Urticaceae Myriocarpa yzabalensis   Elytraria imbricata
Polygonaceae Coccoloba acapulcensis   Hygrophila guianensis1
  Triplaris cumingiana   Justicia pectoralis
Amaranthaceae Iresine angustifolia   Mendoncia gracilis
  I. celosia   Nelsonia brunellodes 4
Phytolaccaceae Microtea debilis   Ruellia metallica
Leguminosae Aeschynomene americana var.glandulosa   Teliostachy alopecuroidea alopecuroidea
  Albizia guachapele3   Trichanthera gigantea
  Cajanus bicolor   Rubiaceae Manettia reclinata
  Calopogonium caeruleum Compositae Ayapana elata
  C. mucunioides   Baltimora recta
  Cassia obtusifolia1   Chromolaena odorata 1
  C. reticulata   Elephantopus mollis
  Desmodium scorpiurus   Fleischmannia sinclairi4
  D. triflorum   Heterocondylus vitalbis4
  Machaerium floribundum   Koanophyllon wetmorei 1
  Mimosa casta   Melanthera aspera
  M. pudica1   Mikania guaco
  Mucuna rostrata   M. leiostachya
  Rhynchosia pyramidalis   Neurolaena lobata
  Teramnus uncinatus   Rolandra fruticosa
  T. volubilis   Spiracantha cornifolia
Oxalidaceae Averrhoa carambola   Synedrella nodiflora
Malpighiaceae Stigmaphyllon hypargyreum   Verbesina gigantea
  Tetrapteris discolor   Vernonia canescens
Polygalaceae Securidaca diversifolia   V. patens
Euphorbiaceae Adelia triloba    
  Phyllanthus amarus    
Anacardiaceae Astronium graveolens    
Sapindaceae Paullinia fuscescens var. glabrata    
  Serjania atrolineata3    
  S. circumvallata    
  S. cornigera    
  S. decapleuria    
  S. mexicana3    
  S. trachygona    
  Thinouia myriantha    
 
2. Flowers and fruits dry-wet
       
Alismataceae Sagittaria lancifolia Flacourtiaceae Casearia guianensis var. guianensis
Gramineae Panicum polygonatum   Hasseltia floribunda
  Paspalum saccharoides   Xylosma oligandrum
Cyperaceae Calyptrocarya glomerulata Lythraceae Cuphea carthagenensis
  Rhynchospora corymbosa Combretaceae Combretum cacoucia
  Scleria mitis   C. decandrum
Piperaceae Peperomia macrostachya   Terminalia amazonica
  P. obscurifolia Myrtaceae Eugenia venezuelensis
  Piper culebranum Melastomataceae Miconia argentea
Lacistemaceae Lacistema aggregatum   M. elata
Moraceae Pseudolmedia spuria Onagraceae Ludwigia leptocarpa
Loranthaceae Phoradendron piperoides Boraginaceae Heliotropium indicum
  Struthanthus orbicularis   Tournefortia bicolor
Amaranthaceae Chamissoa altissima   T. cuspidata
Chrysobalanaceae

Hirtella racemosa (may flower more than once per year)

 Scrophulariaceae Bacopa salzmannii
Leguminosae Dalbergia brownie Rubiaceae Bertiera guianensis
Rutaceae Citrus reticulata   Diodia denudata
Malpighiaceae Heteropteris laurifolia Cucurbitaceae Gurania makoyana
  Hiraea reclinata Compositae Conyza apurensis
  Mascagnia nervosa    
Euphorbiaceae Alchornea latifolia    
Sapindaceae Paullinia bracteosa    
Malvaceae Pavonia rosea    
Guttiferae Havetiopsis flexilis    
  Marila laxiflora    
Violaceae Rinorea sylvatica    
 
3. Flowers and fruits wet season
       
Gramineae Andropogon virginicus Rutaceae Zanthoxylum panamense
  Lasiacis procerrima Malpighiaceae Byrsonima spicata
  Panicum mertensii   Hiraea quapara
  P. milleflorum   Mascagnia hippocrateoides
  Paspalum notatum   Spachea membranacea5
  P. plicatulum   Stigmaphyllon puberum
  Rhipidocladum racemiflorum Polygalaceae Polygala paniculata
  Saccharum spontaneum Euphorbiaceae Acalypha arvensis
  Setaria paniculifera   Croton billbergianus5
  S. vulpiseta   Drypetes standleyi
Cyperaceae Cyperus giganteus2   Margaritaria nobilis
  C. rotundus   Poinsettia heterophylla
  C. simplex   Sapium caudatum
  Rhynchospora micrantha Anacardiaceae Mosquitoxylum jamaicense
Palmae Cryosophila warscewiczii Sapindaceae Cupania cinerea
Cyclanthaceae Carludovica drudei   C. latifolia
  Ludovia integrifolia   Paullinia rugosa
Araceae Caladium bicolor   Serjania pluvialiflorens
  Dieffenbachia longispatha Rhamnaceae Gouania adenophora
  Dracontium dressleri Vitaceae Cissus microcarpa
  Philodendron fragrantissimum   C. rhombifolia
  P. grandipes Elaeocarpaceae Sloanea zuliinsis
  P. inconcinnum Bombacaceae Quararibea asterolepis
  P. pterotum   Q. pterocalyx
  P. radiatum Dilleniaceae Doliocarpus major
  P. tripartitum   D. olivaceus
  Rhodospatha moritziana   Saurauia laevigata
  R. wendlandii Flacourtiaceae Banara guianensis
  Xanthosoma helleborifolium   Casearia arborea
  X. nigrum   C. commersoniana
  X. pilosum Cactaceae Rhipsalis cassytha
Bromeliaceae Aechmea magdalenae Lecythidaceae Gustavia fosteri
Commelinaceae Campelia zanonia Myrtaceae Eugenia coloradensis
  Dichorisandra hexandra   Myrcia fosteri
Pontederiaceae Eichhornia azurea   M. gatunensis
Amaryllidaceae Zephyranthes tubispatha   Psidium anglohondurense
Musaceae Heliconia catheta Melastomataceae Arthrostema alatum
  H. irrasa   Bellucia grossularioides
  H. latispatha   Henriettea succosa
  H. pogonantha   Miconia lateriflora
Zingiberaceae Costus allenii   M. shattuckii
  C. guianensis var. macrostrobilus Onagraceae Ludwigia octovalvis
  C. laevis Araliaceae Dendropanax arboreus
  Renealmia alpinia   D. stenodontus5
  R. cernua5 Myrsinaceae Ardisia bartlettii
  Zingiber oficinale   Parathesis microcalyx5
Marantaceae Calathea inocephala5 Sapotaceae Pouteria unilocularis
  C. insignis   P. stipitata
  C. latifolia Gentianaceae Voyria alba
  C. marantifolia   V. tenella
  C. micans Asclepiadaceae Marsdenia crassipes
  C. panamensis Boraginaceae Cordia panamensis
  C. villosa   Tournefortia hirsutissima
  Ischnosiphon leucophaeus Verbenaceae Aegiphila cephalophora
  Stromanthe jacquinii   A. elata
Burmanniaceae Thismia panamensis   Vitex cooperi
Orchidaceae Brassia caudata Solanaceae Cestrum latifolium
  Dichaea trulla   Lycianthes synanthera
  Encyclia triptera   Solanum asperum
  Palmorchis powellii   S. jamaicense
Piperaceae Peperomia mameiana Scrophulariaceae Lindernia crustacea
Lacistemaceae Lozania pittieri   L. diffusa
Ulmaceae Celtis iguanaeus Gesneriaceae Chrysothemis friedrichsthaliana
  Trema micrantha   Diastema raciferum
Moraceae Dorstenia contrajerva   Nautilocalyx panamensis
  Trophis racemosa5 Acanthaceae Herpetacanthus panamensis
Urticaceae Boehmeria cylindrica Rubiaceae Amaioua corymbosa
  Pouzolzia obliqua   Cephaelis discolor
  Urera eggersii   Geophila croatii
Olacaceae Heisteria costaricensis5   G. repens
  H. longipes   Hamelia axillaris
Rafflesiaceae Apodanthes caseariae   Palicourea guianensis5
Polygonaceae Coccoloba coronata   Psychotria acuminata
  C. parimensis   Alibertia edulis
Caryophyllaceae Drymaria cordata6   P. brachybotrya
Nymphaeaceae Nymphaea blanda   P. deflexa
Menispermaceae Abuta racemosa   P. micrantha
  Chondrodendron tomentosum   P. pubescens
 

Odon tocarya tamoides var. canescens

   P. racemosa
Lauraceae Phoebe mexicana   Randia formosa
Saxifragaceae Hydrangea peruviana Cucurbitaceae Cayaponia glandulosa
Leguminosae Crotalaria vitellina6   Fevillea cordifolia
  Inga hayesii Compositae Clibadium surinamense
  Leucaena multicapitula   Conyza bonariensis
  Ormosia coccinea var. subsimplex   Eleutheranthera ruderalis
  Pithecellobium barbourianum   Erechtites hieracifolia var.cacalioides
  Pterocarpus officinalis5   Fleischmannia microstemon
  P. rohrii   Wedelia trilobata5
      Wulffia baccata
       
4. Flowers and fruits wet-dry
       
Gramineae Andropogon glomeratus Piperaceae Peperomia killipi
  Brachiaria mutica   P. obtusifolia
  Cenchrus brownii Leguminosae Aeschynomene ciliata
  Ichnanthus pallens   Clitoria rubiginosa
  Lasiacis oaxacensis Simaroubaceae Quassia amara
  Olyra latifolia Melastomataceae Miconia borealis
  Panicum grande   Schwackaea cupheoides
  Paspalum microstachyum Convolvulaceae Ipomoea tiliacea
  Phragmites australis Gesneriaceae Drymonia serrulata6
  Polytrias amaura Rubiaceae Borreria densiflora
  Schizachyrium microstachyum Compositae Calea prunifolia
Commelinaceae Callisia ciliata   Pseudoelephantopus spicatus
       
5. Flowers dry; fruits dry-wet
Cyclanthaceae Carludovica palmata Anacardiaceae Anacardium excelsum
Araceae Anthurium littorale Sapindaceae Cupania rufescens
Bromeliaceae Billbergia macrolepis   C. sylvatica1
Commelinaceae Phaeosphaerion persicariifolium   Serjania paucidentata
Orchidaceae Cattleya patinii Malvaceae Hibiscus bifurcatus
  Epidendrum radicans Dilleniaceae Davilla nitida
  Ionopsis satyrioides   Doliocarpus dentatus
Piperaceae Peperomia cordulata   D. multiflorus
Moraceae Pourouma guianensis Flacourtiaceae Casearia aculeata
Chrysobalanaceae Licania hypoleuca   C. arguta 1
Leguminosae Cassia undulata1 Cactaceae Epiphyllum phyllanthus var. columbiense
  Clitoria javitensis1 Melastomataceae Miconia hondurensis
  Dioclea guianensis3   M. impetiolaris
  Enterolobium schomburgkii3 Myrsinaceae Stylogyne standleyi
  Lonchocarpus velutinus Convolvulaceae Merremia umbellata
  Machaerium kegelii Boraginaceae Cordia alliodora
  M. milleflorum   C. lasiocalyx
  Schizolobium parahybum Rubiaceae Macrocnemum glabrescens
Simaroubaceae Picramnia latifolia   Uncaria tomentosa
  Simarouba amara var. typica Cucurbitaceae Cayaponia granatensis
      Melothria trilobata
       
5a. Flowers dry; fruits dry-wet (flowers more than 9 months)
       
Bombacaceae Pachira aquatica    
       
6. Flowers dry; fruits wet
       
Palmae Bactris coloniata Leguminosae Acacia melanoceras
Araceae Anthurium clavigerum   Bauhinia guianensis
  A. friedrichsthalii   Hymenaea courbaril
  A. tetragonum   Inga cocleensis
  Monstera dilacerata   I. fagifolia8
Bromeliaceae Aechmea pubescens   I. thibaudiana
Orchidaceae Lockhartia acuta   Pithecellobium dinizii
Piperaceae Piper arieianum   P. macradenium7,8
Moraceae Artocarpus altilis Rutaceae Citrus aurantifolia
  Brosimum alicastrum   C. aurantium
  Cecropia insignis7   C. sinensis
Nyctaginaceae Pisonia aculeata7   Zanthoxylum procerum8
Menispermaceae Odontocarya truncata7   Z. setulosum7
Monimiaceae Siparuna pauciflora Meliaceae Trichilia verrucosa8
Lauraceae Beilschmiedia pendula Anacardiaceae Mangifera indica
  Ocotea skutchii Sapindaceae Allophylus psilospermus
  Persea americana   Talisia nervosa
Chrysobalanaceae Hirtella americana8 Sterculiaceae Herrania purpurea7
    Theaceae Ternstroemia tepezapote
    Guttiferae Garcinia mangostana
      Rheedia acuminata7
      R. edulis
       
6. Flowers dry; fruits wet (cont.)
       
Flacourtiaceae Laetia procera Apocynaceae Malouetia guatemalensis7
  Zuelania guidonia7,8   Odontadenia puncticulosa
Passifloraceae Passiflora ambigua Convolvulaceae Maripa panamensis7,8
  P. nitida7,8 Solanaceae Cestrum megalophyllum
Caricaceae Carica cauliflora Bignoniaceae jacaranda copaia8
Combretaceae Combretum laxum var. epiphyticum   Tabebuia rosea
Melastomataceae Miconia prasina Rubiaceae Faramea luteovirens8
Theophrastaceae Jacquinia macrocarpa Cucurbitaceae Gurania coccinea
Loganiaceae Strychnos toxifera    
       
6a. Flowers dry; fruits wet (flowers more than 9 months)
       
Araceae Anthurium scandens Passifloraceae Passiflora coriacea
  Syngonium podophyllum Solanaceae Solanum lanciifolium
Piperaceae Piper grande Rubiaceae Psychotria limonensis
Moraceae Cecropia peltata    
       
7. Flowers dry; fruits wet-dry
       
Annonaceae Desmopsis panamensis Sapindaceae Paullinia baileyi
Leguminosae Entada monostachya Myrtaceae Psidium guajava
       
8. Flowers dry-wet; fruits wet
       
Cyperaceae Cladium jamaicense Burseraceae Protium tenuifolium var. sessiliflorum
Palmae Bactris barronis Meliaceae Guarea glabra
  B. gasipaes   Trichilia montana
  Geonoma interrupta Malpighiaceae Byrsonima crassifolia
  G. procumbens Euphorbiaceae Acalypha diversifolia
Cyclanthaceae Asplundia alata   A lchornea costaricensis
  Cyclanthus bipartitus   Jatropha curcas (rarely also bimodal)
Araceae Anthurium ochranthum   Omphalea diandra
  Homalomena wendlandii Anacardiaceae Spondias mombin
  Philodendron guttiferum   S. radlkoferi
  P. inaequilaterum Celastraceae Maytenus schippii
  P. nervosum Staphyleaceae Turpinia occidentalis subsp.breviflora
  P. panamense Guttiferae Tovomita longifolia
  Spathiphyllum phryniifolium Flacourtiaceae Laetia thamnia
  Syngonium erythrophyllum   Tetrathylacium johansenii
Bromeliaceae Aechmea setigera   Xylosma chloranthum
  A. tillandsioides Passifloraceae Passiflora williamsii
Zingiberaceae Costus villosissimus Lecythidaceae Gustavia superba
Orchidaceae Ionopsis utricularioides Myrtaceae Eugenia nesiotica
  Stellis crescentiicola Melastomataceae Clidemia collina
Piperaceae Peperomia ebingeri   C. purpureo-violacea
  P glabella   Conostegia bracteata
  Piper aequale   Miconia affinis
  P. arboreum   M. lacera
  P. darienense   M. lonchophylla
  P. hispidum   M. serrulata
  P. perlasense   Topobaea praecox
  P. pubistipulum Myrsinaceae Ardisia pellucida
  P. reticulatum Apocynaceae Forsteronia peninsularis
Moraceae Castilla elastica   F. viridescens
  Cecropia longipes Boraginaceae Cordia bicolor
  Coussapoa panamensis   Tournefortia maculata
  Maquira costaricana Solanaceae Lycianthes maxonii
Nyctaginaceae Guapira standleyanum Bignoniaceae Cydista heterophylla
Annonaceae Annona hayesii Rubiaceae Alseis blackiana
  A. spraguei   Antirrhoea trichantha
Monimiaceae Siparuna guianensis   Chimarrhis parviflora
Lauraceae Ocotea cernua   Faramea occidentalis
Leguminosae Brownea macrophylla   Hoffmania woodsonii
  Cynometra bauhiniifolia   Psychotria carthagenensis
  Inga pauciflora   P. emetica
  I. pezizifera   P. horizontalis
  I. vera subsp. spuria   P. pittieri
  Lonchocarpus pentaphyllus   Randia armata
  Ormosia panamensis    
  Pithecellobium rufescens    
       
8a. Flowers dry-wet; fruits wet (flowers more than 9 months)
       
Amaryllidaceae Crinum erubescens    
Leguminosae Andira inermis    
  Inga quaternata    
Euphorbiaceae Phyllanthus acuminatus    
       
9. Flowers dry-wet; fruits wet-dry
       
Palmae Bactris major Malpighiaceae Bunchosia cornifolia
Iridaceae Neomarica gracilis Euphorbiaceae Hura crepitans
Orchidaceae Campylocentrum micranthum Flacourtiaceae Casearia corymbosa
Leguminosae Myroxylon balsamum var. pereirae Lythraceae Adenaria floribunda
  Platymiscium pinnatum Apocynaceae Prestonia obovata
Burseraceae Bursera simaruba Rubiaceae Psychotria furcata
       
10. Flowers dry-wet; fruits dry
       
Bromeliaceae Guzmania monostachya Euphorbiaceae Garcia nutans
  Tillandsia monadelpha Apocynaceae Odontadenia macrantha
Orchidaceae Epidendrum difforme Bignoniaceae Arrabidaea chica
Annonaceae Xylopia frutescens   Ceratophytum tetragonolobum
Leguminosae Dalbergia retusa   Macfadyena unguis-cati
  Platypodium elegans   Xylophragma seemannianum
  Swartzia panamensis    
  Tachigalia versicolor    
       
10a. Flowers dry-wet; fruits dry (flowers more than 9 months)
       
Apocynaceae Prestonia portobellensis    
       
11. Flowers dry; fruits dry 1 year later
       
Bromeliaceae Vriesia gladioliflora    
Annonaceae Annona glabra    
Leguminosae Enterolobium cyclocarpum    
Hippocrateaceae Hylenaea praecelsa    
  Prionostemma aspera    
Sterculiaceae Sterculia apetala    
Bignoniaceae Pleonotoma variabilis    
       
12. Flowers dry-wet; fruits dry-wet 1 year later
       
Palmae Scheelea zonensis    
Annonaceae Annona muricata    
Leguminosae Inga spectabilis    
Hippocrateaceae Tontelea richardii    
Apocynaceae Lacmellea panamensis    
  Tabernaemontana arborea    
Rubiaceae Tocoyena pittieri    
       
13. Flowers wet; fruits wet-dry
       
Palmae Geonoma cuneata Malpighiaceae Tetrapteris macrocarpa
Araceae Dieffenbachia pittieri Trigoniaceae Trigonia floribunda
Bromeliaceae Guzmania lingulata var. minor Guttiferae Tovomita stylosa
Haemodoraceae Xiphidium caeruleum   Vismia macrophylla
Zingiberaceae Costus pulverulentus Flacourtiaceae Casearia sylvestris
  C. scaber Loganiaceae Strychnos panamensis5
Marantaceae Ischnosiphon pruinosus Convolvulaceae Ipomoea phillomega
Orchidaceae Catasetum bicolor Verbenaceae Aegiphila panamensis
  Habenaria alata Acanthaceae Mendoncia littoralis
Piperaceae Peperomia ciliolibractea Rubiaceae Coussarea curvigemmia
Moraceae Sorocea affinis   Isertia haenkeana
Annonaceae Annona acuminata   Pentagonia macrophylla5
Leguminosae Erythrina costaricensis var. panamensis   Psychotria granadensis
  Swartzia simplex var. grandiflora   P. uliginosa
      Warscewiczia coccinea5
       
13a. Flowers wet; fruits wet-dry (flowers more than 9 months)
       
Piperaceae Piper marginatum    
       
14. Flowers wet; fruits wet 1 year later
       
Araceae Philodendron hederaceum    
Leguminosae Dioclea wilsonii    
Sapotaceae Pouteria sapota    
 
15. Flowers wet; fruits dry
       
Palmae Socratea durissima Malvaceae Hampea appendiculata var. longicalyx
Araceae Dieffenbachia oerstedii Sterculiaceae Byttneria aculeata
  Philodendron sagittifolium Dilleniaceae Tetracera volubilis
Bromeliaceae Catopsis sessiliflora Guttiferae Mammea americana
  Tillandsia anceps Flacourtiaceae Lindackeria laurina
  Vriesia heliconioides Lythraceae Lafoensia punicifolia
  V. ringens Lecythidaceae Couratari panamensis
  V. sanguinolenta Combretaceae Combretum laxum var. laxum
Dioscoreaceae Dioscorea sapindoides   A. megalocarpon
  D. urophylla   Prestonia ipomiifolia
Orchidaceae Catasetum viridiflavum   Blepharodon mucronatum
  Dimerandra emarginata   Matalea pinquifolia
  Epidendrum rigidum Araliaceae Didymopanax morototoni
  E. sculptum Sapotaceae Chrysophyllum cainito
  E. strobiliferum Gentianaceae Chelonanthus alatus
  Liparis elata Apocynaceae Aspidosperma cruenta
  Maxillaria neglecta   A. megalocarpon
  Peristeria elata   Prestonia ipomiifolia
  Polystachya foliosa Asclepiadaceae Blepharodon mucronatum
  Sobralia fragrans   Matalea pinquifolia
  S. suaveolens   M. trianae
  Trichopilia maculata   M. viridiflora
  T. subulata Bignoniaceae Adenocalymma arthropetiolatum
Aristolochiaceae Aristolochia gigantea   Arrabidaea florida
Connaraceae Cnestidium rufescens   A. patellifera
Leguminosae Acacia glomerosa   A. verrucosa
  A. hayesii   Pithecoctenium crucigerum
  A. riparia   Tynnanthus croatianus
  Adenopodia polystachya Gesneriaceae Kohleria tubiflora
  Bauhinia reflexa Rubiaceae Cosmibuena skinneri
  Dipteryx panamensis   Coutarea hexandra
  Inga punctata    
  Machaerium arboreum2    
  M. seemannii    
  Peltogyne purpurea    
Rutaceae Zanthoxylum belizense    
Burseraceae Trattinnickia aspera    
Meliaceae Cedrela odorata    
  Trichilia hirta    
Malpighiaceae Tetrapteris seemannii    
Sapindaceae Paullinia glomerulosa    
  P. pinnata    
Tiliaceae Apeiba membranacea    
  Corchorus siliquosus    
       
       
15a. Flowers wet; fruits dry (flowers more than 9 months)
       
Orchidaceae Maxillaria crassifolia    
Leguminosae Swartzia simplex var. ochnacea    
Tiliaceae Apeiba tibourbou    
Malvaceae Sida acuta    
Begoniaceae Begonia filipes    
Bignoniaceae Anemopaegma chrysoleucum    
  Callichlamys latifolia    
       
16. Flowers wet; fruits dry-wet
       
Palmae Astrocaryum standleyanum Burseraceae Tetragastris panamensis
  Desmoncus isthmius Meliaceae Guarea multiflora
  Elaeis oleifera Euphorbiaceae Dalechampia dioscoreifolia
Liliaceae Cordyline fruticosa Guttiferae Calophyllum longifolium
Annonaceae Xylopia macrantha   Tovomitopsis nicaraguensis
       
       
       
       
       
       
16a. Flowers wet; fruits dry-wet (flowers more than 9 months)
       
Piperaceae Piper villiramulum    
Bignoniaceae Martinella obovata    
       
17. Flowers wet-dry; fruits dry
       
Gramineae Lasiacis sorghoidea Rhamnaceae Gouania lupuloides3
Orchidaceae Eulophia alta Tiliaceae Trichospermum mexicanum
  Maxillaria powellii Ochnaceae Cespedezia macrophylla
  M. uncata   Ouratea lucens
Leguminosae Desmodium cajanifolium Passifloraceae Passiflora seemannii
  Inga mucuna Begoniaceae Begonia guaduensis
  Machaerium microphyllum Combretaceae Combretum fruticosum
  Mucuna mutisiana Apocynaceae Prestonia acutifolia
Malpighiaceae Banisteriopsis cornifolia Bignoniaceae Amphilophium paniculatum
  Hiraea grandifolia   Arrabidaea candicans
Euphorbiaceae Dalechampia tiliifolia3 Acanthaceae Justicia graciliflora
Sapindaceae Paullinia fibrigera Rubiaceae Calycophyllum candidissimum
  P. turbacensis   Pogonopus speciosus
  Serjania rhombea3    
       
17a. Flowers wet-dry; fruits dry (flowers more than 9 months)
       
Commelinaceae Gibasis geniculata    
Euphorbiaceae Mabea occidentalis    
Apocynaceae Mesechites trifida    
Boraginaceae Cordia spinescens    
       
18. Flowers wet-dry; fruits dry-wet
       
Cyperaceae Hypolytrum schraderianum Lauraceae Nectandra globosa
  Rhynchospora cephalotes Leguminosae Canavalia dictyota
Dioscoreaceae Dioscorea macrostachya (possibly bimodal)   Dalbergia monetaria
Piperaceae Piper carrilloanum   Erythrina fusca
  P. imperiale Anacardiaceae Anacardium occidentale
Moraceae Perebea xanthochyma Bixaceae Bixa orellana
Aristolochiaceae Aristolochia chapmaniana Passifloraceae Passiflora auriculata
Myristicaceae Virola surinamensis Boraginaceae Tournefortia angustiflora
    Cucurbitaceae Cayaponia racemosa
       
18a. Flowers wet-dry; fruits dry-wet (flowers more than 9 months)
       
Bombacaceae Ochroma pyramidale    
Bignoniaceae Pachyptera kerere    
       
19. Flowers wet-dry; fruits wet
       
Palmae Chamaedorea wendlandiana    
Piperaceae Piper cordulatum    
Polygonaceae Coccoloba manzanillensis    
Leguminosae Dioclea reflexa    
  Inga goldmanii    
  I. multijuga    
  I. sapindoides    
  I. umbellifera    
Rubiaceae Psychotria marginata    
       
20. Flowers and fruits bimodally
       
Gnetaceae Gnetum leyboldii var. woodsonianum Vochysiaceae Vochysia ferruginea
Palmae Oenocarpus panamanus Polygalaceae Securidaca tenuifolia
  Synechanthus warscewiczianus Euphorbiaceae Acalypha macrostachya
Araceae Monstera dubia   Hyeronima laxiflora
Olacaceae Heisteria concinna Hippocrateaceae Anthodon panamense
Myristicaceae Virola sebifera Sterculiaceae Guazuma ulmifolia
Lauraceae Nectandra purpurascens Dilleniaceae Tetracera hydrophila
Capparidaceae Capparis frondosa Guttiferae Vismia baccifera
Connaraceae Connarus panamennss   V. billbergiana
  C. turczaninowii Araliaceae Oreopanax capitatus
  Rourea glabra Sapotaceae Cynodendron panamense
Leguminosae Inga marginata Solanaceae Cestrum nocturnum
  Prioria copaifera   Solanum arboreum
Erythroxylaceae Erythroxylum multiflorum   S. argenteum
  E. panamense   S. umbellatum
Burseraceae Protium costaricense Rubiaceae Cephaelis ipecacuanha
  P. panamense   Guettarda foliacea
Meliaceae Trichilia cipo   Psychotria psychotriifolia
       
21. Flowers and fruits all year, no peak
       
Gramineae Acroceras oryzoides Vitaceae   Cissus sicyoides
  Andropogon leucostachyus Elaeocarpaceae   Muntingia calabura
  Bothriochloa intermedia Malvaceae   Hibiscus rosy-sinensis
  Chloris radiata   H. sororius
  Cynodon dactylon   Sida rhombifolia
  Hyparrhenia rufa Guttiferae   Clusia odorata
  Isachne polygonoides Turneraceae   Turnera panamensis
  Oryza latifolia Passifloraceae   Passiflora biflora
  Panicum trichoides   P. foetida var. isthmia
  Paspalidium germinatum   P. menispermifolia
  Pharus latifolius   P. punctata
  P. parvifolius Caricaceae   Carica papaya
  Schizachyrium brevifolium Begoniaceae   Begonia patula
  Streptochaeta spicata Rhizophoraceae   Cassipourea elliptica
Cyperaceae   Cyperus brevifolius   Myrtaceae   Psidium friedrichsthalianum
  C. diffuses Melastomataceae   Aciotis levyana
  C. luzulae   Adelobotrys adscendens
  C. odoratus   Clidemia capitellata
  C. sesquiflorus   C. dentata
  C. tennis   C. ocotona
  Eleocharis caribaea   C. septuplinervia
  E. plicarhachis   Conostegia speciosa
  Fuirena umbellata   C. xalapensis
  Rhynchospora nervosa   Leandra dichotomy
  Scleria pterota   Miconia nervosa
  S. secans   Ossaea quinquenervia
Araceae   Anthurium brownii     Tibouchina longifolia
  A. gracile Onagraceae   Ludwigia decurrens
  Montrichardia arborescens   L. helminthorrhiza
  Philodendron scandens Umbelliferae   Hydrocotyle umbellata
Commelinaceae   Commelina erecta Myrsinaceae   Ardisia fendleri
Pontederiaceae   Pontederia rotundifolia Gentianaceae   Voyria truncata
Smilacaceae   Smilax mollis Menyanthaceae   Nymphoides indica
  S. panamensis Apocynaceae   Allamanda cathartica
  S. spinosa   Catharanthus roseus
  S. spissa   Ervatamia coronaria
Musaceae   Heliconia mariae   Mandevilla subsagittata
Orchidaceae   Ornithocephalus powellii   Rhabdadenia biflora
  Pleurothallis brighamii   Stemmadenia grandiflora
  P. grobyi   Thevetia ahouai
  Scaphyglottis longicaulis Asclepiadaceae   Asclepias curassavica
  Triphora gentianoides   Sarcostemma clausum
Piperaceae   Pothomorphe peltata Convolvulaceae   Aniseia martinicensis
Moraceae   Cecropia obtusifolia Verbenaceae   Lantana camara
  Ficus (all species)   Petrea aspera
  Olmedia aspera   Stachytarpheta jamaicensis
Urticaceae   Pilea microphylla Solanaceae   Capsicum annuum
Loranthaceae   Oryctanthus alveolatus   Cyphomandra hartwegii
  O. cordifolius   Physalis angulata
  O. occidentalis   P. pubescens
  Phoradendron quadrangule   Solanum hayesii
  Phthirusa pyrifolia   S. subinerme
Polygonaceae   Coccoloba acuminata Bignoniaceae   Cydista aequinoctalis
  Polygonum acuminatum   Spathodea campanulata
  P. hydropiperoides Gesneriaceae   Columnea purpurata
  P. punctatum Rubiaceae   Cephaelis tomentosa
Amaranthaceae   Gomphrena decumbens   Diodia ocimifolia
Phytolaccaceae   Phytolacca rivinoides   D. sarmentosa
Portulacaceae   Portulaca oleracea   Ixora coccinea
Menispermaceae   Cissampelos pareira Cucurbitaceae   Gurania megistantha
  C. tropaeolifolia   Momordica charantia
Annonaceae   Guatteria amplifolia   Psiguria bignoniacea
  G. dumetorum Compositae   Baccharis trinervis
Capparidaceae   Cleome parvifllora   Chaptalia nutans
Leguminosae   Aeschynomene sensitiva   Eclipta alba
  Crotalaria retusa   Emilia sonchifolia
  Desmodium axillare var.   Hebeclinium macrophyllum
  acutifolium   Melampodium divaricatum
  D. canum   Tridax procumbens
  D. distortum   Vernonia cinerea
  D. tortuosum    
Malpighiaceae   Stigmaphyllon ellipticum    
  S. lindenianum    
Euphorbiaceae   Chamaesyce hirta    
  C. hypericifolia    
  C. hyssopifolia    
  Dalechampia cissifolia subsp.    
  panamensis    
  Phyllanthus urinaria    
Hippocrateaceae   Hippocratea volubilis    
       
22. Flowers and fruits all year, specially wet season
       
Gramineae   Andropogon bicornis Euphorbiaceae  Chamaesyce thymifolia
  Anthephora hermaphrodita   Croton hirtus
  Axonopus compressus   C. panamensis
  Digitaria ciliaris Vitaceae  Cissus erosa
  D. horizontalis Malvaceae  Abelmoschus moschatus
  Eleusine indica Guttiferae  Symphonia globulifera
  Gynerium sagittatum Myrtaceae  Calycolpus warscewiczianus
  Homolepis aturensis Melastomataceae  Conostegia cinnamomea
  Leersia hexandra Umbelliferae  Spananthe paniculata
  Leptochloa virgata Loganiaceae  Spigelia anthelmia
  Lithachne pauciflora   S. humboldtiana
  Panicum fasciculatum Apocynaceae   Mandevilla villosa
  P. pilosum Convolvulaceae  Ipomoea quamoclit
  Paspalum decumbens Solanaceae  Cyphomandra allophylla
  P. paniculatum   Solanum ochraceo-ferrugineum
  P. repens   S. rugosum
  P. virgatum   Witheringia solanacea
  Rottboellia exaltata Scrophulariaceae  Scoparia dulcis
  Setaria geniculata Bignoniaceae  Phryganocydia corymbosa
Cyperaceae  Cyperus haspan Gesneriaceae   Besleria laxiflora
  Fimbristylis dichotoma   Codonanthe crassifolia
  Scleria eggersiana   C. uleana
Araceae  Anthurium acutangulum   Columnea billbergiana
  A. bakeri Lentibulariaceae  Utricularia foliosa
  Spathiphyllum friedrichsthalii Acanthaceae  Thunbergia erecta
Commelinaceae   Tripogandra serrulata Rubiaceae  Borreria latifolia
Amaryllidaceae  Hymenocallis pedalis   Chiococca alba
Musaceae  Heliconia vaginalis   Genipa americana
Marantaceae  Calathea lutea   Posoqueria latifolia
Orchidaceae  Epidendrum anceps   Psychotria brachiata
  Maxillaria alba   P. capitata
Piperaceae  Piper auritum   P. chagrensis
Amaranthaceae  Alternanthera sessilis   P. grandis
  Amaranthus viridis   Spermacoce tenuior
Nyctaginaceae  Neea amplifolia Cucurbitaceae  Melothria pendula
Phytolaccaceae  Petiveria alliacea   Psiguria warscewiczii
Chrysobalanaceae  Hirtella triandra (flowering 2 or 3 times per year) Compositae  Schistocarpha oppositifolia
Leguminosae   Desmodium adscendens    
  D. axillare var.  stoloniferum    
  Mimosa pigra    
  Phaseolus peduncularis    
  P. trichocarpus    
  Pithecellobium hymeneaefolium    
  Vigna vexillata    
 
23. Flowers and fruits all year, especially dry season
       
Gramineae   Oplismenus hirtellus Phytolaccaceae  Rivina humilis
  Panicum pulchellum Leguminosae  Centrosema pubescens
Cyperaceae  Cyperus densicaespitosus   Indigofera mucronata
  Scirpus cubensis Dilleniaceae  Tetracera portobellensis
Araceae  Anthurium bombacifolium Umbelliferae  Eryngium foetidum
  Stenospermation angustifolium Solanaceae  Browallia americana
Musaceae  Heliconia wagneriana   Markea ulei
Marantaceae  Thalia geniculata Scrophulariaceae  Stemodia verticillata
Orchidaceae  Maxillaria friedrichsthalii Bignoniaceae  Paragonia pyramidata
  Psygmorchis pusilla Rubiaceae  Borreria ocimoides
Piperaceae  Piper dilatatum   Oldenlandia corymbosa
Amaranthaceae  Alternanthera ficoidea    
  Cyathula prostrata    
 
24. Flowers and fruits all year, especially wet-dry
       
Gramineae   Hymenachne amplexicaulis    
  Panicum maximum    
  P. trichanthum    
  Sacciolepis striate    
Orchidaceae  Epidendrum nocturnum    
  Habenaria repens    
Labiatae   Hyptis brevipes    
Compositae  Mikania micrantha    
  Spilanthes alba    
 
25. Flowers and fruits all year, especially dry-wet
       
Gramineae  Paspalum conjugatum Piperaceae  Peperomia rotundifolia
  Sporobolus indicus Moraceae  Poulsenia armata
Araceae   Anthurium flexile Leguminosae  Caesalpinia pulcherrima
Musaceae  Heliconia metallica Vitaceae  Vitis tiliifolia
Zingiberaceae  Dimerocostus strobilaceus Solanaceae  Solanum antillarum
    Rubiaceae  Hamelia patens var.  glabra
 
26. Flowers and fruits all year, bimodal peaks
       
Gramineae   Ichnanthus tenuis    
Leguminosae  Cassia fruticosa    
Melastomataceae  Mouriri myrtilloides subsp. parvifolia    
Rubiaceae  Borreria laevis    
  Sabicea villosa var.  adpressa    
 
27. Seasonality incompletely known
       
Araucariaceae Araucaria excelsa Leguminosae   Acacia acanthophylla
Hydrocharitaceae  Hydrilla verticillata   Cymbosema roseum
  Limnobium stoloniferum   Machaerium riparium
Gramineae   Bambusa amplexifolia   Vatairea erythrocarpa
  B. arundinacea Humiriaceae   Vantanea occidentalis
  B. glaucescens Rutaceae  Citrus grandis
  Chusquea simpliciflora   C. limon
  Digitaria violascens Malpighiaceae   Hiraea faginea
  Ichnanthus brevivaginatus Euphorbiaceae  Codiaeum variegatum
  Saccharum oficinarum   Manihot esculenta
Palmae   Cocos nucifera   Sapium aucuparium
Sterculiaceae   Theobroma cacao
Araceae  Pistia stratiotes Violaceae  Rinorea squamata
  Syngonium sp. Caricaceae  Jacaratia spinosa
Bromeliaceae   Ananas comosus Myrtaceae  Eugenia principium
  Tillandsia bulbosa   E. uniflora
  T. fasciculata var.  convexispica Melastomataceae   Miconia rufostellulata
  T. fasciculata var.  fasciculata Onagraceae  Ludwigia torulosa
  T. subulifera Araliaceae  Polyscias guilfoylei
Smilacaceae   Smilax lanceolata Sapotaceae  Pouteria fossicola
Amaryllidaceae  Amaryllis belladonna Loganiaceae  Strychnos darienensis
Dioscoreaceae  Dioscorea alata Apocynaceae  Forsteronia myriantha
Musaceae  Musa sapientum Convolvulaceae  Operculina codonantha
Orchidaceae  Campylocentrum pachyrrhizum Labiatae  Coleus blumei
  Elleanthus longibracteatus Scrophulariaceae  Mecardonia procumbens
  Encyclia chacaoensis Lentibulariaceae  Utricularia obtusa
  E chimborazoensis Cucurbitaceae  Cayaponia denticulata
  Epidendrum coronatum   Posadaea sphaerocarpa
  E. rousseauae   Sicydium coriaceum
  Notylia albida Campanulaceae   Centropogon cornutus
  Scaphyglottis prolifera Compositae  Clibadium asperum
  Trichocentrum capistratum   Mikania hookeriana
Piperaceae  Piper aristolochiifolium   M. tonduzii
  P. peracuminatum   Pluchea odorata
  P. pseudo garagaranum    
  P. viridicaule    
Moraceae   Coussapoa magnifolia    
Ceratophyllaceae  Ceratophyllum demersum    
Menispermaceae  Abuta panamensis    
Annonaceae  Anaxagorea panamensis    
  Crematosperma sp.    
  Unonopsis pittieri    
Lauraceae  Nectandra cissiflora    
  N. savannarum Acacia    
  N. savannarum    
 
27a. Flowers known; fruits unknown
       
Pontederiaceae   Eichhornia crassipes Chrysobalanaceae Licania platypus
Dioscoreaceae  Dioscorea haenkeana Leguminosae   Desmodium wydlerianum
  D. polygonoides   Inga minutula
Orchidaceae  Coryanthes maculata   I. ruiziana
  Epidendrum stangeanum   Ormosia macrocalyx
  Habenaria bicornis Malpighiaceae   Malpighia romeroana
  Lycaste powellii Euphorbiaceae   Acalypha wilkesiana
  Masdevallia livingstoneana Sapindaceae   Talisia princeps
  Maxillaria camaridii Passifloraceae   Passiflora vitifolia
  Pleurothallis trachychlamys Cactaceae   Epiphyllum phyllanthus var.  rubrocoronatum
  P. verecunda Lecythidaceae   Grias fendleri
  Sievekingia suavis Myrtaceae   Syzygium jambos
  Sobralia rolfeana Ebenaceae   Diospyros artanthifolia
  Spiranthes lanceolata Loganiaceae   Strychnos brachistantha
  Triphora mexicana Asclepiadaceae   Cynanchum cubense
  Vanilla fragrans   C. recurvum
  V . pompona   Fischeria funebris
  Xylobium foveatum   Gonolobus allenii
Proteaceae   Roupala montana Convolvulaceae   Iseia luxurians
Aristolochiaceae   Aristolochia pilosa Verbenaceae   Clerodendrum paniculatum
Nymphaeaceae   Nymphaea ampla Solanaceae   Cestrum racemosum
Lauraceae   Ocotea oblonga Bignoniaceae   Adenocalymma apurense
  O. pyramidata   Clytostoma binatum
 
27b. Flowers unknown; fruits known
       
Palmae   Bactris coloradonis Sapindaceae  Paullinia pterocarpa
Ulmaceae  Celtis schippii Rubiaceae   Chomelia psilocarpa
 
28. Flowers dry; fruits all year
       
Gramineae   Streptochaeta sodiroana    
 
29. Flowers dry-wet; fruits all year
       
Bignoniaceae   Stizophyllum riparium    

 

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