A new species and a new record of oncidioid micro-orchids (Orchidaceae: Oncidiinae) for the flora of Costa Rica

In this contribution seven forms of digenetic trematodes, some parasitic on cattle and some in wild animals, are described. Excepting Fasciola hepatica Linnaeus, 1758, whose presence in cattle and horses in Costa Rica has been cited by Chavarria in 1940, all the species considered are new to the helminthologic fauna of Costa Rica. Specimens of a paramphistomid found in Costa Rica catt1e have been classified as Paramphistomum cervi (Zeder, 1790) Fischoeder, 1901, in view of the presence of a genital sucker, the configuration of the digestive apparatus, and the structure and disposition of the reproductive organs and vitelline glands. Fasciola hepatica Linnaeus, 1758 is also a frequent parasite in Costa Rica, the precise life cycle of this parasite is unknown in the Republic but studies have been undertaken to determine its intermediate mollusean host. Choledocystus intermedius Caballero, Bravo and Cerecero, 1944 has been found for the first time in Costa Rica, in toads Bufo marinus marinus Linnaeus, 1758 thus allowing its range to be defined from southern Mexico to Panama. Gorgoderina megalorchis Bravo, 1948 is another species of trematode parasite of the amphibian host cited above; our specimens differ from those studied by M. Bravo Hollis only in size, the specimens from Costa Rica being larger than those from Mexican toads; the new locality record extends its geo­graphical distribution to Central America. Urotrema scabridum Braun, 1900 has been collected in Costa Rica for the first time, from the bat Eptesicus propinquus Peters which is also considered to be a new host record. Two species of the genus Rhapalias: R. coronatus (Rudolphi 1819) Stiles and Hassall, 1898 and R. horridus (Diesing, 1850) Stiles and Hassall, 1898 are registered from the Costa Rican opossum, Didelphis marsupialis etensi Allen; both species are known to parasitize various South American and Mexican marsupials and the Costa Rican specimens do not differ from these in structural or measurable details. Finally, the trematode Heronimus chelydrae Mac Callum, 1902 has been collected from the lungs of freshwater turtles of the genus Kinosternon. According to the studies of E. Caballero y C. 1940, it is believed that this is the only species in the genus; it has also been reported from turtles of Panama, Mexico and the United States in North America. It has been noted that an early regression of the testes occurs in this species, since these were persistent in only a few of the specimens described.

Hydroquinone derivatives and monoterpenoids from the Neotropical liverwort Plagiochila rutilans

The Shuttle Radar Topography Mission (SRTM) generated one of the most-complete high-resolution digital topographic data sets of the world to date. The elevations generated by the on-board C-band sensor represent surface elevations in “bare earth” regions, and the elevations of various scatterers such as leaves and branches in other regions. Elevations generated by a medium-footprint (� 10 m diameter) laser altimeter (lidar) system known as NASA’s Laser Vegetation Imaging Sensor (LVIS) were used to assess the accuracy of SRTM elevations at study sites of variable relief, and landcover. Five study sites in Maine, Massachusetts, Maryland, New Hampshire, and Costa Rica were chosen where coincident LVIS and SRTM data occur. Both ground and canopy top lidar elevations were compared to the SRTM elevations. In “bare earth” regions, the mean vertical offset between the SRTM elevations and LVIS ground elevations varied with study site and was approximately 0.0 m, 0.5 m, 3.0 m, 4.0 m, and 4.5 m at the Maine, Maryland, Massachusetts, New Hampshire, and Costa Rica study sites, respectively. In vegetated regions, the mean vertical offset increased, implying the phase center fell above the ground, and the offset varied by region. The SRTM elevations fell on average approximately 14 m below the LVIS canopy top elevations, except in Costa Rica where they were approximately 8 m below the canopy top. At all five study sites, SRTM elevations increased with increasing vertical extent (i.e., the difference between the LVIS canopy top and ground elevations and analogous to canopy height in vegetated regions). A linear relationship was found sufficient to describe the relationship between the SRTM-LVIS elevation difference and canopy vertical extent.

One striking paradox in epidemiologic research is the strong association between diet and cancer in ecologic studies compared with the weaker associations reported in many within-country case-control and cohort studies. However, most ecologic studies have relied on indirect measures of dietary intake, such as food disappearance data. The objectives of our study were to assess the feasibility of collecting dietary and biomarker data from individuals living in countries having markedly different dietary patterns and cultures and to examine the magnitude of the between-country variation in their measurement. Adults surveyed in Shanghai (China), Costa Rica and King County (Washington, USA) completed a 24-hr dietary recall, a cancer risk factor survey, and provided a blood sample. We analyzed a subset of the blood specimens for vitamins C, E, carotenoids and phospholipid fatty acids. We observed substantial differences in nutrient intakes and in mean plasma concentrations of dietary biomarkers across the study populations. For example, King County participants had the highest daily intake of vitamin C (mean 78.3 ± 12.2 mg compared with 42.6 ± 38.3 mg in Shanghai and 34.8 ⅔ 43.8 mg in Costa Rica). The mean plasma vitamin C level in King County was also the highest of the 3 study sites: 927.9 ± 43.9 μg/dl in King County, 585.7 ± 35.9 μg/dl in Shanghai and 461.1 ± 33.1 μg/dl in Costa Rica. Plasma trans fatty acids (a biomarker of a diet high in hydrogenated fats) were highest in King County and lowest in Shanghai. Int. J. Cancer 82:28–32, 1999. © 1999 Wiley-Liss, Inc.

Detection of rickettsiae in fleas and ticks from areas of Costa Rica with history of spotted fever group rickettsioses

One striking paradox in epidemiologic research is the strong association between diet and cancer in ecologic studies compared with the weaker associations reported in many within-country case-control and cohort studies. However, most ecologic studies have relied on indirect measures of dietary intake, such as food disappearance data. The objectives of our study were to assess the feasibility of collecting dietary and biomarker data from individuals living in countries having markedly different dietary patterns and cultures and to examine the magnitude of the between-country variation in their measurement. Adults surveyed in Shanghai (China), Costa Rica and King County (Washington, USA) completed a 24-hr dietary recall, a cancer risk factor survey, and provided a blood sample. We analyzed a subset of the blood specimens for vitamins C, E, carotenoids and phospholipid fatty acids. We observed substantial differences in nutrient intakes and in mean plasma concentrations of dietary biomarkers across the study populations. For example, King County participants had the highest daily intake of vitamin C (mean 78.3 +/- 12.2 mg compared with 42.6 +/- 38.3 mg in Shanghai and 34.8 +/- 43.8 mg in Costa Rica). The mean plasma vitamin C level in King County was also the highest of the 3 study sites: 927.9 +/- 43.9 microg/dl in King County, 585.7 +/- 35.9 microg/dl in Shanghai and 461.1 +/- 33.1 microg/dl in Costa Rica. Plasma trans fatty acids (a biomarker of a diet high in hydrogenated fats) were highest in King County and lowest in Shanghai.

Swarming Activity of Polybiine Social Wasps (Hymenoptera: Vespidae: Polybiini)

We studied the intraspecific territorial defense behavior of wintering Willow Flycatchers (Empidonax traillii) in Costa Rica using a randomized playback experiment that exposed male and female birds to recordings of Willow Flycatcher songs and calls, Lesser Ground Cuckoo (Morococcyx erythropygius) vocalizations, and random noise. Flycatchers of both sexes responded most strongly to simulated conspecific territory intrusion, and the agonistic behaviors that we observed were similar to those seen during natural intraspecific encounters in winter. Both males and females engaged in song and aggressive behaviors in defense of territories, and there was no significant difference between the sexes in scored agonistic responses. The similarity between the sexes in intraspecific territorial defense behaviors and aggressiveness may account for both sexes of flycatchers using the same habitats at our study sites in Costa Rica, and wintering females defending territories against males. The Willow Flycatcher, a sexually monomorphic species, differs in this way from a number of sexually dimorphic passerines, in which behaviorally dominant males occur in more optimal winter habitats.

The reproductive ecology of two eastern Pacific zooxanthellate coral species was examined as part of a continuing series of studies relating bleaching/mortality events caused by the El Nino–Southern Oscillation disturbance, and is described for study sites in Costa Rica, Panama, and the Galapagos Islands (Ecuador). This study deals with the sibling agariciid species Pavona varians and Pavona sp.a over a 13 yr period (1985 to 1997). Both Pavona species are broadcast-spawners with some gonochoric, but mostly sequential hermaphroditic colonies. Minimum colony sizes (and ages) at first reproduction were 5 cm (5 yr) and 3 cm (2 to 3 yr), respectively, in P. varians and Pavona sp.a. In the Panama and Galapagos populations, gonochoric colonies spawn eggs or sperm at least monthly. Six fecundity attributes were not significantly different in the two species, but the eggs of P. varians are white to beige and positively buoyant, and those of Pavona sp.a are dark green and neutrally to negatively buoyant. Eggs of both species lack zooxanthellae. Both species are reproductively active year-round, with maximum activity in the dry season in the nonupwelling Gulf of Chiriqui, and in the wet season in the upwelling Gulf of Panama. Spawning is predominantly during full moon, and possibly also at new moon at most study sites. Spawning in P. varians and Pavona sp.a is 12 h out of phase, with the former species spawning ∼1 h before sunrise and the latter about 1 h after sunset. The fecundity of Pavona spp. at Cano and the Galapagos Islands was much greater (19 900 to 27 900 eggs cm−2 yr−1) than at all Panama sites (14 800 to 19 800 eggs cm−2 yr−1). Intraspecific crosses in both species resulted in swimming planula larvae after 25 to 36 h. Recruitment of P. varians was highest in Panama, moderate in Costa Rica, and nil in the Galapagos Islands, matching, respectively, the contributions of P. varians to the pre-1982/1983 El Nino coral-population abundances in these areas. Recruitment success of P. varians at Uva Island was significantly related to maximum monthly positive sea surface-temperature (SST) anomalies that occurred in the year preceding recruitment over the period 1982 to 1996; recruitment failed when SST anomalies exceeded 1.6 to 1.9 C° during the severe ENSO events of 1982/1983 and 1997/1998.

Nectar Availability and Bee-Foraging on Ipomoea (Convolvulaceae)

-We studied the location and site selection of burrows of black iguana, Ctenosaura similis, at Palo Verde, Guanacaste, Costa Rica, in 1989 and 1990. Our study site included the environs of buildings as well as dry lowland deciduous forest. Iguana had burrows on earth banks, and in trees, logs and rocks with a higher frequency than occurred at randomly-selected points. Burrows differed from random points in having less cover over the entrance and within 1 m (both rock and vegetation) than did the random points. Temperatures at the burrow entrance were cooler than at the random points, although air temperatures did not differ significantly. We suggest that the use of burrows and selection of burrow sites by black iguanas may relate to predation pressures, thermal constraints and daily activity patterns. Descriptions of general habitat use are numerous for reptiles (e.g., Heatwole, 1977), although there are fewer descriptions of nest sites (see Van Devender and Howard, 1973; Burger and Zappalorti, 1986, for review) or of nonnesting burrows. Studies of site selection, whereby the authors have shown that the reptiles chose particular sites over all available sites, are even less common. Studies of selection require the location of a sufficient sample of nests or burrows for statistical analyses, and it is often difficult to find reptile nests or burrows. Nonetheless, nest site selection has been studied in alligators (Alligator mississippensis; Garrick and Lang, 1977), green iguanas (Iguana iguana; Rand, 1968), turtles (Burger and Montevecchi, 1975; 430 This content downloaded from 157.55.39.153 on Mon, 19 Sep 2016 04:52:46 UTC All use subject to http://about.jstor.org/terms BURROW SELECTION BY IGUANAS Ewert, 1976; Stoneburner and Richardson, 1981), and pine snakes (Pituophis melanoleucus; Burger and Zappalorti, 1986). Most reptiles do not regularly dig burrows for daily occupation (see Wright and Wright, 1957), although many reptiles use subterranean burrows dug by mammals. However some lizards (Rand and Dugan, 1983) and a snake (Carpenter, 1982; Burger et al., 1988) dig burrows for nests or for use throughout the year. Presumably the location of these burrows reflects evolutionary constraints such as predation pressures, conspecific competition, and thermoregulation. In this paper we examine burrow site selection in black iguana (Ctenosaura similis) at Palo Verde, Guanacaste, Costa Rica. We were interested in whether the same burrows were used from year to year, whether burrow site characteristics varied between years, and whether burrow sites differed from the available habitat. Black iguanas are large iguanid lizards ranging from southern Mexico to Panama. The flesh is edible, and they are often hunted for food (Fitch and Hackforth-Jones, 1983). They range in size from 20-50 cm snout-vent length (SVL), with overlap between the sexes, although males are generally larger (Fitch and Hackforth-Jones, 1983). METHODS AND STUDY AREA We studied black iguanas at the Palo Verde National Park in Guanacaste, northwestern Costa Rica, in March of 1989 and 1990. Palo Verde contains mainly tropical lowland deciduous forest that borders the extensive marshes of the Tempisque River. Further description of the study area can be found in Gill (1989) and Burger and Gochfeld (1991). In 1989 we surveyed the area immediately around the OTS biological station, the deciduous forest within 25 m of the road between the OTS station and the park headquarters (hereafter called the hacienda), and around the hacienda (1 km from the station). We searched by walking transects through each area in a manner that allowed us to see nearly all of the habitat. Burrows included all holes where iguanas lived, but did not include nests or spaces under buildings. We included only burrows actively being used by black iguanas. We used photographs and sketches from 1989 to locate the sites of all burrows for comparison with 1990. To compare overall habitat choice (bank, ground, flat rock, rock or trees, see Table 2) with available habitat, we used a table of random numbers to generate coordinates for a grid used to select points for the entire study site. General habitat types included bank (slope over 15?), ground (flat area), flat rock, rocks or boulders, and trees, logs or roots. We recorded specific site characteristics (see Table 3) from all burrows and from an equal number of random points in each of the three plac s (OTS station, deciduous forest, hacienda). Specific site characteristics recorded included: slope; substrate (ground, rock, bank); percent cover over the opening; percent vegetation cover within 1 m of the point (or burrow entrance estimated); percent rock within 1 m; percent overall cover (vegetation, trees, rock) within 1 m; distance to the closest burrow and closest tree; and temperature of the air and at the mouth of the burrow entrance or spot. We so recorded the height and width of the burrow entrance for all burrows whether they were dug or were in rock crevices. Random points were selected by using a table of random numbers to generate a compass direction and a distance (1 to 4 m) from each burrow. Thus, each random point was matched to a burrow site. We estimated iguana length by noting where an iguana started and ended on the habitat and then measuring this distance, and checked our estimates against iguanas of known length (N = 20 from marked individuals, r = 0.9, P < 0.001). We were usually only 1-3 m from the iguana, so the estimates were possible. In 1989 both general habitat type (bank, ground, flat rock, rock, logs) and specific site characteristics were recorded for random points and for burrow sites. In 1990 we recorded the general habitat types for burrows at the OTS station, hacienda, and forest in between, but we tripled the size of the areas searched in equal proportion to the three types. We also recorded some site characteristics, including location, height and width of burrow entrance, percent cover at the entrance, overall slope of area with burrow, slope at the burrow entrance, nearestneighbor distance, and the length of the burrow. Burrow lengths are minima because we had difficulty measuring beyond the bend if a burrow made a major turn. Indeed it was impossible to determine if the stick was at the end. We used Wilcox X2 tests to determine differences in the distribution of site characteristics between the burrows and random sites, and Contingency x2 tests to determine differences between the general habitat types of random sites and burrows in 1989 and 1990, and between the 1989 and 1990 burrow sites.

The Puerto Rican Common coqui frog (Eleutherodactylus coqui) has a long history as an invasive species in places such as Hawaii. Since its introduction in Costa Rica, scarce information is available to understand why and how the habitat in the Turrialba town is suitable for the species. Our goal was to analyze the habitat selection of E. coqui to identify if there are key habitat features that explained its success there. We measured 9 site variables that may affect the habitat selection of E. coqui in 92 survey units of 10 m radius distributed over a 500 m radius from its introduction point. We registered the presence/pseudo-absence data of E. coqui and environmental variables in each survey unit during eight surveys. We ran occupancy models to determine the influence of the variables on the habitat selection and to estimate its detection probability. We found that sites near the introduction point, containing abundant vegetation, bromeliads, and palms have a higher probability to be occupied by E. coqui. The habitat selection in Costa Rica shares characteristics with the populations of Puerto Rico and Hawaii. But, unlike the case in Hawaii, in Costa Rica this species has maintained a limited dispersal because the potentially higher biotic resistance, as well a sedentary behavior. However, the microhabitat conditions used by E. coqui in the study site are common throughout the country. Therefore, active management in new populations and environmental education programs to avoid human transportation of the species is critical to reduce its dispersal.

We examined the structure of the community of epiphytes growing on Crescentia alata trees in Santa Rosa National Park, Costa Rica. Four orchid species were dominant in this community, showing a distinct order of colonization of the calabash trees, with Oncidium cebolleta colonizing first, Encyclia cordigera second, Brassavola nodosa third, and Laelia rubescens fourth. We were unable to find any differentiation of the locations of these species based on the physical characteristics of the sites. It is hypothesized that the colonization pattern observed is due to the quantity of propagules produced by each species. THE NUMBER OF SPECIES found in a community is determined by five major factors-the time for colonization, the distance over which colonizers must travel, the number of potential colonizers, the size of the area to be colonized, and the biotic interactions between colonizers (MacArthur and Wilson 1967). Two approaches have been used to determine the effects of these factors upon the community organization of oceanic islands (e.g., Diamond 1969, Simberloff and Wilson 1969) and a variety of island-like situations (e.g., Vuilleumier 1970, Brown 1971, Culver, Holsinger, and Baroody 1973, Opler 1974). One approach applied to both plant (Johnson, Mason, and Raven 1965) and animal (Hamilton, Barth, and Rubinoff 1964, Wilson and Taylor 1967) communities has been to use correlative techniques to describe the patterns of species number and distribution from to on a large geographical scale. The second approach, applied exclusively to animal communities, has been to manipulate the five factors experimentally (e.g., Simberloff and Wilson 1969, Addicott 1974, Schoener 1974). In this paper we describe the community structure of plants occurring on a local scale using correlative techniques and suggest that this system is one in which experimental work on the relative roles of these five factors in determining community structures may be performed. STUDY AREA AND METHODS Individual Crescentia alata HBK, the calabash tree (Bignoniaceae), constitute an island-like situation for epiphytic plants. The calabash tree is an ideal epiphyte host because its soft, deep bark provides easy anchorage for epiphytes and recess for humus accumulations (Pittendrigh 1948). The study site, a 10 ha calabash orchard in Santa Rosa National Park, Guanacaste Province, Costa Rica, has been described by Janzen et al. (1980). It is surrounded by dry, deciduous thorn-forest, the dominant vegetation type in this region (Holdridge 1967). No individual calabash tree at this site is located more than 100 m from the edge of the thorn-forest. The spherical canopies of the calabash trees range from 0.5 to 7.5 m radius. Since trees are spaced at least 5 m apart, canopies of individuals rarely intertwine. Each calabash tree was considered an island which may be colonized by epiphytic plants: one cactus species, three bromeliads, and five species of orchids. Nomenclature is that of Janzen and Liesner ( 1980). We measured the canopy radius of a randomly chosen calabash tree and recorded the species of epiphytes present (sample sizes are indicated in figure 1). Canopy radius was used because it was both easily measured and accurate with repetition. The existence of a species-area relationship was investigated by means of regression analysis using a logarithmic curve fit after making a scatter plot of the data. To explore the temporal pattern of colonization, we calculated the percent occurrence of each epiphytic species in one-species, two-species, three-species, etc. communities. To study the role of biotic interactions, we contrasted the relative abundance of each epiphytic species on small trees (less than 2 m canopy radius) with that on large trees (greater than 4 m canopy radius). To obtain these numbers, at least two branches were randomly chosen on a minimum of five inBIOTROPICA 14(2): 137-14

Invasive species alter ecosystem integrity and functioning and are considered one of the major threats to biodiversity on a global scale. The indopacific lionfish (Plerois volitans [Linnaeus, 1758] / miles [Bennet, 1882] complex) is the first non-native marine fish that has established itself in the Western Atlantic. It was first reported in Florida in the 1980s and then spread across the entire Caribbean in subsequent years. In Costa Rica, lionfish were first sighted by the end of 2008 and are now present in all South Caribbean reefs. Lionfish are a major problem for local fisherman by displacing native fish species. The aim of this study was to determine population density, size and diet of lionfish populations at four study sites along the Southern Caribbean coast of Costa Rica. Two of the sites were located inside the National Park Cahuita where regular lionfish removal occurs, whereas the other two study sides do not experiment this kind of management. Total length and wet weight of >450 lionfish individuals were determined between March and June 2011. Three relative metrics of prey quantity (percent number, percent frequency, and percent weight) were compared from approximately 300 lionfish caught with the polespear in shallow waters (<7 m depth). Population density was assessed weekly through visual transect surveys. Our results showed that lionfish preyed mostly upon teleosts and crustaceans. Teleosts dominated lionfish diet in percent frequency (71%) and percent weight (85%), whereas crustaceans had the highest percent number (58%). The top five teleost families of dietary importance were Pomacentridae, Acanthuridae, Blennidae, Labridae and Serranidae. The average total length (+/- SD) of lionfish was 18.7 (+/- 5.7)cm and varied significantly between sites (p<0.001). Mean density of lionfish was 92fish/ha with no significant differences between sites. Smallest fish and lowest densities were found at the two sites inside the National Park Cahuita. Despite management efforts on a regional scale, nationwide efforts are ineffective and lionfish control activities are poorly implemented. We conclude that there is an urgent need to develop an improved institutional framework for local lionfish control that promotes effective coordination among the relevant stakeholders in order to deal with invasive lionfish in Costa Rica.

Flowers of Plumeria rubra, a mass-flowering tropical tree, share many traits with hawkmoth-pollinated flowers, and Plumeria pollen was recovered from the tongues of 17 hawkmoth species in Costa Rica. Rates of visitation and fruit set were exceptionally low compared with other hawkmoth flowers sympatric with Plumeria, apparently because Plumeria flowers produce no nectar reward. Hawkmoths in a flight cage quickly learned to avoid nectarless Plumeria flowers, but readily fed from these flowers after artificial nectar was added. Although no specific model for Plumeria was found in Costa Rica, the presentation of generalized odor and visual cues that mimic typical hawkmoth flowers may facilitate deceitful pollination of Plumeria. FLORAL MIMICRY ENCOMPASSES a diversity of reproductive strategies used by plants to achieve pollination by animals (see reviews of Wiens 1978, Little 1983). Because of experience with a model, pollinators visit a floral mimic expecting a reward. However, the mimic provides no reward and pollination occurs through deceit. Despite the importance of floral mimicry as a reproductive strategy, little detailed information about such systems exists (Wiens 1978, Williamson 1982, Little 1983). Williamson (1982) stressed that floral mimicry should be defined in terms of reproductive fitness. However, showing that fitness of a floral mimic is increased by its interaction with a model has rarely been successful in floral mimicy systems (Boyden 1980, Bierzychudek 1981, Williamson, pers. comm.). The essential difference in selection pressure on floral and insect mimics suggests that floral mimics may be less dependent upon resemblance to a model than insect mimics. Failure of an insect mimic to deceive a predator usually leads to the mimic's death, but failure of a floral mimic to deceive a pollinatory only results in the loss of potential pollination event. Deceitful plants may also attract pollinators by presenting cues that the pollinators innately recognize, rather than relying on strong resemblance to a model. In some instances a specific model may be absent (Dafni and Ivri 1981a, b; Ackerman 1981, 1983). In addition, convergence within floral syndromes may further complicate analysis of floral mimicry systems (Brown and Kodrik-Brown 1979, Williamson and Black 1981, Little 1983). In this paper I describe the floral biology of Plumeria rubra L. (Apocynaceae), a tropical tree apparently adapted for hawkmoth pollination. Plumeria produces no floral nectar and may be a generalized mimic of other hawkmoth-pollinated flowers. The reproductive biology of this species differs from typical examples of floral mimicry in that Plumeria lacks a specific model, is a massively flowering tree with an extended blooming period, and may be the most abundant hawkmoth flower in tropical deciduous forest of Costa Rica during much of its blooming period. STUDY SITES AND METHODS From 1976-1982 I studied Plumeria in northwestern Costa Rica (Guanacaste Province) at Hacienda La Pacifica, 10 km north of Canias, and at the Comelco ranch, centered 10 km north of Bagaces. The lowland deciduous and evergreen riparian forests at these sites (50-100 m elevation) are described by Frankie et al. (1974, 1983), Opler et al. (1980), and Opler (1983). I also made observations near the community of Monteverde (northern Puntarenas Province) from 1979-1982 where Plumeria grows in semi-deciduous, moist forest on the Pacific slopes at 800-1200 m elevation. FLORAL BIOLOGY.-Flowering phenology is based on Frankie et al. (1974) and my observations from 19761982. I studied the floral behavior of Plumeria using the methods of Haber and Frankie (1982) and Frankie et al. (1983). I monitored nectar secretion intensively on bagged flowers of two trees over several days and checked for possible nectar production on about 20 additional trees by probing flowers with microcapillary tubes and inspecting dissected flowers. I counted ovules in dissected flowers and seeds in mature fruits from the lowland site. Fruit numbers were easily observed on leafless trees during the dry season at