K. Greg Murray

Floral neighborhood and pollination success in four hummingbird-pollinated cloud forest plant species.

In a cloud forest at Monteverde, Costa Rica, we examined pollen loads received by self-compatible flowers of two pairs of plant species pollinated by hummingbirds: Hansteinia ble- pharorachis and Razisea spicata (Acanthaceae), and Besleria triflora and Drymonia rubra (Gesneri- aceae). Each pair consisted of one species (Hansteinia or Besleria) pollinated by short-billed hum- mingbirds and a related species (Razisea or Drymonia) pollinated by long-billed hummingbirds. At three different times per species, separated by 1-3 mo, we examined flowers on 28-40 focal plants from a wide variety of floral neighborhoods, ranging from plants isolated from conspecifics, either by distance or by other flowering species pollinated by the same hummingbirds, to plants surrounded by conspecifics. Because short-billed hummingbirds often restrict foraging to areas of high flower density, and because short-tubed flowers adapted for hummingbirds often have similar pollen placement, we predicted that short-tubed flowers isolated from conspecifics would receive fewer conspecific grains and more het- erospecific grains than short-tubed flowers surrounded by conspecifics. Because long-billed hum- mingbirds often forage over large areas and because long-tubed flowers adapted for hummingbirds tend to diverge in pollen placement, we predicted that pollination of long-tubed flowers would be relatively unaffected by floral neighborhood. Effects on pollen loads of floral neighborhood (nearness to or isolation from other flowers) followed few patterns consistent with our prediction or with conventional theory. (1) There were no consistent effects of floral neighborhoods on numbers of heterospecific grains deposited on stigmas; in all four species, regardless of corolla length, effects of particular neighborhood variables (as determined with stepwise multiple regression) were as likely to run exactly counter to conventional models as to corroborate models. (2) In none of the 12 sampling runs did increases in absolute densities of neigh- boring heterospecific flowers adversely affect pollination. (3) However, in two runs, loads of conspecific grains increased with increases in the absolute density of neighboring nonspecific flowers, and/or (in three runs) with increases in their relative density (proportion of conspecifics among neighboring flowers). These runs all involved short-flowered species rather than long-flowered species, tending to confirm our initial prediction, but half the sampling runs, even of short-flowered species, failed to show any density-dependent effects from neighboring flowers pollinated by the same hummingbirds. Flowers frequently received fewer conspecific grains than they had ovules to be fertilized. Therefore, the potential existed for floral neighborhoods to affect seed set and fitness of plants. Nevertheless, even though neotropical hummingbird-pollinated flowers have been cited as examples of species whose flowering peaks are displaced through competition for pollination, competitive effects from neighboring heterospecific plants were only sporadic in the species we examined, and were particularly infrequent in those species with long flowers adapted for long-billed hummingbirds.

SEASONAL FRUIT PREFERENCES FOR LIPIDS AND SUGARS BY AMERICAN ROBINS

Abstract Fruit preference by birds is a complex process based upon the morphology and spatial arrangement of fruits and on the physiological needs and capabilities of birds. In North America, most fruits can be divided into two groups based on nutritional content: those rich in sugars relative to lipids, and those rich in lipids relative to sugars. To investigate how fruit preference may change seasonally and to determine if it is correlated with physiological state, we designed a simple laboratory experiment using American Robins (Turdus migratorius) and artificial fruits. During summer and autumn, we offered eight robins a choice between synthetic sugar-rich and lipid-rich fruits of equal caloric value and then measured food intake and assimilation efficiency for each fruit type. Overall, robins preferred sugar-rich to lipid-rich fruits during both seasons. Robins had a higher assimilation efficiency for sugars than for lipids during both seasons, although assimilation efficiency of lipids increased significantly from summer to autumn. During experiments, robins consumed significantly more sugar-rich than lipid-rich fruits in summer but not in autumn. Coupling fruit intake with assimilation efficiency indicates that in summer, robins had a higher rate of energy gain from sugars than from lipids, but by autumn the rate of energy gain from lipids increased to nearly the same level as that from sugars. Our results suggest that robins prefer sugar-rich fruits because of their simple and fast rate of digestion, enabling higher rates of energy gain, but that lipid-rich fruits become important with the onset of autumn.

DISTURBANCE, POLLINATOR PREDICTABILITY, AND POLLINATION SUCCESS AMONG COSTA RICAN CLOUD FOREST PLANTS'

Cloud forest at Monteverde, Costa Rica experiences frequent natural disturbance. To determine whether species interactions vary spatially due to physical heterogeneity produced by disturbance, we examined relationships between 22 plant species and 11 nectar—feeding bird species in 14 study plots distributed among three patch types: larger landslide—like gaps (hand—cleared areas along a trail), small gaps (formed by recent treefalls), and understory of closed—canopy forest. Species we describe here flowered in two or three patch types. The aspects of pollination we examined varied little with patch type. Mean frequency of pollinator visits varied with patch type in a few plant species but not in most, and there was no significant trend across species. Pollen loads carried by 314 mist—netted hummingbirds did not vary significantly with patch type, either in total number of grains or number of species represented. Cumulative pollen loads that hummingbirds deposited on stigmas of two species of Acanthaceae (Razisea spicata and Hansteinia blepharorachis) did not vary consistently with patch type, except that Hansteinia flowers in treefall gaps received fewer heterospecific pollen grains than flowers in the other two patch types. Frequency of fruit set varied significantly with patch type in three of the four species examined, but the direction of variation in one of these was opposite to the direction of the other two. The absolute frequency with which flowers were pierced by nectar—robbing hummingbirds did not vary significantly with patch type, although the frequency of piercing relative to legitimate pollinator visits did increase in the large gaps. We attribute the latter result to aggregation of the hummingbird Eupherusa eximia, a chronic nectar robber, at dense clumps of long—flowered plant species that occurred in large gaps. Only one feature we examined suggested that patch type might directly affect the nature of species interactions: in two different analyses, the level of variation in frequency of hummingbird visits to flowers declined from large gaps to small gaps to forest. Results suggest that, unless the disturbance initiating a patch is unusually severe or widespread, interactions between the plants and hummingbirds examined are insensitive to patch type. Such species, existing in naturally dynamic forests throughout their recent evolutionary histories, presumably have become accommodated to frequent small—scale disturbance. Results also suggest that those habitat—related contrasts in plant reproductive traits and plant—pollinator interactions documented in other studies, which compare habitats initiated by anthropogenic disturbances with undisturbed patches, may be artifacts to some extent. Anthropogenically generated disturbance mosaics may promote the spread of species whose reproductive traits evolved under very different circumstances from mosaics generated by natural disturbances.

Effects of urbanization on the population structure of freshwater turtles across the United States: Urbanization Effects on Turtles

Landscape-scale alterations that accompany urbanization may negatively affect the population structure of wildlife species such as freshwater turtles. Changes to nesting sites and higher mortality rates due to vehicular collisions and increased predator populations may particularly affect immature turtles and mature female turtles. We hypothesized that the proportions of adult female and immature turtles in a population will negatively correlate with landscape urbanization. As a collaborative effort of the Ecological Research as Education Network (EREN), we sampled freshwater turtle populations in 11 states across the central and eastern United States. Contrary to expectations, we found a significant positive relationship between proportions of mature female painted turtles (Chrysemys picta) and urbanization. We did not detect a relationship between urbanization and proportions of immature turtles. Urbanization may alter the thermal environment of nesting sites such that more females are produced as urbanization increases. Our approach of creating a collaborative network of scientists and students at undergraduate institutions proved valuable in terms of testing our hypothesis over a large spatial scale while also allowing students to gain hands-on experience in conservation science.