Effect of treefall gaps on the patchiness and species richness of Neotropical ant assemblages

Natural formation of treefall gaps plays an integral role in the ecological and evolutionary dynamics of many tropical forests, affecting the spatiotemporal distribution of plants and the animals that interact with them. This study examines the impact of treefall gaps on the spatial and temporal patchiness of ant assemblages in a moist lowland forest in Panama. Using pitfall traps and honey baits, we compared ant assemblages in five 1 to 2-year-old treefall gaps (ca 100 m²) and five adjacent plots (ca 100 m²) in undisturbed forest understory at three different times of year (late wet season, late dry season, and early wet season). We found little evidence that ant assemblages respond dramatically to the formation of treefall gaps and the differences in habitat qualities they produce. Ant abundance, species richness, species composition, and rates of resource discovery did not differ between gaps and forest understory. However, we did find significant differences in numerical abundance related to forest stratum (ground vs vegetation) and resource type in pitfall traps (oil-cockroach vs honey), and significant differences in ant species richness and rates of resource discovery across seasons. While habitat effects by themselves were never statistically significant, habitat and seasonal differences in species richness interacted significantly to produce complex, season-dependent differences among gap and forest habitats. These results suggest that the formation of natural treefall gaps has less of an effect on Neotropical ant assemblages compared to other groups of organisms (e.g., plants, birds) or other causes of patchiness (e.g., ant mosaics, moisture availability, army ant predation). The results of our study also have important implications for the underlying causes of habitat differences in the distribution of ant-defended plants. Received: 3 February 1998 / Accepted: 7 April 1998     

Habitat‐dependent population regulation in an irrupting population of long‐nosed bandicoots (Perameles nasuta)

We used isodars to analyse habitat‐dependent population regulation by long‐nosed bandicoots Perameles nasuta during an irruption and subsequent population crash in three habitats (heath, woodland and forest) at Booderee National Park, south‐eastern Australia. Specifically, we aimed to see whether patterns of habitat‐dependent population regulation matched a priori estimates of quantitative and qualitative differences between habitats. We also tested if habitat preference changed between the increasing and decreasing phase of the irruption as predicted by the reciprocating dispersal theory. Quantitative differences in habitat quality were indexed by the relative abundance of the main food of long‐nosed bandicoots (terrestrial invertebrates), while qualitative differences were indexed by the availability of refuge from predation (vegetation understorey density). One index of fitness, body weight, was highest in forest, and lowest in heath, suggesting an ideal despotic model of habitat selection. Over the entire course of the irruption, there was density‐dependent habitat selection with forest and woodland both quantitatively superior to heath. This reflected the overall abundance of invertebrates with highest abundance in woodland and forest and less in heath. Isodar analysis also revealed that although forest was quantitatively better than heath and equivalent to woodland it was qualitatively poorer than either habitat. Heath had a higher density of understorey than woodland and woodland having a higher density of understorey than forest giving crossover population regulation. When the increasing and declining phase of the irruption were analysed separately, no habitat was quantitatively superior to any other during either phase. The lack of switching in preference between habitats from the increasing to the declining phase of the irruption and the virtual absence of any dispersal by adults, does not support the reciprocating dispersal hypothesis.     

Density‐dependent habitat selection predicts fitness and abundance in a small lizard

Density‐dependent habitat selection has been used to predict and explain patterns of abundance of species between habitats. Thermal quality, a density‐independent component of habitat suitability, is often the most important factor for habitat selection in ectotherms which comprise the vast majority of animal species. Ectotherms may reach high densities such that individual fitness is reduced in a habitat due to increased competition for finite resources. Therefore, density and thermal quality may present conflicting information about which habitat will provide the highest fitness reward and ectotherm habitat selection may be density‐independent. Using ornate tree lizards Urosaurus ornatus at 10 sites each straddling two adjacent habitats (wash and upland), we tested the hypothesis that habitat selection is density‐dependent even when thermal quality differs between habitats. We first tested that fitness proxies decline with density in each habitat, indicating density‐dependent effects on habitat suitability. We also confirmed that the two habitats vary in suitability (quantified by food abundance and thermal quality). Next, we tested the predictions that habitat selection depends on density with isodar analyses and that fitness proxies are equal in the two habitats within a site. We found that monthly survival rates decreased with density, and that the wash habitat had more prey and higher thermal quality than the upland habitat. Lizards preferred the habitat with more food and higher thermal quality, lizard densities in the two habitats were positively correlated, and fitness proxies of lizards did not differ between habitats. These patterns are consistent with density‐dependent habitat selection, despite differences in thermal quality between habitats. We expect that density‐dependent habitat selection is widespread in terrestrial ectotherms when densities are high and temperatures are close to their optimal performance range. In areas where thermal quality is low, however, we expect that depletable resources, such as food, become less limiting because assimilating resources is more difficult.     

The influence of spatio-temporal resource fluctuations on insular rat population dynamics

Local spatio-temporal resource variations can strongly influence the population dynamics of small mammals. This is particularly true on islands which are bottom-up driven systems, lacking higher order predators and with high variability in resource subsidies. The influence of resource fluctuations on animal survival may be mediated by individual movement among habitat patches, but simultaneously analysing survival, resource availability and habitat selection requires sophisticated analytical methods. We use a Bayesian multi-state capture-recapture model to estimate survival and movement probabilities of non-native black rats (Rattus rattus) across three habitats seasonally varying in resource availability. We find that survival varies most strongly with temporal rainfall patterns, overwhelming minor spatial variation among habitats. Surprisingly for a generalist forager, movement between habitats was rare, suggesting individuals do not opportunistically respond to spatial resource subsidy variations. Climate is probably the main driver of rodent population dynamics on islands, and even substantial habitat and seasonal spatial subsidies are overwhelmed in magnitude by predictable annual patterns in resource pulses. Marked variation in survival and capture has important implications for the timing of rat control.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Combined Effects of Host Plant Quality and Predation on a Tropical Lepidopteran: A Comparison between Treefall Gaps and the Understory in Panama

In tropical forests, light‐gaps created from treefalls are a frequent source of habitat heterogeneity. The increase in productivity, through gap formation, can alter food quality, predation and their impact on insect herbivores. We hypothesized that in gaps, herbivores would be less resource‐limited and more predator limited, whereas in the understory, we predicted the reverse. In this study, we investigate the combined effects of food quality and predation on the lepidopteran larva Zunacetha annulata feeding on its host plant Hybanthus prunifolius in two habitats; sunny treefall gaps and the shaded understory in Panama. In bioassays, Z. annulata feeding on sun leaves ate 22 percent less leaf area, grew 25 percent faster, and had higher pupal weights than larvae feeding on shade leaves. However, shade leaves had higher nitrogen content and specific leaf area. In gaps, predation was 26.4 percent compared to 13.8 percent in the understory. Larvae on understory plants traveled greater distances and spent more time searching and traveling than larvae on gap plants. These differences in behavior are consistent with lower predation risk and lower quality food in the understory. Using data from bioassays and field experiments we calculated 0.22 percent and 1.02 percent survival to adulthood for larvae in gaps and the understory, respectively. In conclusion, although these habitats were in close proximity, we found that larvae in the understory are more resource‐limited and larvae in gaps are more predator limited.     

Effects of distinct types of disturbance on seed rain in the Atlantic forest of NE Brazil

In disturbed sites, some groups of seeds might be excluded from the seed rain due to their dispersal modes or seed size, and some groups might be successful as a result of disturbance effects. In the present study, we examined the seed rain in natural treefall gaps and in an area of regenerating forest following an accidental burning, which occurred 4 years before this study. Both of these disturbed areas were compared with nearby forest understorey. The number of seeds, number of species, and proportion of wind-dispersed seeds were compared between these disturbed and undisturbed areas. The treefall gaps have received lower numbers of seeds and species than the nearby understorey, but the number of wind-dispersed seeds did not differ between these areas. The lowest seed number observed in treefall gaps can be attributed to a lower number of animal-dispersed seeds, suggesting that animals may be avoiding treefall gap areas. A higher number of seeds and a lower number of species were observed in the burned area when compared to the adjacent understorey. The high number of small-sized seeds and of wind-dispersed seeds in the burned area was almost surely a consequence of the local production of the pioneer plants established after the burning. In this study, substantial differences were observed in the characteristics of the seed rain at disturbed sites, when compared with undisturbed understorey. However, these two distinct types of disturbance showed quite differing patterns, as treefall gaps received lower number of seeds while the burned area received a higher number of seeds, with a greater proportion of wind-dispersed seeds. The exception was for species richness, which was quite low at both these disturbed sites.     

Influence of gap size and soil properties on microbial biomass in a subtropical humid forest of north-east India

We examined the effects of treefall gap size and soil properties on microbial biomass dynamics in an undisturbed mature-phase humid subtropical broadleaved forest in north-east India. Canopy gaps had low soil moisture and low microbial biomass suggesting that belowground dynamics accompanied changes in light resources after canopy opening. High rainfall in the region causes excessive erosion/leaching of top soil and eventually soil fertility declines in treefall gaps compared to understorey. Soil microbial population was less during periods when temperature and moisture conditions are low, while it peaked during rainy season when the litter decomposition rate is at its peak on the forest floor. Greater demand for nutrients by plants during rainy season (the peak vegetative growth period) limited the availability of nutrients to soil microbes and, therefore, low microbial C, N and P. Weak correlations were also obtained for the relationships between microbial C, N and P and soil physico–chemical properties. Gap size did influence the microbial nutrients and their contribution to soil organic carbon, total Kjeldhal nitrogen and available-P. Contribution of microbial C to soil organic carbon, microbial N to total nitrogen were similar in both treefall gaps and understorey plots, while the contribution of microbial P to soil available-P was lower in gap compared to the understorey. These results indicate that any fluctuation in microbial biomass related nutrient cycling processes in conjunction with the associated microclimate variation may affect the pattern of regeneration of tree seedlings in the gaps and hence be related with their size. This revised version was published online in June 2006 with corrections to the Cover Date.     

Avian distribution in treefall gaps and understorey of terra firme forest in the lowland Amazon

We compared the bird distributions in the understorey of treefall gaps and sites with intact canopy in Amazonian terra firme forest in Brazil. We compiled 2216 mist-net captures (116 species) in 32 gap and 32 forest sites over 22.3 months. Gap habitats differed from forest habitats in having higher capture rates, total captures, species richness and diversity. Seventeen species showed a significantly different distribution of captures between the two habitats (13 higher in gap and four higher in forest). Gap habitats had higher capture rates for nectarivores, frugivores and insectivores. Among insectivores, capture rates for solitary insectivores and army ant followers did not differ between the two habitats. In contrast, capture rates were higher in gaps for members of mixed-species insectivore flocks and mixed-species insectivore–frugivore flocks. Insectivores, especially members of mixed-species flocks, were the predominant species in gap habitats, where frugivores and nectarivores were relatively uncommon. Although few canopy species were captured in gap or forest habitats, visitors from forest mid-storey constituted 42% of the gap specialist species (0% forest) and 46% of rare gap species (38% forest). Insectivore, and total, captures increased over time, but did so more rapidly in gap than in forest habitats, possibly as a response to gap succession. However, an influx of birds displaced by nearby timber harvest also may have caused these increases. Avian gap-use in Amazonian terra firme forests differs from gap-use elsewhere, partly because of differences in forest characteristics such as stature and soil fertility, indicating that the avian response to gaps is context dependent.     

B chromosomes in Brazilian rodents

B chromosomes are now known in eight Brazilian rodent species: Akodon montensis, Holochilus brasiliensis, Nectomys rattus, N. squamipes, Oligoryzomys flavescens, Oryzomys angouya, Proechimys sp. 2 and Trinomys iheringi. Typically these chromosomes are heterogeneous relative to size, morphology, banding patterns, presence/absence of NORs, and presence/absence of interstitial telomeric signals after FISH. In most cases, Bs are heterochromatic and late replicating. Active NORs were detected in two species: Akodon montensis and Oryzomys angouya. As a rule, Bs behave as uni or bivalents in meiosis, there is no pairing between Bs and autosomes or sex chromosomes and also their synaptonemal complexes are isopycnotic with those in A chromosomes.     

Tree seed germination and seedling establishment in treefall gaps and understorey in a subtropical forest of northeast India

Abstract Seed germination, and survival and growth of seedlings of four dominant tree species, Quercus dealbata, Quercus griffithii, Quercus glauca and Schima khasiana were studied in the treefall gaps and forest understorey of an undisturbed mature-phase humid subtropical broadleaved forest in northeast India. Three important microenvironmental factors namely photosynthetically active radiation (PAR), soil moisture and litter depth, were also measured in the forest understorey and gaps and correlated with seedling mortality. Seed germination of S. khasiana was significantly higher in the treefall gaps than in the understorey; among the tree species studied, it had the highest germination. Quercus seedlings were abundant in the understorey and small gaps, while S. khasiana seedlings were more numerous in the large gaps. The survivorship curves for the seedling populations revealed that the three Quercus species survived better in the understorey, while S. khasiana did so in the gaps. PAR and soil moisture were positively correlated with tree seedling mortality, which occurred mainly during the winter months. The Quercus seedlings grew better in the forest understorey and small gaps and S. khasiana seedlings in the large gaps. The differential performance of the tree seedlings to the conditions prevailing in the understorey and gaps of two sizes indicates that different species were adapted to different light environments depending upon their optimum requirements. This could be an effective mechanism for promoting species coexistence in the forest community.     

Consumer movement through differentially subsidized habitats creates a spatial food web with unexpected results

Energy and nutrient flow between habitats, or allochthonous input, can have a significant impact on food web dynamics. Previous theory demonstrated that resource abundance decreases in habitats where consumers are subsidized. Here we examine the effect of subsidies that are available in localized parts of a habitat (such as near the shore in a marine‐subsidized terrestrial ecosystem) with a two‐patch model in which consumers move between patches, resources are stationary, and consumers receive the subsidy in only one of the two patches. In contrast to previous theory, our results show that subsidized consumers can increase resource abundance, though only in the subsidized patch. Furthermore, the total resource population responds positively to increasing consumer movement. These results demonstrate the importance of spatial heterogeneity in food web dynamics and the need for further examination of the role of space in multispecies trophic webs.     

Seeds and seedlings of two neotropical montane understory shrubs respond differently to anthropogenic edges and treefall gaps

Edges resulting from forest clear-cutting and treefall gaps can affect plant populations and consequently the distribution of species across landscapes. These two types of disturbance might interact to exacerbate or ameliorate “edge effects”, a rarely tested possibility. We focused on the effects of distance from forest edge (0–10, 30–40, 60–70, and 190–200 m) and habitat within forest fragments (treefall gaps and intact forest) on the early stages of development of Palicourea gibbosa and Faramea affinis, two common shrubs of montane forests in southwest Colombia. Seed germination and seedling growth did not change with distance from forest edge. Within forest fragments, however, seed germination and seedling growth were higher in treefall gaps than in intact forest understory for both species. In contrast, seed predation was influenced by distance from forest edge and in P gibbosa it depended on habitat. Seed predation was highest in the forest interior (190–200 m from forest edge) and in P. gibbosa this was true only in treefall gap habitats. These results suggest that animal mediated processes such as post-dispersal seed predation are more likely than physiological processes to be affected by anthropogenic edges. Our results provide some evidence that treefall gaps may interact with “edge effects”, however, they are inconclusive as to whether they exacerbate or ameliorate them. Received: 31 August 1998 / Accepted: 18 February 1999     

A comparison of 12 tree species of Amazonian caatinga using growth rates in gaps and understorey, and allometric relationships

1. The relative growth rate of saplings of 12 species from an oligotrophic lowland rain forest were measured in treefall gaps and understorey. Mean relative height growth ( R _H) within treefall gaps was found to be slowest for tall-tree species with branched saplings, intermediate for subcanopy trees and fastest for tall-tree species with unbranched saplings. Most species had similar R _H within the understorey. R _H values were not related to leaf mass per unit area (LMA) or foliar N concentrations.
2. Allometric relationships between the total leaf area (TLA) and height were dependent upon light conditions; in general saplings of a given height had a greater TLA in treefall gaps than in understorey. The species with the largest estimated TLA values in gaps tended to have the greatest R _H values in gaps; no such trend emerged in the understorey. The values of the allometric coefficients were not related to foliar properties.
3. The relationship between stem diameter and height was only weakly dependent on light conditions and the relationship between the growth rates in these dimensions was also weak. The lack of plasticity may reflect the fact that the height–diameter relationship has little bearing on a sapling's tolerance of shade.
4. One way of accommodating the dependence of allometry upon irradiance is to add R _H as a covariate. We derive a relationship between growth rates from this resource-dependent allometric equation and show that it reasonably describes measurements taken in the caatinga forest.     

Consumer versus resource control and the importance of habitat heterogeneity for estuarine bivalves

The relative influence of consumers (top down) and resources (bottom up) on the distribution and abundance of organisms remains a key question in ecology. We examined the relationships between consumer and resource variables along a productivity gradient for a dominant predator–prey interaction in a marine soft‐sediment system. We 1) quantified density and size of the clam Macoma balthica (prey species) in six replicate sites at each of four habitat types (shallow mud, deep mud, muddy sand and detrital mud) in the Rhode River, Chesapeake Bay. We selected one habitat type of high food availability and clam density (shallow mud) and another of low food availability and clam density (muddy sand) for manipulative experiments. Then, we 2) measured M. balthica survival and growth through transplants, 3) measured food availability as sedimentary organic carbon content, 4) quantified predator density, and 5) calculated predator foraging efficiency in the two habitat types. Clam density in the four habitat types differed and was related to sedimentary carbon availability and predator density. One of the habitats, detrital mud, appeared to be a population sink because it only held juvenile Macoma that never survived to reproductive age. Macoma size and growth, and predator (mainly blue crab Callinectes sapidus) densities were positively correlated with productivity and were higher in shallow mud than muddy sand. In contrast, Macoma mortality, local ‘interaction strength’, and predator foraging efficiency were lower in the productive habitat (shallow mud). Thus, predation intensity was inversely correlated with productivity (food availability); consumer and resource effects differed by habitat type; and, at a relatively small spatial scale, consumer and resource forces jointly determined population dynamics in this soft‐sediment marine system.     

Seasonal and habitat differences affect the impact of food and predation on herbivores: a comparison between gaps and understory of a tropical forest

Herbivore populations are influenced by a combination of food availability and predator pressure, the relative contribution of which is hypothesized to vary across a productivity gradient. In tropical forests, treefall gaps are pockets of high productivity in the otherwise less productive forest understory. Thus, we hypothesize that higher light availability in gaps will increase plant resources, thereby decreasing resource limitation of herbivores relative to the understory. As a result, predators should regulate herbivore populations in gaps, whereas food should limit herbivores in the understory. We quantified potential food availability and compared arthropod herbivore and predator densities in large forest light gaps and in the intact understory in Panama. Plants, young leaves, herbivores and predators were significantly more abundant per ground area in gaps than in the understory. This pattern was similar when we focused on seven gap specialist plant species and 15 shade-tolerant species growing in gaps and understory. Consistent with the hypothesis, herbivory rates were higher in gaps than the understory. Per capita predation rates on artificial caterpillars indicated higher predation pressure in gaps in both the dry and late wet seasons. These diverse lines of evidence all suggest that herbivores experience higher predator pressure in gaps and more food limitation in the understory.     

Contrasting bee foraging in response to resource scale and local habitat management

It is hypothesized that two main factors drive the foraging patterns of native and exotic species: food resource availability and habitat composition. These factors are particularly relevant for native bees and exotic honeybees, essential crop pollinators that are sensitive to floral resources and habitat management, and that have recently exhibited alarming population declines. Mechanisms driving native and exotic bee foraging patterns may critically depend on floral resource availability and habitat composition, yet the impacts of these factors on bee foraging have never been simultaneously analyzed. In a coffee producing region in southern Mexico, we investigated the influence of coffee floral resource levels and habitat management on native and exotic bee foraging. We measured the amount of flowering coffee available at multiple spatial scales within two distinct agroforestry habitat types (high-shade and low-shade coffee) and recorded visits to coffee flowers, documenting bee species, visit duration and visit frequency. We observed a significantly greater number of visits in high-shade coffee habitats than in low-shade coffee habitats for both native and exotic bees. In high-shade coffee habitats, native solitary bee and native social bee visitation decreased significantly in response to increasing floral resource availability, exhibiting a 'dilution effect' at the smallest spatial scale. In contrast, in low-shade coffee habitats, Africanized honeybees exhibited a 'concentration effect', increasing visitation significantly in response to increasing floral resource availability at the largest spatial scale. This study is the first to show that foraging patterns of native bees and exotic honeybees contrast in response to floral resource level and scale and that this response is mediated by the vegetation management of the local habitat.     

Dispersal among habitats varying in fitness: reciprocating migration through ideal habitat selection

Current evolutionary models of dispersal set the ends of a continuum where the number of individuals emigrating from a habitat either equals the number of individuals immigrating (balanced dispersal) or where emigrants flow from a source habitat to a corresponding sink. Theories of habitat selection suggest a more sophisticated conditional strategy where individuals disperse from habitats where they have the greatest impact on fitness to habitats where their per capita impact is lower. Asymmetries between periods of population growth and decline result in a reciprocating dispersal strategy where the direction of migration is reversed as populations wax and wane. Thus, for example, if net migration of individuals flows from high- to low-density habitats during periods of population growth, net migration will flow in the opposite direction during population decline. Stochastic simulations and analytical models of reciprocating dispersal demonstrate that fitness, carrying capacity, stochastic dynamics, and interference from dominants interact to determine whether dispersal is balanced between habitats, or whether one habitat or the other acts as a net donor of dispersing individuals. While the pattern of dispersal may vary, each is consistent with an underlying strategy of density-dependent habitat selection.     

Clonal integration is beneficial for resource sharing in a creeping amphibian herb (<Emphasis Type="Italic">Alteranthera philoxeroides</Emphasis>)

Clonal integration facilitates the growth and reproduction of clonal plants by providing the ability to share resources among ramets in heterogeneous environments. The benefits of clonal integration for plant growth may depend on a contrast in resource availability and may encounter costs, especially when a young part of the clone is growing across a border between richer and poorer conditions than the old part. We studied a clonal amphibian plant growing across a border between an aquatic and a terrestrial ecosystem, which typically differ in the availability of resources. We asked whether the young part of the clone is supporting the old part with phosphorus and whether this support has costs. We performed an experiment with Alternanthera philoxeroides where plants grow from water to a terrestrial habitat. The terrestrial habitat had either a low or high phosphorus supply, and the connection between the old and young parts of the clone was either left intact or split. We determined that the young part of the clone growing in a terrestrial habitat supported the old part with phosphorus when growing on a substrate rich in phosphorus. We have found no cost of this resource translocation; on the contrary, whole clones increased not only their accumulation of phosphorus, but also of nitrogen. Our study shows how an amphibian plant may profit from heterogeneous habitats by resource sharing in a clonal network.     

The evolution of life histories in spatially heterogeneous environments: Optimal reaction norms revisited

Summary Natural populations live in heterogeneous environments, where habitat variation drives the evolution of phenotypic plasticity. The key feature of population structure addressed in this paper is the net flow of individuals from source (good) to sink (poor) habitats. These movements make it necessary to calculate fitness across the full range of habitats encountered by the population, rather than independently for each habitat. As a consequence, the optimal phenotype in a given habitat not only depends on conditions there but is linked to the performance of individuals in other habitats. We generalize the Euler-Lotka equation to define fitness in a spatially heterogeneous environment in which individuals disperse among habitats as newborn and then stay in a given habitat for life. In this case, maximizing fitness (the rate of increase over all habitats) is equivalent to maximizing the reproductive value of newborn in each habitat but not to maximizing the rate of increase that would result if individuals in each habitat were an isolated population. The new equation can be used to find optimal reaction norms for life history traits, and examples are calculated for age at maturity and clutch size. In contrast to previous results, the optimal reaction norm differs from the line connecting local adaptations of isolated populations each living in only one habitat. Selection pressure is higher in good and frequent habitats than in poor and rare ones. A formula for the relative importance of these two factors allows predictions of the habitat in which the genetic variance about the optimal reaction norm should be smallest.