Microhabitats in the tropics buffer temperature in a globally coherent manner

Vegetated habitats contain a variety of fine-scale features that can ameliorate temperate extremes. These buffered microhabitats may be used by species to evade extreme weather and novel climates in the future. Yet, the magnitude and extent of this buffering on a global scale remains unknown. Across all tropical continents and using 36 published studies, we assessed temperature buffering from within microhabitats across various habitat strata and structures (e.g. soil, logs, epiphytes and tree holes) and compared them to non-buffered macro-scale ambient temperatures (the thermal control). Microhabitats buffered temperature by 3.9°C and reduced maximum temperatures by 3.5°C. Buffering was most pronounced in tropical lowlands where temperatures were most variable. With the expected increase in extreme weather events, microhabitats should provide species with a local layer of protection that is not captured by traditional climate assessments, which are typically derived from macro-scale temperatures (e.g. satellites). Our data illustrate the need for a next generation of predictive models that account for species'' ability to move within microhabitats to exploit favourable buffered microclimates.     

Thermoregulation and microhabitat use in mountain butterflies of the genus Erebia: Importance of fine-scale habitat heterogeneity

Mountain butterflies have evolved efficient thermoregulation strategies enabling their survival in marginal conditions with short flight season and unstable weather. Understanding the importance of their behavioural thermoregulation by habitat use can provide novel information for predicting the fate of alpine Lepidoptera and other insects under ongoing climate change. We studied the link between microhabitat use and thermoregulation in adults of seven species of a butterfly genus Erebia co-occurring in the Austrian Alps. We captured individuals in the field and measured their body temperature in relation to microhabitat and air temperature. We asked whether closely related species regulate their body temperature differently, and if so, what is the effect of behaviour, species traits and individual traits on body to air and body to microhabitat temperature differences. Co-occurring species differed in mean body temperature. These differences were driven by active microhabitat selection by individuals and also by species–specific habitat preferences. Species inhabiting grasslands and rocks utilised warmer microclimates to maintain higher body temperature than woodland species. Under low air temperatures, species of rocky habitats heated up more effectively than species of grasslands and woodlands which allowed them to stay active in colder weather. Species morphology and individual traits play rather minor roles in the thermoregulatory differences; although large species and young individuals maintained higher body temperature. We conclude that diverse microhabitat conditions at small spatial scales probably contribute to sympatric occurrence of closely related species with different thermal demands and that preserving heterogeneous conditions in alpine landscapes might mitigate detrimental consequences of predicted climate change.     

Tropical forests are thermally buffered despite intensive selective logging

Tropical rainforests are subject to extensive degradation by commercial selective logging. Despite pervasive changes to forest structure, selectively logged forests represent vital refugia for global biodiversity. The ability of these forests to buffer temperature‐sensitive species from climate warming will be an important determinant of their future conservation value, although this topic remains largely unexplored. Thermal buffering potential is broadly determined by: (i) the difference between the “macroclimate” (climate at a local scale, m to ha) and the “microclimate” (climate at a fine‐scale, mm to m, that is distinct from the macroclimate); (ii) thermal stability of microclimates (e.g. variation in daily temperatures); and (iii) the availability of microclimates to organisms. We compared these metrics in undisturbed primary forest and intensively logged forest on Borneo, using thermal images to capture cool microclimates on the surface of the forest floor, and information from dataloggers placed inside deadwood, tree holes and leaf litter. Although major differences in forest structure remained 9–12 years after repeated selective logging, we found that logging activity had very little effect on thermal buffering, in terms of macroclimate and microclimate temperatures, and the overall availability of microclimates. For 1°C warming in the macroclimate, temperature inside deadwood, tree holes and leaf litter warmed slightly more in primary forest than in logged forest, but the effect amounted to <0.1°C difference between forest types. We therefore conclude that selectively logged forests are similar to primary forests in their potential for thermal buffering, and subsequent ability to retain temperature‐sensitive species under climate change. Selectively logged forests can play a crucial role in the long‐term maintenance of global biodiversity.     

Comparison of temperate and tropical rainforest tree species: photosynthetic responses to growth temperature

Little is known about the differences in physiology between temperate and tropical trees. Australian rainforests extend from tropical climates in the north to temperate climates in the south over a span of 33° latitude. Therefore, they provide an opportunity to investigate differences in the physiology of temperate and tropical trees within the same vegetation type. This study investigated how the response of net photosynthesis to growth temperature differed between Australian temperate and tropical rainforest trees and how this correlated with differences in their climates. The temperate species showed their maximum rate of net photosynthesis at lower growth temperatures than the tropical species. However, the temperate species showed at least 80% of maximum net photosynthesis over a 12-16°C span of growth temperature, compared with a span of 9-11°C shown by the tropical species. The tropical species showed both larger reductions in maximum net photosynthesis at low growth temperatures and larger reductions in the optimum instantaneous temperature for net photosynthesis with decreasing growth temperature than the temperate species. The ability of the temperate species to maintain maximum net photosynthesis over a greater span of growth temperatures than the tropical species is consistent with the greater seasonal and day-to-day variation in temperature of the temperate climate compared with the tropical climate.     

Habitat associations of species show consistent but weak responses to climate

Different vegetation types can generate variation in microclimates at local scales, potentially buffering species from adverse climates. To determine if species could respond to such microclimates under climatic warming, we evaluated whether ectothermic species (butterflies) can exploit favourable microclimates and alter their use of different habitats in response to year-to-year variation in climate. In both relatively cold (Britain) and warm (Catalonia) regions of their geographical ranges, most species shifted into cooler, closed habitats (e.g. woodland) in hot years, and into warmer, open habitats (e.g. grassland) in cooler years. Additionally, three-quarters of species occurred in closed habitats more frequently in the warm region than in the cool region. Thus, species shift their local distributions and alter their habitat associations to exploit favourable microclimates, although the magnitude of the shift (approx. 1.3% of individuals from open to shade, per degree Celsius) is unlikely to buffer species from impacts of regional climate warming.     

Facing the Heat: Thermoregulation and Behaviour of Lowland Species of a Cold-Dwelling Butterfly Genus,Erebia

Understanding the potential of animals to immediately respond to changing temperatures is imperative for predicting the effects of climate change on biodiversity. Ectothermic animals, such as insects, use behavioural thermoregulation to keep their body temperature within suitable limits. It may be particularly important at warm margins of species occurrence, where populations are sensitive to increasing air temperatures. In the field, we studied thermal requirements and behavioural thermoregulation in low-altitude populations of the Satyrinae butterflies Erebia aethiops, E. euryale and E. medusa. We compared the relationship of individual body temperature with air and microhabitat temperatures for the low-altitude Erebia species to our data on seven mountain species, including a high-altitude population of E. euryale, studied in the Alps. We found that the grassland butterfly E. medusa was well adapted to the warm lowland climate and it was active under the highest air temperatures and kept the highest body temperature of all species. Contrarily, the woodland species, E. aethiops and a low-altitude population of E. euryale, kept lower body temperatures and did not search for warm microclimates as much as other species. Furthermore, temperature-dependence of daily activities also differed between the three low-altitude and the mountain species. Lastly, the different responses to ambient temperature between the low- and high-altitude populations of E. euryale suggest possible local adaptations to different climates. We highlight the importance of habitat heterogeneity for long-term species survival, because it is expected to buffer climate change consequences by providing a variety of microclimates, which can be actively explored by adults. Alpine species can take advantage of warm microclimates, while low-altitude grassland species may retreat to colder microhabitats to escape heat, if needed. However, we conclude that lowland populations of woodland species may be more severely threatened by climate warming because of the unavailability of relatively colder microclimates.     

Life history strategies of cladocerans: comparisons of tropical and temperate taxa

We review recent works on different life history variables of cladoceran taxa in tropical and temperate freshwater bodies, comparing the strategies that cladocerans have evolved to adapt to contrasting environmental conditions in the two geographical regions. These life-history parameters relate to age and size at maturity, survival, fecundity, life-expectancy at birth, lifespan, gross, and net reproductive rates, generation time, the rate of population increase, peak population density and day of peak abundance. We also discuss the role of photoperiod and temperature on some of these life history parameters. We found a general paucity of experimental work and field data in tropics on cladocerans. There is very limited information on the few Daphnia species found in the tropics. The misconception of low species diversity of cladocerans in the tropics arose due to several reasons including lack of extensive and intensive field collections. Higher water temperatures apparently promote permanent infestation of tropical waters with toxic cyanobacteria, which reduce the zooplankton diversity. In addition to higher temperatures in the tropics, the year-round high predation pressure of planktivorous fish probably causes the tropical species, particularly in pelagic habitats, to reach maturity earlier (< 3 days) than in temperate regions. Species of Daphnia in temperate regions are particularly adapted to living at food concentrations that are much lower and seasonably more variable than those for tropical genera such as Diaphanosoma. This is further corroborated by the more than an order of magnitude higher threshold food concentration (TFC) for tropical Cladocera than for their temperate counterparts. Fecundity patterns differ between tropical and temperate cladoceran taxa: cultured under optimal temperature regimes, tropical taxa have fewer eggs than temperate species of a comparable body size. Predation pressure may act differently depending on the size of the cladoceran neonates and thus on their population size structure. Global warming and climate changes seem to affect the behaviour (migration), distribution, and abundance of cladocerans. Apparently, in direct response to these changes, the possibility of encountering the tropical cladocerans in the northern, temperate hemisphere (bioinvasions) is on the rise.     

Rapid diversification and not clade age explains high diversity in neotropical Adelpha butterflies

Latitudinal gradients in species richness are among the most well-known biogeographic patterns in nature, and yet there remains much debate and little consensus over the ecological and evolutionary causes of these gradients. Here, we evaluated whether two prominent alternative hypotheses (namely differences in diversification rate or clade age) could account for the latitudinal diversity gradient in one of the most speciose neotropical butterfly genera (Adelpha) and its close relatives. We generated a multilocus phylogeny of a diverse group of butterflies in the containing tribe Limenitidini, which has both temperate and tropical representatives. Our results suggest there is no relationship between clade age and species richness that could account for the diversity gradient, but that instead it could be explained by a significantly higher diversification rate within the predominantly tropical genus Adelpha. An apparent early larval host-plant shift to Rubiaceae and other plant families suggests that the availability of new potential host plants probably contributed to an increase in diversification of Adelpha in the lowland Neotropics. Collectively, our results support the hypothesis that the equatorial peak in species richness observed within Adelpha is the result of increased diversification rate in the last 10-15 Myr rather than a function of clade age, perhaps reflecting adaptive divergence in response to the dramatic host-plant diversity found within neotropical ecosystems.     

Thermal Buffering of Microhabitats is a Critical Factor Mediating Warming Vulnerability of Frogs in the Philippine Biodiversity Hotspot

Species may circumvent the impacts of climate warming if the habitats they use reduce ambient temperature. In this study, we identified which frog species from a tropical montane rain forest in the Philippines may be vulnerable to climate warming. To do so, we selected five anuran species that utilize four breeding habitats and identified the sensitivity and exposure of tadpoles and direct‐developer eggs to heat by measuring their critical thermal maximums (CT_max) and the habitat‐specific temperatures they experience. Our study species included two direct‐developer frogs—one species that lays its eggs on exposed leaves, and another that lays its eggs in ferns—and three species that produce aquatic free‐swimming tadpoles—two stream breeders, and one phytotelm (tree hole) breeder. We compared thermal tolerances derived from microclimates of breeding habitats with tolerances derived from macroclimate (i.e., non‐buffered air temperature taken from the rain forest canopy). We also examined whether differences in CT_max existed across life‐history stages (egg, metamorph/young‐of‐year, and adult) for the two direct‐developer frog species. Habitats buffered ambient temperature and expanded thermal tolerances of all frog species. We found that direct‐developers, however, are more vulnerable to increased temperatures than aquatic breeders—indicated by their high sensitivity to temperature, and exposure to high temperatures. Direct‐developer eggs were more sensitive to warming than both metamorph and adult life‐history stages. Thermally buffered microhabitats may represent the only protection against current and impending climate warming. Our data highlight the importance of considering sensitivity and exposure in unison when deciphering warming vulnerability of frogs.     

Thermal tolerance of tropical and temperate alpine plants suggests that ‘mountain passes are not higher in the tropics’

Aim

Tolerance of species to extreme temperatures largely determines their distribution and vulnerability to climate change. We examined thermal tolerance in tropical and temperate alpine plants, testing the hypotheses that: (a) temperate plants are resistant to more extreme temperatures and have an overall wider thermal tolerance breadth (TTB); (b) TTB in temperate plants is wider than TTB in tropical plants during the entire growing season; (c) resistance to frost and heat varies during the season in temperate plants but not in tropical plants; (d) TTB of a species predicts its latitudinal range.

Location

Tropical (Ecuador, Bolivia) and temperate (USA, Austria) mountains.

Time period

Four periods of the growing season (2014, 2016–2019).

Major taxa

Ninety-six vascular plant species.

Methods

We employed the electrolyte leakage method to estimate the temperature resistance, that is, the temperature at which 50% tissue injury (Lt50) occurs in leaves. We used phylogenetic linear mixed-effect models in a Bayesian framework to test for differences between the plant groups.

Results

Temperate and tropical plants do not differ in their temperature resistance. The four hypotheses are rejected since: (a) temperate plants do not have significantly wider overall TTB compared to tropical plants, (b) TTB of temperate plants is wider than TTB of tropical plants only at the end of the temperate summer, (c) seasonal acclimation is observed in both plant groups, (d) the latitudinal range of the plants is not related to TTB.

Main conclusions

The lack of TTB differences between temperate and tropical alpine plants is consistent with trends observed in ectothermic animals, which suggests a general latitudinal pattern in high-elevation poikilotherm organisms. Limited acclimation capacity to cope with long freezing exposures restricts the occurrence of tropical alpine species to thermally aseasonal environments making them particularly vulnerable to climate change.     

Thermal tolerance ranges and climate variability: A comparison between bivalves from differing climates

The climate variability hypothesis proposes that in variable temperate climates poikilothermic animals have wide thermal tolerance windows, whereas in constant tropical climates they have small thermal tolerance windows. In this study we quantified and compared the upper and lower lethal thermal tolerance limits of numerous bivalve species from a tropical (Roebuck Bay, north western Australia) and a temperate (Wadden Sea, north western Europe) tidal flat. Species from tropical Roebuck Bay had higher upper and lower lethal thermal limits than species from the temperate Wadden Sea, and Wadden Sea species showed an ability to survive freezing temperatures. The increased freezing resistance of the Wadden Sea species resulted in thermal tolerance windows that were on average 7 °C greater than the Roebuck Bay species. Furthermore, at a local-scale, the upper lethal thermal limits of the Wadden Sea species were positively related to submersion time and thus to encountered temperature variation, but this was not the case for the Roebuck Bay species. A review of previous studies, at a global scale, showed that upper lethal thermal limits of tropical species are closer to maximum habitat temperatures than the upper lethal thermal limits of temperate species, suggesting that temperate species are better adapted to temperature variation. In this study, we show for the first time, at both local and global scales, that the lethal thermal limits of bivalves support the climate variability effect in the marine environment.     

Are tropical butterflies more colorful?

There is a common and long-standing belief that tropical butterflies are more striking in their coloration than those of cooler climates. It has been suggested that this is due to more intense biotic selection or mate selection in the tropics. We tested whether there were differences in coloration by examining the dorsal surface color properties of male butterflies from three regions of the western hemisphere: the Jatun-Satcha Reserve in lowland Ecuador (tropical), the state of Florida, USA (subtropical) and the state of Maine, USA (cool temperate). We digitally photographed the dorsal wing and body surface of male butterfly specimens from Maine, Florida, and Ecuador. For each photograph, we analyzed the mean and variation for the color-parameters that are thought to be related to colorfulness; namely Hue, saturation and intensity. Overall, the Ecuadorian sample exhibited more varied intensity, saturation, and Hue compared to the other regions. These results suggest a more complex assemblage of colors and patterns regionally and on a butterfly-by-butterfly basis in the tropics. The greater complexity of colors within each butterfly in our Ecuadorian sample suggests that tropical butterflies are indeed more ‘colorful’, at least by some measures. Possible reasons for this include stronger predation pressure selecting for aposematism, greater species diversity selecting for camouflage or warning coloration against potential predators, and easier recognition of potential mates in a species rich environment.     

Rates of metabolism in diapausing and reproductively active tropical butterflies, Euploea core and Euploea sylvester (Lepidoptera: Nymphalidae)

Abstract.  Although the ecology of diapause has been widely studied in the field, the underlying physiological responses occurring in tropical diapausing insects remain virtually unexplored. This is especially the case with rates of respiration in diapausing tropical insect species. The present study compares rates of metabolism, as assessed by measurement of carbon dioxide production, between two species of diapausing and reproductively active tropical butterflies, Euploea core (W.S Macleay) and Euploea sylvester (Fabricius) , independent of temperature. Measurement of metabolism over a day-time/night-time regime confirms that these tropical butterflies display a diurnal rhythm in accordance with many other tropical and temperate insect species, regardless of developmental state. In addition, diapausing Euploea butterflies display rates of carbon dioxide production only 28% lower than those of reproductively active butterflies, and can terminate diapause within days of receiving the correct cues. Maintaining a similar metabolic rate throughout diapause, as well as a rapid termination of diapause, would enable these tropical butterflies to respond immediately to larval host plant resources, without the disadvantage of missing optimum conditions, allowing the species to maximize their reproductive potential.     

Evidence for a freezing tolerance-growth rate trade-off in the live oaks (Quercus series Virentes) across the tropical-temperate divide

? It has long been hypothesized that species are limited to the north by minimum temperature and to the south by competition, resulting in a trade-off between freezing tolerance and growth rate. We investigated the extent to which the climatic origins of populations from four live oak species (Quercus series Virentes) were associated with freezing tolerance and growth rate, and whether species fitted a model of locally adapted populations, each with narrow climatic tolerances, or of broadly adapted populations with wide climatic tolerances. ? Acorns from populations of four species across a tropical-temperate gradient were grown under common tropical and temperate conditions. Growth rate, seed mass, and leaf and stem freezing traits were compared with source minimum temperatures. ? Maximum growth rates under tropical conditions were negatively correlated with freezing tolerance under temperate conditions. The minimum source temperature predicted the freezing tolerance of populations under temperate conditions. The tropical species Q. oleoides was differentiated from the three temperate species, and variation among species was greater than among populations. ? The trade-off between freezing tolerance and growth rate supports the range limit hypothesis. Limited variation within species indicates that the distributions of species may be driven more strongly by broad climatic factors than by highly local conditions.     

Vulnerability to climate warming and acclimation capacity of tropical and temperate coastal organisms

Ecological forecasting on the likely impacts of climate warming is crucial at a time when several ecosystems seem to be responding to this environmental threat. Among the most important questions are: which are the most vulnerable organisms to climate warming and where are they? Recently, there has been debate on whether the tropics or temperate zones are more vulnerable to warming. Vulnerability toward higher temperatures will depend on the organisms’ thermal limits and also on their acclimation capacity, which remains largely unknown for most species. The aim of the present work was to estimate (1) the upper thermal limits (Critical Thermal Maximum (CTMax)), (2) the warming tolerance (CTMax – Maximum Habitat Temperature) and (3) the acclimation capacity of tropical and temperate rocky shore organisms. Differences in biological groups (decapod crustaceans vs fish) were investigated and the effect of region (tropical vs temperate) and habitat (intertidal vs subtidal) was tested. Overall, 35 species were tested. For the assessment of the acclimation capacity, tropical-temperate pairs of closely related species of shrimp, crab and fish were selected. Warming tolerance was higher for temperate species than for tropical species and higher for subtidal species than for intertidal species, confirming that species with the highest thermal limits have the lowest warming tolerance. All species tested presented some acclimation capacity (CTMax_Trial − CTMax_Control), with the exception of gobiid fish, which was not observed to acclimate. The tropical species tested showed a lower acclimation capacity than their temperate counterparts. Given that tropical rocky shore organisms are already living very close to their thermal limits and that their acclimation capacity is limited, it is likely that the impacts of global warming will be evident sooner in the tropics than in the temperate zone.     

Climate change and the effects of temperature extremes on Australian flying-foxes

Little is known about the effects of temperature extremes on natural systems. This is of increasing concern now that climate models predict dramatic increases in the intensity, duration and frequency of such extremes. Here we examine the effects of temperature extremes on behaviour and demography of vulnerable wild flying-foxes (Pteropus spp.). On 12 January 2002 in New South Wales, Australia, temperatures exceeding 42 degrees C killed over 3500 individuals in nine mixed-species colonies. In one colony, we recorded a predictable sequence of thermoregulatory behaviours (wing-fanning, shade-seeking, panting and saliva-spreading, respectively) and witnessed how 5-6% of bats died from hyperthermia. Mortality was greater among the tropical black flying-fox, Pteropus alecto (10-13%) than the temperate grey-headed flying-fox, Pteropus poliocephalus (less than 1%), and young and adult females were more affected than adult males (young, 23-49%; females, 10-15%; males, less than 3%). Since 1994, over 30000 flying-foxes (including at least 24500 P. poliocephalus) were killed during 19 similar events. Although P. alecto was relatively less affected, it is currently expanding its range into the more variable temperature envelope of P. poliocephalus, which increases the likelihood of die-offs occurring in this species. Temperature extremes are important additional threats to Australian flying-foxes and the ecosystem services they provide, and we recommend close monitoring of colonies where temperatures exceeding 42.0 degrees C are predicted. The effects of temperature extremes on flying-foxes highlight the complex implications of climate change for behaviour, demography and species survival.     

Forecasting the viability of sea turtle eggs in a warming world

Animals living in tropical regions may be at increased risk from climate change because current temperatures at these locations already approach critical physiological thresholds. Relatively small temperature increases could cause animals to exceed these thresholds more often, resulting in substantial fitness costs or even death. Oviparous species could be especially vulnerable because the maximum thermal tolerances of incubating embryos is often lower than adult counterparts, and in many species mothers abandon the eggs after oviposition, rendering them immobile and thus unable to avoid extreme temperatures. As a consequence, the effects of climate change might become evident earlier and be more devastating for hatchling production in the tropics. Loggerhead sea turtles (Caretta caretta) have the widest nesting range of any living reptile, spanning temperate to tropical latitudes in both hemispheres. Currently, loggerhead sea turtle populations in the tropics produce nearly 30% fewer hatchlings per nest than temperate populations. Strong correlations between empirical hatching success and habitat quality allowed global predictions of the spatiotemporal impacts of climate change on this fitness trait. Under climate change, many sea turtle populations nesting in tropical environments are predicted to experience severe reductions in hatchling production, whereas hatching success in many temperate populations could remain unchanged or even increase with rising temperatures. Some populations could show very complex responses to climate change, with higher relative hatchling production as temperatures begin to increase, followed by declines as critical physiological thresholds are exceeded more frequently. Predicting when, where, and how climate change could impact the reproductive output of local populations is crucial for anticipating how a warming world will influence population size, growth, and stability.     

Cold-induced vasodilation and vasoconstriction in the finger of tropical and temperate indigenes

While heat acclimatization reflects the development of heat tolerance, it may weaken an ability to tolerate cold. The purpose of this study was to explore cold-induced vasodilation (CIVD) responses in the finger of tropical indigenes during finger cold immersion, along with temperate indigenes. Thirteen tropical male indigenes (subjects born and raised in the tropics) and 11 temperate male indigenes (subjects born and raised in Japan and China) participated. Subjects immersed their middle finger at 4.3±0.8 °C water for 30 min. Rectal temperature, skin temperatures, finger skin blood flow, blood pressure and subjective sensations were recorded during the test. The results showed that: (1) the tropical group demonstrated a lower minimum (T_min), maximum (T_max) and mean finger temperature (T_mean) compared to those of the temperate group (P<0.05); (2) seven tropical indigenes demonstrated a late-plateau type of CIVD pattern, which is characterized by a pronounced 1st vasoconstriction and a single CIVD with a faint and weak 2nd vasoconstriction, whereas no temperate indigene demonstrated the late-plateau type; and (3) the hand temperature at the end of finger immersion was 3 °C lower in the tropical than the temperate group (P<0.05). These results indicate that tropical indigenes have less active responses of arterio-venous anastomoses in the finger and weaker vasoconstrictions after the first CIVD response during finger cold immersion, which can be considered as being more vulnerable to cold injury of the periphery in severe cold.     

The phytogeography ofCladophora (Chlorophyceae) in the northern Atlantic Ocean,in comparison to that of other benthic algal species

About 43Cladophora species inhabit the coasts of the northern Atlantic Ocean. These can be subdivided into seven distribution groups: (a) the tropical western Atlantic group (16 species); (b) the warm temperate Mediterranean-Atlantic group (9 species); (c) the warm temperate North American group (1 species); (d) the Arctic group (1 species); (e) the amphiatlantic tropical to warm temperature group (7 species); (f) the amphiatlantic tropical to temperate group (4 species), and (g) the amphiatlantic temperate group (5 species). These groups agree with general phytogeographic patterns. Thus, the high numbers of species restricted to the tropical western Atlantic region and the warm temperate Mediteranean-Atlantic region are in agreement with the richness and high degree of endemism of these regions. The fact that all species occurring in northeast America also occur in Europe agrees with the high floristic similarity of the boreal areas in America and Europe. The sediment coasts of the Carolinas are an efficient barrier to the south-north dispersal of benthic algae. The temperature bound phytogeographic limits are set in most cases by the species ability to survive adverse temperatures; for “northern” species to survive a high summer temperature in the south, and for “southern” species to survive a low winter temperature in the north. The limits in the Arctic region are all set by the species ability for sufficient growth and reproduction in summer. Conversely, only few northern species have a southern limit which is set by a winter temperature that is not too high for sufficient growth and reproduction. Most species of this group are winter-annuals at their southern limit, and summer-annuals at their northern limit. A comparatively small number of species with a tropical-to-warm temperate distribution have a northern limit at temperate latitudes which is set by a sufficiently high summer temperature for growth and reproduction. A high proportion of this group are lagoonal or quiet water species, which profit by higher summer temperatures in sheltered waters.C. vagabunda is an example.C. rupestris andC. sericea have an amphiboreal distribution and also occur in the southern temperate belt. They probably used a Pleistocene temperature drop to disperse, through the Atlantic along the African coast, from one hemisphere to the other. In the Pacific temperatures were not sufficiently low for this dispersal; and hence these two species reached the Pacific probably by way of the Bering Strait.     

GROWTH RATES OF BIRDS IN THE HUMID NEW WORLD TROPICS

The growth curves of 40 species of lowland neotropical birds were fitted by logistic equation. The birds were mostly from Panama, Trinidad and Surinam. The growth constants of the fitted equations (asymptote A and growth rate K) were compared within and among species, and with previously published data on temperate species. Growth parameters of tropical passerines are about as variable within species as they are within temperate species. In both cases, variation in A and K between broods is greater than it is within broods. Panamanian birds breed during the dry-wet transition and conditions for growth apparently improve as rainfall increases. Asymptotes of growth curves are higher, and mortality within broods lower, as the breeding season progresses. Asynchronous hatching and the reduction of brood-size by selective starvation of young is a prominent phenomenon during the early part of the breeding season. Several instances are reported, however, of young persisting in nests with inadequate feeding and greatly subnormal weights. Slowed development under conditions of poor nutrition may be adaptive in the tropics if periods of low food availability are short and allow the possibility of recovery from undernourishment. As a group, neotropical lowland passerines (30 species) grow 23% more slowly than a sample of 51 temperate passerines. Variation of growth rates among these tropical species is similar to variation among temperate species, and it is related to adult body-size the length of the nestling period. Young of tropical and temperate species attain similar asymptotes, relative to adult body-weight, by the end of the nestling period. Hypotheses are advanced which might explain the slower growth rate of tropical species, and tested to the extent available data permits. (1) Because brood-size can be changed only by adding or removing whole young, changes in growth rate could provide finer adjustment of the energy requirements of the young to the feeding capacity of the parents. This model predicts different means and variances for growth rate within groups of species with different clutch-sizes, predictions not supported by available data. (2) Growth rate is shown to increase the maximum energy requirement of a nestling only if K exceeds some value determined by the energy requirement of the young, growth rate should vary in proportion to the level of basal maintenance metabolism. In a small sample of tropical species, rates of basal metabolism were 25% lower than in a comparable sample of temperate species. These data therefore support the hypothesis, although the cause of the lower metabolic rate of the tropical nestlings is not known. (3) Daily periods of hypothermia could reduce the energy requirement of the young and at the same time reduce their growth rate; but observations of body temperatures of tropical nestlings are contrary to this hypothesis. (4) The short day-length of tropical climates reduces the time during which young can assimilate energy relative to their energy expenditures. This model predicts that tropical nestlings would have less productive energy available, (consistent with their reduced growth rates), but it also predicts that arctic birds should grow faster than temperate species, which is not confirmed by available data. (5) The low nitrogen content of fruits may cause the slow growth of a few strictly frugivorous species (Oilbird and Bearded Bellbird), but among other tropical species growth rate is not correlated with the estimated proportion of fruit in the diet.