Are mountain passes higher in the tropics? Janzen's hypothesis revisited.

Synopsis In 1967 Daniel Janzen published an influential paper titled "Why Mountain Passes Are Higher in the Tropics." Janzen derived a simple climatic-physiological model predicting that tropical mountain passes would be more effective barriers to organismal dispersal than would temperate-zone passes of equivalent altitude. This prediction derived from a recognition that the annual variation in ambient temperature at any site is relatively low in the tropics. Such low variation within sites not only reduces the seasonal overlap in thermal regimes between low- and high-altitude sites, but should also select for organisms with narrow physiological tolerances to temperature. As a result, Janzen predicted that tropical lowland organisms are more likely to encounter a mountain pass as a physiological barrier to dispersal (hence "higher"), which should in turn favor smaller distributions and an increase in species turnover along altitudinal gradients. This synthetic hypothesis has long been at the center of discussions of latitudinal patterns of physiological adaptation and of species diversity. Here we review some of the key assumptions and predictions of Janzen's hypothesis. We find general support for many assumptions and predictions, but call attention to several issues that somewhat ameliorate the generality of Janzen's classic hypothesis.

[1]  D. Janzen Why Mountain Passes are Higher in the Tropics , 1967, The American Naturalist.

[2]  G. Gilchrist Specialists and Generalists in Changing Environments. I. Fitness Landscapes of Thermal Sensitivity , 1995, The American Naturalist.

[3]  Raymond B Huey,et al.  Behavioral Drive versus Behavioral Inertia in Evolution: A Null Model Approach , 2003, The American Naturalist.

[4]  Joseph H. Connell,et al.  On the Prevalence and Relative Importance of Interspecific Competition: Evidence from Field Experiments , 1983, The American Naturalist.

[5]  Dawn M. Kaufman,et al.  LATITUDINAL GRADIENTS OF BIODIVERSITY:Pattern,Process,Scale,and Synthesis , 2003 .

[6]  Theodore Garland,et al.  Physiological Differentiation of Vertebrate Populations , 1991 .

[7]  Peter Kareiva,et al.  Biotic interactions and global change. , 1993 .

[8]  M. Bertness Predation, Physical Stress, and the Organization of a Tropical Rocky Intertidal Hermit Crab Community , 1981 .

[9]  R. Huey Latitudinal Pattern of Between-Altitude Faunal Similarity: Mountains Might be "Higher" in the Tropics , 1978, The American Naturalist.

[10]  W. Heyer A Herpetofaunal Study of an Ecological Transect Through the Cordillera de Tilarán, Costa Rica@@@A Herpetofaunal Study of an Ecological Transect Through the Cordillera de Tilaran, Costa Rica , 1967 .

[11]  W. Porter New Animal Models and Experiments for Calculating Growth Potential at Different Elevations , 1989, Physiological Zoology.

[12]  Dawn M. Kaufman,et al.  THE GEOGRAPHIC RANGE: Size, Shape, Boundaries, and Internal Structure , 1996 .

[13]  E. Pianka Latitudinal Gradients in Species Diversity: A Review of Concepts , 1966, The American Naturalist.

[14]  D. Bradford,et al.  Thermoregulation of Lizards and Toads at High Altitudes in Peru , 1976 .

[15]  R. Dekker,et al.  Predation and the western limits of megapode distribution , 1989 .

[16]  B. Brattstrom,et al.  Thermal acclimation in anuran amphibians as a function of latitude and altitude. , 1968, Comparative biochemistry and physiology.

[17]  D. M. Gates,et al.  THERMODYNAMIC EQUILIBRIA OF ANIMALS WITH ENVIRONMENT , 1969 .

[18]  D. H. Morse Niche Breadth as a Function of Social Dominance , 1974, The American Naturalist.

[19]  R. V. Bovbjerg Ecological Isolation and Competitive Exclusion in Two Crayfish (Orconectes Virilis and Orconectes Immunis) , 1970 .

[20]  H. Heatwole,et al.  Some Aspects of the Thermal Ecology of Puerto Rican Anoline Lizards , 1969 .

[21]  F. Allendorf,et al.  Blackwell Publishing, Ltd. , 2003 .

[22]  Kevin J. Gaston,et al.  Macrophysiology: large‐scale patterns in physiological traits and their ecological implications , 2004 .

[23]  Andrew R. Cossins,et al.  Temperature Biology of Animals. , 1989 .

[24]  J. Roughgarden,et al.  Effect of species interactions on landscape abundance patterns , 2005 .

[25]  G. Walsberg,et al.  Physiological Diversity and Its Ecological Implications , 2001 .

[26]  J. Connell The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus , 1961 .

[27]  G. S. Bakken Measurement and Application of Operative and Standard Operative Temperatures in Ecology , 1992 .

[28]  Anthony R. Ives,et al.  An Introduction to Phylogenetically Based Statistical Methods, with a New Method for Confidence Intervals on Ancestral Values , 1999 .

[29]  P. Jones,et al.  REPRESENTING TWENTIETH CENTURY SPACE-TIME CLIMATE VARIABILITY. , 1998 .

[30]  M. Feder Environmental variability and thermal acclimation of metabolism in tropical anurans , 1982 .

[31]  D. Schemske,et al.  THE EVOLUTION OF SPECIES' DISTRIBUTIONS: RECIPROCAL TRANSPLANTS ACROSS THE ELEVATION RANGES OF MIMULUS CARDINALIS AND M. LEWISII , 2005, Evolution; international journal of organic evolution.

[32]  P. E. Hertz Adaptation to altitude in two West Indian anoles (Reptilia: Iguanidae): Field thermal biology and ph , 2009 .

[33]  F. H. Berkum EVOLUTIONARY PATTERNS OF THE THERMAL SENSITIVITY OF SPRINT SPEED IN ANOLIS LIZARDS , 1986 .

[34]  A. Wallace Tropical Nature and Other Essays , 1972 .

[35]  T. Dobzhansky,et al.  Evolution in the tropics , 1950 .

[36]  A. Shapiro,et al.  Geological barriers and restricted gene flow in the holarctic skipper Hesperia comma (Hesperiidae) , 2004, Molecular ecology.

[37]  Richard Field,et al.  ENERGY, WATER, AND BROAD‐SCALE GEOGRAPHIC PATTERNS OF SPECIES RICHNESS , 2003 .

[38]  J. W. Patterson Thermal Acclimation in Two Subspecies of the Tropical Lizard Mabuya striata , 1984, Physiological Zoology.

[39]  C. Rahbek The Relationship Among Area, Elevation, And Regional Species Richness In Neotropical Birds , 1997, The American Naturalist.

[40]  R. Huey,et al.  Cost and Benefits of Lizard Thermoregulation , 1976, The Quarterly Review of Biology.

[41]  Kevin J. Gaston,et al.  Elevation and climatic tolerance : a test using dung beetles , 1999 .

[42]  V. Markgraf,et al.  High altitude tropical biogeography , 1988 .

[43]  M. McPeek,et al.  The community context of species' borders: ecological and evolutionary perspectives , 2005 .

[44]  J. Terborgh Bird Species Diversity on an Andean Elevational Gradient , 1977 .

[45]  D. Lieberman,et al.  TROPICAL FOREST STRUCTURE AND COMPOSITION ON A LARGE-SCALE ALTITUDINAL GRADIENT IN COSTA RICA , 1996 .

[46]  Fredrica H. van Berkum,et al.  LATITUDINAL PATTERNS OF THE THERMAL SENSITIVITY OF SPRINT SPEED IN LIZARDS , 1988 .

[47]  R. Jaeger,et al.  COMPETITION LEADS TO AN EXTINCTION‐PRONE SPECIES OF SALAMANDER: INTERSPECIFIC TERRITORIALITY IN A METAPOPULATION , 1998 .

[48]  M. Chappell Behavioral Factors in the Altitudinal Zonation of Chipmunks (Eutamias) , 1978 .

[49]  A. D. Bradshaw,et al.  Evolutionary Significance of Phenotypic Plasticity in Plants , 1965 .

[50]  R. Andrews,et al.  Geographic variation in field body temperature of sceloporus lizards , 1998 .

[51]  R. Huey,et al.  Physiological Consequences of Habitat Selection , 1991, The American Naturalist.

[52]  David F West,et al.  Breeding structure of three snow pool Aedes mosquito species in northern Colorado , 1998, Heredity.

[53]  S. Herzog,et al.  The elevational gradient in Andean bird species richness at the local scale: a foothill peak and a high‐elevation plateau , 2005 .

[54]  J. Freyhof,et al.  The Alps as barrier to dispersal in cold-adapted freshwater fishes? Phylogeographic history and taxonomic status of the bullhead in the Adriatic freshwater drainage. , 2004, Molecular phylogenetics and evolution.

[55]  G. Thomas,et al.  Genetic variation and population structure inOryza malampuzhaensis Krish.et Chand. endemic to Western Ghats, South India , 2001, Journal of Genetics.

[56]  Arthur E. Dunham,et al.  Interfaces between Biophysical and Physiological Ecology and the Population Ecology of Terrestrial Vertebrate Ectotherms , 1989, Physiological Zoology.

[57]  R. Huey,et al.  Plants Versus Animals: Do They Deal with Stress in Different Ways?1 , 2002, Integrative and comparative biology.

[58]  R. Huey,et al.  Compensation for Altitudinal Changes in the Thermal Environment by Some Anolis Lizards on Hispaniola , 1981 .

[59]  R. Dudley,et al.  Into thin air: Physiology and evolution of alpine insects. , 2006, Integrative and comparative biology.

[60]  R. Jaeger Competitive Exclusion as a Factor Influencing the Distributions of Two Species of Terrestrial Salamanders. , 1971, Ecology.

[61]  Roy Haines-Young,et al.  Biogeography , 1992, Vegetatio.

[62]  R. Jaeger Moisture as a factor influencing the distributions of two species of terrestrial salamanders , 1971, Oecologia.

[63]  M. Taper,et al.  Interspecific Competition, Environmental Gradients, Gene Flow, and the Coevolution of Species' Borders , 2000, The American Naturalist.

[64]  D. Sc. Prof. Dr. M. S. Mani M. A.,et al.  Ecology and Biogeography of High Altitude Insects , 1968, Series Entomologica.

[65]  R. Brandl,et al.  Ecological and genetic spatial patterns of Urophora cardui (Diptera: Tephritidae) as evidence for population structure and biogeographical processes , 1994 .

[66]  J. Felsenstein Phylogenies and the Comparative Method , 1985, The American Naturalist.

[67]  Kevin J. Gaston,et al.  Thermal tolerance, climatic variability and latitude , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[68]  J. Terborgh,et al.  Interspecific aggression and habitat selection by Amazonian birds , 1995 .

[69]  K. Gaston,et al.  Hemispheric Asymmetries in Biodiversity—A Serious Matter for Ecology , 2004, PLoS biology.

[70]  P. Jones,et al.  Representing Twentieth-Century Space–Time Climate Variability. Part I: Development of a 1961–90 Mean Monthly Terrestrial Climatology , 1999 .

[71]  J. David,et al.  Chromosomal inversion polymorphism in Afrotropical populations of Drosophila melanogaster. , 2002, Genetical research.

[72]  J. Lawton,et al.  Making mistakes when predicting shifts in species range in response to global warming , 1998, Nature.

[73]  M. Feder,et al.  EFFECTS OF LATITUDE, SEASON, ELEVATION, AND MICROHABITAT ON FIELD BODY TEMPERATURES OF NEOTROPICAL AND TEMPERATE ZONE SALAMANDERS' , 1982 .

[74]  T. Garland,et al.  Evolutionary Physiology , 1926, Nature.

[75]  Robert D. Stevenson,et al.  The Relative Importance of Behavioral and Physiological Adjustments Controlling Body Temperature in Terrestrial Ectotherms , 1985, The American Naturalist.

[76]  M. Bertness Competitive Dynamics of a Tropical Hermit Crab Assemblage , 1981 .

[77]  C. Navas,et al.  Patterns of distribution of anurans in high Andean tropical elevations: Insights from integrating biogeography and evolutionary physiology. , 2006, Integrative and comparative biology.

[78]  O. Reichman,et al.  Physiology on a Landscape Scale: Plant-Animal Interactions1 , 2002, Integrative and comparative biology.

[79]  C. Navas Herpetological diversity along Andean elevational gradients: links with physiological ecology and evolutionary physiology. , 2002, Comparative biochemistry and physiology. Part A, Molecular & integrative physiology.

[80]  J. Lawton,et al.  Individualistic species responses invalidate simple physiological models of community dynamics under global environmental change , 1998 .

[81]  R. Briers Range limits and parasite prevalence in a freshwater snail , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[82]  Joyce S. Tsuji,et al.  Thermal Acclimation of Metabolism in Sceloporus Lizards from Different Latitudes , 1988, Physiological Zoology.

[83]  J. H. Carothers,et al.  Altitudinal zonation among lizards of the genus Liolaemus: questions answered and unanswered questions , 2001 .

[84]  R. Verheyen,et al.  Altitudinal variation of the thermal biology and running performance in the lizard Podarcis tiliguerta , 1989, Oecologia.

[85]  G. Rogowitz Evaluation of Thermal Acclimation and Altitudinal Variation of Metabolism in a Neotropical Lizard, Anolis gundlachi , 1996 .

[86]  H. G. Andrewartha,et al.  The distribution and abundance of animals. , 1954 .

[87]  Paul R. Martin,et al.  ECOLOGICAL AND FITNESS CONSEQUENCES OF SPECIES COEXISTENCE: A REMOVAL EXPERIMENT WITH WOOD WARBLERS , 2001 .

[88]  Kevin J. Gaston,et al.  The structure and dynamics of geographic ranges , 2003 .

[89]  C. Navas Metabolic Physiology, Locomotor Performance, and Thermal Niche Breadth in Neotropical Anurans , 1996, Physiological Zoology.

[90]  G. Graves,et al.  Multiscale assessment of patterns of avian species richness , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[91]  Gregory K. Snyder,et al.  Temperature Adaptations in Amphibians , 1975, The American Naturalist.

[92]  Robert D. Holt,et al.  On the evolutionary ecology of species' ranges , 2003 .

[93]  H. Lüddecke,et al.  Are Tropical Highland Frog Calls Cold-adapted? The Case of the Andean Frog Hyla labialis1 , 2002 .

[94]  L. Rieseberg,et al.  Gene flow between cultivated and wild sunflowers , 1994, Theoretical and Applied Genetics.

[95]  E. Pianka Latitudinal gradients in species diversity , 1989 .

[96]  M. Feder Environmental Variability and Thermal Acclimation in Neotropical and Temperate Zone Salamanders , 1978, Physiological Zoology.

[97]  J. H. Carothers,et al.  Thermal characteristics of ten Andean lizards of the genus Liolaemus in central Chile , 1997 .

[98]  R. Huey,et al.  Thermal Biology of Anolis Lizards in a Complex Fauna: The Christatellus Group on Puerto Rico , 1976 .

[99]  R. Ruibal,et al.  Eurythermy and Niche Expansion in Lizards , 1970 .

[100]  W. Heyer A Herpetofaunal Study of an Ecological Transect Through the Cordillera de Tilaran, Costa Rica1 , 1967 .