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Biodiversity and metacommunity structure of animals along altitudinal gradients in tropical montane forests

  • Michael R. Willig (a1) and Steven J. Presley (a1)

The study of altitudinal gradients has made enduring contributions to the theoretical and empirical bases of modern biology. Unfortunately, the persistence of these systems and the species that compose them is threatened by land-use change at lower altitudes and by climate change throughout the gradients, but especially at higher altitudes. In this review, we focus on two broad themes that are inspired by altitudinal variation in tropical montane regions: (1) dimensions of biodiversity and (2) metacommunity structure. Species richness generally decreased with increasing altitude, although not always in a linear fashion. Mid-altitudinal peaks in richness were less common than monotonic declines, and altitudinal increases in richness were restricted to amphibian faunas. Moreover, gradients of biodiversity differed among dimensions (taxonomic, phylogenetic and functional) as well as among faunas (bats, rodents, birds) in the tropical Andes, suggesting that species richness is not a good surrogate for dimensions that reflect differences in the function or evolutionary history of species. Tropical montane metacommunities evinced a variety of structures, including nested (bats), Clementsian (rodents, bats, gastropods), quasi-Clementsian (reptiles, amphibians, passerines) and quasi-Gleasonian (gastropods) patterns. Nonetheless, compositional changes were always associated with the ecotones between rain forest and cloud forest, regardless of fauna.

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S. J. ANDELMAN & M. R. WILLIG 2003. Present patterns and future prospects for biodiversity in the western hemisphere. Ecology Letters 6:818824.

R. P. ANDERSON , E. E. GUTIÉRREZ , G.J. OCHOA , F. J. GARCÍA & M. AGUILERA 2012. Faunal nestedness and species–area relationship for small non-volant mammals in “sky islands” of northern Venezuela. Studies on Neotropical Fauna and Environment 47:157170.

J. A. BARONE , J. THOMLINSON , P. ANGLADA-CORDERO & J. K. ZIMMERMAN 2008. Metacommunity structure of tropical forests along an elevational gradient in Puerto Rico. Journal of Tropical Ecology 24:110.

W. BARTHLOTT , W. LAUER & A. PLACKE 1996. Global distribution of species diversity in vascular plants: towards a world map of phytodiversity. Erdkunde 50:317327.

A. BECKER , C. KÖRNER , J.-J. BRUN , A. GUISAN & U. TAPPEINER 2007. Ecological and landuse studies along elevational gradients. Mountain Research and Development 27:5865.

J. G. BLAKE & B. A. LOISELLE 2000. Diversity of birds along an elevational gradient in the Cordillera Central, Costa Rica. The Auk 117:663686.

G. BREHM , D. SÜSSENBACH & K. FIEDLER 2003. Unique elevational diversity patterns of geometrid moths in an Andean montane rainforest. Ecography 26:456466.

J. H. BROWN 1971. Mammals on mountaintops: nonequilibrium insular biogeography. American Naturalist 105:467478.

C. A. BRÜHL , M. MOHAMED & K. E. LINSENMAIR 1999. Altitudinal distribution of leaf litter ants along a transect in primary forests on Mount Kinabalu, Sabah, Malaysia. Journal of Tropical Ecology 15:265277.

M. W. CADOTTE , J. CAVENDER-BARES , D. TILMAN & T. H. OAKLEY 2009. Using phylogenetic, functional and trait diversity to understand patterns of plant community productivity. PLoS ONE 4:e5695.

J. CAVENDER-BARES , K. H. KOZAK , P. V. A. FINE & S. W. KEMBEL 2009. The merging of community ecology and phylogenetic biology. Ecology Letters 12:693715.

L. M. CISNEROS , K. R. BURGIO , L. M. DREISS , B. T. KLINGBEIL , B. D. PATTERSON , S. J. PRESLEY & M. R. WILLIG 2014. Multiple dimensions of bat biodiversity along an extensive tropical elevational gradient. Journal of Animal Ecology 83:11241136.

B. D. COLEMAN , M. A. MARES , M. R. WILLIG & Y. H. HSIEH 1982. Randomness, area and species richness. Ecology 63:11211133.

J. H. CONNELL 1980. Diversity and the coevolution of competitors, or the ghost of competition past. Oikos 35:131138.

N. J. CORDEIRO 1998. Preliminary analysis of the nestedness patterns of montane forest birds of the Eastern Arc Mountains. Journal of East African Natural History 87:101118.

A. W. DIAMOND & A. C. HAMILTON 1980. The distribution of forest passerine birds and Quaternary climatic change in Africa. Journal of Zoology 191:379402.

S. DÍAZ & M. CABIDO 2001. Vive la différence: plant functional diversity matters to ecosystem process. Trends in Ecology and Evolution 16:646655.

L. M. DREISS , K. R. BURGIO , L. M. CISNEROS , B. T. KLINGBEIL , B. D. PATTERSON , S. J. PRESLEY & M. R. WILLIG 2015. Taxonomic, functional, and phylogenetic dimensions of rodent biodiversity along an extensive tropical elevational gradient. Ecography 38:876888.

L. FAHRIG 2003. Effects of habitat fragmentation on biodiversity. Annual Review of Ecology, Evolution & Systematics 34:487515.

D. P. FAITH 1992. Conservation evaluation and phylogenetic diversity. Biological Conservation 61:110.

J. E. FAUTH , B. I. CROTHER & J. B. SLOWINSKI 1989. Elevational patterns of species richness, evenness, and abundance of the Costa Rican leaf-litter herpetofauna. Biotropica 21:178185.

G. W. FERNANDES & A. C. F. LARA 1993. Diversity of Indonesian gall-forming herbivores along altitudinal gradients. Biodiversity Letters 1:186192.

J. C. T. GARTEN , W. M. POST , P. J. HANSON & L. W. COOPER 1999. Forest soil carbon inventories and dynamics along an elevational gradient in the southern Appalachian Mountains. Biogeochemistry 45:115145.

A. A. GIARETTA , K. G. FACURE , R. J. SAWAYA , J. H. DE M. MEYER & N. CHEMIN 1999. Diversity and abundance of litter frogs in a montane forest of southeastern Brazil: seasonal and altitudinal changes. Biotropica 31:669674.

H. A. GLEASON 1926. The individualistic concept of the plant association. Bulletin of the Torrey Botanical Club 53:726.

G. GONZÁLEZ , E. GARCIA , V. CRUZ , S. BORGES , M. ZALAMEA & M. M. RIVERA 2007. Earthworm communities along an elevation gradient in northeastern Puerto Rico. European Journal of Soil Biology 43:S24S32.

G. L. GRAHAM 1983. Changes in bat species diversity along an elevational gradient up the Peruvian Andes. Journal of Mammalogy 64:559571.

G. L. GRAHAM 1990. Bats versus birds: comparisons among Peruvian volant vertebrate faunas along an elevational gradient. Journal of Biogeography 17:657668.

P. J. GRUBB 1977. Control of forest growth and distribution on wet tropical mountains: with special reference to mineral nutrition. Annual Review of Ecology and Systematics 8:83107.

J.-A. GRYTNES & C M. MCCAIN 2007. Elevational trends in biodiversity. Pp. 18 in S.A. Levin (ed.). Encyclopedia of biodiversity, Academic Press, Waltham.

L. R. HEANEY 2001. Small mammal diversity along elevational gradients in the Philippines: an assessment of patterns and hypotheses. Global Ecology and Biogeography 10:1539.

A. HEMP 2006. Continuum or zonation? Altitudinal gradients in the forest vegetation of Mt. Kilimanjaro. Plant Ecology 184:2742.

P. HERNANDEZ-ALCANTARA , D. A. SALAS-DE LEON , V. SOLIS-WEISS & M. A. MONREAL-GOMEZ 2014. Bathymetric patterns of polychaete (Annelida) species richness in the continental shelf of the Gulf of California, Eastern Pacific. Journal of Sea Research 91:7987.

H. HILLEBRAND 2004. On the generality of the latitudinal diversity gradient. American Naturalist 163:192211.

U. HOFER , L-F. BERSIER & D. BORCARD 1999. Spatial organization of a herpetofauna on an elevational gradient revealed by null model tests. Ecology 80:976988.

U. HOFER , L-F. BERSIER & D. BORCARD 2000. Ecotones and gradient as determinants of hepetofaunal community structure in the primary forest of Mount Kupe, Cameroon. Journal of Tropical Ecology 16:517533.

M. A. HUSTON 1997. Hidden treatments in ecological experiments: re-evaluating the ecosystem function of biodiversity. Oecologia 110:449460.

J. E. JANKOWSKI , A. L. CIECKA , N. Y. MEYER & K. N. RABENOLD 2009. Beta diversity along environmental gradients: implications of habitat specialization in tropical montane landscapes. Journal of Animal Ecology 78:315327.

D. H. JANZEN 1967. Why mountain passes are higher in the tropics. American Naturalist 101:233249.

M. KESSLER 2000. Elevational gradients in species richness and endemism of selected plant groups in the central Bolivian Andes. Plant Ecology 149:181193.

K. KITAYAMA 1992. An altitudinal transect study of the vegetation on Mount Kinabalu, Borneo. Vegetatio 102:149171.

M. A. LEIBOLD & G. M. MIKKELSON 2002. Coherence, species turnover, and boundary clumping: elements of meta-community structure. Oikos 97:237250.

M. A. LEIBOLD , M. HOLYOAK , N. MOUQUET , P. AMARASEKARE , J. M. CHASE , M. F. HOOPES , R. D. HOLT , J. B. SHURIN , R. LAW , D. TILMAN , M. LOREAU & A. GONZALEZ 2004. The metacommunity concept: a framework for multi-scale community ecology. Ecology Letters 7:601613.

D. LIEBERMAN , M. LIEBERMAN , R. PERALTA & G. S. HARTSHORN 1996. Tropical forest structure and composition on a large-scale altitudinal gradient in Costa Rica. Journal of Ecology 84:137152.

T.-S. LIEW , M. SCHILTHUIZEN & M. BIN LAKIM 2010. The determinants of land snail diversity along a tropical elevational gradient: insularity, geometry and niches. Journal of Biogeography 37:10711078.

C. LÓPEZ-GONZÁLEZ , S. J. PRESLEY , A. LOZANO , R. D. STEVENS & C. L. HIGGINS 2012. Metacommunity analysis of Mexican bats: environmentally mediated structure in an area of high geographic and environmental complexity. Journal of Biogeography 39:177192.

R. H. MACARTHUR & E. O. WILSON 1963. An equilibrium theory of insular zoogeography. Evolution 17:373387.

P. H. MARTIN , R. E. SHERMAN & T. J. FAHEY 2007. Tropical montane forest ecotones: climate gradients, natural disturbance, and vegetation zonation in the Cordillera Central, Dominican Republic. Journal of Biogeography 34:17921806.

M. A. MARTÍNEZ-MORALES 2005. Nested species assemblages as a tool to detect sensitivity to forest fragmentation: the case of cloud forest birds. Oikos 108:634642.

N. W. H. MASON , C. LANOISELÉE , D. MOUILLOT & C. ARGILLIER 2007. Functional characters combined with null models reveal inconsistency in mechanisms of species turnover in lacustrine fish communities. Oecologia 153:441452.

C. M. MCCAIN 2005. Elevational gradients in diversity of small mammals. Ecology 86:366372.

C. M. MCCAIN 2009. Global analysis of bird elevational diversity. Global Ecology and Biogeography 18:346360.

C. M. MCCAIN & R. K. COLWELL 2011. Assessing the threat to montane biodiversity from discordant shifts in temperature and precipitation in a changing climate. Ecology Letters 14:12361245.

M. MEYBECK , P. GREEN & C. VÖRÖSMARTY 2001. A new typology for mountains and other relief classes. An application to global continental water resources and population distribution. Mountain Research and Development 21:3445.

G. G. MITTELBACH , C. F. STEINER , S. M. SCHEINER , K. L. GROSS , H. L. REYNOLDS , R. B. WAIDE , M. R. WILLIG , S. I. DODSON & L. GOUGH 2001. What is the observed relationship between species richness and productivity? Ecology 82:23812396.

C. MORITZ , J. L. PATTON , C. J. CONROY , J. L. PARRA , G. C. WHITE & S. R. BEISSINGER 2008. Impact of a century of climate change on small-mammal communities in Yosemite National Park, USA. Science 322:261264.

R. NANIWADEKAR & K. VASUDEVAN 2007. Patterns in diversity of anurans along an elevational gradient in the Western Ghats, South India. Journal of Biogeography 34:842853.

S. M. D. NOR 2001. Elevational diversity patterns of small mammals on Mount Kinabalu, Sabah, Malaysia. Global Ecology and Biogeography 10:4162.

O. F. PALIN , P. EGGLETON , Y. MALHI , C. A. J. GIRARDIN , A. ROZAS-DÁVILA & C. L. PARR 2011. Termite diversity along an Amazon–Andes elevation gradient, Peru. Biotropica 43:100107.

B. D. PATTERSON 1982. Pleistocene vicariance, montane islands, and the evolutionary divergence of some chipmunks (genus Eutamias). Journal of Mammalogy 63:387398.

B. D. PATTERSON & A. ATMAR 1986. Nested subsets and the structure of insular mammalian faunas and archipelagos. Biological Journal of the Linnean Society 28:6582.

B. D. PATTERSON , D. F. STOTZ , S. SOLARI & J. W. FITZPATRICK 1998. Contrasting patterns of elevational zonation for birds and mammals in the Andes of southeastern Peru. Journal of Biogeography 25:593607.

S. PAVOINE & M. B. BONSALL 2011. Measuring biodiversity to explain community assembly: a unified approach. Biological Reviews 86:792812.

O. L. PETCHEY & K. J. GASTON 2006. Functional diversity: back to basics and looking forward. Ecology Letters 9:741758.

O. L. PETCHEY , K. L. EVANS , I. S. FISHBURN & K. J. GASTON 2007. Low functional diversity and no redundancy in British avian assemblages. Journal of Animal Ecology 76:977985.

E. PINEDA & G. HALFFTER 2004. Species diversity and habitat fragmentation: frogs in a tropical montane landscape in Mexico. Biological Conservation 117:499508.

S. J. PRESLEY , C. L. HIGGINS , C. LÓPEZ-GONZÁLEZ & R. D. STEVENS 2009. Elements of metacommunity structure of Paraguayan bats: multiple gradients require analysis of multiple ordination axes. Oecologia 160:781793.

S. J. PRESLEY , C. L. HIGGINS & M. R. WILLIG 2010. A comprehensive framework for the evaluation of metacommunity structure. Oikos 119:908917.

S. J. PRESLEY , M. R. WILLIG , C. P. BLOCH , I. CASTRO-ARELLANO , C. L. HIGGINS & B. T. KLINGBEIL 2011. A complex metacommunity structure for gastropods along an elevational gradient. Biotropica 43:480488.

S. J. PRESLEY , L. M. CISNEROS , B. D. PATTERSON & M. R. WILLIG 2012. Vertebrate metacommunity structure along an extensive elevational gradient in the tropics: a comparison of bats, rodents and birds. Global Ecology and Biogeography 21:968976.

C. RAHBEK 1995. The elevational gradient of species richness: a uniform pattern? Ecography 18:200205.

C. RAHBEK 1997. The relationship among area, elevation, and regional species richness in Neotropical birds. American Naturalist 149:875902.

M. A. REX 1981. Community structure in the deep-sea benthos. Annual Review of Ecology and Systematics 12:331353.

B. A. RICHARDSON , M. J. RICHARDSON & F. N. SOTO-ADAMES 2005. Separating the effects of forest type and elevation on the diversity of litter invertebrate communities in a humid tropical forest in Puerto Rico. Journal of Animal Ecology 74:926936.

M. L. ROSENZWEIG & E. A. SANDLIN 1997. Species diversity and latitude: listening to area's signal. Oikos 80:172176.

R. J. ROWE 2007. Legacies of land use and recent climatic change: the small mammal fauna in the mountains of Utah. American Naturalist 170:242257.

K. SAFI , M. V. CIANCIARUSO , R. D. LOYOLA , D. BRITO , K. ARMOUR-MARSHALL & J. A. F. DINIZ-FILHO 2011. Understanding global patterns of mammalian functional and phylogenetic diversity. Philosophical Transactions of the Royal Society London. Series B. Biological Science 366:25362544.

V. SÁNCHEZ-CORDERO 2001. Elevation gradients of diversity for rodents and bats in Oaxaca, Mexico. Global Ecology and Biogeography 10:6376.

S. M. SCHEINER & M. R. WILLIG 2005. Developing unified theories in ecology as exemplified with diversity gradients. American Naturalist 166:458469.

S. M. SCHEINER , A. CHIARUCCI , G. A. FOX , M. R. HELMUS , D. J. MCGLINN & M. R. WILLIG 2011. The underpinnings of the relationship of species richness with space and time. Ecological Monographs 81:195213.

B. B. SIMPSON 1974. Glacial migrations of plants: island biogeographic evidence. Science 185:698700.

S. A. SMITH , A. N. MONTES DE OCA , T. W. REEDER & J. J. WIENS 2007. A phylogenetic perspective on elevational species richness patterns in Middle American treefrogs: why so few species in lowland tropical rainforests? Evolution 61:11881207.

M. J. SPASOJEVIC & K. N. SUDING 2012. Inferring community assembly mechanisms from functional diversity patterns: the importance of multiple assembly processes. Journal of Ecology 100:652661.

D. S. SRIVASTAVA & J. H. LAWTON 1998. Why more productive sites have more species: an experimental test of theory using tree-hole communities. American Naturalist 152:510529.

R. D. STEVENS & M. R. WILLIG 2002. Geographical ecology at the community level: perspectives on the diversity of New World bats. Ecology 83:545560.

R. D. STEVENS , S. B. COX , R. E. STRAUSS & M. R. WILLIG 2003. Patterns of functional diversity across an extensive environmental gradient: vertebrate consumers, hidden treatments and latitudinal trends. Ecology Letters 6:10991108.

R. D. STEVENS , M. R. WILLIG & R. E. STRAUSS 2006. Latitudinal gradients in the phenetic diversity of New World bat communities. Oikos 112:4150.

R. D. STEVENS , M. M. GAVILANEZ , J. S. TELLO & D. A. RAY 2012. Phylogenetic structure illuminated the mechanistic role of environmental heterogeneity in community organization. Journal of Animal Ecology 81:455462.

N. G. SWENSON & B. J. ENQUIST 2009. Opposing assembly mechanisms in a Neotropical dry forest: implications for phylogenetic and functional community ecology. Ecology 90:21612170.

J. TERBORGH 1971. Distribution on environmental gradients: theory and a preliminary interpretation of distributional patterns in the avifauna of the Cordillera Vilcabamba, Peru. Ecology 52:2340.

J. TERBORGH 1977. Bird species diversity on an Andean elevational gradient. Ecology 58:10071019.

J. TERBORGH 1985. The role of ecotones in the distribution of Andean birds. Ecology 66:12371246.

D. TILMAN , J. KNOPS , D. WEDIN , P. REICH , M. RITCHIE & E. SIEMANN 1997. The influence of functional diversity and composition on ecosystem processes. Science 277:13001302.

M. G. TURNER 1989. Landscape ecology: the effect of pattern on process. Annual Review of Ecology and Systematics 20:171197.

H. D. VANCE-CHALCRAFT , M. R. WILLIG , S. B. COX , A. E. LUGO & F. N. SCATENA 2010. Relationship between aboveground biomass and multiple measures of biodiversity in subtropical forest of Puerto Rico. Biotropica 42:290299.

R. B. WAIDE , M. R. WILLIG , C. F. STEINER , G. MITTELBACH , L. GOUGH , S. I. DODSON , G. P. JUDAY & R. PARMENTER 1999. The relationship between productivity and species richness. Annual Review of Ecology and Systematics 30:257300.

D. M. WATSON 2003. Long-term consequences of habitat fragmentation – highland birds in Oaxaca, Mexico. Biological Conservation 111:282303.

D. M. WATSON & A. T. PETERSON 1999. Determinants of diversity in a naturally fragmented landscape: humid montane forest avifaunas of Mesoamerica. Ecography 22:582589.

P. L. WEAVER & P. G. MURPHY 1990. Forest structure and productivity in Puerto Rico's Luquillo Mountains. Biotropica 22:6982.

C. O. WEBB 2000. Exploring the phylogenetic structure of ecological communities: an example for rain forest trees. American Naturalist 156:145155.

C. O. WEBB , D. D. ACKERLY , M. A. MCPEEK & M. J. DONOGHUE 2002. Phylogenies and community ecology. Annual Review of Ecology and Systematics 33:475505.

J. J. WEINS , G. PARRA-OLEA , M. GARCÍA-PARÍS & D. B. WAKE 2007. Phylogenetic history underlies elevational biodiversity patterns in tropical salamanders. Proceedings of the Royal Society B 274:919928.

R. H. WHITTAKER 1960. Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs 30:279338.

M. R. WILLIG , D. M. KAUFMAN & R. D. STEVENS 2003. Latitudinal gradients of biodiversity: pattern, process, scale, and synthesis. Annual Review of Ecology, Evolution and Systematics 34:273309.

M. R. WILLIG , S. J. PRESLEY , C. P. BLOCH , I. CASTRO-ARELLANO , L. M. CISNEROS , C. L. HIGGINS & B. T. KLINGBEIL 2011. Tropical metacommunities and elevational gradients: effects of forest type and other environmental factors. Oikos 120:14971508.

B. J. WILSEY , D. R. CHALCRAFT , C. M. BOWLES & M. R. WILLIG 2005. Relationships among indices suggest that richness is an incomplete surrogate for grassland biodiversity. Ecology 86:11781184.

R. D. WILSON , J. W. H. TRUEMAN , S. E. WILLIAMS & D. K. YEATES 2007. Altitudinally restricted communities of schizophoran flies in Queensland's wet tropics: vulnerability to climate change. Biodiversity and Conservation 16:31633177.

H. WOLDA 1987. Altitude, habitat and tropical insect diversity. Biological Journal of the Linnean Society 30:313323.

Z. WOLDU , E. FEOLI & L. NIGATU 1989. Partitioning an elevational gradient of vegetation from southeastern Ethiopia by probabilistic methods. Vegetatio 81:189198.

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