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• www.clas.ufl.edu/users/mbinford • Click on class link. BIOGEOGRAPHY GEO 4300/5305

The Science and History of Biogeography Lecture 2 – 8 January 2015 Last Time • Introduction – Biogeography? – The Science of Biogeography • Major questions • Methods – Why Geography instead of Biology? • About spatial heterogeneity • Significant overlap but different perspectives • This Class – the Semester – Objectives – Logistics – Evaluation – Schedule • Reading: Chapter 1 & 2 in Lomolino et al.

This Time 8 January

• Who are You? • History of Biogeography; • Organization of Life • Systematics and Biological Nomenclature • Perhaps: The Environmental Setting Who are you?

• Name • Undergrad/Grad • What are you studying? (and thesis/dissertation topic for grad students) • Background in geography • Background in biology • What do you want to do when you graduate? Primary Question of Biogeography

• How and why does biological diversity vary over the surface of the Earth?

Topics and Subdisciplines of Biogeography

• Natural History Zoogeography • Spatial modelers • Vegetation geography • Spatial ecology • Floristic phytogeography • Animal dispersal • Macroevolution paleontology • Speciation • Panbiogeography • Population genetics/evolution • Cladistic/vicariance • Landscape ecology biogeography • Wildlands/conservation • Systematist/phylogenetic • Climatology biogeography • Paleoecology • Island biogeography • Ethnobiology • Biodiversity • And anything else that is • Metapopulation biology concerned with the distribution and abundance of organisms on Earth’s surface.

Biogeography in the News Biogeography in the News

• The biggest factor determining species diversity and distribution on islands is not size and isolation, as traditional island biogeography theory states, but economics. Simply put, the more trade an island is engaged in, the more boats visit it, and with more boats comes more hitchhikers.

A study published in Nature this week examines the species distribution of Anolis lizards throughout the Caribbean islands, finding that their pattern of colonization is exactly the opposite of what traditional island biogeography theory would predict. Theory of Ecology is Biogeography

• One view: In ecology, the scientific domain is spatial and temporal patterns of distribution and abundance. Eight fundamental principles attend this domain: 1. organisms are distributed heterogeneously in space and time 2. all organisms interact with their biotic and abiotic environments 3. the distribution of organisms and their interactions are susceptible to contingency 4. environmental conditions are heterogeneous in space and time 5. resources are finite and heterogeneous in space and time 6. birth and death rates result from interactions with abiotic, biotic environments 7. ecological properties of populations are the consequence of evolution 8. individual variation predominates

History of Biogeography

Overview Time • Accumulation of Species • Pre-13th Century Distribution Information • The Age of Exploration • Theory of Evolution • Biogeography of the 19th • Plate Tectonics Century • Theoretical, Empirical, and – Four British Scientists Technological Advances – Other Contributions of the 19th – Island Biogeography - 1967 Century – Cladistics/Vicariance • The First Half of the 20th – Biodiversity interest Century – Rise of Conservation • Biogeography since the 1950s – Remote Sensing/GIS – Phylogenetics and distinct evolutionary lineages Early

• Pre-15th Century – Greek • Aristotle (384-322 BCE), Theophrastus (c. 370-287 BCE), Virgil (70-19 BCE - Aeneid ) – Roman • Roman Empire – administration, agriculture, written records

http://www.alexandria.ucsb.edu/ The Age of Exploration, Age of Discovery • European: Written accounts of travels to Asia – Giovanni da Pian del Carpine (c. 1180 – 1252) journeyed to Mongolia and back from 1241– 1247. – Marco Polo (1254-1324/1325) journeyed throughout Asia from 1271 to 1295, in the court of Kublai Khan of Cathay. – Trade with the “East” led to exploration by both land and sea – reports of fantastic beasts and strange environments

Travels of Marco Polo 1600 - 1850 "Age of Reason" Linnaeus, Buffon, Lamarck, Lyell

• Linnaeus (1707-1778): Noachan deluge Plants and Animals spread from Mount Ararat (Turkey) • Elevational Zones of Ararat are origins of "biomes"

• Georges-Louis Leclerc, Comte de Buffon (1707- 1788): spread from the Arctic • Buffon's Law: distant regions with similar climate (& similar-appearing vegetation) have different animal species – Mediterranean climate - biome – Monsoonal climate - biome – Climate and Species are changeable

1600 - 1850 "Age of Reason" Linnaeus, Buffon, Lamarck, Lyell • Sir Charles Lyell (1797 - 1875): author of The Geological Evidence of the Antiquity of Man in 1863 and Principles of Geology (12 editions). • Presently observable geological processes were adequate to explain geological history. Uniformitarianism, not book of Genesis! • Vast time scale for Earth's history. • Major influence on Charles Darwin • Jean-Baptiste Lamarck (1744-1829) • Changes in the organic, as well as in the inorganic world, being the result of law, and not of miraculous interposition. • Forerunner of evolution. • Also author of discredited theory of evolution by inheritance of acquired characteristics. The Age of Exploration

• Johann Reinhold Forster (1729 - 1798) Cook's 2nd Expedition 1778 • Global biotic regions (plants) • Higher species diversity in tropics • Species diversity correlated with island size

• Alexander von Humboldt (1769- 1859): Plant Vegetation types strongly correlated with local climate • Elevational Vegetation Zones (Andes) • Latitudinal Belts of Vegetation 1850 - 1900 Evolution by Natural Selection, but pre-Plate Tectonics • Charles Darwin (1809-1882) Evolution through Natural Selection: The Origin of Species in 1859. • Theoretical Framework for Biotic Patterns in Space and Time • Fundamental to all Biology • Also a barnacle, coral, and earthworm expert. 1850 - 1900 Evolution by Natural Selection, but pre-Plate Tectonics • Alfred Russel Wallace (1823-1913) – Biotic Regions similar to Sclater's – Originator of Zoogeography • Distance not equal taxonomic similarity • Integrated geological, fossil, evolutionary information • Considered paleoclimate influences distributions

17 Biogeographic Principles Advocated by Alfred Russel Wallace • See Lomolino et al. page 33, Box 2.1 • Still the basis of much of biogeography today, but with the addition of plate tectonics and genetics. Hooker and Sclater • Joseph Dalton Hooker (1817 – 1911) – Asst. Surgeon and Botanist on Ross Expedition to southern hemisphere (1839 – 1843) – First proposed “breaking up of a continuous tract of land” – Director of Kew Botanical Gardens – Drew analogy between montane and island floras

• Phillip Lutley Sclater (1829-1913) Five Terrestrial biotic regions (for birds) – Palearctic – Aethiopian – Indian – Neotropical – Nearctic – Australasian • Six Marine regions (marine mammals)

Nineteenth Century “Name" Rules (Laws) • C.W.L. Golger (1833) within a species, individuals in moist climates are darker • C. Bergmann (1847) for warm blooded animals, those in colder climates are larger • J.A. Allen ( 1878) for warm blooded animals, those in colder climates have shorter limbs and appendages • E.D. Cope (1887)(orthogenesis vs. G.G. Simpson) groups tend in one direction, e.g., larger body size with time • Guthrie-Geist (20th Century '85 '87) for larger mammals, more food yields larger animals (island dwarfing)

Late Nineteenth Century

• C. Hart Merriam (1884) – Life Zones – Elevation and Latitudinal (cf. Alexander von Humboldt) – Arizona, S. Idaho • Asa Gray (botanist) – disjunctions: taxonomically similar groups distantly separated

Merriam's Life Zones

See also Fig. 2.12 in Lomolino et al. Biogeography after 19th Century • The big question was: How did the world get this way?

The First Half of the 20th Century • C. Raunkiaer (1934) - Ecological classification (vs. taxonomic) – Therophytes – Geophytes – Epiphytes • A. L. Wegener (1910) ; Drummond Matthews and Fred Vine (1963) – Continental Drift (Plate Tectonics) • Ernst Mayr (1904-2004) - genetics – Biological species concept (potentially interbreeding to produce fertile offspring) – Allopatric speciation (arising through geographic isolation) • Centers of Origin - current patterns – George Gaylord Simpson (1902-1984) Paleontologist – Philip J. Darlington (1904-1983) Zoologist

Biogeography since the 1950s (technological revolution, ecology, paleontology) • Plate Tectonics – Magnetometers; deep sea drilling – sonar, submarines • L. Croizat (1958) vicariance biogeography: disjunction of multiple species due to the growth of barriers • R. H. MacArthur and E. O. Wilson (1963) Island Biogeography • Technological Advances – radiometric dating – GIS and Remote Sensing – The entire Earth can be imaged synoptically – Genomics – evolutionary linkages

Need for Biogeography in Conservation, Climate Change • Species Diversity function of overlapping species ranges.

Ladle, Richard J., and Robert J. Whittaker, eds. Conservation Biogeography . Chichester, UK: Wiley, 2011.

• As climate changes, environmental conditions change. Organization of Life

• Levels of Functional Organization • Biological Systematics and Taxonomic Levels Hierarchical Levels of Functional Organization • Subatomic Particles • Population • Atoms • Community • Molecules • Ecosystem • Cell organelles • Landscape • Cells • Biosphere • Tissues • Earth • Organs • Organism Taxonomic Levels of Biological Organization - Linnaean TAXON (plural TAXA) Binomial • Domains* Nomenclature • Kingdom • Phylum (Division for Pinus elliottii plants and Fungi) • Class Homo sapiens • Order • Family Pan troglodytes • Genus (plural Genera) • Species (singular AND Genus name + plural) specific epithet Carolus Linnaeus (1707 — 1778) = Species name Systema Naturae published 1758 *Not Linnaean Domain and Kingdom

Three Domains: Archaea, Bacteria (Eubacteria), and Eukaryota (all eukaryotic groups: Protista, Plantae, Fungi, and Animalia) Purves et al., Life: The Science of Biology, 4th Edition. Sinauer Associates Taxonomy of Humans • Domain: Eukarya (organisms which have cells with a nucleus) • Kingdom: Animalia (with eukaryotic cells having cell membrane but lacking cell wall, multicellular, heterotrophic) • Phylum: Chordata (animals with a notochord, dorsal nerve cord, and pharyngeal gill slits, which may be vestigial or embryonic) • Subphylum: Vertebrata (possessing a backbone, which may be cartilaginous, to protect the dorsal nerve cord) • Class: Mammalia (endothermic vertebrates with hair and mammary glands which, in females, secrete milk to nourish young) • Cohort: Placentalia (giving birth to live young after a full internal gestation period) • Order: Primates (collar bone, eyes face forward, grasping hands with fingers) • Suborder: Anthropoidea (monkeys, including apes, including humans; as opposed to the lemurs, lorises, and tarsiers) • Infraorder: Catarrhini (Old World anthropoids) • Superfamily: Hominoidae (apes, including humans) • Family: Hominidae (great apes, including humans) • Genus: Homo • Species: Homo sapiens (high forehead, well-developed chin, gracile bone structure) Kingdom: Animalia Phylum: Chordata Why Scientific Names? Subphylum: Vertebrata Class: Actinopterygii Order: Amiiformes • Amia calva L. Family: Amiidae • Bowfin in most of U.S. Genus: Amia Species: Amia calva L. • beaverfish, blackfish, cottonfish, cypress trout, freshwater dogfish, grindle, grinnel, John A. Grindle, lawyer, marshfish, scaled ling, speckled cat, and western mudfish. Choupique is a common name used in Louisiana that was derived from the Choctaw name for bowfin. http://www.flmnh.ufl.edu/fish/Gallery/ Descript/Bowfin/Bowfin.html What is a Species? • Species are the fundamental taxonomic units of biological classification. Environmental laws are framed in terms of species. Even our conception of human nature is affected by our understanding of species. • Ernst Mayr: all the individual organisms of a natural population that generally interbreed at maturity in the wild and whose interbreeding produces fertile offspring. • But there are problems with this definition. • After thousands of years of use, the concept remains central to biology and a host of related fields, and yet also remains at times ill-defined. • "Right now we can only guess that the correct answer for the total number of species worldwide lies between 2 and 100 million," Michael Rosenzweig. What is a Species? Systematics and Biological Nomenclature • Rules set by International Commissions • International Commission on Zoological Nomenclature (ICZN) – http://www.iczn.org/ • International Code of Botanical Nomenclature – http://www.bgbm.org/iapt/nomenclature/code/Sai ntLouis/0001ICSLContents.htm

Some general rules for nomenclature 1. All taxa must belong to a higher taxonomic group. 2. The first name to be validly and effectively published has priority. 3. All taxa must have an author. When you see a scientific name such as Homo sapiens L., the L. stands for Linneus, who first described and named that organism. Most scientists must have their names spelled out, for example Libopollis jarzenii Farabee et al. Next Time

• Environmental Setting: – Energy – Atmospheric Circulation Patterns – Climatic Regions of the World – Soils • Parent Materials • Soil Formation – Aquatic Environments – Microenvironments