PLANT SPECIATION AND EVOLUTION LECTURES
Biogeography
I. Plant distributions
A. Factors influencing plant distributions
1. Wind and ocean currents [Cox & Moore 1993, p. 83]
2. Physical barriers and land masses
3. Latitude and elevation--different communities distributed along a latitudinal gradient and also along an elevational gradient (1000 m elevational change roughly equals 10 degree latitude change) [Raven et al. 1992, p. 661]
4. Annual precipitation and temperature cycles
5. Natural processes--e.g., substrate weathering on mountain slopes, riverbank ice-scouring, wind deposition of sand in dunes, fire in savannas and prairies, drought/rain cycles in deserts, drawdown in coastal plain marshes
B. Vegetation patterns
1. Regional/continental biomes--e.g., desert, mediterranean scrub, arctic tundra, temperate deciduous forest [Raven et al. 1992, p. 658]
2. Floristic regions and elements--e.g., Palaeotropical vs. Australian regions [Cox & Moore 1993, p. 209]
3. Autochthonous vs. allochthonous species
a) Autochthonous--evolved to present-day form in an area, e.g., island endemics
b) Allochthonous--evolved elsewhere and arrived to area subsequently, exotics
4. Indigenous (native) species vs. exotics (introduced or persistent after cultivation); e.g., latter would be dandelion (Taraxacum officinale) and purple loosestrife (Lythrum salicaria) in the U.S.
5. Frequent range patterns and disjunctions
a) Eastern Asian-eastern North American, e.g., tulip-tree (Liriodendron)
b) Circumpolar (Northern Hemisphere), e.g., linear-leaved sundew (Drosera linearis)
c) Gondwanan (eastern South America-western Africa), e.g., Brocchinia and nearest relative
d) Bipolar (high latitudes in both hemispheres), e.g., sweet-cicely (Osmorhiza)
e) Austral (southern South America-southern Australia/South Pacific), e.g., southern beech (Nothofagus)
C. Widespread vs. rare species
1. Cosmopolitan species--very widely distributed across continents, e.g., dandelion (Taraxacum)
2. Evolutionary and climatic relicts
a) Evolutionary relict--scattered survivor of a once more dominant and widespread group that largely died out, e.g., magnolias
b) Climatic relict--distribution greatly impacted by climatic or sea level changes, e.g., redwoods
3. Rare species--some may have always been rare, some are rare because of human impacts or natural environmental change
II. Biogeographic issues
A. Types of biogeographic analysis--three different approaches emphasizing different types of data: phylogenetic, cladistic and panbiogeography
B. Vicariance vs. dispersalism
1. Vicariant ranges within continents determined by emergence of physical barriers, intercontinental disjunctions determined by continental drift-->poor to no seed dispersability; e.g., cycads, southern beech (Nothofagus)
2. Dispersalist ranges determined by active movement of disseminules (seeds or plantlets or fragments) by dispersal agents such as wind, water, birds-->high dispersability; e.g., perhaps most plants
3. Hotly debated, but most species surely disperse at least moderate distances at least rarely, and some disperse very long distances, e.g., intercontinentally--extremes of a continuum
4. Example of trouble with strict reliance on vicariance alone--molecular phylogenetic and biogeographic study of mistletoes (Arceuthobium) in the Western Hemisphere
a) Investigators suggested that substantial molecular divergence in nuclear ribosomal DNA sequences between a northeastern North American and Hispaniolan species was result of vicariance, with 20 million year old disjunction occurring by over-land migration of mistletoes when the islands were semi-connected to North America
b) However, mistletoes have highly developed dispersal mechanisms that squirt seeds with sticky goo onto birds' legs; birds must carry such seeds hundreds (thousands?) of miles
c) Numerous bird species overwinter in and south of the Caribbean every year-->long-distance dispersal far more reasonable (researchers didn't bother to explain how the mistletoes exhibit a North American-European species disjunction with considerable lower molecular divergence)
d) Great sequence divergence can be explained differently, too--one or more species not sampled in the study would "fill in the gap" between the Hispaniolan and North American mistletoes, or the northeastern population is not very closely related whereas southern populations would be, and should be sampled (might close the time and sequence divergence gap dramatically)
C. Centers of origin, centers of endemism, and refugia
1. Center of origin--"birthplace" of a lineage, e.g., "southern Gondwanaland" for southern beech (Nothofagus)
2. Center of endemism--"hotspot" for diversification, with lots of narrowly restricted endemics; a high incidence of endemics in a region also indicates this area may be a center of origin for the group, e.g., southern Patagonia (=Argentina and Chile), SE Australia and southern Pacific islands in present-day Nothofagus, but fossils known from Antarctica
3. Refugium--place where different groups survived environmental catastrophes such as Pleistocene glaciation, from which they later dispersed, e.g., Queen Charlotte Islands hypothesized for Tellima and other Pacific Northwestern species [more next Wednesday]
D. Phylogeography--molecular data for populations within a species or among closely related ones clarifies dispersal patterns, helps illuminate major geologic/geographic events and their relative sequence in a species' evolutionary history
1. Canary Island lizards (Gallotia galloti)
a) Mitochondrial DNA study of three lizard species including different island populations of one clarified the inter-island migration of the one species
b) Allowed interpretation of variant traits on different islands, some with unique origin and reproductively related, others convergent (e.g., body size changes)
2. Pacific Northwest Tellima grandiflora
a) Restriction site variation in chloroplast DNA separated northern and southern populations of Tellima grandiflora; northern type presumed to survive at a refugium (isolated spot remaining snow-free during glaciation) on the Queen Charlotte Islands
b) Surprise!--an evident ancient hybridization event led to the "capture" of the chloroplast of a different genus, Mitella, in the northern Tellima populations
c) North-south division has since been revealed in many other Pacific Northwestern plant species, indicating a region-wide floristic division during the Pleistocene
3. Human evolution [Cann et al. 1987, p. 34]
a) Restriction site study of mitochondrial DNA from people around the world were analyzed to infer the pattern of continental diversification and migration
b) Genetically most diverse populations were African; fossil evidence suggested that the "root" of the tree was also in Africa, and phylogenetic relationships of remaining human populations reflected repeated colonization events into other continental areas from Africa-->became known as the "Out of Africa" theory
c) These earlier investigators proposed that all humans arose from a single female ca. 200,000 years ago-->the "Mother Eve" hypothesis
d) Much debate and controversy, and another theory developed ("Multinational Origins") with at least three simultaneous origins of modern humans (polyphyly!); further population genetic studies discounted "Mother Eve"
e) More recent extensive re-analysis of earlier mitochondrian data and more careful evaluation of fossil and anthropological data support the "Out of Africa" theory
4. Phylogenetic units in conservation--definition of "monophyletic" and geographically distinctive units (=sets of populations) by phylogeographic approach, to identify "worthy" populations for conservation; e.g., tiger beetles (Cicindela)
Bibliography
Avise, J. C. 1994. Molecular markers, natural history and evolution. Chapman and Hall, New York, New York. 511 pp.
Cox, C. B. and P. D. Moore. 1993. Biogeography: an ecological and evolutionary approach, 5th ed. Blackwell Scientific Publications, Oxford, United Kingdom. 326 pp.
Malhotra, A., R. S. Thorpe, H. Black, J. C. Daltry, and W. Wuester. 1996. Relating geographic patterns to phylogenetic processes. In: Harvey, P. H., A. J. L. Brown, J. M. Smith, and S. Nee (eds.), New uses for new phylogenies. Oxford University Press, Oxford, United Kingdom. pp. 187-202.
Moritz, C. 1996. Uses of molecular phylogenies for conservation. In: Harvey, P. H., A. J. L. Brown, J. M. Smith, and S. Nee (eds.), New uses for new phylogenies. Oxford University Press, Oxford, United Kingdom. pp. 203-214.
Raven, P. H., R. F. Evert, and S. E. Eichhorn. 1992. Biology of plants, 5th ed. Worth Publishers, New York, New York. 791 pp.
Soltis, D. E., M. S. Mayer, P. S. Soltis, and M. Edgerton. 1991. Chloroplast-DNA variation in Tellima grandiflora (Saxifragaceae). American Journal of Botany 78:1379-1390.
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