Reid's Paradox
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Reid's Paradox of Rapid Plant Migration or Reid's Paradox, describes the observation from the paleoecological record that plant ranges shifted northward, after the
last glacial maximum The Last Glacial Maximum (LGM), also referred to as the Last Glacial Coldest Period, was the most recent time during the Last Glacial Period where ice sheets were at their greatest extent between 26,000 and 20,000 years ago. Ice sheets covered m ...
, at a faster rate than the
seed dispersal In spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
rates commonly occur. Rare long-distance seed dispersal events have been hypothesized to explain these fast migration rates, but the dispersal vector(s) are still unknown. The plant species' geographic range expansion rates are compared to the actualistic rates of seed dispersal using mathematical models, and are graphically visualized using dispersal kernels. These observations made in the paleontological record, which inspired Reid's Paradox, are from fossilized remains of plant parts, including needles,
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
,
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
, and
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s, that can be used to identify past shifts in plant species' ranges. Reid's Paradox is named after
Clement Reid Clement Reid Fellow of the Royal Society, FRS (6 January 1853 – 10 December 1916) was a British geologist and paleobotany, palaeobotanist. Life Reid was born in London in 1853. His great uncle was Michael Faraday. His family circumstances m ...
, a paleobotanist, who made the principle observations from the paleobotanical record in Europe in 1899. His comparison of
oak An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
tree seed dispersal rates, and the observed range of oak trees from the fossil record, did not concur. Reid hypothesized that
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
was not a possible explanation for the observed paradox, and supplemented his hypothesis by noting that birds were the likely cause of long range seed dispersal. Reid's Paradox has been subsequently documented across Europe and North America.


Dispersal kernels

Dispersal kernels are statistical models that represent the probability of seed dispersal from the source tree. Realistic biological data is required to complete the models. These data are used to accurately fill in variables such as seed number, seed size, and reproductive age. Depending on the plant species, the variables in the equation will change. In the years since Reid hypothesized the methods for seed dispersal, the models have gained more complex elements which attempt to resolve Reid's Paradox. The dispersal of seeds from a parent tree are initially occurs as a
normal distribution In probability theory and statistics, a normal distribution or Gaussian distribution is a type of continuous probability distribution for a real-valued random variable. The general form of its probability density function is f(x) = \frac ...
, as predicted by a standard
diffusion equation The diffusion equation is a parabolic partial differential equation. In physics, it describes the macroscopic behavior of many micro-particles in Brownian motion, resulting from the random movements and collisions of the particles (see Fick's l ...
. However, biological phenomenon complicate the diffusion equation by adding biotic vectors of dispersal such as
blue jay The blue jay (''Cyanocitta cristata'') is a passerine bird in the family Corvidae, native to eastern North America. It lives in most of the eastern and central United States; some eastern populations may be migratory. Resident populations ar ...
s and eastern grey squirrels, species which possess caching behaviors, and abiotic agents of dispersal such as high velocity wind storms. These additional vectors of seed dispersal make the dispersal kernels have a "fat-tail", or a large
kurtosis In probability theory and statistics, kurtosis (from , ''kyrtos'' or ''kurtos'', meaning "curved, arching") refers to the degree of “tailedness” in the probability distribution of a real-valued random variable. Similar to skewness, kurtos ...
. This means that the probability of a long-range dispersal event is higher than that of the standard diffusion dispersal kernel. In order to resolve Reid's Paradox, the vector(s) of seed-dispersal, which give the dispersal kernel a fat-tail, must be identified.


Possible explanations for Reid's Paradox


Animal dispersal

Long distance seed-dispersal events due to animal-seed interactions (such as caching or endozoochorous dispersal) would fatten the tail of the dispersal kernels. To fully explain Reid's Paradox, these rare animal induced seed-dispersal events must have been more important during migration events than recognized or recorded currently.


Cryptic refugia

Small populations of plants may have grown closer to the ice sheets in microhabitats that possessed the habitat characteristics needed for growth and reproduction. This would minimize the actual post-glacial dispersal distance. Such hypothetical populations would not be abundant enough to leave fossil evidence, so have escaped detection. In North America, there is some genetic evidence of cryptic northern refugia for
sugar maple ''Acer saccharum'', the sugar maple, is a species of flowering plant in the soapberry and lychee family Sapindaceae. It is native to the hardwood forests of eastern Canada and the eastern United States. Sugar maple is best known for being the ...
and
American beech American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, p ...
.{{Cite journal, last1=Cruzan, first1=Mitchell B., last2=Templeton, first2=Alan R., date=December 2000, title=Paleoecology and coalescence: phylogeographic analysis of hypotheses from the fossil record, journal=Trends in Ecology & Evolution, volume=15, issue=12, pages=491–496, doi=10.1016/s0169-5347(00)01998-4, pmid=11114435, issn=0169-5347


References

Forest ecology Conservation biology Environmental modelling Paleontology