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High-net-worth individual
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Not to be confused with Ultra high-net-worth individual.
Parts of this article (those related to documentation) need to be updated. Please update this article to reflect recent events or newly available information. (January 2017)
High-net-worth individual (HNWI) is a term used by some segments of the financial services industry to designate persons whose investible assets (such as stocks and bonds) exceed a given amount. Typically, these individuals are defined as holding financial assets (excluding their primary residence) with a value greater than US$1 million.[1][2][3]
However, there are distinct classifications of HNWI and the exact dividing lines depend on how a bank wishes to segment its market. For example, an investor with less than US$1 million but more than US$100,000 is considered to be "affluent", or perhaps even "Sub-HNWI".[1] "Very-HNWI" (VHNWI) can refer to someone with a net worth of at least US$5 million.[4]
By 2007, the expansion of HNWI assets led to the creation of a super class of HNWIs, known as ultra-high-net-worth individuals (UHNWIs), i.e. those with US$30 million in liquid financial assets according to the Capgemini and Merrill Lynch World Wealth Report 2006[3][5] or with a disposable income of more than US$20 million.[6]
At the end of 2016, there were just over 13 million HNWIs in the world. The United States of America had the highest number of HNWIs (4,400,000) of any country, whilst London had the most HNWIs (357,200) among cities as based on data from the Knight Frank Wealth Report.[7]
Contents
[hide]
1 United States: SEC regulations
2 Annual World Wealth Report
3 Capgemini figures
3.1 Banking and finance
3.2 Retail
4 Knight Frank Wealth Report
5 Academic studies of asset management trends
6 High-net-worth individual's taxable portfolios
7 Liability insurance
8 See also
9 References
10 External links
United States: SEC regulations[edit]
See also: Accredited investor
The U.S. Securities and Exchange Commission requires all SEC-registered investment advisers to periodically file a report known as Form ADV.[8] Among other things, Form ADV requires each investment adviser to state how many of their clients are "high-net-worth individuals." The Form ADV Glossary of Terms explains that a "high-net-worth individual" is an individual with at least $1,000,000 managed by the reporting investment adviser, or whose net worth the investment adviser reasonably believes exceeds $2,000,000 (or who is a "qualified purchaser" as defined in section 2(a)(51)(A) of the Investment Company Act of 1940). The net worth of an individual for SEC purposes may include assets held jointly with his or her spouse. Unlike the definitions used in the financial and banking trade, the SEC's definition of HNWI would include the value of a person's verifiable non-financial assets, such as a primary residence or art collection.
Annual World Wealth Report[edit]
The World Wealth Report was co-published by Merrill Lynch and Capgemini, previously known as Cap Gemini Ernst & Young who worked together since c. 1993, investigating the "needs of high-net-worth individuals (HNWIs are individuals with more than $1 million in assets excluding primary residence)" in order to "successfully serve this market segment." Their first annual World Wealth Report was published in 1996.[9] The World’s Wealth Report[10] defines HNWIs as those who hold at least US$1 million in assets excluding primary residence and ultra-HNWIs as those who hold at least US$30 million in assets excluding primary residence. The report states that in 2008 there were 8.6 million HNWIs worldwide, a decline of 14.9% from 2007. The total HNWI wealth worldwide totaled US$32.8 trillion, a 19.5% decrease from 2007. The ultra-HNWIs experienced the greater loss, losing 24.6% in population size and 23.9% in accumulated wealth. The report revised its 2007 projections that HNWI financial wealth would reach US$59.1 trillion by 2012 and revised this downward to a 2013 HNWI wealth valued at $48.5 trillion advancing at an annual rate of 8.1%.[11]
The 2013 World Wealth Report[12] was jointly produced by Capgemini and RBC Wealth Management and included, for the first time, the Global HNW Insights Survey produced in collaboration with Scorpio Partnership. The inaugural survey represented one of the largest and most in-depth surveys of high-net-worth individuals ever conducted, surveying more than 4,400 HNWIs across 21 major wealth markets in North America, Latin America, Europe, Asia-Pacific, Middle East, and Africa. Scorpio Partnership have established themselves as a market leader in the supply of HNW insight having spent over 15 years conducting private client interviews, collecting business intelligence and working with over 100 clients who range from universal banks, domestic retail banks, specialist private banks and fund managers, to family offices, high-net-worth clients and regulatory bodies. The partnership is independently owned and managed by Sebastian Dovey and Cath Tillotson and carries out global assignment overseen from its base in London's West End.
Largest HNWI Population, 2012[13][14]
Country
HNWI Population
Percentage of total population
United States 3.44 million 1.09
Japan 1.9 million 1.49
Germany 1.0 million 1.24
China 643,000 0.05
United Kingdom 465,000 0.73
HNWI Wealth Distribution (by Region)[14]
Region
HNWI Population
HNWI Wealth
Global 12 million $46.2 trillion
North America 3.73 million $12.7 trillion
Asia-Pacific 3.68 million $12.0 trillion
Europe 3.41 million $10.9 trillion
Latin America 0.52 million $7.5 trillion
Middle East 0.49 million $1.8 trillion
Africa 0.14 million $1.3 trillion
Capgemini figures[edit]
The World Wealth Report published Capgemini has estimated the number and combined investable wealth of high-net-worth individuals as follows,[15][16][17] using the United States Consumer Price Index (CPI) Inflation Calculator.[18]
Year
Number of HNWIs
(millions)
HNWI Wealth
(trillions USD)
In 2012 USD
(trillions USD)
1998[15] ? 17.4 24.3
1999[19] 6.0 21.6 29.5
2000[20] 7.0 25.5 33.7
2001[21] 7.2 27.0 34.7
2002[22] 7.1 26.2 33.2
2003[23] 7.3 27.2 33.7
2004[24] 7.7 28.8 34.7
2005[25] 8.3 30.8 35.9
2006[5] 8.7 33.3 37.6
2007[26] 9.5 37.2 40.9
2008[27] 10.1 40.7 43.1
2009[11] 8.6 32.8 34.8
2010[16] 10.0 39.0 40.7
2011[17] 10.9 42.7 43.2
2012[14] 12.0 46.2 46.2
2013[28] 13.7
2014[28] 14.7
2015[28] 15.4
2016[28] 16.5 63.5
Banking and finance[edit]
Main articles: Private banking and Wealth management
Most global banks, such as Credit Suisse, Barclays, BNP Paribas, Citibank, Deutsche Bank, HSBC, JPMorgan Chase and UBS, have a separate business unit with designated teams consisting of client advisors and product specialists exclusively for UHNWI. Because of their extreme high net worth and the way their assets were generated[further explanation needed], these clients are often considered to have characteristics similar to institutional investors.
By 2006, asset managers working for HNW individuals invested more than £300 billion on behalf of their clients. These wealth managers are bankers who in 2006, earned multimillion-pound salaries and owned their own companies and equity funds.[29] In 2006, a list of the 50 top investment bankers was published by the Spear's Wealth Management Survey.
Retail[edit]
Brands in various sectors, such as Bentley, Maybach, and Rolls-Royce in motoring, actively target UHNWI and HNWI to sell their products. In 2006, Rolls-Royce researchers suggested there were 80,000 people in ultra-high-net-worth category around the world. UHNW individuals "have, on average, eight cars and three or four homes. Three-quarters own a jet aircraft and most have a yacht."[6]
Knight Frank Wealth Report[edit]
Knight Frank has been publishing an annual wealth report since 2005. The following is a list of the cities with the most number of HNWIs as of December 2016 according to data compiled for the Knight Frank Wealth Report by New World Wealth.[7]
Rank
City
Number of
HNWIs
(2016)
1 London 357,000
2 New York City 339,000
3 Tokyo 280,000
4 Hong Kong 228,000
5 Singapore 217,000
6 San Francisco Bay Area 180,000
7 Los Angeles 173,000
8 Chicago 135,000
9 Frankfurt 128,000
10 Beijing 122,000
Academic studies of asset management trends[edit]
The Wharton Global Family Alliance whitepaper was released in 2008 to study the investment strategies of single family offices in the United States and in Europe.[30] The research was segregated into sub-groups representing those with less than $1 billion in assets and those with assets above $1 billion. The study found that U.S. families reported a more aggressive attitude toward investment objectives than their counterparts in Europe. One recommendation of the WGFA study advised the advisors and family offices serving this niche to avoid complexity in the structure of portfolios.
The authors cite that the more complex the portfolio and number of holdings, the more difficult the job of performing adequate governance, reporting, and education. The Institute for Private Investors, a peer networking organization for wealthy families and their advisors, suggested a similar theme to its membership in 2008 with a conference themed, "The Return to Simplicity".[31] Kotak Wealth Management and CRISIL Research, published a report on the Ultra High Net Worth Individuals in India titled "Top of the Pyramid Report".[32]
High-net-worth individual's taxable portfolios[edit]
Since 2009, the Gannon Group has managed wealth for exclusively HNW and UHNW individuals and families at Morgan Stanley's Portfolio Management Group.[33][34] In his book entitled Investing Strategies for the High Net Worth Investor: Maximize Returns on Taxable Portfolios,[35] Gannon Group's executive director Niall Gannon investigated asset management strategies for high tax bracket investors on taxable portfolios. Gannon advises taxable investors in terms of long-range estate planning decisions and assists them in making more accurate estimates of their expected total portfolio value. He studied the S&P 500 index on an after-tax basis and found the return to be 6.63% after paying taxes at the top prevailing federal tax rate and a constant average of 6% for state taxes from January 1, 1957, through December 31, 2010.[35]
Additional results from Gannon's research found that a portfolio of municipal bonds out-performed the re-invested S&P 500 index in 17% of the rolling 20-year periods since the inception of the index. Gannon rejects the use of historical returns for future asset allocation modelling for high-net-worth investors. He argues that an observation of stock portfolio earnings yields (the inverse of the p/e ratio) are more indicative of the future return potential of the portfolio when modelled with the impact of taxation on performance.[35]
Liability insurance[edit]
Main article: Umbrella insurance
High-net worth individuals have unique risks to their wealth through legal liability and tort law. Therefore, they commonly purchase umbrella insurance policies which further protect them from expensive lawsuits. These "umbrella" policies are designed in an open-ended way to cover risks which fall through the "gaps" of other insurance policies. They are also commonly specifically designed to have higher limits for covering lawsuits resulting from expensive vehicle accidents, home accidents, defamation, and fiduciary liability arising from nonprofit board volunteering.[36] In the United States, a trade group called the Council for Insuring Private Clients was formed in 2012 to focus on this market.[37] Companies known to focus on the market include Chubb,[38] ACE Limited and Privilege Underwriters Reciprocal Exchange (Pure).[39]
As a cheaper alternative, a "personal excess" policy may be purchased instead, which is not designed to cover the gaps in coverage the way that an umbrella policy does.[36]
See also[edit]
Billionaire
Ultra high-net-worth individual
Millionaire
Middle Class
Global assets under management
Plutonomy
Politically exposed person
Swiss bank
Wealth
References[edit]
^ Jump up to: a b Investopedia Definition of "High Net Worth Individual (HNWI)"
Jump up ^ "Capgemini 2007 World Wealth Report" (PDF). Capgemini. 2006-06-12. Retrieved 2007-07-08. World Wealth Grows to $33.3 trillion Says Merrill Lynch
^ Jump up to: a b Nivedita Chakravartty (18 Jan 2007). "For A Few Dollars More". The Times of India.
Jump up ^ Stefano Caselli; Stefano Gatti (2005). Banking for Family Business: A New Challenge for Wealth Management. Springer. ISBN 3-540-22798-9.
^ Jump up to: a b 2006 World Wealth Report (PDF) (Report). Capgemini. 2006. Archived from the original (PDF) on January 2, 2010. Retrieved 11 September 2013.
^ Jump up to: a b Ray Hutton (5 November 2006). "Rich spurn ultra-luxury cars". UK: The Sunday Times. Retrieved 10 September 2013.
^ Jump up to: a b http://www.knightfrank.com/wealthreport
Jump up ^ SEC Form ADV
Jump up ^ 2003 World Wealth Report (PDF) (Report). Capgemini. 2003. Archived from the original (PDF) on January 2, 2010. Retrieved 10 September 2013.
Jump up ^ 2009 World Wealth Report (Report). Thought Leadership. Capgemini. 2009.
^ Jump up to: a b 2009 World Wealth Report (PDF) (Report). Capgemini. 2009. Retrieved 11 September 2013.
Jump up ^ 2013 World Wealth Report (PDF) (Report). Capgemini. 2013. Retrieved 10 September 2013.
Jump up ^ World Wealth Report 2013. capgemini.com
^ Jump up to: a b c "World Wealth Report 2013". Capgemini.
^ Jump up to: a b 1998 World Wealth Report (PDF) (Report). Capgemini. 1998. Retrieved 10 September 2013.
^ Jump up to: a b 2010 World Wealth Report (PDF) (Report). Capgemini. 2010. Retrieved 11 September 2013.
^ Jump up to: a b 2011 World Wealth Report (PDF) (Report). Capgemini. 2011. Retrieved 11 September 2013.
Jump up ^ "Consumer Price Index (CPI) Inflation Calculator". U.S. Bureau of Labor Statistics.
Jump up ^ http://www.capgemini.com/m/en/doc/WWR99.pdf
Jump up ^ http://www.capgemini.com/m/en/doc/WWR00.pdf
Jump up ^ http://www.in.capgemini.com/m/in/tl/pdf_2001_World_Wealth_Report.pdf
Jump up ^ http://www.capgemini.com/m/en/tl/pdf_2002_World_Wealth_Report.pdf
Jump up ^ https://web.archive.org/web/20100102085510/http://ml.com/media/14076.pdf
Jump up ^ http://www.hr.capgemini.com/m/hr/tl/World_Wealth_Report_2004.pdf
Jump up ^ http://www.hr.capgemini.com/m/hr/tl/World_Wealth_Report_2005.pdf
Jump up ^ https://web.archive.org/web/20100102074059/http://ml.com/media/79882.pdf
Jump up ^ 2008 World Wealth Report (PDF) (Report). Capgemini. 2008. Archived from the original (PDF) on November 23, 2008. Retrieved 11 September 2013.
^ Jump up to: a b c d "The proliferation of high-net-worth individuals". The Economist. 4 October 2017.
Jump up ^ Rivkin, Annabel (12 December 2006). "How I make the rich richer". London: The Times. Retrieved 10 September 2013.
Jump up ^ Wharton Global Family Alliance. "Benchmarking the Single Family Office: Identifying the Performance Drivers".
Jump up ^ Institute for Private Investors. [1].
Jump up ^ Ultra HNI segment set to treble: Report. Indian Express (2011-06-07). Retrieved on 2013-07-18.
Jump up ^ "The Gannon Group at Morgan Stanley Portfolio Management Group". Morgan Stanley.
Jump up ^ Paul Sullivan (22 February 2013). "Reasons to Avoid Buying Stocks, and Why You Should Ignore Them". New York Times. Retrieved 9 September 2013.
^ Jump up to: a b c Gannon, Niall Joseph (December 2009). Investing Strategies for the High Net Worth Investor. New York: McGraw Hill. ISBN 978-0-07-162820-4.
^ Jump up to: a b "Comparing Stand-Alone Personal Umbrella Policies | IRMI.com". www.irmi.com. Retrieved 2016-01-16.
Jump up ^ "New Trade Group to Focus on High-Net-Worth Segment". Retrieved 2016-01-16.
Jump up ^ Ruquet, Mark E. (5 March 2007). "High-Net-Worth Clients in an Insurance World All Their Own". Insurance Office of Central Ohio. Retrieved 2016-01-06.
Jump up ^ "Feature Story: Big Fish, Bigger Pond". PropertyCasualty360.com. Retrieved 2016-01-16.
External links[edit]
2013 World Wealth Report (WWR) (PDF) (Report). Capgemini. 2013.
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Forensic footwear evidence
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Forensic footwear evidence can be used in legal proceedings to help prove that a shoe was at a crime scene. Footwear evidence is often the most abundant form of evidence at a crime scene and in some cases can prove to be as specific as a fingerprint. Initially investigators will look to identify the make and model of the shoe or trainer which made an impression. This can be done visually or by comparison with evidence in a database; both methods focus heavily on pattern recognition and brand or logo marks. Information about the footwear can be gained from the analysis of wear patterns which are dependent on angle of footfall and weight distribution. Detailed examination of footwear impressions can help to link a specific piece of footwear to a footwear imprint as each shoe will have unique characteristics.
Contents
[hide]
1 Types of footwear evidence
1.1 Footwear outsole impressions
1.2 Footwear insole imprints
1.3 Footwear trace evidence
2 Detection of footwear evidence
3 Recovery of footwear impression evidence
3.1 Lifting
3.2 Casting
3.3 3D scanning
3.4 Three-dimensional imaging
4 Examination of footwear impressions evidence
5 Limitations of footwear evidence
6 Footwear databases
7 Footwear Software
8 High profile cases
9 References
10 External links
Types of footwear evidence[edit]
Footwear evidence can come in at least three forms, footwear outsole impressions, footwear insole impressions and footwear trace evidence.
Footwear outsole impressions[edit]
Footwear outsole impressions are impressions left on an object that was caused by contact with a piece of footwear. These can be left on the ground or raised surface by persons treading over it, left on doors or walls by persons attempting to kick or climb over a wall or even left on other persons after being kicked or stomped on.
Shoeprints left on a dusty surface
There can also be latent impressions not easily visible to the naked eye, on many different surfaces such as floor tiles, concrete or even carpet. Detection may require the use of additional specialized light sources such as portable ultraviolet lighting. Recovery typically includes photography as well as lifting with "gel" or "electrostatic" dust lifters.
Footwear insole imprints[edit]
Imprints of a person's foot can be left on the insole's inside footwear. The size and arrangement of foot and toe imprints can be used to determine whether a person has worn a piece of footwear. The analysis and comparisons of foot imprints is part of the discipline of forensic podiatry. The insole will show a virtual image of the bare foot print of the wearers foot. This can be compared to the actual barefoot print of the shoe owner to gain a match. Useful in forensic evidence casting and identification. H Farmer 2017
Footwear trace evidence[edit]
Footwear trace evidence is trace evidence that is recovered from footwear. Types of trace evidence that could be recovered include skin, glass fragments, body hair, fibers from clothing or carpets, soil particles, dust and bodily fluids. The study of this trace evidence could be used to link a piece of footwear to a location or owner. DNA can be one of the contributing factors in forensic footwear evidence.
Detection of footwear evidence[edit]
Footwear impression revealed by oblique lighting.
Footwear impression details revealed with fingerprint powder
Footwear impressions can be detected with a variety of methods including:
Visually using natural or artificial lighting.
Using artificial light sources to provide oblique, coaxial, and polarized light for detection of visible and latent impressions.
Using electrostatic lifting devices to lift dusty impressions.
Using fingerprint powder to develop latent impressions.
Using physical or chemical enhancement methods to develop or enhance faint impressions.
Recovery of footwear impression evidence[edit]
Footwear evidence occurs most often as either footwear impressions left in a soft surface, such as mud or as dust deposits, which are difficult for the human eye to detect. At violent crimescenes footmarks can be left as a result of a person standing in blood and subsequently trailing it as they move around the scene.
Lifting[edit]
Footwear impressions can be lifted from surfaces with tools such as adhesive lifters, gelatin lifters or electrostatic lifting devices.
Casting[edit]
Evidence left via impressions can generally be recovered utilizing a plaster cast. Initially the impression is isolated by framing the area with a solid boundary. Following this a plaster mix can be gently poured inside the frame; it is generally considered not best practice to pour directly onto the impression. In some cases where the surface is not ideal for casting prior techniques can be utilised to gain a better cast of the impression. Sand can often be fixed in place by applying an aerosol resin or glue although hair spray is often used. Wet mud impressions can be dried using a combination of pipetting water from the surface and applying hot air, often in the form of a hair dryer.
3D scanning[edit]
Footwear evidence can be recovered with use of 3D scanners.
There are two types of 3D scanners used in forensics:
crime scene scanners able to capture a large overview map of the scene;
‘close-up’ 3D scanners able to capture individual objects in high resolution and full colour.[1]
Advantage of 3D scanning over casting is that 3D scanners can scan the object without touching or affecting it – most 3D scanners use lasers to capture 3D information. Some scanners can also capture the colour surface, [null producing] a visually [null accurate] replica of the object. To create a complete and highly detailed 3D model of a footprint less than 15 minutes is required.
Three-dimensional imaging[edit]
Recent approach takes advantage of digital photography. A set of digital images of a footmark or target area of interest, taken in different viewpoints is required to create detailed and accurate 3D model. Appropriate software allows to visualise a 3D model in different ways, make accurate measurements from a 3D trace or perform a comparison of multiple traces.
Examination of footwear impressions evidence[edit]
Footwear impression can reveal information that may be of use to forensic investigators. Analysis of impressions found at a crime scene may provide the following information:
Number of people at a crime scene: Different footwear impressions left at a crime scene will indicate more than one person was present at the crimescene.
Approximate height of the wearer: There is statistical correlation[2] between foot/footwear size and height and also stride length and height. The dimensions of the footwear impression and the distance between impressions can be used to provide an estimate of height.
Activity of wearer when impression was made: Plastic footwear impression left on soft surfaces can reveal whether a person was walking, running or carrying a heavy load when the impression was made. An impression made by person running will make impressions that are deeper in the heel and toe sections. A person carrying a heavy load such as a body will leave imprints that are deeper than when not carrying a heavy load.
Manufacture, model and approximate size of footwear: Footwear impression can show the design elements (shapes, patterns, arrangement) that form the outsole design. These can be compared with a footwear outsole database to identify the model of footwear that made the impression. Knowledge of the model of footwear that made the impression can assist in narrowing down the list of suspects and can also be used to link multiple crime scenes to the same perpetrator(s).
Establish link between crime scene impression with a specific piece of footwear: Comparison of crime scene impressions with a specific piece of footwear recovered from a suspect can determine if there is any relationship. A worn piece of footwear will gradually acquire unique wear and tear on the outsoles. These randomly acquired characteristics will be unique to that specific shoe and may show in crime scene impressions.
If class characteristics and size matches: investigators can determine that footwear could have made the impression, but other footwear with similar design and size could also have made the impression.
If class characteristics, size match and randomly acquired characteristics found in the footwear outsole can also be found in the crime scene impressions: Investigators can determine that the specific piece created the crime scene impression. This relationship can be used as evidence to prove that the footwear's owner was at the crime scene.
Latent footwear impression found at a crime scene
Latent footwear impression after treatment with fingerprint powder
A suspect's footwear 1
A suspect's footwear 2
Suspect's footwear class characteristics
Suspect's footwear with unique characteristics labeled
Comparison of crimescene impression with suspect's footwear (labeled)
Comparison of crimescene impression with suspect's footwear (closeup)
Limitations of footwear evidence[edit]
The Unabomber, Theodore Kaczynski, was known to keep shoes with smaller soles attached to the base in order to confuse investigators about the size of the suspect's feet.[3]
Footwear databases[edit]
Forensic investigators can use computerized footwear databases to quickly compare the class characteristics such as tread design and possibly identify a manufacturer, brand, and model to the ones stored in the database.
Everspry http://www.footprintmatcher.com/products/products_01.htm
[forensity] https://forensity.com/index.php/en/product-en
Foster and Freeman http://www.fosterfreeman.com/trace-evidence/356-sicar-6-solemate.html
Raven technologies http://raven-technology.com/products/footwear/
Footwear Software[edit]
DigTrace http://www.digtrace.co.uk/
High profile cases[edit]
This section is empty. You can help by adding to it. (January 2015)
References[edit]
Jump up ^ "3D Scanning: A New Tool for Cracking Tough Cases". Forensic Magazine. 2009-02-01. Retrieved 2016-12-01.
Jump up ^ Davis, Kevin T. The foot length to stature ratio: a study of racial variance. Diss. Texas Tech University, 1990.
Jump up ^ "FOXNews.com - Newly Released Unabomber Evidence Offers New Insights". Fox News. 2006-11-29. To evade authorities chasing him, Unabomber Theodore Kaczynski kept shoes with smaller soles attached to the bottom in his reclusive Montana cabin, according to evidence released 10 years after his capture
External links[edit]
Scientific Working Group on Shoeprint and Tire Tread Evidence (SWGTREAD)
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Biological dispersal
From Wikipedia, the free encyclopedia
Jump to: navigation, search
This article is about biological dispersal in ecosystems. For other forms of dispersion, see Dispersion (disambiguation).
Wind dispersal of dandelion seeds.
Biological dispersal refers to both the movement of individuals (animals, plants, fungi, bacteria, etc.) from their birth site to their breeding site ('natal dispersal'), as well as the movement from one breeding site to another ('breeding dispersal'). Dispersal is also used to describe the movement of propagules such as seeds and spores. Technically, dispersal is defined as any movement that has the potential to lead to gene flow.[1] The act of dispersal involves three phases: departure, transfer, settlement and there are different fitness costs and benefits associated with each of these phases.[2] Through simply moving from one habitat patch to another, the dispersal of an individual has consequences not only for individual fitness, but also for population dynamics, population genetics, and species distribution.[3][4][5] Understanding dispersal and the consequences both for evolutionary strategies at a species level, and for processes at an ecosystem level, requires understanding on the type of dispersal, the dispersal range of a given species, and the dispersal mechanisms involved.
Biological dispersal may be contrasted with geodispersal, which is the mixing of previously isolated populations (or whole biotas) following the erosion of geographic barriers to dispersal or gene flow (Lieberman, 2005;[6][7] Albert and Reis, 2011[8]).
Dispersal can be distinguished from animal migration (typically round-trip seasonal movement), although within the population genetics literature, the terms 'migration' and 'dispersal' are often used interchangeably.
Contents
[hide]
1 Types of dispersal
2 Costs and benefits
3 Dispersal range
3.1 Environmental constraints
3.2 Dispersal barriers
4 Dispersal mechanisms
4.1 Plant dispersal mechanisms
4.2 Animal dispersal mechanisms
4.2.1 Non-motile animals
4.2.2 Motile animals
4.2.3 Dispersal by dormant stages
5 Quantifying dispersal
6 Consequences of dispersal
7 See also
8 References
9 Further reading
10 External links
Types of dispersal[edit]
Dispersal from parent population
Some organisms are motile throughout their lives, but others are adapted to move or be moved at precise, limited phases of their life cycles. This is commonly called the dispersive phase of the life cycle. The strategies of organisms' entire life cycles often are predicated on the nature and circumstances of their dispersive phases.
In general there are two basic types of dispersal:
Density-independent dispersal
Organisms have evolved adaptations for dispersal that take advantage of various forms of kinetic energy occurring naturally in the environment. This is referred to as density independent or passive dispersal and operates on many groups of organisms (some invertebrates, fish, insects and sessile organisms such as plants) that depend on animal vectors, wind, gravity or current for dispersal.[9][10]
Density-dependent dispersal
Density dependent or active dispersal for many animals largely depends on factors such as local population size, resource competition, habitat quality, and habitat size.[11][12][13]
Due to population density, dispersal may relieve pressure for resources in an ecosystem, and competition for these resources may be a selection factor for dispersal mechanisms.[14]
Dispersal of organisms is a critical process for understanding both geographic isolation in evolution through gene flow and the broad patterns of current geographic distributions (biogeography).
A distinction is often made between natal dispersal where an individual (often a juvenile) moves away from the place it was born, and breeding dispersal where an individual (often an adult) moves away from one breeding location to breed elsewhere.[1]
Costs and benefits[edit]
Epilobium hirsutum - Seed head
In the broadest sense, dispersal occurs when the fitness benefits of moving outweigh the costs.
There are a number of benefits to dispersal such as locating new resources, escaping unfavorable conditions, avoiding competing with siblings, and avoiding breeding with closely related individuals which could lead to inbreeding depression.[12]
There are also a number of costs associated with dispersal, which can be thought of in terms of four main currencies: energy, risk, time and opportunity.[2] Energetic costs include the extra energy required to move as well as energetic investment in movement machinery (e.g. wings). Risks include increased injury and mortality during dispersal and the possibility of settling in an unfavorable environment. Time spent dispersing is time that often cannot be spent on other activities such as growth and reproduction. Finally dispersal can also lead to outbreeding depression if an individual is better adapted to its natal environment than the one it ends up in. In social animals (such as many birds and mammals) a dispersing individual must find and join a new group, which can lead to loss of social rank.[2]
Dispersal range[edit]
"Dispersal range" refers to the distance a species can move from an existing population or the parent organism. An ecosystem depends critically on the ability of individuals and populations to disperse from one habitat patch to another. Therefore, biological dispersal is critical to the stability of ecosystems.
Environmental constraints[edit]
Few species are ever evenly or randomly distributed within or across landscapes. In general, species significantly vary across the landscape in association with environmental features that influence their reproductive success and population persistence.[15][16] Spatial patterns in environmental features (e.g. resources) permit individuals to escape unfavorable conditions and seek out new locations.[17] This allows the organism to "test" new environments for their suitability, provided they are within animal's geographic range. In addition, the ability of a species to disperse over a gradually changing environment could enable a population to survive extreme conditions. (i.e. climate change).
As the climate changes, prey and predators have to adapt to survive. This poses a problem for many animals, for example the Southern Rockhopper Penguins.[18] These penguins are able to live and thrive in a variety of climates due to the penguins' phenotypic plasticity.[19] However, they are predicted to respond by dispersal, not adaptation this time.[19] This is explained due to their long life spans and slow microevolution. Penguins in the subantarctic have very different foraging behavior than the subtropical waters, it would be very hard to survive and keep up with the fast changing climate because these behaviors took years to shape.[18]
Dispersal barriers[edit]
A dispersal barrier may mean that the dispersal range of a species is much smaller than the species distribution. An artificial example is habitat fragmentation due to human land use. Natural barriers to dispersal that limit species distribution include mountain ranges and rivers. An example is the separation of the ranges of the two species of chimpanzee by the Congo River.
On the other hand, human activities may also expand the dispersal range of a species by providing new dispersal methods (e.g., ships). Many of them become invasive, like rats and stinkbugs, but some species also have a slightly positive effect to human settlers like honeybees and earthworms.[20]
Dispersal mechanisms[edit]
Most animals are capable of locomotion and the basic mechanism of dispersal is movement from one place to another. Locomotion allows the organism to "test" new environments for their suitability, provided they are within the animal's range. Movements are usually guided by inherited behaviors.
The formation of barriers to dispersal or gene flow between adjacent areas can isolate populations on either side of the emerging divide. The geographic separation and subsequent genetic isolation of portions of an ancestral population can result in speciation.
Plant dispersal mechanisms[edit]
Burs are an example of a seed dispersion mechanism which uses a biotic vector, in this case animals with fur.
Main article: Seed dispersal
Seed dispersal is the movement or transport of seeds away from the parent plant. Plants have limited mobility and consequently rely upon a variety of dispersal vectors to transport their propagules, including both abiotic and biotic vectors. Seeds can be dispersed away from the parent plant individually or collectively, as well as dispersed in both space and time. The patterns of seed dispersal are determined in large part by the dispersal mechanism and this has important implications for the demographic and genetic structure of plant populations, as well as migration patterns and species interactions. There are five main modes of seed dispersal: gravity, wind, ballistic, water and by animals.
Animal dispersal mechanisms[edit]
Non-motile animals[edit]
There are numerous animal forms that are non—motile, such as sponges, bryozoans, tunicates, sea anemones, corals, and oysters. In common, they are all either marine or aquatic. It may seem curious that plants have been so successful at stationary life on land, while animals have not, but the answer lies in the food supply. Plants produce their own food from sunlight and carbon dioxide—both generally more abundant on land than in water. Animals fixed in place must rely on the surrounding medium to bring food at least close enough to grab, and this occurs in the three-dimensional water environment, but with much less abundance in the atmosphere.
All of the marine and aquatic invertebrates whose lives are spent fixed to the bottom (more or less; anemones are capable of getting up and moving to a new location if conditions warrant) produce dispersal units. These may be specialized "buds", or motile sexual reproduction products, or even a sort of alteration of generations as in certain cnidaria.
Corals provide a good example of how sedentary species achieve dispersion. Corals reproduce by releasing sperm and eggs directly into the water. These release events are coordinated by lunar phase in certain warm months, such that all corals of one or many species on a given reef will release on the same single or several consecutive nights. The released eggs are fertilized, and the resulting zygote develops quickly into a multicellular planula. This motile stage then attempts to find a suitable substratum for settlement. Most are unsuccessful and die or are fed upon by zooplankton and bottom dwelling predators such as anemones and other corals. However, untold millions are produced, and a few do succeed in locating spots of bare limestone, where they settle and transform by growth into a polyp. All things being favorable, the single polyp grows into a coral head by budding off new polyps to form a colony.
Motile animals[edit]
Main articles: Animal locomotion and Animal migration
The majority of all animals are motile. Although motile animals can, in theory, disperse themselves by their spontaneous and independent locomotive powers, a great many species utilize the existing kinetic energies in the environment, resulting in passive movement. Dispersal by water currents is especially associated with the physically small inhabitants of marine waters known as zooplankton. The term plankton comes from the Greek, πλαγκτον, meaning "wanderer" or "drifter".
Dispersal by dormant stages[edit]
Many animal species, especially freshwater invertebrates, are able to disperse by wind or by transfer with an aid of larger animals (birds, mammals or fishes) as dormant eggs, dormant embryos or, in some cases, dormant adult stages. Tardigrades, some rotifers and some copepods are able to withstand desiccation as adult dormant stages. Many other taxa (Cladocera, Bryozoa, Hydra, Copepoda and so on) can disperse as dormant eggs or embryos. Freshwater sponges usually have special dormant propagules called gemmulae for such a dispersal. Many kinds of dispersal dormant stages are able to withstand not only desiccation and low and high temperature, but also action of digestive enzymes during their transfer through digestive tracts of birds and other animals, high concentration of salts and many kinds of toxicants. Such dormant-resistant stages made possible the long-distance dispersal from one water body to another and broad distribution ranges of many freshwater animals.
Quantifying dispersal[edit]
Dispersal is most commonly quantified either in terms of rate or distance.
Dispersal rate (also called migration rate in the population genetics literature) or probability describes the probability that any individual leaves an area or, equivalently, the expected proportion of individual to leave an area.
The dispersal distance is usually described by a dispersal kernel which gives the probability distribution of the distance traveled by any individual. A number of different functions are used for dispersal kernels in theoretical models of dispersal including the negative exponential distribution,[21] extended negative exponential distribution,[21] normal distribution,[21] exponential power distribution,[22] inverse power distribution,[21] and the two-sided power distribution.[23] The inverse power distribution and distributions with 'fat tails' representing long-distance dispersal events (called leptokurtic distributions) are though to best match empirical dispersal data.[21][24]
Consequences of dispersal[edit]
Dispersal not only has costs and benefits to the dispersing individual (as mentioned above), but it also has consequences at the level of the population and species as well.
Most populations have a patchy spatial distribution. Dispersal, by moving individuals between different sub-populations, can increase the overall connectivity of the population, helping to minimize the risk of stochastic extinction, since if a sub-population goes extinct by chance, it is likely to be recolonized if the dispersal rate is high. Increased connectivity can also decrease the degree of local adaptation.[25]
See also[edit]
Competition (biology)
Disturbance (ecology)
Dormancy ('dispersal in time')
Gene flow
Habitat fragmentation
Island hopping
Landscape ecology
Metapopulation
Oceanic dispersal
Phoresy
Population modeling
Population distribution
Population ecology
Species distribution
References[edit]
^ Jump up to: a b Ronce, O. (2007). "How does it feel to be like a rolling stone? Ten questions about dispersal evolution". Annual Review of Ecology, Evolution, and Systematics. 38: 231–253. doi:10.1146/annurev.ecolsys.38.091206.095611.
^ Jump up to: a b c Bonte, D.; Van Dyck, H.; Bullock, J. M.; Coulon, A.; Delgado, M. D. M.; Gibbs, M.; et al. (2012). "Costs of dispersal". Biological Reviews of the Cambridge Philosophical Society. 87 (2): 290–312. doi:10.1111/j.1469-185X.2011.00201.x. PMID 21929715.
Jump up ^ Dunning, J. B. J.; Stewart, D. J.; Danielson, B. J.; Noon, B. R.; Root, T. L.; Lamberson, R.H. & Stevens, E. E. (1995). "Spatially explicit population models: current forms and future uses" (PDF). Ecological Applications. 5 (1): 3–11. doi:10.2307/1942045. JSTOR 1942045.
Jump up ^ Hanski, I.; Gilpin, M. E., eds. (1997). Metapopulation biology : ecology, genetics and evolution. San Diego: Academic Press. ISBN 0-12-323446-8.
Jump up ^ Hanski, I. (1999). Metapopulation Ecology. Oxford: Oxford University Press. ISBN 0-19-854065-5.
Jump up ^ Lieberman (2005). "Geobiology and paleobiogeography: tracking the coevolution of the Earth and its biota". Palaeogeography, Palaeoclimatology, Palaeoecology. 219: 23–33. doi:10.1016/j.palaeo.2004.10.012.
Jump up ^ Lieberman, Bruce S. (2005). "Geobiology and paleobiogeography: Tracking the coevolution of the Earth and its biota". Palaeogeography, Palaeoclimatology, Palaeoecology. 219: 23. doi:10.1016/j.palaeo.2004.10.012.
Jump up ^ James S. Albert; Roberto E. Reis (2011). Historical Biogeography of Neotropical Freshwater Fishes. University of California Press. p. 308. ISBN 978-0-520-26868-5.
Jump up ^ Maguire Jr., B. (1963). "The passive dispersal of small aquatic organisms and their colonization of isolated bodies of water". Ecological Monographs. 33: 161. doi:10.2307/1948560. JSTOR 1948560.
Jump up ^ Nathan, R. (2001). "The challenges of studying dispersal" (PDF). Trends in Ecology & Evolution. 16 (9): 481. doi:10.1016/S0169-5347(01)02272-8. Archived from the original (PDF) on 2013-09-28.
Jump up ^ Johst, K. & Brandl, R. (1997). "The effect of dispersal on local population dynamics". Ecological Modelling. 104: 87. doi:10.1016/S0304-3800(97)00112-9.
^ Jump up to: a b Bowler, D. E. & Benton, T.G. (2005). "Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics". Biol. Rev. 80 (2): 205–225. doi:10.1017/S1464793104006645. PMID 15921049.
Jump up ^ Fernández, L. (2015). "Source-sink dynamics shapes the spatial distribution of soil protists in an arid shrubland of northern Chile". Journal of Arid Environment. 113: 121–125. doi:10.1016/j.jaridenv.2014.10.007.
Jump up ^ Irwini, A.J. & Taylor, P.D. (2007). "Evolution of Dispersal in a Stepping-Stone Population with Overlapping Generations" (PDF). Theoretical Population Biology. 58 (4): 321–8. doi:10.1006/tpbi.2000.1490. PMID 11162790.
Jump up ^ Groom MJ, Meffe GK, Carroll CR (2005). Principles of Conservation Biology. Sunderland (MA): Sinauer Associates, Inc. ISBN 0-87893-597-5.
Jump up ^ Martin TE (1998). "Are microhabitat preferences of coexisting species under selection and adaptive?" (PDF). Ecology. 79 (2): 656–670. doi:10.1890/0012-9658(1998)079[0656:AMPOCS]2.0.CO;2.
Jump up ^ Turner MG, Gardner RH, O'Neill, Robert V (2001). "Landscape Ecology: In theory and practice, pattern and process". New York: Springer. ISBN 0-387-95123-7.
^ Jump up to: a b Tremblay, Yann (April 11, 2003). "Geographic variation in the foraging behaviour, diet and chick growth of rockhopper penguins" (PDF). Marine Ecology Progress Series. doi:10.3354/meps251279.
^ Jump up to: a b Forcada, Jaume; Trathan, Philip N. (2009-07-01). "Penguin responses to climate change in the Southern Ocean". Global Change Biology. 15 (7): 1618–1630. doi:10.1111/j.1365-2486.2009.01909.x. ISSN 1365-2486.
Jump up ^ http://ngm.nationalgeographic.com/2007/05/jamestown/charles-mann-text/2
^ Jump up to: a b c d e Chapman, D. S.; Dytham, C. & Oxford, G. S. (2007). "Modelling population redistribution in a leaf beetle: an evaluation of alternative dispersal functions". Journal of Animal Ecology. 76 (1): 36–44. doi:10.1111/j.1365-2656.2006.01172.x. PMID 17184351.
Jump up ^ Zhou, Y. & Kot, M. (2011). "Discrete-time growth-dispersal models with shifting species ranges". Theoretical Ecology. 4 (1): 13–25. doi:10.1007/s12080-010-0071-3.
Jump up ^ Miller, T.; Shaw, A. K.; Inouye, B. D. & Neubert, M. G. (2011). "Sex-biased dispersal and the speed of two-sex invasions". The American Naturalist. 177 (5): 549–561. doi:10.1086/659628. PMID 21508603.
Jump up ^ Radinger, J. & Wolter, C. (2014). "Patterns and predictors of fish dispersal in rivers". Fish and Fisheries. 15 (3): 456–473. doi:10.1111/faf.12028.
Jump up ^ Lenormand, T. (2002). "Gene flow and the limits to natural selection". Trends in Ecology and Evolution. 17 (4): 183–189. doi:10.1016/S0169-5347(02)02497-7.
Further reading[edit]
Ingold, C. T. (1971) Fungal spores: their liberation and dispersal Oxford, Clarendon Press 302 p. ISBN 0-19-854115-5
Lidicker, W. Z. and R. L. Caldwell (1982) Dispersal and migration Stroudsburg, Pa. : Hutchinson Ross Pub. Co 311 p. ISBN 0-87933-435-5 (Dispersal of animals)
Bullock, J. M.; R. E. Kenward and R. S. Hails (editors) (2002) Dispersal ecology : the 42nd symposium of the British Ecological Society. Oxford, UK : Blackwell Science 458 p. ISBN 0-632-05876-5 (Animals and plants)
Lieberman, B. S. 2008. Emerging syntheses between palaeobiogeography and macroevolutionary theory. Proceedings of the Royal Society of Victoria 120: 51–57.
Turner M. G., Gardner R. H., O'Neill. 2001. Landscape Ecology: In theory and practice, pattern and process. New York (NY): Springer Science.
Levin S. A. (1974). "Dispersion and Population Interactions". The American Naturalist. 108 (960): 207. doi:10.1086/282900.
Clobert, J., E. Danchin, A. A. Dhondt and J. D. Nichols (eds.)(2001) "dispersal", Oxford University Press, Oxford.
Cousens, R., C. Dytham, R. Law (2008), Dispersal in plants: a population perspective, Oxford University Press, Oxford
External links[edit]
Fruit and seed dispersal images at bioimages.vanderbilt.edu
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