lecture 1 - introduction to geometric morphometrics

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Department of Geological Sciences | Indiana University (c) 2012, P. David Polly G562 Geometric Morphometrics Rychlik, L., G. Ramalhinho, and P. D. Polly. 2006. J. Zool. Syst. Evol. Res. 44: 339-351. An introduction Geometric morphometrics P. David Polly Department of Geology, Indiana (Biology and Anthropology) University, 1001 E. 10th Street, Bloomington, IN 47405, USA [email protected] http://mypage.iu.edu/~pdpolly/

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Page 1: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Rychlik, L., G. Ramalhinho, and P. D. Polly. 2006. J. Zool. Syst. Evol. Res. 44: 339-351.

An introductionGeometric morphometrics

P. David PollyDepartment of Geology, Indiana(Biology and Anthropology)University, 1001 E. 10th Street, Bloomington, IN 47405, [email protected]://mypage.iu.edu/~pdpolly/

Page 2: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Course websitehttp://www.indiana.edu/~g562/

Page 3: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Spring 2014 Syllabus

Date Topic Assignments17#Jan#14 Introduction0to0Geometric0Morphometric0Methods0(GMM)

24#Jan#14 Applied:0Introduction0to0Mathematica Assignment01

31#Jan#14 Aopplied:0First0GMM0Analysis Assignment02

7#Feb#14 No0Meeting READING01

14#Feb#14 Applied:0Procrustes,0Principal0components0analysis0and0morphospaceAssignment03

21#Feb#14 Discussion:0Multivariate0tests0and02B#PLS0 READING02

28#Feb#14 Applied:0Multivariate0tests0for0GMM0(proposals0due) Assignment04

7#Mar#14 Discussion:0Randomization0and0Bootstrap READING03

14#Mar#14 Applied:0Bootstrapping0and0randomization0tests Assignment05

21#Mar#14 No0Meeting:00Spring0Break

28#Mar#14 Discussion:0Outlines0and0EDMA READING04

4#Apr#14 Applied:0Outline0analysis0and0EDMA Assignment06

11#Apr#14 Discussion:0Phylogenetic0trees0and0morphometrics READING05

18#Apr#14 Applied:0phylogenetic0trees0and0morphometrics Assignment07

25#Apr#14 Discussion:0evolutionary0and0growth0trajectories READING06

2#May#14 Applied:0Monte0Carlo0simulations Assignment07

5#May#14 Projects0Due:00Monday,0May05

Page 4: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Computers and Mathematica®Laptops should be brought to class every week. The software we will use is compatible with both Mac and Microsoft operating systems.

General software: If possible, please install the following on your system: ImageJ (http://rsbweb.nih.gov/ij/ ), Adobe Creative Suite (especially Photoshop and Illustrator, http://iuware.iu.edu/ ), Microsoft Office or Mac iWork or OpenOffice (for spreadsheet programs, http://iuware.iu.edu/ ).

Mathematica®: Most of our work will be done in the Mathematica which is a mathematical and statistical application that does efficient computations, is flexible for customized analysis, has powerful graphics capabilities, and is easy to learn (compared to R, for example, which is an equivalent application). You can purchase a student license for Mathematica 8.04 at the IU Stat/Math Center at 410 Park Avenue for $30 (cash or check only). They will provide licence code and download link for installation.

Page 5: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Participation 20% Assignments 20% Reading reports 10% Project proposal 10% (due 7 March) Project report 40% (due 5 May)

Grading

Page 6: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

ProjectsThe focal assignment for the course is to carry out an original morphometric study that you will present as a written project paper. Ideally your project will be related to your broader research interests. During the first half of the course you will develop your study, with advice if you need/want it.

Project proposals are due mid semester. Proposals consist of a one to two page description of the aim and subject of your project, the source of your data, the question your study will try to answer, and the tools you’ll need to collect your data. Please feel free to consult me as you develop your proposal.

The final project report should be modelled on a scientific journal paper, with introduction to the problem, materials and methods, results, discussion, conclusion, references (cited properly in the journal format of your choice). Target length is 10-15 pages, including figures, tables, and references.

Page 7: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Supplementary texts

Hammer, Ø. and D.A.T. Harper. 2006. Palaeontological Data Analysis. Blackwell Publishing, Oxford, United Kingdom.

Zelditch, M.L., D.L. Swiderski, H.D. Sheets, and W.L. Fink. 2004. Geometric Morphometrics for Biologists: a Primer. Elsevier Academic Press, San Diego, California.

MacLeod, N. 2012. Palaeo Math 101. A series of essays published in the Palaeontological Association Newsletter. http://www.palass.org/modules.php?name=palaeo_math

Page 8: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

What are morphometrics?

Any quantitative measurement and analysis of morphological traits

Mandible Length (cm)

Man

dib

le H

eig

ht

(cm

)

Shrew

Marmot

Page 9: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Geometric morphometricsthe quantitative representation and analysis of morphological shape using geometric coordinates instead of measurements

Component 1 Com

ponent 2

Shrew

Marmot

Page 10: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

The Major Goal of Morphometrics:Measuring morphological similarity and difference

Page 11: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Types of geometric representationOutlines (2D or 3D)Semilandmarks, sliding landmarks

3D surfaces

Landmarks (2D or 3D)

Polly, P. D. 2008. Adaptive Zones and the Pinniped Ankle: A 3D Quantitative Analysis of Carnivoran Tarsal Evolution.  Pp. 165-194 in (E. Sargis and M. Dagosto, Eds.) Mammalian Evolutionary Morphology: A Tribute to Frederick S. Szalay. 

Springer: Dordrecht, The Netherlands.

Polly, P. D. 2003. Paleophylogeography: the tempo of geographic differentiation in marmots

(Marmota). Journal of Mammalogy, 84: 369-384.

Page 12: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Definitions

Landmark – any point described with cartesian coordinates (x, y, z) used to represent the shape of a structure.

Landmark (2) – any point that can be placed on a biologically or geometrically homologous point on the structure.

Semi-landmark – a point that is placed arbitrarily using an algorithm, often by defining endpoints at biologically homologous points and placing a specified number of semilandmarks between them.

Page 13: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

0 10 0

10

5

5

X-axis

Y-ax

is

(2,5)

(4,8)

(5,2)

(6,8)

(6,2)

Landmarks

Landmarks are coordinate points used to represent a shape

They are quantified as Cartesian coordinates (x,y[,z])

At least 3 are required (two points make a line)

Example analyses: Relative Warps (PCA of landmarks), Euclidean Distance Matrix Analysis (EDMA) of distances between landmarks

Page 14: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

0 10 0

10

5

5

X-axis

Y-ax

is

Outlines

Outlines are perimeters delimited by many points

They are quantified as Cartesian coordinates (x,y[,z]), often converted to angles

Many points are required to represent a shape

Example analyses: Semilandmarks, sliding semilandmarks, Eigenshape (PCA of outline), Fourier analysis

Page 15: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Polly, P. D. 2008. Adaptive Zones and the Pinniped Ankle: A 3D Quantitative Analysis of Carnivoran Tarsal Evolution.  Pp. 165-194 in (E. Sargis and M. Dagosto, Eds.) Mammalian Evolutionary Morphology: A Tribute to Frederick S. Szalay. 

Springer: Dordrecht, The Netherlands.

Surfaces

Surfaces are the 3D surface of an object

They are quantified as Cartesian coordinates (x,y,z)

Many points are required to represent a shape

Example analysis: Eigensurface (PCA of surfaces), sliding semilandmarks

Page 16: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Advantages to geometric representation

Results can be presented visually as a “shape” than tables of numbers

Data are easily collected from digital photographs

Size is mathematically removed from the analysis to focus on pure shape

X-axis

Y-ax

is

Page 17: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Y-ax

is

X-axis

Y-ax

is

Vector Plot

Spline Plot

GMM results can be presented graphicallyTwo examples of graphically showing those differencesDifference in shape of mandibles

of shrew and marmot

Page 18: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Marmot

Shrew

Traditional morphometrics mixes size and shapeThe size of the animal affects all measurements so that primary morphometric difference between two taxa is size rather than shape

Marmot

Shrew

0

5

10

15

20

25

Mill

imet

res

H L H L

Page 19: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Marmot

Shrew

Geometric morphometrics removes size by rescalingShapes are enlarged or reduced to achieve a standard, equal size

Coordinates of rescaled landmarks show difference in relative position only

X-axis

Y-ax

is

Page 20: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Disadvantages of geometric representationSize is completely absent from the analysis, and size may be biologically relevant

Only single rigid structures can be easily analyzed

Page 21: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

-1.50

-1.00

-0.50

0.00

0.50

1.00

1.50

-4.0 -2.0 0.0 2.0 4.0 Log body mass (kg)

Log

met

abol

ic ra

te (O

2 /hr

/g)

data from Eisenberg, 1981

Size is biologically importantand it may be of interest in a morphometric analysis

Page 22: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Polly, P. D. 1998. Variability in mammalian dentitions: size-related bias in the coefficient of variation. Biological Journal of

the Linnean Society, 64: 83-99.

Size and shape may behave differentlySize or shape data may be appropriate for different analyses

Page 23: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Only single rigid structures can represented with geometric morphometrics

Okay Not okay

Page 24: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

A short history of geometric morphometrics....

Page 25: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Albrecht Dürer (1471 -1528)

Page 26: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

On Growth and Form, 1917

D’Arcy Thompson (1860-1948)

Page 27: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Francis Galton (1822-1911)

1891: starts biometric laboratory at University College London

Biometric approach to genetics: regression & correlation

Composite portraiture: photographs of different subjects combined (through repeated limited exposure) to produce a single blended image

Anthropometry & differential psychology: quantitative analysis of fingerprints

Page 28: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Modern Geometric Morphometric Methods (GMM)Development of landmark geometrics was driven by Fred Bookstein (long of University of Michigan, now Washington and Vienna)

Joined very productively by F. James Rohlf (Stony Brook)

Ian Dryden, Kanti Mardia, Les Marcus, and Dennis Slice have been important names in developing techniques and theory.

Bookstein was originally intent on creating a truly quantitative way of producing d’Arcy Thompson transformation grids.

Bookstein

Page 29: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Steps in a geometric morphometric study

Study design

Data collection

Data standardization

Analysis

Results interpretation

Page 30: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

How do you choose landmarks (or outlines, or surfaces)?1. The data must reflect a hypothesis

2. The data must represent the shape adequately

3. Landmarks must be present on all specimens

Measurement Error and Sample size

1. Measurement error (ME) always exists in any collection of data, but ME doesn’t matter if it is substantially less than the differences you want to measure.

2. Sample size required for a particular study depends on the within-group variation relative to differences between groups.

Page 31: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

How many specimens do I need?

• Depends on the question being addressed

• Depends on the error in your data

• You need more specimens when the differences you want to measure are small compared to the variation within your group (natural or due to error)

• For sexual dimorphism in skulls of humans or other primates, 10 individuals of each sex might be enough

• For differences in genetic strains of mice where the mutation doesn’t obviously affect the skeleton, 50 individuals of each strain is more realistic

• For species that belong to different families or orders, 1 specimen per species is almost always sufficient

Page 32: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

What morphometrics can’t answer for you..

• Morphometrics does not tell you what ‘large’ or ‘difference’ or ‘shape’ mean(These are definitions you must supply and your results depend upon them)

• Morphometrics does not tell you whether you unwittingly have two unrecognized groups in a single sample(Although comparison with known groups may help such an endeavour)

• How to identify cladistic characters(For the first two reasons combined)

Page 33: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Examples of available software

Digitizing landmarks and outlines: tpsDIG, ImageJ

Superimposition: Morpheus (plus integrated in some below)

Outline analysis: Eigenshape, PAST

MANOVA: Statistica, PAST

Discriminant functions, CVA: Statistica, PAST

Principal components analysis of landmarks: tpsRELW, PST

Construction of trees: PHYLIP, PAUP, NTSYSpc, PAST

Simulations: Mathematica, R

Links and downloads at SUNY Stony Brook morphometrics site: http://life.bio.sunysb.edu/morph/

Page 34: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Equipment: 2D outlines and coordinates

High-quality digital cameras

(resolution doesn’t matter as much as the possibility of lens distortion: test your camera first by photographing a piece of graph paper and looking for “fish eye” distortion)

Calipers or scale bar

Page 35: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

Equipment: 3D outlines and coordinates

Reflex Microscope for collectingthree-dimensional landmarks, outlines and measurements (good for objects the size of a cat skull down to things about 2-5 mm long)

Microscribe robotic arm for collecting 3D landmarks and measurements (good for objects the size of a human skull down to a rat skull)

Page 36: Lecture 1 - Introduction to Geometric Morphometrics

Department of Geological Sciences | Indiana University (c) 2012, P. David Polly

G562 Geometric Morphometrics

3D surfaces

Microscan Laser scanner for scanning surfaces (good for objects the size of a cat skull down to ones about 2-3 cm long)

NextEngine laser scanner (good for objects the size of a horse skull down to a single tarsal bone)