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2<HTML>
3<HEAD>
4<TITLE>fitch</TITLE>
5<META NAME="description" CONTENT="fitch">
6<META NAME="keywords" CONTENT="fitch">
7<META NAME="resource-type" CONTENT="document">
8<META NAME="distribution" CONTENT="global">
9<META HTTP-EQUIV="Content-Type" CONTENT="text/html; charset=iso-8859-1">
10</HEAD>
11<BODY BGCOLOR="#ccffff">
12<DIV ALIGN=RIGHT>
13version 3.6
14</DIV>
15<P>
16<DIV ALIGN=CENTER>
17<H1>FITCH -- Fitch-Margoliash and Least-Squares Distance Methods</H1>
18</DIV>
19<P>
20&#169; Copyright 1986-2002 by the University of
21Washington.  Written by Joseph Felsenstein.  Permission is granted to copy
22this document provided that no fee is charged for it and that this copyright
23notice is not removed.
24<P>
25This program carries out Fitch-Margoliash, Least Squares,
26and a number of similar methods as described in the documentation
27file for distance methods.
28<P>
29The options for FITCH are selected through the menu, which looks like
30this:
31<P>
32<TABLE><TR><TD BGCOLOR=white>
33<PRE>
34
35Fitch-Margoliash method version 3.6a3
36
37Settings for this run:
38  D      Method (F-M, Minimum Evolution)?  Fitch-Margoliash
39  U                 Search for best tree?  Yes
40  P                                Power?  2.00000
41  -      Negative branch lengths allowed?  No
42  O                        Outgroup root?  No, use as outgroup species  1
43  L         Lower-triangular data matrix?  No
44  R         Upper-triangular data matrix?  No
45  S                        Subreplicates?  No
46  G                Global rearrangements?  No
47  J     Randomize input order of species?  No. Use input order
48  M           Analyze multiple data sets?  No
49  0   Terminal type (IBM PC, ANSI, none)?  (none)
50  1    Print out the data at start of run  No
51  2  Print indications of progress of run  Yes
52  3                        Print out tree  Yes
53  4       Write out trees onto tree file?  Yes
54
55  Y to accept these or type the letter for one to change
56
57</PRE>
58<P>
59</TD></TR></TABLE>
60<P>
61Most of the input options
62(U, P, -, O, L, R, S, J, and M) are as given in
63the documentation page for distance matrix programs, and their input format is
64the same as given there.  The
65U (User Tree) option has one additional feature when the N (Lengths)
66option is used.  This menu option will appear only if the U (User Tree)
67option is selected.  If N (Lengths) is set to "Yes" then if any branch
68in the user tree has a branch length, that branch will not have its
69length iterated.  Thus you can prevent all branches from having their
70lengths changed by giving them all lengths in the user tree, or hold
71only one length unchanged by giving only that branch a length (such
72as, for example, 0.00).  You may find program RETREE useful for
73adding and removing branch lengths from a tree.  This option can
74also be used to compute the Average Percent Standard Deviation for a
75tree obtained from NEIGHBOR, for comparison with trees obtained by
76FITCH or KITSCH.
77<P>
78The D (methods) option allows choice between the Fitch-Margoliash
79criterion and the Minimum Evolution method (Kidd and Sgaramella-Zonta, 1971;
80Rzhetsky and Nei, 1993).  Minimum Evolution (not to be confused with
81parsimony) uses the Fitch-Margoliash criterion to fit branch lengths to each
82topology, but then chooses topologies based on their total branch length
83(rather than the goodness of fit sum of squares).  There is no
84constraint on negative branch lengths in the Minimum Evolution method;
85it sometimes gives rather strange results, as it can like solutions
86that have large negative branch lengths, as these reduce the total
87sum of branch lengths!
88</PRE>
89<P>
90Another input option available in FITCH that is not available in KITSCH
91or NEIGHBOR
92is the G (Global) option.  G is the
93Global search option.  This causes, after the last species is added to
94the tree, each possible group to be removed and re-added.  This improves the
95result, since the position of every species is reconsidered.  It
96approximately triples the run-time of the program.  It is not an option in
97KITSCH because it is the default and is always in force there.  The
98O (Outgroup) option is described in the main
99documentation file of this package.  The O option has no effect if the
100tree is a user-defined tree (if the U option is in effect).  The
101U (User Tree) option requires an unrooted tree; that is, it require
102that the tree have a trifurcation at its base:
103<P>
104&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;((A,B),C,(D,E));
105<P>
106The output consists of an unrooted tree and the lengths of the
107interior segments.  The sum of squares is printed out, and if <EM>P = 2.0</EM>
108Fitch and Margoliash's "average percent standard deviation" is also computed
109and printed out.  This is the sum of squares, divided by
110N-2, and
111then square-rooted and then multiplied by 100 (n is the number of
112species on the tree):
113<P>
114&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;APSD = ( SSQ / (N-2) )<SUP>1/2</SUP> x 100.
115<P>
116where <EM>N</EM> is the total number of off-diagonal distance measurements that
117are in the (square) distance matrix.  If the S (subreplication) option
118is in force it is instead the sum of the numbers of replicates in all the
119non-diagonal
120cells of the distance matrix.  But if the L or R option is also in effect, so
121that the distance matrix read in is lower- or upper-triangular, then the
122sum of replicates is only over those cells actually read in.  If S is not in
123force, the number of replicates in
124each cell is assumed to be 1, so that <EM>N</EM> is <EM>n(n-1)</EM>,
125where <EM>n</EM> is the number
126of species.  The <EM>APSD</EM> gives an indication of the average percentage
127error.  The number of trees examined is also printed out.
128<P>
129The constants
130available for modification at the beginning of the program are:
131"smoothings", which gives the number of passes through
132the algorithm which adjusts the lengths of the segments of the tree so
133as to minimize the sum of squares, "delta", which controls the size of
134improvement in sum of squares that is used to control the number of
135iterations improving branch lengths,
136and "epsilonf",
137which defines a small quantity needed in
138some of the calculations.  There is no feature saving multiply trees
139tied for best,
140partly because we do not expect exact ties except in cases where the branch
141lengths make the nature of the tie obvious, as when a branch is of zero
142length.
143<P>
144The algorithm can be slow.  As the number of species
145rises, so does the number of distances from each species to the others.  The
146speed of this algorithm will thus rise as the fourth power of the
147number of species, rather than as the third power as do most of the
148others.  Hence it is expected to get very slow as the number of species
149is made larger.
150<P>
151<HR>
152<P>
153<H3>TEST DATA SET</H3>
154<P>
155<TABLE><TR><TD BGCOLOR=white>
156<PRE>
157    7
158Bovine      0.0000  1.6866  1.7198  1.6606  1.5243  1.6043  1.5905
159Mouse       1.6866  0.0000  1.5232  1.4841  1.4465  1.4389  1.4629
160Gibbon      1.7198  1.5232  0.0000  0.7115  0.5958  0.6179  0.5583
161Orang       1.6606  1.4841  0.7115  0.0000  0.4631  0.5061  0.4710
162Gorilla     1.5243  1.4465  0.5958  0.4631  0.0000  0.3484  0.3083
163Chimp       1.6043  1.4389  0.6179  0.5061  0.3484  0.0000  0.2692
164Human       1.5905  1.4629  0.5583  0.4710  0.3083  0.2692  0.0000
165</PRE>
166</TD></TR></TABLE>
167<P>
168<HR>
169<P>
170<H3>OUTPUT FROM TEST DATA SET (with all numerical options on)</H3>
171<P>
172<TABLE><TR><TD BGCOLOR=white>
173<PRE>
174
175   7 Populations
176
177Fitch-Margoliash method version 3.6a3
178
179                  __ __             2
180                  \  \   (Obs - Exp)
181Sum of squares =  /_ /_  ------------
182                                2
183                   i  j      Obs
184
185Negative branch lengths not allowed
186
187
188Name                       Distances
189----                       ---------
190
191Bovine        0.00000   1.68660   1.71980   1.66060   1.52430   1.60430
192              1.59050
193Mouse         1.68660   0.00000   1.52320   1.48410   1.44650   1.43890
194              1.46290
195Gibbon        1.71980   1.52320   0.00000   0.71150   0.59580   0.61790
196              0.55830
197Orang         1.66060   1.48410   0.71150   0.00000   0.46310   0.50610
198              0.47100
199Gorilla       1.52430   1.44650   0.59580   0.46310   0.00000   0.34840
200              0.30830
201Chimp         1.60430   1.43890   0.61790   0.50610   0.34840   0.00000
202              0.26920
203Human         1.59050   1.46290   0.55830   0.47100   0.30830   0.26920
204              0.00000
205
206
207  +---------------------------------------------Mouse     
208  !
209  !                                +------Human     
210  !                             +--5
211  !                           +-4  +--------Chimp     
212  !                           ! !
213  !                        +--3 +---------Gorilla   
214  !                        !  !
215  1------------------------2  +-----------------Orang     
216  !                        !
217  !                        +---------------------Gibbon   
218  !
219  +------------------------------------------------------Bovine   
220
221
222remember: this is an unrooted tree!
223
224Sum of squares =     0.01375
225
226Average percent standard deviation =     1.85418
227
228Between        And            Length
229-------        ---            ------
230   1          Mouse             0.76985
231   1             2              0.41983
232   2             3              0.04986
233   3             4              0.02121
234   4             5              0.03695
235   5          Human             0.11449
236   5          Chimp             0.15471
237   4          Gorilla           0.15680
238   3          Orang             0.29209
239   2          Gibbon            0.35537
240   1          Bovine            0.91675
241
242
243</PRE>
244</TD></TR></TABLE>
245<P>
246</BODY>
247</HTML>
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