Phylogenetic Analysis

Martin Cavalluzzi martin at VIMS.EDU
Sun Mar 2 10:48:05 CST 1997

Here is the compilation of the responses that I received regarding
my inquiry on phylogenetic analysis packages.  I thank the 13 individuals
who responded in a professional manner.  Their comments regarding
the four packages that I asked about, as well as others that are new
to me, are much appreciated. I also received one response "flaming" me for
using TAXACOM instead of the Web (strange but true).

URLs that have information on phylogenetic packages, analysis, and related

Comments regarding programs:

PAUP: latest released version was 3.1.1, date 1993, released by the
Illinois Natural History Survey.  This is now 'out of press' and
unobtainable.  A new version PAUP* is in beta test and will be published by
Sinauer sometime: maybe later this year if Dave Swofford can just get the
last few bugs ironed out?

PHYLIP: latest release is available from Joe Felsenstein's website at
University of Washington. cost zero.

Hennig86:  first and only version is numbered 1.1, date 1988.  It still is
available from Steve Farris or several of his authorised distributors. cost

MacClade:  I think the latest is version 3.0.6.  It is published by
Sinauer. cost something like $50-100 US.

Merits of each package:::
Well, MAcClade is not strictly a PHYLOGENETIC ANALYSIS package, its a
tree-exploration/ character mapping on trees/ etc, package.  Its for Macs
only.  It includes a rudimentary tree-search algorithm using an NNI branch
swapper and operating on only one tree in memory at one time, but if you
want a package to do cladistic tree searches this is not the one for you.
However, if your scene is exploring alternative trees, mapping characters
on trees, and generally indulging in what Willi Hennig called reciprocal
illumination (ie, between intiial tree estimates and a developing data
set), then MaClade is recommended.

Phylip has cheapness on its side, and it does most things that the other
phylogeny packages do.  It's a bit clumsy to use because the package
consists of three-dozen or so programs each for a specific task and you
have to keep track of infile and outfile names.  ALso, each program
requires its own type of data: eg, parsimony on binary characters is
separate from parsimony on protein characters is separate from parsimony on
dna sequence characters, etc. (It does not do parsimony on multi-state
characters, if ordered you can recode to additive binary, if unordered you
may be stuck.)  You can't do total-evidence cladistic analyses with Phylip,
combining different types of data.  The branch swappers are equivalent to
PAUP 3.1.1's medium-weight SPR swapper but slower, meaning tree search is
fairly effective for smallish data sets but gives out much earlier than the
more powerful search engines, generally at about 30 taxa.  Phylip includes
a good range of phenetic tree-building tools including such standard
methods as UPGMA and NJ.

PAUP 3.1.1 is strictly a cladistics package, and for Macs only.  It does
not offer phenetic tree-building.  The data format is the same as for
Maclade. (Well, more or less; at least the same file can be used.  PAUP
actually accepts a wider range of input format than does MacClade, eg
character weights in range 0-32767 as compared to 0-9999, maximum 32 states
per character as compared to 26.  But for ordinary analyses the same file
can be used in both programs.)  There's a full range of character-type,
character-weight, etc options, mixed analyses can be done as easily as any
other analyses, there are many options for tree searching and the branch
swappers are fast and effective. Heuristic searches up to 75 or so taxa
need no special search strategies, analyses with several hundred taxa have
been attempted. Branch-&-bound (ie, complete, non-heuristic) tree-search
operates up to about 16-20 taxa (depending how messy your data is) but is
not as fast as in PAUP* or Hennig86.  PAUP will do tree searches under
topological constraints, and it's the only program which will do so.  It
therefore is the only vehicle for techniques such as the FAITH T-PTP test
for the significance of groups on trees.  PAUP 3.1.1 is an EXCELLENT
all-round phylogenetic cladistics package.  Get one if you can, or get into
collaboration with someone who has an authorized copy.

Hennig86 is a PC cladistics package.  It is far, far, FAR less user
friendly that PAUP 3.1.1, and the 'manual' is totally incomprehensible
except to someone with a degree in c++.  Data matrices cannot have more
than 999 characters (unlimited in PAUP).  Only 10 states per character,
symbols 0-9 (32 in PAUP, symbols arbitrary).  Character types can be
individually declared  but only as either ordered or unordered (many other
options in PAUP, or invent your own data type via a cost-stepmatrix
function). Character weights integer 0-9 (0-32767 PAUP). Branch-&-bound
algorithm faster than PAUP 3.1.1, about the same asin  PAUP*, with it you
can get a complete list of every most-parsimonious tree for a 25-26 taxon
data set in around 3 days. (As you probably know, the time taken for
branch&bound rises very steeply with the number of taxa: 10 taxa 1 minute,
15 taxa 5 minutes, 20 taxa 10 minutes 25 taxa 3 days, 26 taxa 3 weeks, 27
taxa 3 months, 28 taxa 10 years ...).  Hennig86 does not do
random-addition-sequence searches or bootstraps at the press of a button
(PAUP does, and lots more beside).  Add-on shells which call Hennig86 as a
subroutine, such as Mark Siddal's package 'Random Cladistics', are
available for doing some but not all of these things.  If you have large
and messy data with lots of homoplasy, Hennig86 sometimes will find shorter
trees than PAUP 3.1.1 -- but just as often PAUP 3.1.1 will find shorter
trees than Hennig86.  This is because the two programs have different
starting tree and branch swapping routines.  When I'm working with a large
and messy morphological data set of >50 taxa, if PAUP starts to give me
multiple islands of trees I generally run the matrix once through Hennig86
to check it can't find something shorter.

PAUP* is the *Ferrari* of phylogenetics packages.  It combines the
capability of PAUP 3.1.1 with faster algorithms all-round (including the
all-important branch&bound).  It has oodles of new features, data
manipulations, tests relating to input (eg, partition homogeneity, PTP) and
tree output (eg, bootstrap, T-PTP).  Also its no longer just a cladistics
package, it offers all the phenetic methods found in Phylip PLUS many
others: eg, you can apply any sequence evolution model ever invented (not
just Kimura 2- or 3- parameter, but also many more) at the press of a
button.  It does maximum likelihood analysis too -- and it is both fast
(1000 times faster than Phylip) and more effective (because the likelihood
search is operating in conjunctionn with the PAUP TBR swapper).  Run all
sorts of tests, explore your data, exploreyour evolutionary model, evaluate
your assumptions, do anything!   Finally PAUP*, like PAUP 3.1.1 and
MacClade, will give a quality tree-print.

PAUP is undergoing revision, and is to be marketed soon by Sinauer; at
present, it is unavailable, except by "borrowing" from those who have it.
It only does Parsimony-type analysis, as the name implies (Phylogenetic
Analysis Using Parsimony), but it has a very nice interface (on Mac) and
is easy to get going with.

PHYLIP you can get free from Joe Felsenstein, via the net at:
It has versions for all platforms.  YOU MUST DOWNLOAD ALL THE
DOCUMENTATION! Otherwise you'll have a hell of a time running things.  It
has a variety of arcane details... i.e. it is NOT user friendly.  But I
really like it because it is general and open-ended, much more flexible
than PAUP (maybe the new edition of PAUP will be broader) and can do
pretty much all sorts of phylogenetic analysis, can apply bootstrapping to
any method, do consensus trees, etc.  Many of the routines are VERY fast,
at least on PowerMac.

MacClade is also marketed by Sinauer.  It is a file-editor and tree
manipulator and a variety of other stuff.  It doesn't do phylogenetic

Another free web-item is TreeView, for Mac or Windows from :
It is very fast and useful way of looking at trees from Nexus treefiles.

Thank you for your time.


Marty Cavalluzzi

martin at

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