FORUM

Cladisticc ( 1990)6:369-- 3 72

COMBINABLE COMPONENT CONSENSUS

Kare Bremer’

‘ Department o f $fycternatzc Botany, yptala I‘nzuei r i ~ y ,

Box 541, S-75121 C‘ppsala, Sweden

Page ( 1989a) distinguished between strict consensus trees and Nelson consensus trees.
T h e former were originally used by Nelson ( 1979) and Schuh and Polhemus ( 1980), and
named strict consensus trees by Sokal and Kohlf( 1981).Before Page’s distinction, strict
consensus trees were frequently termed Nelson trees, first by Schuh and Farris (1981).
For example, Mickevich a n d Platnick ( 1989: 42) stated that “a Nelson consensus
includes only those subtaxa exactly replicated [=consistently repeated I in tlic
cladograms being compared”. Following Page ( 1989a), this is the drfiriition o f a strict
consensus tree, whereas a Nelson tree includes “replicates [=replicated subtitxa]” n.r
well as “components [ = subtaxa] combinable [ = compatiblej with replicates and with
each other” (Nelson, 1979: 10; Nelson arid Platnick, 1981) .
For comparison of two cladograms (e.g. Schuh arid Farris, 198 1 ), the strict and the
Nelson consensus trees are often identical, but they may becomc dillercnt if‘ morr
cladograms arc included. I will elaborate on this below. I will arguc that thrrc is anothcr
related type of consensus tree available, and that this type may bc a useful tool in
systematic studies.
Consider the cladograms (top row) and the consensus trees (bottom row) in Fig. I .
They are partly taken from Page (1989a: fig. 12) and are based o n an examplr of
Indo-Australian biogeography from Nelson and Platnick (198 1 ) . T h e components of
the cladograms and the consensus trees are indicated with dots arid numbercd 1L12.
T h e strict consensus tree contains only those components occurring in all the
cladograms, i.e., component 6. Combinable ( =compatible) componerits present only in
some of the cladograms are riot included in a strict consensus tree. Component 7 is
present in the butterfly and bird cladograms but absent in the bat cladogram. I t is
combinable with the components of the strict consensus tree, but discarded.
Nelson trees are built in two steps. First, all replicated components are combined.
Replicated components are those repeated in at least two, but not necessarily all of the
cladograms compared. I n Fig. 1 the replicated components are 2, 3 , 4, 5, 6 and 7. Note
that of those, components 2, 3, 4, 5 and 7 are replicated in two o f t h e cladogrnms only.
In Fig. 1 and in Nelson’s (1979) original example all replicated components are
combinable. Nelson did riot compare cladograms with different, conflicting sets of
replicated components. Page ( 198%) dealt with this problem, and fbrmalized
construction of Nelson trees from cladograms containing non-combinable sets (cliques)
of replicated components. We d o not need to consider this complication further here. It
affects construction of Nelson trees only.
I n the second step of constructing a Nelson tree, all non-replicated components,
combinable both with the replicated ones a n d with each other, are added. In Fig. 1
components 1, 8, 9, 10, 11 and 12 are unique to single cladograms, i.e., non-replicated.
However, since they are all non-combinable either with the replicated components or
with each other. none is added to the Nelson tree.
370 ti. KREMISR

Butterfly Bird Bat

clodograrn clodogram clodograrn

I I

2 3

3 4
4 5

5 2
6 6
4
7 7 7
8 8
9 9

Combinoble
Nelson Adarns component Strict
consensus tree consensus tree consensus tree consensus tree

I I

2 2
3 3
4 4

5 5

6 6

7 7
8 8

9 9

Nelson’s rationale for accepting the rcplicated componeiits as “true” was the alleged
low prol)al)ility of component rrplication d u c to chance alone. Nelson further argued
that non-replicated components, which are non-combinable with the replicated “true”
oms, then can bc discarded as false, whereas the combinable oiics may be added to the
roriscnsus tree, incrrasiiig. its information content.
Nelson trrrs, like Adams trees (Adams, 1972) and majority r u l r consensus trecs
(Margush arid Mchlorris, 1981; I’enny rt al., 1982), may be ill conflict with thc
cladograms, since they m a y contain components that are non-combinable with the
components of some of‘the cladograms. I n the Nelson tree in Fig. I , components 2, 3 , 4
and 5 arc rion-combinable with components 8, 9, 10, 11 and 12 in the cladograms.
Majority rulc consensus trees according to Margush and McMorris ( 1981 ) contain
only those components that occur in more than halfofthe cladograms (slightly difkrent
dchnitions are applied by other authors, e.g. Penny et al., 1982). For the cladograms in
Fig. 1 , the majority rulc consensus tree is identical to the Nelson tree.
‘\dams (1972) consensus trees essentially work by pulling down unstable taxa and
components with diflercrlt positions in the cladograms to the first node suinmarizing the
dif'erent positions. For the claclograms in Fig. 1 , taxa 1 a r i d 2 a s well ;is c o n i p o ~ i e l i t s3 , 5
and 6 arc placed a t the basal iiodr in the resulting Adanis tree.
Since Nelson trees, Adams trees and majority rule cmisensus trees fi-cqucntly contain
components in conflict with some of thc cladograni components, these c o n s c n s ~ ~trees s
ma). be urisatisfirctory fbr tcmatic purposes. Instead, tcmatists oftcn rcsort to t h t .
more restrainrd, o r cautious, strict conscnsus trees, in in prcting their results. At [tic
s m i e time tlicy lose infbrniation from comltiriahle comporicrnts n o t prtwnt i n a l l of' thc-
cl:tdogrums. I n Fig. 1 the conittinaljlc component 7 is lost in tlic strict consciisus trcr.
There is ;in at ternativc consensus trcc ai,ailable, summarizing (111 comliinablc
coniporicnts. In 1;ig. 1 it is constructed b y adding componcnt 7 t o the strict consciisus
tree. 'l'lir strict coiisensus tree difkrs b y summarizing a sultset of' thc coin1)inaItle
components, viz. all the completely replicated OIICS. 'I'lic altcrnative tree, t l i c consensus
t r w of'cornbinable components, includes a largcr set ofcomponcnts. Hence, i t is more
informative than the strict tree, yct i t is not in conflict with a n y ofthe c l d o g r m i s , which
m a y Iic thr c;isv with Nclson trees or Atlams trees.
11' only two c1adogr;ims arc compared, the Nclson tree becomes idciitical with the
consciisus tree ofcorritiiri;i tde components, riot with the strict trc 1s suggested by r a g e
(1989:~)and McMorris ct al. [ 1983). 'Iliis i s because the two fitrmcr trees incliidc a11
comltinable components, whereas the strict tree rejects all components not present i n
both cladograms.
i2 host of conscnsus techniques has been dcvcloped for biogeographic analysis fscc c.g.
\Vile)., 1988), b u t i t is n o t my purpos' to discuss thrm her(,. For phylogcnctic ;inalysis,
Miyamoto ( 198.5j and Carpenter (1988) have ;rrgued that conscnsus trees should tic
iL\:oid r d, sinct t h cy ii rr 1r s s in fcirrn a t i ve i d h e n ce ti avc lcss ex pl ;in a t ory powe r t 11;I t i
individual cladograms. However, in plaili csamination of multiplr solutions from
p;irsirnoiiy malyses, coriserisus techniques are u s e f d tools. Strict coiisciisus trees a r c
commonly usrd fbr this purpose, b u t they arc often much collapsed and uiiirifi)rrri;itivc.
T h e consensus tree of cornbinable componcnts m a y then hc an alternative.
Although the rationalr for using consensus trccs in comp;tring multiplc cladograms
from a single data set may be viewed clifferently from that fix coniparing single
cladograms from several data sets (as in Fig. I ) , I would arguc that the combinable
componcrit consciisus may be preferable t o the strict corise~is~is in Itot ti cascs. I n thr c;zsv
with independent data sets, some ofthose (in Fig. 1 from I)uttcrflies and birds) provide
support for comhinatlle components (number 7 ) that can he addcd t o the strict
coiisensus trce, increasing information without conflict. I n thc former case multiplr
cladograms result from diferent interpretations of' the same data. 'llit- more r e s o l \ d
cladograms provide combinable components that, similarl)., caii be added t o the strict
consensus tree, increasing information without conflict. Even though these added
components are supported only by somc ofthe equally parsimonious ititc.rl"etatioiis, the
resulting comttinable component coIiscnsus tree may bc preferred, since it is more
informative (cf. the argument by Miyamoto, 1985; Carpenter, 1988).
Standard programs like Hennig86 (Farris, 1988), PAUP (Swofford, 1989), or
C O M P O N E N T (Page, 19891,) all compute strict conscnsus trees (in Hennig86 undclr
the Nelsen option) a s well a s Nelson tree (COMPONEKT) and Adams trrcs (PALJI'].
I n future versions, options for computation o f the combinahlc component conscnsus
would be welcome (presently incorporated in the mast recent 1990 version of' PAUP).
For systematic purposes, a consensus tree of all cornbinable components seems
conceptually more logical than a strict consensus tree of' replicated components only.
372 K. BREMER

Acknowledgments
1 th;ink i\rnr :\nderlwrg, Birgitta Bremer, James Carpenter, Christopher Humphrias,
1’c.r Ola Karis, Mari Kiillcrs-jti, Brent Mishler, Gareth Nclson, Koderic Page arid two
anony-mow rc.\.icwers for uscful suggestions.

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(RPceizied f o r publication 15 June 1990; accepled 29 July 1990)