ANALYSIS OF SOME PHYLOGENETIC TERMS,

WITH ATTEMPTS AT REDEFINITION

OITO HAAS AIW 001l.0 GAYLOIlD SiJdPSOII
CO:-!Tf:NTS
""'trac.. . . . . . .. . . .. . . . .. . . .. . ., .. , .... , ... .. .
In 'n>1uo ..... . . . .
Ht>n><>Io,:. ulosy. """""' ... Y
Oriai nd hi"ory, . ..... , .. " .. , . .. , .... , .. , .
(4) Horoolocy." by <>-n . . . . , ...
.. Iou , _
(.) H<Moo ".y. homopla.y _ , __ ... . __ ... _ . . _.
(I). 'flu: .
Do6n,...,.,. and <Ii.,,""'..,... . , .... ... ,
(.o ) H"""""", , , " . . ... . . .. ,
A"."",. vo, """-,",,,, . _ .
(,) A .. "",y . _. ___ ... __ _ . _. ,
(I) HonqoIuy .. . ... " .. __ .. .
(.) H"mo'o&), , ... h"""",Ia ... ,
s,......,.m nd .-eI .. <d =0. _ ....... __ .. _.
(.o) 'Ionoolory _ , . .. __ ..
(.) lionqolu . ....... ... . . , . , . ,
"" .. , .. ,1 ..... """""",...,. .
or..; nd Hi"...,. . . .. ______ _ .
(.o) . . .
M ParoUeli..,., . . , .. . . . . .. ,
l<) Co."""''''''''' ... . , ... . .
lIdinitiun ...:I <Ii,,''''''''''''_
(.) Geoenl . .. ,. ...... ,. _ .. .
0; .... '11....,., " ,
- -- -- . -
(,) Para",I"m d ... , . . .
s..n. ... ",pIeo'" """,l ldiom . nd
. . __ __ __ . . . ___ .. . .
Ori, .. , <l<6";ti.,... .. , ...... ,
K<d,6,,; , .... _ __ ____ _ __
v, . .umkry .. . . . . . _. _ ... . . .
Syo.;.n ... , phon.,;"'II. "mila, t<rm., . , , , , ... .
,Dd l..-.ob"",,,,, ...,dlr.c..,ioo .
(.,.. .nd poly.
'"
'"
'" m
""
'"
'"
'"
'"
'"
'"
'"
m
'"
'"
'"
'"
'" m
'"
,.
'"
'"
'"
"'
"'
=
"' w
..

'"
...
....... tic <Hirio __ . . . ,. . ... . .... ... . . . ..... -iiI
Time "" .. ," ... ... _ . . . __ ... . ... .
"'
H""""""",ic """".-en<:<. i .... ';... . J<J
Sum ..... . . _ , .... ... . . . . .. . .... . .I
Kelet<fIOtO, ......... , ... , . . . . ... , . . . . . . .. , ... , .. .lf5
[ad",o/ ..... Ioo!_.1< , ...... "00<1 , ,
Aosn .. tT
The ....... ""'-Ioc. (homoreny) ",'on. Ioo.nopiao.,
po",I"""". ""''''.0""" time
,; .... tu .... ,0<1 ;t<n.,ion ...
,.-.=I bocIt to 'hel, .... , ... nd crhically .... Iy<ed.
_. .
obo<u", . '"
"".;v,"nt to. nor don
"""",,pI> o_lap_
Neitber """"",Iaoy. 1\01" ...... lleIism. """ """'<'iC_
. nd bOO><om<l>h. , .. ply , binl '" _ .... pon,..;,y
01 ,b. oi.rular ""0 .... , ,II ""'"' "'",_""u mo. 1M: <itllet
"",orocIo,.,,,,.,, or ''''''''_
. oocItronoo. rotlv<I'""",.nd homoomor .... y mo 'n
_QJI ...... d to poIypIt.ietk ....,.oboloeicall ...... . oo
.,....,. hiOb ... ni ...
l4<qut. te ..... 'z., ..-.....-b'ldunO' ... 01 z.;tbou.ti .. '.
Of. io<=I. hi> "Z.ltoigna," .. n" (tim< oi .... ' .... )
.d.ru,t<d ... d<oco-ipti .... "'" pta .. ..,.... term ,,, desio
.... wtain ....,. ". ioucltruoutn ...... "" <On .....
-
1I.ocu ... """ 01 morobolooic '''-'''y 1M: ",...". hom<-
"""""",1<""""""", or ."rH .... b .. to Ko/to, "i _tive
Artctt>ilduOi:" (it ..... ,'" I"" ... ,;... <JI >pe<ieo _ !Ie>o,"n.
i,..,.,,,,,,,).
[n oddl,.,.. ""ho _ ..... , ... ..1 mony ou..r.
..., di><u.-i or .... nUoo>ed. All phy",""",,,", _..-
...
INTKODUCT. ON
TJUS paper hal grown out of a st udy uoo.,r
taken a lew yea .. ago by one of its autho ...
OF ' '' pH"-O..". _ ","" oocr .... VOl. 90, .. 0. 5. 0<CtI .. .... lH6
'"
0Tf0 BAAS GEORCE C .. \YWRD SIMPSON
(H ..... ). 't,mulated by a by Cloud (1941).
Th," 'tOOy by 1-1 ......... late. dioc,,1IWed with
Slmpoon "nd it .. as decide<.! to upand ito IOI>J)<
and to modify it oomewha t. "''h,,," the expanded
, t .... y Wall in pnpar.otion. d'IICUM"",' of homol
OIlY and analogy we", publithe<.l by Boyden
( 1943) and by Hubbo ( 19-14). Thoir v;...,.,.s w .....
diec..-d id by Boyden. Hubbo.
and Simpoon, and in con_tion by fIoyden,
........ , add Simpoon. Bo)-dm .... kindly IrWIe
.",Habit to H ..... and Simpoon....., written but
(a t P<"""nt) unpublished Mnlrb "" thi'lUbject.
Ou. uudy " difi<:rent in 1OOp<.r>d in of
view from tl>ow: by lIoyden and by Hubbo and
it oomewhat d ifferent eon<:!u.ion. 011
ce.tain of t he pnjntll that have been treated by
all of u.. It tberefon: _ n\fd advi .. ble to con,
plete and to publioh the ....... k originally un<J.e.
t.keeI by Haaa ... ith furthH modification in the
li.&lh t 01 this di"' .... on. EKept for the
fim t ... o par3ir"3pb, of thU inttodllCtion.
noo:u explidtly ltate<! to be by Simpooon. the
diocuooion 011 t/M, bu .. of the phenomenon
u ...... lIy called poralk-liom (p. 336). and " few very
brief p' .... elge .. be"'. the .. 0((\101 t he
paper hav<: been wrilten by flu . ... ho
formed the eXlen""" lleBn:h involV<!d.
The wbole IIndy h"". howe"" been repeatedly
dillCuued with $impoon. aoo H .... and Simpoon
an: in all:,.,.,ment "" to a lmott all the opinion,
0:" 1. At H ..... ' n:que-t. Simpoon il
a..nated with him all c:cHuth.or in the .. _
that "" ohatH """",, .. bility for the C1lI>d' ''''1
reached. rathe, than in t he .. n., that he .....
oha ..... "'luaJ]y in the baJic reooearcfl or in tbe
act ..... 1 writinll: of thi. pa ... ' , The", remain twO
minor pointl rtya rding ... hich our detailed opin.
1onI_ not prt"Ci..,I) Ihe IIOme, but lhe ... have no
important bearinll: "" the cooclu",ool
reache<.!. On each of thete pointl tlul main tut
..... been left all written by H ..... and a note by
Simpoon II,.. been added.
The int.....,.t hao been foc"-! pi-
marily on the hitherto nthH amhill:u<>mJy and
iddiocrimi .... tely ....... tennl parallelism, con
a nd """-'morphy. Homoplaay had
to be induded linu it t urned OOt ttl be the lUg"'"
""hich <:()fTtI! b(l1h pandJelilm and
converEence. Thu. homolOil:)". ftOl)'which ""'"
cept homoptasy had oooe been tpUt . <:arne into
play, dnwing aIonll:;1I old counterpart analogy.
MQI"<I(I""" many Other tenno ha ... had 10 be
diotu...d, or at leut ,.,..ntiened. mootly for tM
uk" of d .. ti"ll:\liohi"K them from the tennI ""
which ",tention ;" mainly focuoed. SiDCe m..
cullion and mention 01 u- Ie. important
1...-mI h .... had to be O(:att...oo throullhmtt lbe
pa ..... ' a. OttasiOIl aroo., it hal_me<! deoinble
to 'pp"nd an alphabetic indu 01 atl the phylo
lenni involved in thilltud).
Tlu-ouelwlut the invt'ltill:ation. it i, firO! at-
temp(<<i to t,.,.,., tM lorml under di-.:uooion
back 10 tMi. -oriiin, not beca_ of oome eon
"'"ptina 01 priori t y .. bich would here be out of
place, but beau.. insiihl inlo t be qnal usage
of U ell" '00 is bel ... """ to be helpful for ,II
..opt<" undentandinlt;. Then, the further evoIu
tion of the ....... ept is .. far all _m, to
be ....,nh while. a critict.l "naira' io attempted,
and. where it _mo neceuary. amended defini.
lions are propooed. Finally Iynonyml of the
tc,mlltudied. wbe.., ouch ex"l. are l .. ted. Tile
clarification of phyloge...,tic ttrmir>OloiY by
tliminau"K a hoot 01 1)'_)11111 or oo.oIete or
othHw'" ",pertlu",," tenn. may be ..,...idered
one of the pUJ IX of thU otudy.
IIOMOLOGV (HOMOGEl'n
ANALOGY. !IOMOI'LASY
ORI GIN MiD 1!15TORV
(0) lI"",oloO,'" CDIO<riPed OuR
Tail (1928: ]53). bel ..... ' the eeomelnc..1
meaninll: of th" tenn to be the orieinal one. cor
'eopondinl memben 01. "milar
"hornoIosou"" Thu. it re...-ntll to him a
....... word cond" ...... tinn of edroy St. Hilaire'l
"pinciple 01 connexiodo." It may, howe_.
be doubted .. het hero...-, ... hen ddiniilg (1M3 :
379) "hon>nlog",," BI ' '"The An", OIPn in
diff.",nt anima!. under e....-y va ..... ty 01. form
and function," had Ihi, i"Ometrical in
mind.'
Soon afterward" thio term Waf to unde'll:o a
conoiderablc amount of amplification on tlot <:me
hancJ ,,00 ,,,Wi .. ;.;.,., on tho other. In 1&46
Owen 0 847: 1lS-176) dlotinguiehed th_ kidd.
01 hon>oIoJY: ''The fint io that .bcwe
viz. the c:orrnpondeflcy of a or orran. dete,
mined by i tl ... !atl"" position alld conntiono.
with .. or OtiaD in a diff_nt .nimal: Ihe
dtttnnination of .. hich ltomoIory indica"" that
,ueh .nimal, are constructed on a common
ty ....... " nil .. Owen'l 1J>i4l /ocmDIoo
It wilt be notice<.! at once that in the 6 ... 1 place
'0.. ... (tl6(;. XU ) h'-dl poi ... "'" ''''1 "!'rio. 1<1
tau !be ..... 1_" ... l ... d bttft i ..... boo, v....,.ty
.. _.ty." .od q_ I"'_p r ..... Goof"", St.
Hiloire (lIn) i. ",bicII "h._+ ....... od " ...........
__ ;.0 , 1< b ",Io00d
ANALYSIS OF PHYLO(;EN'TIC TKMS
considen.tOom of topographic and comparative
anatomy.", made for this kind of
hornolOK}', although mention of homoIogues as
being "c<K\lItructM on a """,mon type" may
imply K &light forebodina: of common origin.
"A hi&her relalion of homoIOiY i. deJined by
(I. riI. ) a. "That in which a par! or ...nel
of par!1 stand. to !he fundamental or Ij:eneral
type;" this is !ermM hy him ,.JUraI Jum,cIoD
Thu. the ron-eopondence of the basilar proceso
of the human occipital hone ,,;th the "",.;...,.,;Ipi.
tale of a fish or crocodile I. cited ao an example of
'!"'cia! homology, but the character of that J>rO-
t"e$S .... the "oentrum or "hod)" of the ""'t
<;Tanial u one of general hnrnoIogy.
Finally, rmal Iwm<>/()CJ i. defined ao the .. rial
repetition or "'p.--ntative relation in the
oewment. of the same okeleton, and t h..e .. nally
related or repMted partsa ... <;8. lled
This diviokm Rnd distinction are repeated with.
out any _ntia! change by 1866 (pp.
XII , XIII).
M can he readily seen from thi d>eme, both
general and -enal hornolO&'Y, as there deline<!, a,.,
phwomena which do not pI"I'IIlPP""" two Or
IllOI"f> organisms to be c<>mpaced with each other
and, therefore, lie ou!&ide the original orope of
homoloirY, as co"""ived by o...en in 1M3. Thu.
!hey may well be eliminated Ir"\lIIl the lurt!>er
eou..., 01 this invcotigation, a. may other """'.
proJ>OI:d by later autbon for cat<;gori .. 01 "bo_
moloirY" apart from special bomology' FoIlow-
ina:, in !hi. _peel;, Boyden'. (19U: 233) pro-
pooal, and contrary to Hubbs (19: 293), the
"""" I>omoJogy i. h ...... nafter '*" within the
_tricl iom, l hough nol with the definition, of
Owt!n, 1M3, i. e. as equivalent to his .pecial
homology of 11147. This .....,m. tn be in acconl
a""" with mool COlltempnrary usage. and we have
dispeofJe<l ,,ith adding "r. IIr.' or like to this
term to indicat.! that il io not meant to indude
Owen'. general and llerial homologies. The twn
latter term. may serve a us.ful purpooe in c<>m
parative ana!<>lny and li"netico (ef. Borden,
1943: 234) , but it i. q"".tinned whether they
(III'}.!, 13). _. withi8 tho _ 01
Iol. "AU ......... ..... icIt 10 o\Ot .... ""'100' to
o..u., _I alb .,...........,. ....... ,h.
_ ... liDo homotypy. The......, homo>dl""'my 10 .. ed
by Georea.bo..r to """" _ 0..."., _I."" . ., I ... ,
10. port. oerioI homo\ocy.
'lblo .ppI"" '" lioK .. bo. (1m, lJI elob<n ..
I , = .. ....u., to """'y 01 tho n dl.,... .. " e>., .... ""
diatloopiob<d by MI .. rt "$-119), ... Iy _ 01
... bdt """'" with .. tho..".. r:J 0-".,
have any "r-aionn in the ... a!m 01 pbyl_
ogeny.
(6) A...u..V'
The hiotnr} 01 this term before Owen hat been
t raced back a. lac ... to Aristotle by Boyden
(19U: 2J1-232). but Owen'. delinit;"n nf 1M3
(p. may he", be chooen a. pOint 01 depac-
ture, "Analogue. A par t nrorglin in nne animal
which hal the .. IDe function a. another part nc
organ in a different animal." Owen himself
.,,,,,,. later to n<>lice<! Ihat the word
"anotber i. dioturbing in this definitinn beca""'.
... ben "'pealing it in 1866 (p. XII ), he ",placed
the laO! eighl word. by "a pan or orp;an in an
animal. "
) H""",,, .. y, lIomopiru,
!\/!er the advent of evolutionary idea_ll
but ridieuled by Owen a. late a. 1866 (p.
XXXVI)'-Owen" C<lncept nf homology proved
00 longer quite ... lacking, as it did,
any pbylogenetic implicatio..... 11 was thio need
tor h""""n;";ng Owen', pr-e-evolutionary term
witb t he new ..,hool nf thoullht that convinced
Lanke<Jter" (1810 .. , 36) of t he n"""""'lf "10 have
two lerm. in place of the one 'homologue,' and
to broadly diMingu"", the nature 01 tbe resem
blances to which they are applied.' T hu. he
prop< ' f 10 call ".truetu .... which are i"netic-
ally .... Iated, in on far as they have a single "'1',.,.
.. ntalive in a common ancestor: hOlll"KennUI
and the relalinn botween them homogtny. and
to '_ak of one as the h<KItOgen of the other."
In the pallSaiC just quoted. Lank .. t ... clearly
started nut to establish a dichotomowi dininctinn
but be muddltd the sub;.ct immtdiately after-
wards anaWffing his own (ibid., 38):
"What i. the other quantityoovered by the term
homology over and above In
.toad of dearly defining the Ot\oer limb of hi.
dichotomy, al !>e had the one, he gave a wordy
and by no mea ... unequivocal e"Planari"" nl
what he had in mind. ' " He mad. thing. even
...... tho hypothetical ...... """,pUr...! by tho
_ ..... " 01 Lo.....-dr.. Do ........ 1'0'.1,""", oDd o<hen..
_tQ! '" R<arly .... ilar '" .. vifoon.
"""'to. ad on ''''' or ".",... PO"" 01 aD "'P"iom winch are
."",,<iy or n ... ely olik., .... '-'I ........ ".lifH:otio ... 01 ....
nriouo po ... .. ill "" ....."Iy '" nearly F....u...-, II
io .... d of """br "" ... in tho ...... OIJ&niom, ... "'p_
.... .... f.,.".. to 0<1 "" PO'" I ...... o;qoooiom . ...
"",,,'''' '" oeariy >llke.nd _ .... h_
,"'- tho .--.1,i<Ic ,.11ed I.,...., I. the
"" ... in tho ,...., "'Pn;"'" .. ill ho ... riy (Jf .><&<111 alik<.
,....... "';U bo, I imqioo, ... Iciod r:J otilficuity to th. noIu_
OTTO HAAS :\:-" U GEORGE G,W I-OKD SI W 'SO;':
worse by indudini' and even hi. ex_
amples from oerial homology, and o:onclud<'<l
by propooing "to cal! thi. kind of BgrK"lll<:nt
honlOplaJi. or homoplaoy" (Ioc. <iI.). La nk_
estt'r. word. "which part. are exactly or near!)-
alike and r<mUtt ..... homogenet ic" (italics OUrl)
cle ... ly 11 ... 1 hom"",,"y was to him not
an ......"tial prerequisite of homoplaoy. "Ibe
same holds l rue for hi. lurtl>cr attempt. (ilM ..
42) al defini"i homoplasy a. "dependini' on a
,",,,nmon (l<Olion of ""okinK cau""" or moulding
on .uch hom<>genou. partl, M on
paris u'ltkh fer oIIur r ...... "'" offer a lib ... ., <if
"",Inial I<J 1M, ;" ...tth (italics ours), N<:VeI"the-
1_, hi. oltng,ll..". oboeure """",mati"" misled.
o.Wm (l9021J' 261) inlo tbe bt,l ie/ that "borne>-
pla.y ah.'ay. invol."" ",-hite parallel-
iom and convergence mayor may no( involve
honlOlogy" and to publish a chart of "Analogy
in Ewlution" in which homoplaoy w"'" COn-
.idered a phenomenon distinct from both paral_
Ieli"", and converg<!nC<! ,' Ho"""ver. in a let ter
published by o.bom in 1901 (19()11J: 23S-2.W).
Lank .. ter "demurred lrom Osborn'. interpreta-
tion of h;" def,nition of 'homoplas)'. '" 'lating
that "Homoplasy d""" tIIA demand an element
of bomolngy" and that he "recogni.ed (as
hitherto unde< one term 'homology')
homogeny-or hereditary quality-and
homoplasy 0.- moulded non_heredity quality."
T"" "lilre""", of material to begin with" of 1810
(!lee above). a ll uded to in Osborn', paper 19024.
i. now explained by LanL:eater at "eilJur a
liken""" 01 true hornog<:ny (Inat i. of form and
relation inherited) er a likenes. of ';milacity in
material, in position. oc in initial fonn-flot due
to d0ge hon>O!l""y-but pOssihly a lilren .... of
SllCh a general a. the nf ma_
t erial ' (not of elaborated form and p;tcUj in two
epidermal .urfaces: " Thereupnn o.botn
,;0..;0< <0; .... _t 01 Mr. Il_t Spenr'. writi .... in
ad.rutti"" ,be ...,." p.upoo;,iont;.0.:1 it;'
with , "" printipk 00' lonh tbat oerial homoloo;ie>.nd
owoeh .... whKt.. t.,..th<r,.;th .. ba,;. ...... di ..
hu '-" ino;!.d<d .n<i<r lIomoIoiy may ...
u,,-;ned" (;hiI . 39) .
.. nu. t..d. 01 dadty in Lonkeo .... "' ..... tion ..,."..
"" b.o"" """tribuoed '0 """" mi .. OO"' .. <>dine> 01 til<
""""pt "homopb.," by Spomann (t9U, I? $.2), K.-,ly
attacked by SMe..ttOOo (l91S) .
' StiU .. ,r;.". (1<>00, Oabon> h04 OJ>Obn 01 ......
1 ... ,01 .,.rolloli ... '" homop.Io.y" .. i,bout any
.cion betw_'- "'-0 """"" .. (<I, o.t.:.no, 356,
fro...,..).
'T,;, (1928; 159. lbO) _ inclined '" ............ ,
Lo....".... ..,.". . ba, ..... d<d ,be oropr ol tb" t<rm Joomo.
J>b.oy .. b .. 10_ Iii., bu, i, may -rl be .... t be hod.n tloio
(190;,,; 228) concluded that "Lankestec. IJomo.
PlMy is 10 "",u"to,,", """,..no... '"
p.uGiklism, '" < .... _:<"".:. Two yea ... earlier.
Osborn (1905; 959) had already induded parallel
and COIlV<:li:tfIt adaptation. in the analOl[ou.
oms tadni'; ''To these processes and ",.ulu 01
similar n.odcllng Lankester hao the
6tting t..-mo homnplaay a nd homoplattic."
(4) Tiu Modern MIa"'''t of Homol<lD'
When "!,Iitting O ... "n'. concepl of homol<>&y
inln l>omogt1ty and homoplaoy. Lankcotcr
(1870..) clearly intended to ,..,place homology
by the one co"'''''Pt or the ot""r, ao the case
might be. and explidtly stated as his 're.unn for
not retaining 'homology'" tnat "It ;" Je,,& likely
to cause confusion if we have a new tenn than if
WI' an.end a n old ""e."
Stransely enough, just what Lankest .. wanted
to avoid turned out to be the ",."ltol the lutthet
evolution of this terminological problem: "Ho-
mology" did not dioappear, U ""as hi. inten tion.
nor. in thooe ca..,. wbere it i. e<tuivalenl to
Lank ... ter'. "homogeny," was il "'J>Iaced by lhe
lalter tt'rm which ne""," .... cceeded in becoming
lI"n ...... Uy or even ",-idely adopted. In.tead.
"homology" quite gradually B$lumed t"" COlI-
notation that "homogeny" w a.! intended by
LanL:eater to <:On""'y. It;" true that Sourne as
late as 1910 (p, 38) complained not only tnat
"the term homogeny has not bt...,n "" seneraliy
and many, if nol moot """,Iogi.to ha"",
prelerred to retain lhe old word homoloio" ," but
al"" that "in "" doillj!:. . man}' 01 them have
failed 10 di,tinj!:uisit bel"""'" the two quantit""
;" mind . , ,be O/<ry ".,,"', but failed <0 .. ..- i, w;tII
ouffiOont clarity. I n h .. 1<,,,,,. quowi ob<w<. I>< .......
to toke (I>< pooition <bat "" ;" ........ Iy intoetpci ... hi. POP""
oi. 1370. rbi. p>aitioo ... "'. to be .... par"d by hi.
<0 Mi ...... (13706).,..b<re he """"'von <0 mak _
d;"i""tion be,_ hornopIuy <ban in hi.
.... y. bot 0100 b> ,"" .... .....,. 01 ,.., _y iueU
(Uro." I), "H"""",looy iDel ..... . n ""_ oi. """" '-">-
bb"", oi. f""" ...t>ich or< not ... "",bl< '" h"""""",y . U
-.01 a..---, "'" "n"".""
_dly "'"""c<""'''. "' .... U .. in ."""tu,.,. Ioo,i", no

'Obviously "':,bou, kno.lodro 01 _ t. .... , ....... ""
oi. Ooboroo II ... , .. (1910, .19) f""nd d;lI'Iaolty ;. d;l!er
ca. iatioc be,_ 00/>0<0" "*ltf: .. ieo (01 (90l) .nd
"",. l<d .... (;hiI . W), " , .. l.onk<s.."., .. rm h-..Ja,.,
",..,...U til< <0 ... :' Similorly, Willey ( ' 9", X_XII ).
"t..n qu" .. loninl o.t.:.no . _ 01 1!>O1 . on. " ... ..
01 tho 10'_ ..... """ 01 "","ion that bod tn<n "'"'" in tho
__ nti"",_
'F", .,.".,rJe", .. tioo, 01 "'" chona< 01 ....
uN _ by til .......... f"" tho o.,,,i.i,,,,_
Sporn .. n. 19f 5; 6&--71, " .. d Ins: lJ.6-l5&.
ANALYSIS OF SO\lt I' HYLOGEr.< ETIC
[I] contained ... ithin the "n,le term. 01 "'hkh the
dif("",,,,,,,,,.....,... .., dearly pointe<.! Ollt in Lank_
.. ter'._y [of 18701."
Ho_ver. in til<! meantime til<! term homology
had to go a Jona: way to ",pla.tt tho one
of iUl ",placers. It had OOon , tute<.!
in thi. dirtion. even a Ie .. ' }'t'a'" before Lank-
tlter' o publication, by Owen. him .. lf, ..,ho ad
mitte<.! (11!6(i : XXXVI ) that. "The moot intel
ligible idu of homologou. pam in I Uch series il
that they are due to inheritance." About t he
end of the nineteenth century Gegt,nbaur (1898 :
22) empha.oized that "h<>mology i, governed by
d .. -ent ; homologou. mgano are of <;(Immon
or;,i""" (translation from the G.nnruo). and
Osborn conduded his I"pe' of 1902a (I'. 270)
with thill .. ntenee: ' The only decioive teot of
homolol(y i. hi,tori< commilnity of derivation
(i. e., homov;t'ny)." This coneeption _ m' to
have been rather generally """"pted about
fill""" yea", later. when DeDliy in hi. glos$ary
(11123: XXVII) definoo h<>mology aa "_ntial
.truetural identity due 10 community of
ocent." Thus """n Royden_Ito late.- (1943)
adVlXate<.! the ret urn to Owen', original m"'n
ingo of homolOgy and anaIOKY-admit<! in an
eariier paper ( 19JS : 448) : " It i. K"""",lly aKreOO
.. . that homoklgy is due to community of
d"""""t." Quit<! cwmtly Hubb6 (1944: 290)
summarized thi' development aa follOW", " H.,.
moIOgy haa thus"""", 10 .;gnify an ",,,,,,m0111 in
<'Volulionary derivation. . . . The icleao of
Iwmolov;y and community or origin ha"" become
intimately ao.odated. almost to the point of
oy""nymy. "
It mult nol be overlooked. however, that oomc
otudentll Itill adh,,", to the oppooite behel. Thu.
Kalin (1946: 136-137) quite cwmtiy pleaded for
t ho morpholOgic COClcqltion of h<>mology as 01"
I" 5; j 10 the phylogo:netic """. quoting Herlwig
(1906) for hi. fIOl nt of v;..w. Thu, he join,.
though unMrt$ciOO4ly, Boyden (lIlU) in the
battle ay "Bad: to [ HUl)."
Spemann (11115: 81), on t he other hand,
advocate<.! .till another terminology that would-
against Lankester'o intention-retain homology
... a n""tral temI. with I>omOi[eny and homo-
pl,..y. both unde .. tood as ori.<I:inally defined by
that author .... oubdivi.ion ..
Not only because we have to "i"'" with the
historical fact that Lankest...-. t ... m I>omov;eny
hao gradually been relegated to me", oynonymy
.. Wh ... otUdyinl .... wort. o.t..n. made a
not< KI p_ It, "Spociol ... b tho ............. y.
01 Lanlo<o",,:'
with homol"lY. par t 01 "'hkh it "'all in
tended to ", place. bUI ,,130 for the oloje<:ti""
rea..".. ""pte " -d helo ... , "homolOgy" i. u.oed. in
this paper too. with the meaning Lankeole.- ga""
hi. term "homogeny."
.. ND
(< lIomoloD
Thu> homology (Owen. 11343. _ spe;:ial homol
ov;y, Owen. 1847) is !.ere defined a. a oimilari 5Y
bet w""" partll, organ . or otrucU,res of different
orp.ni ...... attributable to common ancestry.
It ,,il1 he noticed that-----<lespite Hubb', (N-I4 :
292) adVOC"-ting " that "'e t hink and write of
homolOgOtlo function. in the same way that
homologous . tmetu ..... ha"" been treatet!"-
funclion. are nol included. in I"" above delini
tiQn. in the enumeration of similar
It is believed that the function of mga!U may
or may not be the same in different mganismo.
but that functions ... ""I. are never homologous
the one 50 the other. What Hubbo ",aliy
_ m. to have in mind is to u ... in c""ain caaeo,
idt.ntity of function ao a criterion of homology
(ibid . 291) . There i. certainly no objection
"iain.t thio. but it must be kept in mind l hat
idtntity of function hao been consldc,....]. from
the very beginning of t he terminology under
discussion. to be 1M charactorilti.:: of art.1l<>gy
rather than homology.
N .... 1 $;", p"",: Tho preeedinJ potOliraph Tepre
.. n" the ,-iow d H...... I do no. qui{<o .ilTe aI.
IhoOilh the dioa,5ftm<ont i. 01 _y oIi,h. impor'an
and it in 1>"<1 m..-.ly. mattor <>I _bal preoaion.
Hobb', point "'AI to . ,,.,.. .h. diffioulty ud. at
tim ... the und .. inbili.y <>I o:o"';d.rini "rue' u", and
lunotion .. po.rotdy. Similarity QI luncti<m i. not
doaively charackri.,;c of . ",Ios;y. It it dten.
indeed ..... Uy. charaoteri.tk <>I homoloi]y. aloo.
I" lact. Imm one poin' of vie .... "alocy and hom"'.
. ... t..-Q . Ikrnath-e 'h<OTelicol plan.tiOOIO of
airnilarit y both in lunotion ond in .tructu.... In
. och oa_. I can _ no objection '0 ,_kinJ of
hom"'osou. funotioPl>. u an .. p ..... "" 01 opin;""
to the origin 01. dynamic complex iocludin, the
IUneti"" Ionl with .he .Iruclo .... ""norm;n, .hem .
That "' unction. a aTe n.vor tho
OlIO to the othu" i. a matter of d.finition. ""Iy. I
ai]T<' that .uch a d. hitioD it d ...... ble if ph",oed
tha' ' onction conllid..-.d .. ob.tnoetiono and .ith.
oul """aidora""" fo.- the otructUTe& WI ""rform
tho .. function .. Mould not b< 'pOteen 01 .. hom"'.
"i0o,. bu. that hom"'OIIY ... "tly involve. both
function and .teocture and i. thi, combined oenoe
it it proper to '""ok 01 ,h. oomoiogou, fonction>
""rforn".d by hom%tou. olructo_ I t i . 0100
OlTO HAAS A;Io'1) GEORGE GA\' toRI) SIMPSON
If .... bu, 1_ COIn ...... , 'Ilot the al bomoI
"""'" .... _y "'" be """"I""", ..
Moment', (1945) recen' propooal. to ]um[>
epecial ... rial. and i:eneral homologi.,. and tven
to diecanJ the unn homol"i)' al'<IIIether, re-
i t by the all too "oimibrity," an:
decide<lly rejectc<l. Hornolog:y . ... under
'Iood, io Ail! a ,,-dl usable w-m.
PanicuWiy unfortunale and. " 'fcre it to be ao:.
oep(od. bound greatly 10 ;,",re_ the """'ailing
Itt"JIunological confuoion il 1...-tM.
PlOPOW] of no f""".. than ",x "I\n"" I.,...... fot the
ditliOlCUon of "".-iou. kind. of timilarit;"', ..-t
of .... hich al'fc preoccupied. ... ith a diffe",nt
meaning, in the preYinu. lil ..... 'u"' on the oub-
ject."
(b) A "'*0 vl. lIo",op/<uy
Rtf.,... aHempti .... defini,iom of nlher anal.
0f0' or homoplaoy ot both, ;t "ill ha, ... to be
examine<! ot r>OI , he. ,...., t.,...... are
'1nony_ nu. q,,"lion has veKed . tudentl
",nee bo.nopIny he )'0\1"11"1" tenn, ""'.
...... "' 5i d in 18;0. La nh-lter. him .. lf, (18100:
404]) the quer:lion a. foll"",-.:
I, may be oaid tbo, I.,m ...... lorY, .. al.-eody
I" u". i uffici,"1 to indi .. ,. ",hat i. hor, tennod
"homopl y"; but .... I"'y h ... wider .. tion
rP"ttI to i. in which it iolouad vory uodul to ,,,,ploy
h, l ad it "",,1<1 ..." be u...:f .i,h amy _uracy ia
place 01 bomopluy. ,1., t wo orpm hlvi_, the
.. me I"""tion 1'" 1 .... """"' ........ "'" ck.<Iy ....
nd! otbe< i. !.hoi. ''''''''u", and .-ellti""
100 ...... PI' ..... _ : aad i, i ..-.11 to ... 1Ii. ,be word
i. ,bat "ide .... Ie.
Dendy (1923). a lthough lliv'''g in his G","",,-ry
(PI', XVII! and XXVll. retpec:tiVl'ly) different
definition. for a nalOiY" On 'he 0 .... hand a nd
" H-..Iynanoy (.djo<ti ... , _y"''''''') ___
I _I .... "".""' ..... ,oJO-'-. "-,py (_u ...,.,
,N-!, ZIP) ) .OO<I _ r"",,", by Abo! ( '9011: 211---lJP))"
.. _-..,. (f, ' -, .5) _ .. , _ ....... ""u .. <ic
__ .. ;,10 t>-r w.-t- H".:",,,,, _ by_.
_. (,19S, 456) OD<! to r-. .. " ..........
i: ... - h - ::-. i:!- _ ..... :>
... t ,'troo) .......... "" .
11_ .... r .. _ (19:11 : It !.) o:od Da<quol
(!9U, _b.d,. w. ,,",y 1M".,.. pilI"""""
.... il\co ...... _ ..... M"""" .. , ,""oX 5:: Iboz", .. _tlc:o!
fI .. .. "', yIe<ic wo. in ,,,,, Gtnnan
'0"""" .. bom.>pt.yl," ..... ,5y 5Ut (p_ ! 7), by l'ab-Mn,
by FPlrMn,.. (5m: 1522) . nd ",._, .nd '"
botlfta _ by "'u_ (5"': .. ;, ill """aU<p&tt
"',...",.,yPotio. Ao po"UIoi "". botlow. 1000 ," rry 10 ""'"
od G"':'r" 0/ '- $1 , .1Id. '_""" 0/
: ... ,." -..
'"$00 .... 1' 5,_ j._ ..... , ..
honIopIIIY" on It.. olbn, """aks (ibN 2%) of
"Tile (emil homology and analoKy (or homo-
platy)" and ,d...cribco (;/M . 258) or
homopla"ic organ, " indiocriminately.
Tait (1928 : 159--160) explicilly dillC ...... thI:
question, .tadni: lhal "Ihe varion, in,
n an_ of anaq-y are ju&1 of [thel kind" of
reliinbla_ to 9"h;d. the l erm homopbsy ;.
applicable, and eloewhere (ibN" 1611) .t.o..,....b
of "hotnopIuio ot ana!0IY."
Hubbt (1944: 296) ani"" al lb. """",Iusion
tbal Lankoe.ter" "merefy ..,I>.Iitoll 'loomori:eny'
for PlPtCial homology and 'lw:In.opIuy' lor anal
<lilY." T hus he . too, .,quat.,. homoplalY
anal"i)' ,
In our opinion, analogy and homo-
pl.:u)' Ire: nol synonym .
If "analovy" io to ",uin, btyolld ito rather
v"I"" eeneral oi.gnilicanoe, any l echnical "",;aning
in """,parati"" a natomy and pbyqen)'. it can
bt 001 one implying luo>ction or u... 11: it
.. the ...... n'nggi......, to tlU. trrm by 0-.. (1843)
"",intaine<! in the bioIorical lcitlK'ft. thotJjj:b
....i lh _tain qualiliraliona. lince. E......,
lIubb. (1944: JO(i) .... m. to ...-""" ... i,h 1loy<kn
(I<)04J) in con.wenng "ag"",menl .. ..... [ita li<;.
0111'11 of charactera (.trueIU .... or funct iona) "
of dil!inCI evo)"tio"",)' origin u _ntial lor
"a modern concept of analogy."
If u"" or function i. impil.cd by Ihe 'ffm anal
"i)', then this t""" :Ioeo not connote, >U 00 bot h
looonolocY and loomopbsy,
i(. LI ,blana: (<1. lanh-ltrr-, 18700: 41). 'n-.
at lout. partl, ""Ianl, or I\tUCIU_ of
'-.o'Oor Ll>Or1'orr-miomo may _ the u" or
have tl:.t """" l unclion ... ithout oImi la.,
On t ile otho-r hand. no m .... n;"' ... 10 _ or
fUOIClion io implied in the term homnp/MY. It
be rallr"r te'nplinr to have to do with
one dichotomy only: homol"i)' (,;milarity d ""
to common anculry)---a",, 1ogy ("milarity I>Ot
due 10 common a""",tryl. H()lto' e,'rt', a ller
clariflc.;;p, ion of the ..... kLp .. i">"OI>'I the nttd
lor ouch a oimple dichotomy -.no to be fulfilled
btlter by the one conl .... 'i"' and
homoplaAi<: oimilarit>e., anakivY bftng- on a
dHf..,-en1 .. 11"" ........ 1 level from homology and
homopIuy i. , a
!eatt primarily, an Plot !"" bringing
aboul homoplrurlic wlre __ ""al<lllY
il from the oulllCI a ",la,i"" btl ...... " l"'"OOraan'
it:n. ot p;r.rtr: th,-.t.
ANALYSIS OF PIIYLOGNETIC TERMS
'"
Th .... honK>l<!gy and a llaJoty IlOl. mutually
ex.clu.;....,. and Boyden'. 231) of
homologu ... ",hich are at the .. me ,i_ analog.
ouo. or of "",,homolog,," wbith are not anal
orou. all {he same. are o,ill coowde"'" valid.
S; .. Th .. tho _ond of
POill," .... "hie" H . .. and 1 .'" ...,. in ..,.., pI.t<:
1,_"", ... . . ... ". al .... "'" i . aot
lun<lomenulaad oJ<- not alleet tho __ importan'
"""d ........... at "'" d.'"i , ...... oIl1omo1o&y .""
ho..,puy.
.... .1 ...... it boot d<'_ .. Ii",;
lorily in'ffJIf<',I u d .. > """ ..... '_'ry.
Homoptuy ... .110 al_. I. boot defiMd as limi
llri,y (or at ,"dudinl 0, Ph'" . " Ieod;nl '0 .. milo'
i'y) tha, i. no' .xplici,ly In,.,p ... ted to con,
mon ... .,..try. Both .... m. toth., ,han hoi., pu,.ly
d_ripti,'. (lO in Boyd . .. ', .... 01 homolos,.
lor lu' ..... ) ptao on opiniott ...... """tiv. and
0 ... nopt..... H""""",,y .,,1" ...... <>pi_"'" ..
'" bow .ho .. milarity 11'<*. Ho.>opIaoy
u oP ...... u to how . ...... ilari,y tlU ..." an...
i . 'bat i, did l ot ari .. by 110 ... ,,1017. but i.-.
not "' .. an <>pi" ..... u to how ,ho oimilarity tlii
.ri ...
I do ...... in qui'. tll< ...... "1)' " H .... _ tboot
... " ...... tiy ... , th< ...... c., .. ori<al k>'d. Til<
ot' il not """riv ....... and "b .... mu,uaUrad"oi y.
O&"lori . .. bu, i" dicho,,,m, 01 ...... nd "not - ....
U .. d., "n"t _ a" i . i. " ill poooibl. '0 hay. a .. qu.nco
01 . ........ ri""' ouch "b," . ,' ...... that " .
,-i,;v ...... coOco on t h ....... Ie,'" ... "a," Quit.
dearly by a I ... "w.h"", _1>(\ , l>OIy by "'ny
.... aklcy' .... ........ an ol' ..... riv. 01
"b." ... , ...... ,h ... "'" ., ..... tiooo. pthtnd by Hou
......... i. haaol .. n '-" ..... lor " ..... _ari.o<I he_
0 .y_y'" ol """""" .. y.
Itaao ..1 I ...... ha, oq .... iooo ol o",,1orY .. i.h
.. y .. undeoi"bI . ,he """oit\ ..... bIt
""';lht oC u .. w..u .. it omito ,II< ".m .... of
lo",,';on. which h;.toricall y bolo" i ' i" the ('On""p'
of ... IOlY and "hieh "ill tin .... I t leaot. t hat C()n-
copt in the mind. 0I1E\OO' " .... oC .he ' .. m. 1 1",1.
ho ............ hot oim;lority in "'wo'.'" ,Ionl ",i.h .",i.
l.ri .y i. lun<ti.o<t .. oq ... ny .. __ .t;"! pan of ,II<
"""o<p' 01 ..... Iou ... , h erm .... 01 ...... aI.a,.
'-"....... M"""", ... tlte i ",plica' ;'" i ...... lly __
e '" ...... d"l ..... a"" i. h .. oo ..... i ..... '-'" .,.pIic.
illy ... ,od , b. <hoe .I",,,"ural limilori.y hoo.<ein q .....
"Of! i . IIot d ... . o ho .. oIocY buI i. _ ... Ia\.!:d .itloo
"""'''unify of fu"". iooo .. Of>!><*d.o """,muify of
'n_'ry. It i . ill ,hl ... not th .. on.Ioc, i. a "-
oh.", .. iv. (bu' no, the only ''' ... nati ) .o homolot:y
... poo;,;ye ca'''ItOfY on the ... m. " b"
ory .... th .... than I "no, -o" .. tq<>ry .... "'m"
,h'''l On dille ... n. In .. l '''Oi h... Tha, i .
... Iot:Y. wh I&0O<I in thlo woy. upo ', . """tive
.,poioR . .... I.eo.y. tbot "."utol rnembla_ 10
wi.h l.n<tiOll. j... ..... .....y .,.po " , .'
"'" vie. tbat it;' .. ",. .... 1 .nth ...... ""'" an_try.
U" lik. hoonopluy, .""Ion 011 ................ Ii, ..
'''-Y .. to .he !>a., of .h< ,-",hlonoo i. qU<'ltion.
I, I. In one "' Y' of """"",l .. y. d"'I'
like homopla.y. a limi l.ri.y that is "'" i .. ,.,
.,,,, d ., homok>iY bot oddinl to thl. pu,", n<ia
definition or hom""l,.., .h. opinion that tho
"mil.ri.y i, '0 be in""p",tod in .'IOth., ".y. AI
.hou,h po,,.Ueli ..... ...... """.' . .... "". mIIy 0100 _.
_tly be <01111 lubdirioioo. or h ............ y. , h" <loot
..." pl_ .hem ..... lev .. "i, h a.lJoey. They bo:-
..... , i dil....,"t trqoriQol _rot.", boa ... ,My
In' (ao ......... 6 ..... ) ""'y. "hile AftalorY
."" i ..... f (bu, oftly oq: i .ely).
i.terproti ,'C.
(el A..aJoV
AnalOJ/Y il thus here defined a. t he relat i""
bel ........ parto. '''It:anl, or " f di ller""t
orpnw... .. hich ...... the oame ute (It' have the
.arne lunetioo. "
(<I) H_iJ<u,
Homoplaoy. <>II t he other hand. may bedmned
to compnle an """'utionary poc- . bringing
about ';milari';"" '''It:ani,,,," or thoir
pan OIV'n . .... otructw"," ... hich ,,,,,",,t d..e to
common an"","y, but to independent acqui';ti<>n
of tile oi milar chamc..,,,. AhhouRh hom"plaoy.
if IrRnolaled lil .. rally. indical'" primml, a pfO.
cao. tbe lenn ha, ... nco: Lankttler ( 1870..).
eqMlt y been applird to lite .. rnilariliot. prOtlucrd
by th. poox-. i. ... 10 the rd..otion hr-
t........ the oimilar organisma or hoonopI ... to."
HomopiallY thlll in.dudm ooth p.molJ.eliom and
I" be deaJ t .. ith under the next
"-'ing. Thi. mighl rai .. the que.tion whe,)"'"
Ie<m homoplaoy might nOl bediopenoed with
the t " .... Other. being well eotahlished
and even better Im""'n and. ao i t """m . at lcut
hitMrto ltatuontly uoed ,hall homopluy.
HowtVet', il _ml advio.:oblt: t" retain ,I.e laot
te<m ao the ..-.. toml""hen';vt oo>e aU the oa .... ,
,II ,I.e MOt'<' 10 Ii""" the boundarit. bot- .....
paralleliom and oon''''we<''''' an: quite in.ddinik
and it i, f""l ..... tly difficult to dt:citk which of
thtIe fl)t'cial term .... pplieo (c/. below. pp. 333 If.).
f...u".....uou of the similar char
..,t .... in both or all concemtd is thlll a
ch"racteriotic of hornopluy ,.. ... e ll ... of pIOnl.
Idil m and <""vergence. It may be added right
he-re tha t th;" esoential criterion ...,n to have
"Soo .. 0<1,.. _ by Sint ..... ,
"n.o .. _ .... .;n boo _ .-; . ....
.................. ....t _ .... e e 1oo prilllOrilr iod""t
... jIO' : ",tho< ........ -..
'"
ana H,\AS AND GEORGE GAYLORD SIMPSON
been =<>gnizru by Darwin in a later" edi.
tion of hi. of Sp.cies (l909, II: 221).
'J'h.,re, in tiill<u .. ing hi. "analogical ",.nations"
(cn.lfIiJered 3 opeclal ca!le of homoplasy I><-Iow) .
I.e !ofH>ke <>( "tl>l: a(.]uirement through Batura]
.. !clion of parts or organ_, &trikingly like each
other, independently of their di=1
from a common pr<>gcnitor"
n.. thought, though expre-.:i in other
word .. may b. found in Cope'. (1811: 343)
""nteoC<"' "Fmm the!!e and many analogous
ca ..... the law may be dodua.od that
identical modificati<>n' of .tructure, ron5!i(U{inll'
c,-oIurion of ty(>O&. h,."" oupervened on d;"tinct
lin.,. of descent." There cannot be any doubt
that "'hat C""" he", <Je.crihe . apI"'rently with.
out knowing Lo.nke.ter'. 187(} paper, i. homo_
plasy" or, mo", orecifitally. paralleli."".
Scott, when diocuosing p"Talielism and conver-
Renee in hi. "Oboervations on the Modes and
Futon of Evolution in the :\fammalia" (1891 :
364 H.), repeatedly cmphasiu-s the independent
a"qui.ition of the oimibr feature; by the corwe r-
gt:nt or parallel li nes. "" he d""", a few yea",
bter (18%: 56). in hi. definitiom 0/ paralleliom
and conWI"g<'IlCe.
Similarly. Buckman (11'01: 132, 238). in di.
c""'ng paral\cli.rn a nd homeomorphy. define<
the lonner as " t1>o tendency of different
to<;b to p.lSS. 'luite independently, through
limilar pl",_ of rlevelopmt. ... t and points out
that two lineages of terebratulid. "develop
.,mibr multiplicatiOl"" q"ite independ_
ently."
as well as in "p<'Cial caoea parallel_
i!m and convc'l(cnce do "ot imply anyl hing ;lS
to conwmpo<aneity of the ";mil"r chang .... in
11>< two or more li neages concerned. For
t;nclinn bet"""n hOIno"I",,)". paralleli.m, and
COnve'l::enre in moxe or Ie"" conl empo<anoous
form. "" the one hand and in non-<:orlt,"",po<-
aneouaO",," on other, ref"",n"" it made to a
followi"g "",...ring.
By way 01 appendix, Tait', 1928 (p. 171) COn'
cept of phyiog.,nel k pia.,. here hneAy be
"n.. edition whOch ..... <he"", '" <DO ... io ,hi, ...
"", . ,-ailah": by l"fff<..,,;, ",.ot h ..... tw..n <he
I" ."h (,&6(;) <J< <he fiftb ( I WI): ,he followin. q.-..''''''
f""" , hellmer;.,an edition vi 1'IIl").
" n.. ...... bold ",. for li,,,,I". (1&95: ( 4) a.",O\(I' to
phi" " ... Si"", <he ....... <> __ pnxI<><e
<he .. "", eII ... not oo ly i. ,b, Loan; ...... W<rll. bu, in
_ni ....... wrll. ';"';lar forma_ 01 """,no ...cur in
deed ;D _y rlilfrren. animak."" .. I .... if ,hey hove bn
.. to "" .. "'" ".tt" fu!---. .. t><ci>ll}' to <he..,...
_n;",1 a<,.,...... (Tra"''''lon from ,M Gomuon.)
m"ntinned; .... ithout properly defining it. Tait
n.cogni .... in it "Th,..,., <J< four elementtt. (vi . ]
(I) the power to tran!mit by h<re<lity the
temporarily exi.ting condition of .tructore; (2)
adaptive modification of ex;.tini; structure; (3)
tile invention of ne ... structures; to which (may
t... added). whet her an independent thing or not.
(4) the ,. ,.".sional dropping of ex;'ting char.
acleR." Later (;bid . 172) Tait deal. ,,ith
pIa:;;, "" .... ith a synonym of adaptation. Any_
way, Tail'. term ooenlO to have heen intended a.
t be hi.<:het" cat"l'lory indoding homoplasy, but it
d""" not SOCnt to ha", met with wide accptanre.
(c) lIDml)/()O \"5. llD"'DpWsJ
Di.tinction 01 homologous and homopla.tic
oimilantic. i. not always ea.y and i. ",ndcred the
more difficult by the a .. "mption. recurnng
again and again in lit""'lore. of oome inherited
latent <J< potential predisposition of more Or 10M
clo.dy relau:<1 groUp" to undct)(O the same Or
oimilar evolu.;""","y modification. independ_
ently of each olher.
On t h;" very assumpt ion Osborn (1901a.
e"J'l'Cially p. 2611) baoed hi. "law" of latent or
potential hnmology;' involved in homoplasy.
Quite Simpoon (1945 ,9) sugg<-sted a
genetical mechanism for .. mte manil""tat;ons 01
t his pllenomen"", stating : "Homologous gt:ne
tend to mutate in the !arne way."' Of CO""",
thi. nggt:stion i. not new. In<ked, the fi"'t to
maJ.:<, i ".,m. '0 ha", been Darwin who. ,,hen
explaining hi. analogo". "arutio ..... in the
example 0/ the pigeon (1860 : 14.1-144). expresoed
the finn belief "' hat all .uch "nologou, variati"".
are due to the ..,,,,ral ra",," of the pigeon havin.<:
inh ... ited lrom a common parent the .. ,me con-
.titution and tendency to variation. when ""ted
on Ly ';milar unknown inAllenee.... Denling
with the ,""me suL;""'. in a later edition of the
ON';" (..,., footnote 17) , he wrote
(19O'},voI.II:121):
.. the m.mbe .... 01 .he .. me ola ... al.hoUKh only
di,"'n,ly allied, have in h<rit"" 00 much in >mmon
in 'beir "" ... ,i.u.ion. tho h<Y . ... ' p"o vary .nder
similar exoitiug cauoeo in .,milar manner; a nd .hi.
.......lrl oL"i<>uoly oid in .he oequif"Omen' ,htQ"ih
nat u",) .. leotion 01 .... '" oriu, . " rikioi(y like
"Lik< "lOfty 01 hi. ronmoonrido. Ooborn i.ovioJ>ly
.oed 'M ... oro . .... ... """td ptd ... ,I>< "on!
"pI _ . . . ,_ . ,,-", " '
'" ... or __ , . n_, .
" . I'd othor "h;d, ........... Iy
oIooom-.d ia<to-m;gh, I>< ci,<oJ ... port;rula,(y .nmi,i
P "'" eM< 01 "'10 Ia,,mokinc ,..-cy 01 ,to., perkod.
,\N . .>,LYSIS OF SO;\IE PHy u:x:.ENET IC TERMS
eacb oth.r, of thol, di_t inheritance
from a common pfOKt"ilO<.
A similar thought i. expre.....,<:J hy FUrhringer
(1888, 1112), according to "'h"", "a mor-phol_
<>Rical similarity of a hiKh.r degr ... (i..,morphy)"
pre ..... in moot of """" that the
genealop;ical diocrepancy het,,-een the animal.
he not too wide. Ahoolutely he"'rogenetic
animal. rarely attain per/eet ionmor-phy (at
feasl in vcrtelmo.le phylum) ," [fran.:iation
fn>m the German.)
Scott (18%: .\.8) "the .. atement [tkat
num ... OU. and dO!le of olnlclure
are prim<! of relalionship to bel
true, even though the resemblances have hoen
independently 8cqUIrro, ... because t he more
nearly any Iwn nrganisms are, the mon:
likely are they to undergo similar modifications."
Osborn, aft.r his paper of 190!. quoted ab<:>ve.
retumo again and again to this idea: In .tudyinK
the origin nl new characlers (1907b: 747), he
arriv .. at the condusions "thaI ouch origin.
[Ihough aru.;IlR independenlly] are
mined by hereditary kin.hip" and that "thi,
prnIel ... minalion i. due tn ",milarity 01
hereditary potential." i n l it" nut year (19084:
150) he "ateo that "he<edit)' appears to domi-
nate the ongin nf new cn.pules in the (nf
Titanoth.....,.] bocau .. they ari .. in Ih. form of
... controlled by anceotral af_
finity. Tkat i., the fa .... """,Its appear in_
dependently in deocendants of the ..,mt an0;e8-
tors" In a ,""",what later 1"""" Osborn
(1912a: 25J) oboe.ves "both in ",spe<::1 to new
CUSI'S on tlto teelh and new horn rudimenlll on lhe
skull [of Titanothereol . .. what in our ;gnor.
may be C.111ed an anlftiral pn.<\ispoo:ilion
to the gen .... of ";milar recliKradation.. Thi.
predispooil ion betrays the uiotence of law in the
origin of certain new characters. .. In hi.
theoMica l consideralion. concluding this paper
.. _n to b< , oy ....... 01 "'"''''fI'''y be""'.
nit 'erm of (lobon .. < ... ...:1 ,00 6 .... _ t.
1\105 (p. 9(1) '0 iOOi ..... "o.,;.i",'" ... ".,..,t ....... tic: vari.
. 0.:-." dio<u.-l ...... ;n t901 (1901.: 2211-1./9). ...J..,.
0\0:6...,;1 .. ", qu.t; ... ,i"" duO". , he i<neoi. of, "' ... du,
.., ...... lin _,.adiotinttioot to ,ho .11otne!ro<>. witi<h ..
", .. ti ... ti"" ""' ......... ....,.... 01 ne .. propor,lorut in
,n .. ;.,i",duoo:"") in 19t2 (t9124: 250),.- .. , loU
"i"'in tho 0tp0 of u.i. po .... , __ do<o ..
with .. hich ;, i , _y n.. 10, ... term ....
ocroNi .... ,. E.i_ ('3\13: t9, 100""'" 1), by
Witholm lWIcu i. tl!\ll (or , .. .,......,...... .. , coiled
"boo';"""t I"rid>'ot. En'wid<lun," ( .. di<ecwd
.,....10<100.1 by E:i ..... and __ aootpt<d by ,ho lat..... (q,
Fen_. t9J I : 114, M.l
Osborn (ibid .. 271) "",,umet "that certain
naI predisposition. 10 the formation of new char
act .... , conMcte<i, as Da .... in ronje<:turt<l, in
oome way with romn,unity of d.,..,nt, are onl)'
evoked under ".,.\ain ""matic and environmental
ronditions, wi l hout which they appear to be in a
lalent, potential or form... .. I "
hi. paper on "Tet rapla'y .... Osborn ( 1912b, 292)
",vi"", hi , 1902 concept nf "polenlial homo!
OIl:y," caJUnii' it "one of l he most
manile. tations of the for".. of heredilY, ,,hieh
predi.".,..,. d...,.,ndanu nf a similar sl ock, ho ...
ever remote, 10 gi"" rioe to similar new charsc"''''
through heredily"; !at .... (ibid" 305) Ihe principle
of potential homology is Mid to be "based upon
tht oboervation U,at anima!. nf similar kin.hip
do nol rontinuoooJy evolve in certain direction.
but "",,,,ly l ran...,it a ";n,ilar potentiality in the
genesis nf n ...... charo.cte ... "
Schindewolf (1936, 11), when attrib"'ing
rertain ca._ of parallelism It> the
\'Erbgut'l in oomJDOn to the lincageo concerned:'
adhereo In tht """'" idea; he uplairut (ibid., )
Ihe "...,pealed and _minglr independent forma_
l ion nf triangular con<:ru. in three Ii,.... of Cly.
mcnid. by the lact lI ... t all th_ fines "are
cIototly ",Ialed, derived from a COmJDOn an"""lral
KrouP, and inherited from it ""Ientia l ft)["
the nf Iriangulac ohape." (Tran"_
lalions from the ("",noan.) A ";milar way of
thought see"", 1;0 be alluded to in a more 1"<'<;I'nt
paper of the ... me author {1944 : 2791.
Finally, HubLs (1944: 302) , in discuMing hi.
"independent homologieo," h<:rc belie""" to be a
..,..,ial case nl h<)rnoplasy. after Italing that
"the .. n .. or a llimilar modification of I>omologou&
atlnbm"" may O<:cuc ... in d;"tantly a ....,lJ
,.. in d"""ly related groupo" add., "E ... idence
keeps accumulating, Ioo ... ""c. to indk ate that
ouch homologies are particularly
charactenlttc nf clo.ely related lineaR .... " In
..... Slm"""" Iuo ,100 "'nll ""'. ,.m <> ... ""'Y in
""Iv< ,....,jc """"'" (root tb< I.,..
""""""*'-....... '0 "' ...... in the .. "", .... y. Some,.,....
.. ..... ,ho" ."'" ""ty "Ddt< en.,;""' ......... t
<t><><Iitiono .... io """nUon ... _ devtlopmen .. 1
(oc,'"' (I .. i_nee . .. _. "" ... in ..,.. oi".;'
b. tho <qani_!. In ouch _ ....... may. ind>d . be
Lottn' ..... notyPlc c .... <a<t< .. ,ha, ._, .. """,ted ;n
dill<.-, Ii .... 01 doo<m, bu, that ' " ........ tho_ due to
,ho ;"!.ori\aftot of tb< .. to< _ "hi<h .-d not ... ""
mU""N oi""", ,ho """'''''''' .""",.,. 1<1. p . .lJ6),
Tit .. """" 01 OoI>of,t', ( t 9OSo. t 9116). t .. korl to """""
pI&oy by ,ho O<ted haU .. bich both ,_ Wtlpoond
,...",., ha"" , n """""'"'. do<o '"" """" .... 'hin 'he Kt>pt 01
thit ' lUdy.
" c.-""", ,ho ._. lrom 1884.
OTTO UAAS AX' D GEORGE GAYI..QIW SL\t f'S()S
cri,..;" all al ulrineic faaon a ... probably
,"""",,,bIc for Ih," drcun .. tance."
In OUr opinion. 110""""<,,. IIomololl!' ohould be
l'<!CO ni w;! only whe-re ,. cleM oblCrvalw",,1
evide....., of a common of the si milar char
acten: wherev"r new ch .. racten a ... indepen
dently aoquired in di fferenl lineagea, no assump-
of _ inheriled laknt or potential JI'"C-
W "milar dun&""" ohould be deemed
.... fficicnl 10 ronsido.-r 1_ "","rae,.,.... h<;Imolq:;.
QUI inRead of mc:rd y hornopIutic... In ... p-
porting this it iJ 1>01 nINnl to deny the
poIIIibitily Ihal tomatic cha",ctcn may
inliependent!y bauoe of the cornmon inheri_
lenee 01 an basi.. 10e in{en
tin" il In the aubject;ve clement in ap-
plkatioo 01 u.- phylol;r<::ncl k (emu by placing
\hoem "" fully .u r-aible on an oo..rvational
pbne. 10e relationohip the "'-cved
phcnot)'p<'S and l he renntypes . .. hieh cannot be
d iA!Ctly oJ- . [ ,..,.. and ",hich an: _U) unk...,.....
in the po nt"13&O' of reoc.>...:h, iJ ... bje<:t to
"pArate ron,.;&,nl ion and ,,",I nol be inbc.crt l
in the uoageo oIt"" cern .. hc", The
,e<>etic ptocew l h<-re in quettion aloo hI"" a
bearinr on the oubjeN of I ... <alleli"", and will be
mentioned again in thai conn""tion (oee bel_,
p. lM).
S'J<O)lVK. ANO '!!lATeo TnKS
(0) lI_<Ho
II has ....... pointed out abo, ... thaI , .. """
t.omoIOf!:)' in ita mod ..... meIOnin, hal"""'" to
,ndie",e juot what LankKler .,..",.,.... 10 <::Ill
the latter term might .. ell bedropped
u a I)"fl onym of th forme-r,
T"ie, (1928 ; 162) vie .... that ... ;"
not a thins: of moonohine. ,. the
hnrn<Ig<>ny i. likely 10 be cOOlluoed .... ilh homo-
... hkh has a m<>ch """", ink ; thi.
app!;'" in an e-o..,n higher dealt<! 10 1M adj""l;ve
hom0l"netlc. Equally &uperAUOUI iJ ita coun-
terpart heWOf!:"""tit. All uoed. e. R. by B""""'an
(180$: 456, 451). whoec meanin,;' rendcft<I by
"homoplall ic'" ju", al well. Aecooding to FO,
brinK"" (I888: 1122. footnote I), W. K. Parker
""'" tM 'oymmorphy"' inot .... d of hontol-
011', but it teeml hardly eVet to 11.>"" been llIood
by ol her aulhon.
Synon)",,"ul with IIomogt!ny and, theref"",.
.. \\1m..,. ('9!1, 119. 91 ) om .... It ..-.tially tb< .....
__ '[' Mol" ' F'UUC:_"'''''
01 L V=-tic: ........... 10 ... ' i .. 'ordUoal _ .....
...... (if. btbr.
a nother lI)'nOQym of I>nmnIorY 10 homopltyly.
which -.nl to ha"" t-n uoed Ji. ... t by Paltmn
(1884 : 17; cJ. FOrbringer. 1888: 1122) in the
lorm "homophyl" lloth thi. ad_
jec\i,,<, and "homophyletic .. are IhUilynOO)"m.
of "homOIogoUI."
(6)
n.e 'l .... ti,," whether or <101 analon and
bomnpluy ..., Iynonrm- hal t-n uamin"" a"d
e,..,nUtaJly an .. ..,;d in the abo,,,.
Thul 1>tIrnc>pI.u)" .. a "Y".
OD)tn 0/ . ""\0&)"; nor i. anaJocy ynon)"m 01
homopialY, but there ",any " the.... In_
dM<!. "' ...... ')""""yml of !hil term can be found
in the 'i terature t han 01 any olhcr here dealt
... ith. Thi, mOl Y be d .... , ttl a ",rui n to
the lacl lhat original upla .... t ion 01
tlw: " ho""npwy' "' ... <>01 """'ily unde,
.Wl<bblo.
II.,....,,,,,,", 1M first of [he t......-no hen: 10 be
diocllllOd. Da ..... in. is
<>01 euetly a IYnonym of hornopl".y. In the
6,..1 edition of the oneill.' Sp.d" (1860: IU ...
144) Darwin Ihil ter'" wi th the ",n_
tence alJowo (p. 326)," which IIllYI.,..,.nti
a ll y the ... ,.. (18700:1: 39) in hi.
...pla""tlon 01 homopla.y at 't he """'" force<
[actinr) on ""rta in 10 .. 0 organi .... l , which parts
are <"UCliy or .... arly alike and _tim.,. hom<>-
If''''tCic'' with the
aIled rorth in lbe ,"",,,,,,I ""nl in 1M 1_ corpn_
;..... [beingl """rly or e"",,,ily alik .. : Osborn
(1!IOla: 261) limd Da .... in'l 'A""kJgou. V"ri_
.tion' among hi. varian, COl ... of ""'naIOflY:
thousIh ktoeping it apart from Lank,,"I""' homo-
pluy, at Ihal Ii"", not yet COO"rt!Ctly un<k ... t.ood
by Osborn. In cont",<l i. tinctioo 10 homoplasy.
",hid [ i' ''mply conc." n.,.j willt ';milarity that iJ
01<0/ (regardleM of what it iJ). ... ""1-
oroul Varialion io an a l wnpt to uplai .. homo-
plasy, corninc lairlr c ...... 10, t hou,b toOl exactly
", Ih parallel; .... <_ beto ..... p.. 337.
(_noc .. 53). or ..... y be consid..,;d a IUbdivision
01 homopI""y. ad<Iing a t hotOr)' as wau,""." AI
", M""",,". _ 01 nlooonop/lylrtic:". _
._. p. J14. 1-...... ll.
.. 1:10)'<1<0, (I 11<1, 1 H ) io t.t,,,,,",,, lioo [It.o [ .... tblo .I'"
plita.;o.. ,"" , ........ q,- " I .... to ",,, 0,.0\<1 ...... .. Itioh
.. _ ""ly opoo;ioll, lout 01_ iokotital
be q ... ' ioo.<d ...... ..::It .. ;[ .. tho .... rio'""'" ... hido
...... _WJ[ col .... Ioc-. b, 0.,...;0 bo, _ llloo orpno
... PO'" .lIoc:rod.
"N_ by s;., ...... ' It .. tlou, .
wittom "I 10-,.. 1O ... .. 1 ........ to ... tloio_ .
ANALYSIS OF SOME PHYLOGENETIC TERMS
though occasionally mentioned by lat .... authors,
Da ..... in'. lenn h,.. Dev .... be<;ome generally
adnptfll: too, t""- younger, more neutral,
and, at the ... me time, wider tenn homoplaoy i.
gi,",n Pl"ferrnce.
Another which aloo antedat ... Lankes-
t..-'. paper of 1810 i. luWol4u, by
(1868: 2S1) to designate "what s-.,.,.l.......,
and othen called 'analOgy' a. di.tinKuished from
affinity." Thi. very definition proves Cope .
Inn 10 be synonymou, with h<>moplasy, as here
understood. It may con.idered entirely
ot..olele.
Of I",er term. 1oom..",,,,pIoy" ;" conoi<kr<>d-
decidedly by (1888 : 1122). doubt
fuUy by Osborn (190la : 26l }--a .ynonym of
homoplaay, whkh it oeem. indeed to be. Thi.
term was, bowev .... , nol coined by Filrbringer,
a. eegenbaur (1898: 25) , Osborn (/. cit.),
Bourne (1910 : 40), and Hubbs (1944: 301) be--
lie,"" bul i. credllffl by FUrbringec (I. m. ) 10
the Finnish naturali" J. A. Palmlm ... ho ""'" it
in .. pa!",cof 11184 (p. 17), along with homophyly,
in the followinrr oenten",: " ... inw;ewt"it
Formen v<>n Orgalll'1l und Organe ""Ibot. die wir
jet. t homolog nennen. in de, Tat
entotanden, al"" "-/>11)'/ oind, od<:c ob sie nur
ar. I::ndresultate tiell
entwide"lt ". To judge by !hi. sentence.
homomoq.hy may _II be considered a synonym
of t ank ... I ... , homoplasy." Filmringec (10<.
riO. however, ' ejoe," both lerm. lor- rea""". of
COR"""ien"" and f>WI>OOCS to u.., W. K.. Par-k.,..'.
'";'_/>IIy.' or "i ...... "'(JMmI," "ao uoed in the
morphological field," instead. Thu> ioomorphy
(IoomO<"phi"",) may here be dispoeed 01 an as
oth.r synonym of homoplasy: it ;" <><xasi<>n;olly
(e. II"-> Hower, 1941 : 12, 49) u.oed .ynonymously
.. ith homeomorph}" al.... In both meani ngs the
u"" of Ihis term ohould be diocouraged 101" merely
etymological reuono since it indicate, lull like
n_ 01 .hape, harrlly ever round in nature, not
even between individual. of the "'II>< .ptti.,..
" 11"""""""' .. it .. , ....... lU'ed .. lnady ;"
0-.,., (I!on, JX G"-o'Y. bu, ... it ...... 1d _ m I"""
.. pia ... , .... "Of ,;mi"'r ''''''',.. ,..;,b no _Rico)
_Inc
.. "_Id "'" be.o<w .. ined ,,""tlou.,.. no! 'b;, .... toneR
ron ..... 1""-1 U>o (adjocti .... l) uoo both .. ...........
ph..I"',mo;I by Pal ...... .
.. 1.tbe'i .... ...... l_.od 1945, .. ..
_ .... o1y ....... """"'-phi." ... n"""_ i .....
".....t papn-o 11"""-..... '-' ... IM to fiod
.... t wba' """'ninl II ;" b.,...,....,.".y ("botno-
morpb;"'} 10 N.mkoll (l9J6) .,;_ <hi, pap ..... not
. vai!&ble.
Eimer (1898: 27) listo amonR hi. "deven la ...
of evolution' "the law of independent similarity
01 development, or- 1wmO<lr.......u, .. which otate.
that the orune direction. 01 evolution may be
operative in dilf..-ent and not immediately reo
laled lorm. and may lead to quite simil ac mor
phological ","ults." From Ihi. de/inition it
may readily be conduded that Eimer. term-
u<Cd by Grabau (1913: 916 If.) and, evrn n>O<"e
amply. by (1921 : 245-2M; 19J5: 131-
loW) in d ooe connection with but
olhe ..... ;.., not wi<J.ly auoepted-i. ju", anothe r
. ynonym of homopla.y. Eimer (10<. <".) add.
lhal waS "alterwardlo demon
.lrated by Hyatt in ArididM a. mOt"phologi.
cal eQuivalenu, and by eumples from
Ihe shell. of
All a matter 01 lact, Hyatt lonnulate<l hi. con
""pI 01 "MM/>IIolotk<J &Juiooknu" fi ... t in 18-84
(p. 361), deriving it lrom hi, "law of organic
equivalence' a. 10110 ..... :
Th. la .... or o'o.ic t"i,olo. may be d<lcrip.
uv<ly "oted .. 10110 ..... tho aetion of physical
eh.angn tako ffoct .pOn an irritable O'1:ani, .. ,
.... hich ."", ... rily .... pOnd. to the ""lernal .tim.
Lanl>, by ao int ..... l ..,.etio. '" <:fIor!. Thi, .ction
lrom ... ithin ul",n tk. paTto 01 tit. o'1l, niom modilied
thoir f.,.."", by the prOd.ction of n ....
VO .... th. ot" which ther.,fore. adaptod
or ouitablo ..., the condiuon, of tho habitat, .,.. Iho
phyai01oy;ieal tuinl " of Ih. phy';"" "on" and
lorce. lrom .... hich th.} dir:tJy Ot" indirectly origi .
nalod. I n"" I.r th. n .. "". _ and habita
aimilaT thoy produ"" "'pn .. ntauon Ot" morpkological
tui.alon"" in ..,rio, and ... I.r ao thoy
dilf .. oot tltey prod.,," the dilf.nmtiat. whieh di,
ting.i oh th ... ..,rie. from oach othor.
Later (1889 : VI II) Hyatt ...,..phraOO'd Ih;" con.
""pt ,.. foll owo: "M"'/>IIolotk<J EquiDak"u.-
In tbe diHerent """'" of a Iyr>'
lrom one .tocl<: there ;" 3 perpetual
Teeu""nC<! of similar lorm. in similar .,,,,cession,
which are usually "ailed t cpt_Mati"" and often
laloely dasoified together, though they really
belonr to di""'"lr"nt, genetic oerieo." adding
(ibid., IX): Th_ fo,ma and th.ir similar char.
acler;"tico are not deri'-"'<l by direct inheritan""
rrom the common ancestor, in .. hieh all tbe
form. are n""""",,cily similar and primitiv<:, but
originate everywhere independently 0/ hereditary
"5".I1;nc of tho E...,I;,h "" .. Ia, ...... ..",.. !G
hioo Cor ... n po ..... F.i ...... Op<11o the _ .. ;V oM, .,
Su..:. lIy.,,, "c;..",m .... tbr An.tidae' in
tit .. -'>, to imply tIto.t imor, '''"'' "" ... bo.dr."
Ieoo,to ItS&.
(Tno HA.,\S AND GEORGE GAYLORD snIPSO'"
in the diffe.rent toen"", anti al.., in all
formation< independently of chron<.>l<>g;cal 0<
dtOrolngical distnbution." Th ... Hyatt's ""rlier
deJinition, although called deotnpti"" by him. is
"""n 10 be explanatO<)' in a Lamarddan sen"",
whkh ill intentionally omitted from our rotIcep'
of hOJn()pla.y. Ho"..,,,,r, from his laler defini-
tion it may. in Our opinion, well be inferrro thai
hi.t mo<pIJolngical !uivalents a", nOlhing el""
than the r,,"ult of homop/aAlic inlluences. This
Il:nn, which h"" not b ;ome widely acttplro
either," should, the",f_. a.loo indudod. in a
broad ..,0"", in the oynonymy of Lank .. ter's
older and "'orter term homoplasy.
Finally. Hubb.' (1944: 3QO-.l()2) "IIIdtPtlldtIU
alon turn out to be nothing doe but
special c.aseo 0/ homoplasy, as he hi,n.df indio
cateo with ,he wOrtJ.: "They....,..., apparently
prominent in mind ,,hen he fin'
(1870) propo!I<!(l the term homoplruty to desiRnate
the similar but independent ",odificalion 0/
homolO!'looS ,""etm..".: His stalemenl that
"Ihe same 0< a similar modification of homolog-
ous attnbutes OCCur"" convel"l:ent or di
VM"!' nt as well at on ""rallejlines of evolution,
and in distantly as wen as in clooely relaled
groups" IUpporto the aix.>w' vic"'; for all the.:
modification. corne under the wid .... COftpt of
homopIasy_ Huht.. i. certainly n&ht in ern
phasi%ing that the boundanes belw""n analogy
and homology an: rather bUI O.hom,
(1902a-not 1907, at quoted by Huht.-p. 270)
.. nten"" ". . I think " 'e have 10 rleal with
homology or. more .trielly. ",ilh a principle in_
tenna]iate homology and analogy"
d.,.. not ....,m ,...,11 in point, dating. as it doeo,
from Ihe time when o..born had not yet
at a undentaIXl ing of Lankestcr'. term
homoplasy (..,. ahove. p. 322). T he "uamples
of Identical but in<.iep"nden' morl ification of
homologous "ruetu",," taken by Hubho (ibUJ.,
301) fro", the of _tain groupo 01
noh ... an: to WI ca_ of- heterochrnnOOl ...... par-
allelism or, in on" ca .. al least in ",hich "the
end r-ewl'- of soch parallel evolu,ion may be
.... arcely distinguishahle On any morphological,
phy";ological or developmental grourKt.,' even
if not homeomorphy. Huhbs'
ca ... give us, on the ot"'" haoo, ""'" examples
of morphological, if not horne<)morphlc =ur
renre (verh. ; "repeated!}'," "again and agaia")
in the Clupeidae. Thus fOOl r or five
11;>, . J . .--.p<at<d,y u...-l by Buekmo.n.;n h. po "...
on a...dow,p,d (1\lO6). ;n tho a'*""<via""
"""""" .. """ph", ""u;v.I<n.:'
terms may well be aPJ>lied 10 Ihe phenon .. na
which Hubt.. has in mind; there is "",lIy no need
for a Jiflh or sluh. Ihe less eo";.....e the phenom-
ena nan>ed by him "iodepe""ent homol<>gies"
are, in our opinion. independent , it is m e , but
not homoIORie .
DIVERGP.NCF.. P.-\RALLEI.lSM, CO:'VERGENCE
A.'!O
{al Di_r'''u
Although lhe "",Ii'alion of II>< phenomenon of
divergent evolution i. Iraced by Oohorn (1902b:
35J) "" far hac k as to Lamarck. t he creation 0/
the phylogenetic lerm "diveryenc<:" appan:ntly
has to he to Dan<in "'ho maoterly ex
plained it in hi. Oriti" of under the head.
ing "Divergence of Charnet ... ' (1860 : 103 ff.)
and illu.trated il by. dasoieal diagram. Start
ing out, all ... ually. from experienc .. in
domeotie ani",als. h. wnteo; "H....." thell. ",e
...... in man's production the action of wbat may
he called the principle of divergence, CaUolnll"
diff ..... """', at lim harely aJ>Pl"l'clabJ<, steadily
to increa"", and til<- breeds to divel)(c in char_
acter hoth from each o.h", and from ,heir
common pa",nl."
It _m. that the term hrut been need ever
oi""", will,oUI any pen:<'vlible deviation . in the
o<iginal ",caning given it by Darwin_
(b) P(Jf(JJk#,,,,
This tcrm I. younger than the p<ecWent one,
and may be <aid 10 h""" into "'orl<l
"""" or '- incidentall y. W. are unable to
clanfy it. origin fu lly. It oeems to have been
u....-d fint by Cope (1868, 1872: po .... ''''), but as
Osborn ( 1902a: 260, Cope
gave il two differe nt m""nillj(S: On on. hand. rut
an equivalcol-of "",,,,,,Iulali,,,, (of the previous
phylogeny in the OJIlogtny), a. ddina] in 1812
(1887: 7) and. in aO emended fnrm, in 1896
(p. 20). The oruoe meaning io implied in the
headin& for Chapler I I I of Pari I of hi. PrimM,
FadI1rl of Orll""'" (1896 : 175-210;
note, es'-;alJy, on p. 1;6: "the ""mlleli,m he-
taJlonomy, ""tog""y, and phylogeny")."
On the oth ... h""d. COD' used, lOOulI'h heoitantly
as il "",n:. thi . l erm in the modern meaning 0/
para!lel i"", of .>'Oluti"" aloo. Then . ho",'.ver,
be spoke moolly of "the parollds:' using the
.. Tho. ... 01 ' ..... l .. 'i"" till '",nd .. ;n 1907
;. D.pb-., (p. !-3, "u I<M d'lhoch' ",loti"" ,u pora'!<"; ....
du Mv<lopp<ment "",,,,,,,,,,,iqu. <t du .... ,
palOont<>l<>Plu< d<o w...").
ANALYSIS OF SO}'lE TERMS
I\OOn "poora!leIism" ",,!y t..ice (1887: 98. 11)2).
It thu, IeJtrnI that to Cope paralJt.liorn .... a
technical UNn. meant rtelpitulaoon and t .... t
he u-t the word in it. modem meaning only<>e-
caeionallyand incidentally."
The ... t, a t !em in America. to u .. "p;uallel
iom" in it. modem .... aning and to
dofi ... it po .. ' dy ..... W. B. Scoll in hJ..-
important paptn of !89! (pp.. 362 fl.) and 13%
(p. S6). For. critical analyw;. 01 bJ. defi.niciono
m elco io to tM nexl htadinlr;.
Some yean la ___ ntei- "pa<alJdiorn"
aa I ... 11 phyJog .. ",tic UNn. hei,,&"
u-t, fM enmrk by Buckman (!90I : 231, 232,
at "pa .... lll'liMn of ").
(.) c-.." ......
'Tho pI..ce of on,in 01 thO. term .. found in tM
1.1_," ... MOil of Carl V",I. Prof...,.. of
GeoJory ond ZooIof!y in Cmo-va. 186J (p. 2M).
I);ec:u"'''' It... family t_ 01 t he prirnateo. he
...oct :
0..- Typus Afie rOhn"'" .. hl .... che.o Ridltun",_
.uoei ... oda-: Of tcir . oido nenl I ....... H"uptbw.
Allo" do, al,u. All.., do. " ...... \\'eI,. Jod..- do-r
U .. ptll " " .. b. ,.. .. t.u , .. t. ""d Z ..... die oteU
meh' a .. oi""nde " r.h_ I(h.ino.. Aher di<
V ....mo ..... 8nl Z"';I' .. it ill S,.. ...
...... 1', ....... 04... Auo don 00 "u"dwnchiod ..
.... do< Gibboo .. do< M.hken unod do<
p",.;,,, ".orid<.r" wei> die drei _oclotna),'id>c-o
All ... , die doudo ..... MonCO co ................ Cbrak
"'" .;_..tor bodtuttod db .. ,_, all Gn. ...
"' ..... .s..- Spiw.. ole oich bdi ..... " (ital;co 0.'-).
Hunl (I&6t: (63) tranolated lhe two Iut
Itnt<:...,.., not quile ... lollowo:
lI ul '" .k "'1' , ..... O{O;.
...... otk .., th.t from ,h. fundam<1ltalJy
di ldn"1 l.mi1i .. 01 \.b, , iboo .... Macari, and 1>&_.
... developed the , h_ uthropold 'po&, whid. by
a "umi>tf 01 00lIl""", <:ha_ ........ od "".MIorably
.'''TH r.ach otloet u-.. t he IroII", 01 "'hick u.ey "'"
the I..,ad. (italics "" .. ).
Th .. vny .-.are 01 lIu"t', IIllnIIatioo J.
quoted by Oaniill in Tu lNJuO/l -I M (1871.
I: 211- 222), ..-htn "" decb. .... il to be
poooiblo, toou,h I., lrom ,,",bable, that u..: .. .,Iy
Pl"OCt1lilon of man milh. a' han
muth i" cha",cu<. thoy becam. mOO"< u"lib
Hob other tbaa any ,Ili.ti". run, bu, .hat , ....
_oilY . .. ' ou<1I<tl V.". /My _tOJ i.
....,oan. Wltoil ....... 1I'cto fOOl' .1It .. "'. obj..:. tho
0/1""' "1 01 .wo dilli"" ....,;n. he """"tin>eo i,,
d"_ u la, ... ,eneral . P1'" ........ i. COft,,..,r
""noid.",br. "moun' 01 < ...... ,.. " (it. Ii" ou .. ) .
A. -" lrom this quotalion, Darwin J>M<,
6ut to the phcnomeoon mentioned by
Vort, which u ron ..... , ... oce in Itchnica! .. n .. ,
if 10 <nate confuoion from lhe very
ou\ltt....;m ........ tely aflWard. par.olleli>H it
with tM oombinuion 01 the eha.raettt'5 01 t_
dilunc:t eptc". in tbeU ODmmort oIJ..,nnr aa "
fftljlt of h)'bridization brought about by man,
... hich is J<ol conwrgenoe. 'Tho nOll" "con......--
l[tIlee" i. applied only 10 the latl.er phenomenon.
but .. nee Darwin appliea the Vt'rb "to Cllllve<ge"
to the former . juolly CTfllilillJ VOCt with it.
6nt dttcriplioo. t... may """II he c:oruKlered tb..
crutor oI .. """ ..... ,g<...,. .. all a term
(aa distinct from the original . _' Klric meaning
01 tM word). n
Evoea tltoouch u.-, f at"t& clearly prove the
Sw;"' Britialt oricin 01 the term, il oeem. 10 .........
_mI a definitely German fbvor aJ'ter<nrd. ,
btcau .. it .. u ""'nl .....d n-.: ren
erally by German otoo.,nta than by loo.c of ol lter
nation,. Thu. Ooborn (19020: 200) apeak. 0>/
" the 'con_genz' 0>/ writ<:lI," even
. peJ!ina- the word the German way, and Traquai
(1905: 382) OItribut"" .....ain oimilaritirs .. '"
pa.ition of tY"" in on the ....., h.and
and certoin primiti... MhH on the otbn' to
"wbat the v.,rmanoo "'" a 'ConYefi('ru-i'::...:hci
nuna:"
That the lerm w;u in the late nineties iAdeftl
quile familiar to Genn;o.n ICholan. but mud!
leOIliO to En,lioh and American onee," rnay he
from a cornparioon of the texU of
Zittel'. (1895: 14) and Eul man',
finl Enilit/t ",,'tiart (1896 : Il): Whereas
.....d the tenn (in tM ODmpound form " Konvcr
la"lOl'l)l\ by German
ot\ldt<lu, c/. below, p . .JJ8), Eastman .. .,.. reluct_
ant to ifl'rp< ..... t<: it in hio _ t ond P"lIphrued
Zit!d', ClIp.ui ... ; ... trad....;n the mio:klle 01 an
otherwiae fairly faithful tranaiation-by tM
lither cumoo ... ,'" of word, "phenom.
ena of p"",llditm, or rteurrin, 'c:ommon typeI
of Itruc: tUl't. '" Only in 1t... IO.'C01Wi En,Ii""
.di t.ion (l911: Il ) <tid Ea man adop' ,he tenn
1. 1I ,.I'l'l) I' llt 01
II ....... . io. <he
OTTO HAAS ,\ND GEORGE GAYLORD SIMPSON
"oonvergenCf'," but .till tI.,.m.,.J it nccns.'F)' to
I'"t it int" 'Ill"teo anti to atIti the term "(>O.ral"' l.
ism" whi<;h he obviouoly Ih<>ught I" be more
familiae to hi' r .... de.... Thus " Konve'!:.nzer
ICheinungen' ;" n(IW translated by
of parallelism or 'convergence."'"
Sl .... ngely a .. arch of the German and
Au.trian ]ite .... ture of the I""t decad .. of the
nineteenth and t he nn;t of mi. cenlUF)' d;,t not
yidd any (>0.1"'" .. discuosing or denning
eithce (>O.calleliom or convergence. It """". that
both ,,id.ly uoed on Eu"-,poan continent
in teaching .... well a. in paper$ dealing with
certain groupo. but not made the oubjects of any
analytic studies. T .. i. imprmon i. reinforced
by .... t..J'. (1909: (222 complaint thaI "the
teTm 'cunverw:n",,' has recently been used ""F)'
f"",uentl y a nd ruined Q catchwor<:l. a8 Mve ...
many tenns in the field of evolution. but that the
vru-iou. stutlentll could not ani"" at any uniform
uae of this designation." He add. that "in
addition to t he term 'CQfI""'Ol("flce' the .x:.;gna
tion. 'I"'r.l.llelism and 'divergence' ha"" come
to be used more and more.. .. The ""Iy
author credited by Abel ,,ith "having. a. t he
n"t, attempted a sharp analYl ical ""paration of
the concepts: (>O.rallcJi..". conve<gence and di.
vergen",," i. - H. F. Osborn (190.9. (T<anola
tion. from t.he German.)
In the meantime Scot! had introduced "con
vergen",," ao well "" "I"'ra lleli.", >0 in the Ameri
can literatu ... (IS91, 18%), clearly de6"ing- them
and cmphapzing the imponance of thege phffiOftl.
cna lor evolution in generaL Under hi.
Osborn also uoed and d iocus.....:l th .... , in ""me of
hi. papers "''''ting to <>f phyl
ogeny (l90la, 1905. 1097a: <1. Abcl 's ",{crence
<l uoted abc=).
lkpl!,..,i (1901, 22J lin....,,,,,,.. docs n(>t
make any such clear distinctions as Scott and
Oobo", belween parallelism and convergence.
He attrihuw. the de CQfl.vc,!:ence"
(OOviou.ly a translation of the German "K.on
vergt:n..,,,,,,heinunKffi ") to "un" ""-'e de nlarche
01 t he cvolu.ionaey motli6cations of
""etai n grouJlS, thu. producing ap/>t<>"'UI <>f
mutual approac" e""n to confuoion_
Once (il>id .. 234) he m""tion. "ca. paral.
"In , .... ro ... , Ena-t>oh O<!itlon, tho to ... , .. D....,.
.... . .. in .., ....... b.t! to _ .ft<>th<r by ;.he.it
. moo." fol\owioa- ,he word< "meuni .. '""""""" t)'1><O 01
"""<t""';" mad< ...... 11_. ,hu. """Uvo .eo'eo<
-. unf..-t""",ely Ief, .",,1_ iD , t.. *""'''' edition.
it """trt to .. pI><""' ...... 01 .... 0.- '"""",r-
..
... of uncoiling in nautiloid. on the one hand
"nd ammoniteo on the other. but a ltogether he
t"", .. ... . phenomena here differentiated as
parnJleliorn. convergence. and homeomorphy
under heading con,,,,!:,nce <>oly.
fn the cou,.. of the decade of t he t""nti.
eth centur y t"e ",rm "cmt""rgence" oeems to
have becumc more familiae '0 English 'tudents
as a. to American "Del. In 19 11 A. Willey
publiohed I.;' book on Co_.""",;,. J:;roIrdi",, _
Since 0011. ter",s parallel is", ant! convergence,
though not their de6nitions. ap"",r to be 6rmly
by then. their history a. ouch need
not be f"Uowed any further. Ho,.." ver. tt..
opinions of late,. authors as weU "" of thooe
hi tbcr to wiU have t" be di..,ull/!Oed
under the f,,((owing heading.
DEf,slTWsS AND DI!IT,"ctWsS
( .. ) c .... rol
.... U three of the term" here discu.....:! Mve their
orIgIn in geometry. It i. bebcved that their
usage in phylOitCny wiU be clarified if their
original i'CODletric meaning i.kept in mind_
If this general c""",,plion be adopt..-l, it f"l
lows that each of th""" term, should be uoed to
indicate only the mutual relati"n of t wo (or
nlOU) definit" duri n, a certain ."",tch
(If l ime. Hence it folJ",,o fur thee:
(I) That tbe same t wo lineages may be div.e
gent, pacalfc!, and con"",!:"nt in con$eCuti""
time period even rep<'.atedly ".,. at illustrated,
though in a ",mewhat """r-simpli fied way. by
Willey'. (1911: 70, SO, 112) fLgur.,. 2, 5. and 11.
This i. obvi"u.ly the mea ning of that author',
(il>id_. 69) ",..,ming .. that con""r
gence d"I"'OO. 6ntly on diverw:nce and oecondly
on pacalldism ...
(2) T ha t a certain .t"'teh of the cvololjon of a
given lineo.ge may be divergent in relation to a
""""nd but at the oame time ""rallel to
a third, and """n c"nvergent ,,i.h a four th. To
make this clear, Ict u. imagine that'" the fo",_
li"""ges cnding up in the fom" wH. yH. v". a nd
.", in the upper ri<:ht corner of Darwin', (18()(l.
betw.,.n 1'1'. lOS and 1(9) diageam illustratin<:
di""rgence the y. and v.lineages exhibit I"' rallel
ism in one character, and t he v and .lineages
convergence in another character, between each
other. Then the vlineage wonld be ""nver
gent with Ihe .Iin<'.age. parallel t" the y.li",,
.. A " mib.. <:<><><rp<ion .... y bo roood , i. _ .. ot ....
_ 01 $chiN' ''oU', (19<14: 11 .... 280) di_...,. 01
.. "" ... lle1nihen" (po .. tt.I'..n..j.
ANALYSIS Of SOME PHYLOGENETIC TERMS
aee. but OliU dih' ient in ",lation to II..,
.... 1Dc ... and yllneai"'" would t...
""pecti,,,,)y. CODVCIlIent with. and paral"'l to
t he vlineage. but di""'icnt in ...,Ialion to the
t wo otlle",. w. lineai'). howcVN. would
ti ll t.. d;"" ... i", from all thn!ll 01 tile ot h ......
(6) Di_,."".
Of all the conccpI' ui'K\er thia
_ ........ to loa"", been subject to thr Jmst
.""""iI of d.,...,.;o" and hardly
any rnodemizalion. Schi...w,wolr. (III: 213,
fie. 3) diagr,un .. an u",l!;,nt ide.>
01 the t.uonOInic implication. 01 thi. """""pt ....
conceiV<:d by modem otandardo,
b UI il doeo not oho ... any CMIlnt;"1 chan,e. u
comp.red to Darwin' ..
Simpooo'. (I 'M5: 12) ..,.,tonce: "If t ho "",Iy
mo:mbc.s 01 two ,moll'll an: more alike than u..-
bter, thei. later dia.imilarit;'" an d;vn-gcnt;'
may .."..., "" an up"to..date ddinitioa 01 di...".
".ooe.
Otbom. ;n propooing his " La .. of AdaJil;...,
Radiation" (1902b: 354). cau. it a "law of
I4J tul41>/i.,. ,ad;,mol< or incipient di',CliCl>te
on omaller OCIlle in a oink: locality" and "".
plai"" it later (1910 : 71) ... "an applicalion of
paleont%iY of t he idea of dive......, t evol" l ion."
(e) C.......,lt'.n
NOIwithotandi,.. Oaborn. (19026: 3.\.4) lUte-
......,1 that bomoplaoy. paralk:lism. and CODYCr'
"""" ... are "by DO mcaRllYROoymouuen n.
or identical 1'" c . ' " thenl hu been, oince
theM ........ -.., born, plenty 01 01
opinioa, not to .. y """fullion, .. to t hei. dc/in;.
tioo and their from eao:h nthe<.
Thil coofusion bea ... , 10 ... ith, on the
hierarchic order ",,,,,,mi n. Ihe i"Clation. tH, t....,.n
them: From Lanknter'. point 01 view, as con
by Boorne (1910: 010) and, 1905
(po 228). by Oobom (_ po JU), I>omc>.
pluy ;. the higho-r ,,"tq:ory. including '-h
"" ... 1"'6.." and 000.'1."1.......... Thioo is 1M <>pin.
ioQ adher<:d to in thi.Mooy. Bourne (100<. cII.).
'-ever. panJldi ... a men: .ynonym
ollo.on>oplasy, and, opoakil\i of"a oontrnot tJo..
: ...... n homoplasy and lea"", UI in
doubt '6'hether theM! t wo ,,",,<1i .
nated or if t he oecond il oobor<! inatcd 10 the
6"'t. Willey (1911), in turn. conoideno. ra th..,.
OU' prWnlly, """....,......,ce the higher category.
includinc homoplaoy," mimicry, and paralltl.
.",o.u.. :' Ill' ;, _ .......... ".It ._ ... __
10 _' ,"."Y" ( ..... ,
lam,- Ihu.evm I mvi", u. .. pa .... llel ooo-
verre ....... (ibi4 .. 78) which. by the ... ay, ioo one
01 the Ii,..,.t nampa 01 a ,f>OiI.odOUi<>'II adjtdiJ
to be encountered in ocicnti6c literatu...,,
Even ... .,... appearo t o be t he CO!1fulion a. to
... 01 the varioul t!>em.,l"", .
With _y few oexceptionl the delinitlona given
roo- the one art applicable to the other equally
or even better, or t hey may even COYf."I" tbe
other but not thr one tMy .'" inte .. tI.!d to
dmnt."
n ... Zittel', 01 " KonY'Cfl[f:n .....
qUOted (p. 326. footnote
18), defineo homoplaooy rather than oonVM'i[1'n.,.,.
WIman', oentenC<." in t he ...conol Miliiih iitlon
of Zittd'. textbook (1913: 13). al q"'ltcd on
POOl<' 332, lum", t ... o PI<C L I I (llOrallclioom and
... hich al"C distingulaMb!e aft..,. all.
'I"be wordo in the 6rM En,gliah iiti""
(1896: 13 ..... p. 331) a'" ttlUinly r""'"
"""" .. tWactory ;til a definition of the phe;l(lmo;:na
of COlhCO,CI '"' which they _ 10 "'"
Ewn , .... otudents ... ho make a dear the-
oretiuJ tIiotinction bctwecoI panllelitm and <;Oil .
'"<1"J"noo: emphuioe the difficulty 01 h. pracl i<.aJ
applicuiool. Scott (1891: 343) points out:
" The diotinction het ... """ tht t ... o d" 01
p/M:no",cna i. obviouo1y one of dtgrec .... thcr than
kind." (194$: 9) allO call. "the dio;.
tinction . , fa. from .&.Iin,: '''I"be
two phenomena inleoi",de COCIlinoouo1y and
..... oft .... indiootinlUiohabie in praclicoe. ".. Scott'.
(I. dI.) ODDCluoioD that "it -ill therefore be
oon ...... icnt to oonMder them locetf>er" ;. valid
lor :he ...... nt INdy aloo. n<)I. only for 1M
"" ..... pointed out by him. but allO for thOle di ..
CIIII<ld in the p.-coedini ""...,.....ph ..

ul).
"00";.', (, 1(00, I U - I " ! pia _ ...... q_
.-( ... _y -U ....... to d;tIioolti..
01 dioti ....... i>ot; ........ 0 rh.. I , .. In lI<I<olco. A. "-
it ";PI.""",' '"" & ,ypbl ..... 0/ panllol;"" (1oo-
t_ .... _ ro<ta). lout 1 .. .. 0/ """.... '" . ..
. ... .. tit< IDi"","", WOfd ... In 1m .. o.r.
........ 01 .... ,*""",-," ... ,,;" y 0/
_ ..,..r.o 01 _ ___ 01 .... <hatw: ..... 01 ...
. _ Of ...... (p. "00 ......... tI"y 0/ _ opo<Woo
........ , II. ;" _ douocwoo .",.., --." (p.
IU)."_ ...... i , ...... _o/ .......... , . 0/ ....
__ " tp. 146).
OTTO 1r. ... AS .... ND GEORGE GAYLORD S I)'IPSOC\'
Hen:, at in similar cue. , a c!ear ,Iiflt:rentiation
betll'een the two neighboring C01l""l'1lo i. th.,
be.t "'ay to arriw at . uitablc definition .
Appa"",tly the fi ... t to at tempt . ""h a differ
entiation ... :to Scott (1891 : 362): "1/ t he vario".
opecies ol the It"nUlJ acquire the
independently oleach other," there ill
parallelism; " if the ol widely d iffe ... ",t
genera , . . gradually a .. urne a commOn lik.,.
ness," there i. convergence. In hi. Wood',
Hole l.ture of 1895 Scot! (1896: 56) ga'"e the
f<.>lI()Wing definitions:
By p",..IIe1i, m il meant .he indepe ndent a""u;';.
lion 01 similar ",,,,,ture in lorm .... are them
.. Iv,," . nd ...,h .<.
Qui oi ';"" in f<><m 'hieh a", no. cIo",I}' ... la ted nd
.hu. in one or more reopecto come 10 be mor< n ..
.lik. than wo re 'heir 'nceoton.
It will be that the laO! part of lhc latter
definition indicates the geOITl<'lrical meani n, of
tbe word as an ."",ntial characteristic of con
""rgence, but the dijfernolia jpu;fiaJ hetwttn .he
two pr""""""" is .till tbe degree of rda tionship
betoo""" the lineag<:o involved.
In hi. paper (1S91: 3<>2) s.:ot t em
phasi...d "the deceptive character" <.>f thcoc
meaning that tltey oimnlatc a rela-
tionship which i. not reall)' I"""I"' nl (u all
phenomena of homoplMY """"ssarily do), and
pointed out that their decepliven .... "may !'n-
crally be unma$ked upon careful u.aminalioo.
though even the c_s 01 1Iw: Ammonite.
.. MOW the need 01 extreme caution."'" In
hio oecond paper (1896 : Si) he add. that "Pal<-
ontology enahles ... accurately 10 diocriminate
bet"..,... rescmblanceo wh;';h are due to
affinity and th""" whi.ch ..., ... 11 from parallelism
<)T COO"'-"lr"nce"----<>r, in u.., terminology here
pro...-:i, between homologoul and homoplastic
similaritie<l."
.. ..... ou< d "_<>1 I' . ... o.li"" it .... y .. ",til
mo ..,., ........ , D.pfto,- "bQ <10<0 oot ... m "" hlo",
""-n So:ott'. "ri<lop ..... " I the ..... ,oo.,hto witll
.I1noo, <he .. me .....n.: " (Of Woo
f"";Io, ""...." .rrn.-... da .. <Ito iT""_
d;"illrto,' 000_ "",iii ... "-" eomp\Cte """.
trQrnP<r l"oboen .... u' .. (1901 , US). "rr-,ur "",joo,..
n .. , f .,;I. au .... 'u ... I .... d< dt', ""1'.>0' COl ..... ...-;..
trompOU_ on f.i.u' .ppeI ' l"..-.bl. "" 1''''P .... tIoo ..
. 136).
" q , lIuck"",n. 191), 9, "It it by . ... 1,..,. 0/ chane""
. .. ,bo, ,I\< d<o<;o., and the pooicion 0/ f . ... \ieo
.nd "" .. to I>< "",--->I by .-b .... m!'<iooo.
d atiioi,y /""" ,.... .. 1 .amilori'y. "hich toO 0("", may I><
...... Iy ,,-!>'ioo''', Sirnpoon. 1945:
"1l""",iocY" .I .... yo ""litln-KIm"" d allinity. PafOllel
Osborn (19024: 261) adopted Scott', Jil!t:r.
<' ntiatOon , huing it m=ly on the antilbesi.:
relatod-umdatod, thn. proVilking thE following
criticillm by (1911 : XI) ; "Th_ denni
t10m lea"" n, in u.. dark as to whal dcKrees of
re Lationohip "",,uld entitle a ca .. to be
da...w at one of parallel;'m or of convergence."
In hill chMt of 1905 (I'. 960) , reproduced hy
Abel ( 1\109 : (223, Osborn diolinguished "anal-
ogou_ adaptat ions, i. e. simil3r charnet,,", arising
independently in "",iUJr ". rtlaltd ,,",!tIDiJ ".
"."' .... . causing a similar evol ution, and ",""Iting
in j>aralkli,m," and "simi lar adaptations arisi ng
independently in diJ"",iUJr ", unrtlalttl anima/,
.r .. p .... , caumng a secondary simil3rilY or ap-
proKiOlatioJJ of type, re.ulting in
Adding similarity and dissimilarity at! a oecond
pair of chamcten to rdalion.hip and non_..,Ia
liooship was aPl"'["('ntly intended 10 lIive a
brooder bMe to the di$tinction, but it may Iw:
doubted if it it any d"artr.
.... bel (1909) a te<.mpted to ;mp<o"" theoe
d.finitions; to him u.., _ntial d ifference 1>/,.
to .. ren pa",lleliom and conwryenre i. gi,-en by
the fact lhat the former brings about t he oame
", . .. It of t",nsiortn.ation in both forOls concerned,
the latter a "'"'" or Ie .. similar one. For par_
alldi,m the "homOOynam;" function of h<.>rOol _
<>gou. organ." i. considered eMCIltial. for con-
vergen"" that of hetet<Jgenelic ones. Thus,
Oobom'. r<"<)uirement uf non-..,lationship is
mainl3ined fOO" co"'''''gen"", bUI parallelism is
&aid to <>xur bet ...... n nichl
Haher verwandtcn Arten .peeieo, nol
",latOO to any considerable extenl).
D"""ui< (1921 ; 253) repeat.< AbE!".
conception, slightly modifying the abo,.., "'-
Quirement of par>.lIe1ism 10 "'veTllChied"""n.
nieht imrner """"",nd,,,,n Arten (various, not
al",ayo ...,Lated speeies)"; on the other hand. he
not only admits in d<>oely relaled
opecies ""d but find. them to be tbe
mo.l ones (Wid., 251). In ad
dilion 10 Abel'. conrept of convergcoce, he lists
two more which n,ight be called constitutional
convergen"" and accidental reo
'peeti""ly (ibid . 251, 261- 262) . All of them
iom;, 1<00 dimot and b.n i,;, . 1", v. 1id
"ithi. _ .. ha, _ Iimi .... It .... y)e.d '" (IVU-
..ti ...... of de .... d atl;.i,y. but it i, .... 1ik<ly to ind..,.
beli<f ift "boIIy I>Ioe .tI;nity. eoo'"l<_, ho.,cv.",
moy be wI><>Ily ..u.Ioea<Ii .. , and p.;nd ... 1 P"'blem 01
.-pboloii<:al ol>osioa 'ion "". pilvloo!<...u.: 10' il the
.. k<tion 01 eh> .. ", ... ,bo, .... Of ..... 11.1 and
no< ."..,....."",:.
A."AlYSIS OF SOME I' H\ ' u:x.:ENI;TIC TERMS
'"
como within homopiuy, .. her-r undtr-
oto::Jd. In his latH book (19J3: 124. UI- I40)
Dacq..t-..toretum to tM nrl"r distinction.
.1 made by Scott; if 1M ;n; tial rpeciH or g=era
are related, IMre i . "more or ""'" p.arallel.
thoua:h oIowly a>nwrgi!lj[ tranofonnati"" (horne-
i f they are not. t.hHe ia "genuiM
01' .. ttic repotition
10'011:" "i')..... (Tronolatlolld""" wGonna.n..)
EYe'll thou"" ;n lhill pa at .. in nwly otbon,
Dacquf dot.hill '-I 'Oconto_ the urmi, ..... ".
panlltliom and con",,,,,",,,,, may Mill bo: reo-
ornizood ,.. the wo 01 thil antitheoi.. A
few P"iulattT (ibid . US). he '"'1",,10 " Pa,.,.lltl_
formt nbildung" to Eimtr. ortoo.
homeogo:neoi. and diltinguiorn.. it from
COfI'o"tT(tnce by the following ;nternal criterion:
i. oo.io:I to be baled "" "primarily the
oune buic orpniution. producing Me 'nly
the f(nUtion 01 orran .. t. ,. for the fal-
flJl"",nt 01 the """" "'"'Iuitoemtntlol lift,
conH.",...,. doa not ariM from. typon-
hafte Sindunr' (_ inmnal
tion), but oo:u ... ,..lwn like condil ...... tr.mofonn
btinp 01 ,..,tirely J,..., ora;aniulion."
AnaJog""", tr.moformalionl 01 . ... tk
form . which may be ieochl'OtloUI or Mter()(hrt>Q.
001 ...., claimed to be ...... yo con""rgence.
from Ike German.) Ot>.cure aI
theoo conu" .... q are. they Item alill to reinforce
the ......",ption t:h.o.t Dacqut in 19J5 ba...J the
diatinction primarily 0<1 relationohip. Howe_.
ke Mill admiq, .. he did in 1911, <:On'"'KU""
bel. """ related fonno (ibid., IJ2. US); on the
othtr hand. he lea .... the q\Jt8lion undecided
.. hetller to ttp!:ai" lhe ""nted """"I<>pment 01
th41 t)'K 01 co...., d .. an;ma!., be tlwy
vnphibia"* or TtJamrrW by con""ri"''':e or by
paralleli .... (im4., 135).
..... indico.red. abo"", Willey (1911), probabl y
the 6nt author to write an tnti..., book on con.
'tl iCJ>Oe. ci""" thill conc<'pt a far broader
than usu.al. Hio definition i, ... followl (ilM..
52) :
The t ... m i. appliod 10 ..... mbl .noet
.mont.t .. 1ma" whick an: Aot due to direct ... Ia
tionMip or tuetic affinity; in o.h .... wO<d .. ... hick
.... 1>01 d"';v! by in htri\aJloe ftom co ........ an_
-.. but .. ....wt I ...... iodtptlldent / .....
tioaaJ odoputio<t 10 .. ,..; .... ..,d ..
n.it definition, .. it Oland., ill 01 no .... tance
for the diatinctioo ullder COOloidaation. Tbt
... mupPIiH 10 Dendy'. definition (1m: XXll):
Tho """.i'; rion. by dill."'nt orpnlo", .. 0/ Ii"';J ..
lorm Of "ruct.", tho ",".h 01 ... milaT mod. of
Iii anrl n<" ....... ",".11 01 inktri.anoe ' tom rom_
moo 'ntHw..-
Annandale'. (1924 : 2) definition 01 con""'"l"ttt
evoI"tion." although rathor wordy and alto-
",the.- unl3tiofactory for .... r P"I'JIOII!.;. 01. in ...... _
to u. t..ca .... itadrtlill the p...;bililyol con.
WIltflCe be'WUll related ora;anlsrno..
Eva! ....:h a "",ndan! work &. Di.rne<'1
'"' Bi6.trolirn>pllu. (l9n: 3-9) deals
.... ith con"""""n"" only (under the name "Kon_
""""""""nl odop"'!ion).
without even ,""nlionin& p.a .... lleliw; thus it
cannot , ivt UI any due to the ""ution "r the
.. _ut problem.
Schindotwolf. in an "'te. tll"" paper 01 lWO.
R2rq out (p. 223) to giYe a prclirninary. "UD.
mtGI" definition 01. <:On', Ci.enoe ..
in ..... pt wt.ich haw.n.:n on different organiu_
lional buN and, therefort:, are not butd on
imlnfdiate relation.hip." It will bt clearly_"
that thit definition CO'<tB the whole ""Ope of
hornopluy. u defined o.I>ove (p. J 2$). In di .
tinguiohing conv<:rgence from pa .... l!c:li .... on the
0fIe hand and from repotition 01 aI_
......)'1 tke ...". evolutionary cou ....... (. itera_
tion lleurlt!n) on the othe . he """"ted. to de_
clart : "A !nil: ""'." .. &ent evolution can"", "".
"'. within one and tho: ...... p/lyr"m [StammJ.
-.i!hin the ." .. n" f"nd_w orv:aniution" .nd
'"
OTTO HAAS AND GEORGE GAYLORD
to suW"! that the t<:nn "onve<gen<:e I .. ",.
.. rvro "to t he Co:J.",'S 01 mutual app,<>a<:h in
.hape betw"".. membe ... of di/ft'Tenl phyla
ISt1I. mmej in ,,hich ,he .imilariti .. alleet merely
analogon. of very different origin and
. truet ore." (Tr anslation. lTOm the c",rman. )
Her<:, then. &on'. difl"",nt;ation , [>arnlkl;"rn
be tween related 10",,0. converge .. .,., bet ween un-
relaled ones, i. _n at itt [lCak.
Simp80n (1945: Y) also though mlleh
less tleeKJedlr. to tha I differentiation. il./ter
di,tinguishing homology from "OII"'-"'"\'nce by
a ttri buting t he reoomLlanc..'S t<> inherited like
character. in the former phen<>r""non, bUI to
inrlel"'nrlent.ly ac<juire<! lih d,aracters in the
latler. he contino,," .tMing that
,hinJ _t 0/ proee .. ... producn llimilari,i ... :
p;>ralleli.m. T he term i. dellCrip'ive ""I,.,
."plan.to')' a nd <de ... to th .. di,tinct
of COnln'lQn vriVn f ... qu.ntly .volv. io much
Ih ... dir"".ioo ah.r .h. dillC()fl.;nuity b.' weon
t b.m has ari .. n, 110 .hat at I.'. r 0\.>K" .h. porI>
",ay h. ," in comm.". that w.", not
i,;t,le in til< oonlmoo ancettry but that '.nd, neve<-
t h.I ... , to b. mon or I ... in p_rion to .h.
n ... r n ... 01 anc .. 'ry. This p!'OJ>Ol'tional ... d_
.noy di,ring"ish .. ""ralleli.m rrom ronv ..... n<e,
On the other hand, Simpoon that "{he
maximum pooo.ible 01 conwl'g<'nu tend
to be in prop<>rtion {o ncameso 01 affinity" and
al"" does ju",ice {o {he "!r"Ornctric" concept 01
""ralJ.lisln and COIlvergcnce, "" he", . dv0C3.too,
with d", fol lowing di. tinction :
II the .arly m.mb. D 01 two trO"P' a ... more aM=<
than the I . .. " t hei' la , di";milariti ore dive .... n.
If .11< urry m.mb. .. . re I ... . ,h';r la, .. ';mi
lariti<1t .... ronv ..... nt. If the urly memb.rs ar.
mu<h .ri ao,j the lat .. m.mb.ro h.ve rurth ..
.. H I , the ...... ""ron. r.
Moret , in It;" 01 paleont<>l<>gy (1940 :
27) admits con",,.gcnce ,,-ithin the
same group."
Finally, an indication of what h.u been c3lk.<.J
above th .. con.cept 01
i. found in Krocber (1943, 108),
In ohort, it i. bau .. the h';, ""ttern. diller th.,
the "milarity i. oooveri<nt. Th. hi , lOti .. or. un_
lik . the .. condary .... ull . . ... lik . .. tho"lrh til<
lib .... ulto n.-l DOt b. ,u,,"rfici. l or .';vial- in fact
may b. accompanied by "",,,allive modifi",tioo, .,{
the Qrlrani , n,."
k " "", [de ""' .... . . <01 .. poooui .....
_ _ I'i".n;..r .. I"0Il""."
In a p<!i ....... 01 K"",bor' , (;oM.), 100...,.....,..
.... foc.>u, .nd ( ... in the u .... 01 ,h,.
paper, ... m to be '1iP,ly """'u .... .
From the p<eoeding ,u,,",y of opinions it would
appear that Scott', original di.tinction bet"..,.n
paralkli.m and hased on the dcgJ'/:
of relationohip ""tween lineages i,
"ill the dominant one. The following conoidera-
tion. make t hi, fact .... ell ,mdcrstandable:
On the one hand, paralleli.m occun, it i, true,
between more or I .... related group". The con
"'pI that ciearly "nder,ies the use 01 paralleliom
in the oen"" of Scott and of moot, but not all,
later authors i. that this p<ocess or phenomen<)f'
in>'OI"". charac",,,, that an"" a fwr .. paration
from a common ancestry hut t hat neve"heie ..
arc in some way due {o the of that
corum,," ity of inhentance. The
""lief in such a connection ha.. been .. d
on previous paf's (pp. 326-321) and one of
u. (Sim"""n, 1945: 9-10) has emphaoi<e<l that
{he ... are at ica", 'OiO known genetM: proce_
that would lead '0 the .......... It. without recau"""
to such or. metaphysical ide,.. as phylo_
genetM: "predetermination" or " latent'- char _
ac"'..... Th"", i. li ttle doubt of the n>ality of
th""" phenomena and the .. 11',,1 phen<>mena cor_
r .. pond with the """""ption of parallelism a. a
..".t of oomol<>gy, but a sort that dOl,. not read,
pht-notypic until after tho> oepM:ltion
of the two or ",ore concerned. Thu. there
is a real validity in the idea 01 paralldi$m in the
sen .. of Scott and of th""" who have foll"""ed hi.
definition, a lt houglt the basi, lor th;" validity
has un ally I><"n obscured. A r<:Otateme!lt in
;>ettptable modem ternlS ha5 nnt Iteen made
a lthough that 01 Sin'p""'n (194-5) may perhaps
be conoi<kred a. an approach tow .. ... 1 th;", The
",ain diff,culty h"" been reteT1tlon of tlte ra\r.cr
crude and hiS!orically out-dated criterion of
ncarness or distance of affinity a. thc distinction
bNWttl1 parallelism and , ,,",,,,Wnce, The in_
t<11ded d istinct ion would Ita".. been validated if
it had carlier been in genetic :rm .
Redefined in . uch term_, parnlMism woukl he
similarity in .... uctu"' due to a COIn","" genet;".
basi. (and"" far n...,mblinl( ho",ology) but 1>'"
reaching morphological ""pression unt il after 'he
.. ""ration of the \wo or more line.
(and in thi. d iffering from homoIORY). Thi$
""""'. really to he the concept toward which
earlier stu<.J;,n .. "..,re prog"""";ng, gropingly in
the ahs<:noo 01 clearer genetic kn"""lool?", fmm
two different d it'\lC{ions: wben they .poI<e of
parallelism in the .,,1"'0 of Scott a. similantr
arising in ne.uly T'\'l:tted Ii""" (i. e., with a con
oiderablc cornmon basi,), ru .. j when {her
'pOke 01 !a\"", or pol<:ntial homolo\ly and
ANALYSIS O. SOME PHYUJGENETIC TERMS
like (i. e . of a common genelic 00.;. for char-
acters not morphologically evidl.1l1 until a later
in phy""," ny)."
Tit;" phenomenon i. 8 opKial c_ of hom"..
pla.y and i. logically on a le",,1 with homology
bocauoe it oIkn an altemati"" explanation of
. imilarities. whklt homoplasy d""" not do. It
's. clearly. not the only altemative to homolOf(Y.
(Analogy in lhe senoe of Simpson. but not of
Haas, is another and there are .I ill mher possibi_
lit;"o.) Thi. "'"tatement makes it clear thaI the
phen()lllenon is not to be de fi ned enl irely ,n t"",,"
0( nea.....,.. or d;"tane<: of C<l mIDOtl ancestry.
although thi. factor io im[>Ol"tanl in il. Even
very d;"tant!y ",late<! animal . Ita"" oome genelic
fact ..... in wmmon. but are more numeroul
the '''''$ the phylogenetic position. 0( the
animal. <:<>n<:erned. T.......,fore the inl"".ily and
frequency of thi. phenomenon will tend to vary
with nearneos <Ji relationship.
So far. the posit ion seem. clear. I I i. possible
to Ralher together hroadly d i""...., ... om
a nd idtal <Ji the paot and to ",late them to a
defini te and modem "'"tate"",nt <Ji a phyl.,.
g ... etic ",ineiple. It remain however. ro de-
cide whether to define or n:<lefine parallel,,,,,, a .
the term applicable to t h;" principle. On
hi.toric;tl ground. this would be justified and
perhape preferable bocaU!le this gt:netic delin;t'on
is cl""". to lhe"""i" of Scott and of most later
authon than i. the strictly geon><tric definit ion.
N"""rthel .... we haw decided againot suc h a
proposal (very ",Iuctantly. "" eone<:ms onc of
uo) becau .. <Ji the inappropriateness of the term.
ParalielilU1l does haw a wrnacular. or at lea" a
non_phylogenetic !lenoe. ",hich ;" applicablo to
tlte phenomenon in question onl y in a deocrip.
tive way .... bereas the .--ne. <Ji genetic defini_
tion would be ro . upply a theory oropinion as 10
t he cau .. underlying the deocriptive situation.
Thi. genetic principle i. not the only one that
can produce a oequen"" pa",11e1.
Mor"" ........ the re."lt of Ih;" genetic facio<" is not
""" "<ily paralleli3m in a deocriptive .. noe.
It can. i""o-ed. be COIlve'll.nc" in this ..,noe
although eonwrgene<: has alm""t al ... a)"l been
"""side"'" ... _li,..!ly d;"l ;nCI from panllel-
isn,. One of the reason. fn .. this vie ..... aud pr0b-
ably tM one. hal t-n that eonvergl.1lce
may be fouud between groupo f3r distant from
each other in tit" zoological ')"Item. a. in the
g Datwin ...... Ioiw> """",;'-.' f.U. .oder
thio """"'Po. . I,""'.h;, .. 00\ """ ...,.;"".nth , ... coo.
_t ........ .,.. ODd of """ .. d;d "'" Inv""" of
the medlaniom of ....... i' y (if b<rvo. p. 3n).
cla,..;",,1 eAAmple of tbe DeV<lnian coral CakeoJa.
the Permian brachiopod RidJlI4j.nia and the
Cretaceous pelecypod all of which
OMume. under ';milar conditions of en";,..,.._
ment. the ""me homlike shape and closely ap.
",,,,, ;mate each othc. cven in somc detail. <Ji
intemal .Iruetu"" such caoes a","" conspicuous
and .trik;ng all to foe". on themoelve. the at-
tenlion of natu",list. Sludy;ng c""vergence.
It cannot be doubted. howcver. thai it may
al .. occur bet ... een ",Iated II"I"OOPO. even between
."""ies of clooely ",Iated genera 00" even of the
oame genu... AmonR the beat k"""' n exam pi""
.'" lhe various lineagel of the Upper Jura .. ic
group of A ... ,,,,,,,ikI ",uta/>UiJ Sow. (nowada, ..
",fe!"ted to the Rcn ... P",p/anuJikI Teisoe)"re by
Roman (1938: 266). but to the genu. &S"";"
Salfcld by A!"kell (1946 : 12. 15 which COIlWtge
to ouch a d"8"'" ... to become homeomorph 3.
d"""ribed by WUrtenberger (1880: 85-86. 106-
101). Hc", it i. only the Ottloge>' y of the form.
compared by that authOO" which. thanb I;<>
la ... <Ji recapitulation. betra)'ll their different
origi n." A oimilar case of different ammonite
speci"" assumi ng in malurity the ""me characte ...
"" "" to inl o O"e voIyphyletic opecie. is
that of lhe Lower Cretaceous C.w.:wu duvaU
a . deotribed by Nol3n (1894). Brink_
mann (1929) 8100 report:f CaIOe" <Ji ","all""t con-
""'1t"nceo among his cROmoe<:ratid. (el. Dacquf.
1935: 1I8). Another cue of intragencric ron_
wrgence i. reported. under t he deaignaUon h""",_
omorphy. by Buckman ( 1\101: 238-239) to occur
bet",...,n " two nearl y related foooil l 1<Ji the
I nf....-ior Oolite). namely: T .... brat"'" pliwIa and
T. flmlwilJ" both of .... hich cle'"eiop oi milar multi
plication of ri bo. On a o<>mewhat higher tax0-
nomic level 'of several difTe",nt
R"netic .. ties into the genu. llaculiU. wa. as-
Mlmoo by Hyatt (189(: 519) . just as W. D.
Smith (1905: 653) found "tltat a number 01
'morphological 10 uoe Hyatt'o
.. To th< tim,. f.,.."" ... mod abo ... Dac.!.of (1921: 252.
6o:. 91).dded. bo ..... id <>h;l.oib"" ....,.y ,,;m;!ar ,hape of
;" 011.11.
.. H .... _ ood i>o<a"", 00 .i"'; .. , to ... <1>
_ <hot ........ 1 of e>'<o ;!ed ;"to.
opecin, 000 .... nUgh' eve<> ,_l of. polypbyl<tio
<II <he ..,,;e.. $Om;"r .. tilt ","'" " borlo 01 .. ch ..... .
10/;, /"""" may _. to -=II o,h.,.. the;nner voIu,i .. .
ano the '""'" ' 00 di_ boo, ..., dearly th<
0/ otic;. 0/ Uod;riI .. h wltO:h ho"" ........ t.... n
lumpe<l;"to 00< '''''';'' : th. , mel> poIypbyletic .".....
, ...... OIl. to "" """,Iy . ....... ' ... 1 oornpout>d of diH .... n'
'hiop to _'lainy by .... _ booed on
th<......., th<lonna: (T ... DOb.tioo from <he
G.nn..o.: W.r ... ' Ull: l06-t01.)
OTIO ANI) GEORGE GA\'LOKD SIMPSON
u .... ..: .... , b.m grouPf'd tojIetheruspoc;a
01 one Il"nul: that th" 15.:ap.au,1 il
poIyPllyletic, and not monophyletic, a. it hal
been treated hcrotof...... . .. " H .. OJ>inion wao
mared by (1907: 117) and Nowak
( 1912): the laner _plit the old ienul 5.:apMln up
into th...., gmera: Il.u:op.ilu, Ac,,,ot4D=.
p/tUu, and dfrivinll each of thc.e
"""'oer"&ing .. n" from anot her ancostralll'l""u"
viz. HoIuJkp/ID' ..... and
Thai all the al>oYe .... mpIH an: taken from
the may nOt be purrly incidental
but "";ng to the f""" that kardly any other group
of lendo it.oetf.o -.ull to Itudi<l8 of
evolution in general." Scott (1896: 57) nli..oo
among the "", lebo-awl c.un "w""", all tt.e.e
IIl'UCtu",. Ii. e. """th. okult or reel) ha"" btrome
alikt through the operation of ei l boer parallel or
development." though adding: "A
moni the tboe cue is different ."
although be k_ well, and .. .,r .... :d
w the.,..... of (OD'egcnoe llOMiicd by WIII"teI>
bell'" and Hy.o.tt, Scott (1891 : 366-361; 1896:
57- 53) failed to da ... the oew;o..l conclusion.
that aloo cloooely ",lated li ... may
II admitted by Oacqu.\ and Morel (_ abm..,
Quotalion.).
Since, therefore. a hii;h de,..ee <>f taxonomic
diltance bet ... ...,n the initial "",mben of 1M
conwrgiJlg linu(tfS il. in our opinoo. e5-
.. nlial for con"","","",,". we <IecidM to
.. pport otrir:t.ly i ..... trio; definition .. ,-lz. of
poNJlklisrlt u homopIatcic modifM::ootions in 1_
01 more 1"- producinr simil&ritios .. hich do
not lncrHoo with evolution. and 01 ,_!...u
u ... elI modification. prod",,;n, similaritieo
... hich do inc:reaoo ... ith e"Qlution.
A. in the ca."" of homOf' lalY, common u""",
appl ... th..., term. to 1M of t ....... evoIlI"
tion:uy ,,"OOCl' iO u "'<l11 al to the plOtt ....
t be",oel__ II"",.."...".. the fortt>er" Gemoan
term "Kon,,.,-gen"" oo:heinunJtft''' (_ pberoom-
ena of convUiC"a: 01. """" prtciIcly, phenom
ena due w look that lI i.tinction
fully into a<:wunt.
It nl llit be arlmitled that the ooI"too <:t>o-en
above lea"", IInoettled Ih. problem of lenni n
oIoR:y for the more tloo:or"ical Ind "",,,tical
principle to ",hid> "'.., pn>poIC denying the 3.p-
pellalion of pU2.llclisnt. In.pte of ill .,.,......
... ha, metaphysic . l undotrwoot. the U... 00
"LJtcnl homology-' might DOt be inappoopoiu".
bul th" is tIu: more Mrictly n ... j>I>oloJi.
(:JIl-phylotil .... tic """"'" 01 ou .. dioao..,;,onl and we
Ire not P"'1"In:d t<o make a definite ",op<..:I at
thil time.
From tboe al:oove ",eomttric" conOi'plion <>f
and it follOWli that only
l be latter but not the former may lead to home-
omorphy. to be m.ou""",, under lboe """t headini.
SOIut KXAIIPLIOI or PAot.U.UIU SM
AND
So far only nampia 01 """"" h.a." bttn
Ii""". in m_ 01 which (CGk ..... Ridl/l#jni4-
JJippwri/u; group <>f A",_il.. ",uI4l>ili.s;
C"",ora' d.....u; .&<oJik.) IUCh a high d"i'""" of
io rtacilt:d Ihat lhey might .. ""
exnmploo of hO/ttOOlllorphy II .. .ll.
An ."""llent illllStration of po.ralleliom- be.
IW<!en a l><:ne and a both of the
Middle Eocme-ia Ii"",, in Oobom. (19086)
lieu'" J. ohooIu., how ...... cuopuleo riot inde-
pcndmtly in both. This ia. in ou .. opinion. an
example of but not of con'ell'"'''''.
boca .... the appear;u><:<: <>f theoo n,.... culpulcs in
both liooo does not bring ,bern mo.-pholoska.lly
clooer logel ...... than they wert boerore .
Another 8000 u.amplc of a!on
taken from the evoIutiQl'l of the mammo.l .. is
Ii""" by Senti (1891: 364). lie citeo the in
dependent toequisition of "t"" prismatic cement
coycred moI;u- in many lonna: t. 1 ... ....,.-,.1 of the
ruminanu. certain pip. lhe """ of the
rhlnocoe' (111 the elephants,
many roo]o,nto e tc.'- Scotl add.: "1". , JD.
blanoc of the molar 01 Elou_ ..... ,.'" to thaI <>f
the hone .. Itrilring. but ill fundamen tal
plan ;. rbinocfl"Otio and not equi .... " Th"
.. indicale. thai. deopite tM appearana:
of the _ rnodific.atiQl'l in both. tM ORe is , till
a hone tooth, and the other . till a rhinoce",s
tooth. Thclf fundamental plane ha,.., nol a p-
proached each other .... hich i, I criterion
lor th .. modification bcinl but not
-"" ...........
ANALYSIS OF SOME PHYLOGENETIC TERMS
'"
Among inYffttbrateA, Buckman', (1906),
plate XLI, although "",ani to illustrate "Brachi
opOd Hom"""morphy," .h""" a beautiful c.ut
of parallelism in dir>hyoid terebrat ulida, bxh
ioochronous (all far"" the three Portlandian-
Neoromian """"- in relation to each other are
roncertlM) and hetenxhl'ODOU' (there heing, in
addition, a I i.we .. riel), The r>a",Uellineageo
undergo the """'" changea, but their "",!>"Ctive
memben do not thereby hec<>me similar
then their pced""""""n, although it lias 1;(1 be
admitted that even at the earliest alageo there
alrndy a ""ry high degree of similarity.
On the ",hok this elm,t "f Ruckman', illn.trate.,
in OUT opinion, parallelism, n"t converg<:ncc or
homc<KI1o'1'hy,
A oomewhat different r>icture i. f""nd in
&hindewolf', (1936: 41) figure 15, iUu. tr.o.ting
the indepM<U:nt ",,<>Jution of triangular _bell
ehape in three li"""ges "f Upper Devonian
CIymmjiu. Sdlindewolf calis t hese th""", serie>
r>araUel , ",hich they are indeed in tbe lower half
of the diagram, after liaving di\'t't"ged al ito hue.
In i to ufJPet half, howeV<1", the lin .. II and III
(line I doeo not reach up lhat hiJ(h) become con
"Vt1"i<'nl, thill C"OfI""rgetI"" \tadin, to home
omorphy in the end ""'mbert .... hieh can, ac
rording to the author (ibid., (3) be distinguished
men:ly hy their suture lines. Thus this ""oe
yields a good illu,\ralion for the statement mad<
at the very beginning oJ th. preceding seclion
(p. 332) that the <arne two lineages may be di
' .. ,,,eut, parallel, and con""""nt in """"''(:utive
time period ...
I{OMJ;OMORPI{Y"
OIlGIN, PltEVIOUS PEFINITIONS
Buckman, after havini uoed the adjectival
form "h()tn<)<;OmOTr>hous" in an earlier paper
(1895: 456), defined in 1901 (pp. 231- 131) hi.
tenn "hornoe<>morphy" no "the ph<1>ornenon of
OPOCles nearly alike 10 far ....... perficial appear
ance ;., concerned, hut unlike when particular
ItructurnJ detail. a..., closely oxarnined:' and
it from "the phenomenon known as
of development."
In the decad"" to follow, the Ierm Wnl uted in
this meaning by Buckman himoelf as well a. by
other r>a1eontoIO\li.ta (oee CI"ud, 1941: 900). "
Cloud, in a paper on "Homwmorphy and a
remarkable inu,vation" (1941: 899), redefines
homeomorphy "as the phenomenon of
superficial ..,..,mblanc<: but cli"';milarity in
""rticular structu""' " thn. making Buckman'.
original definition, "ithout any ..",tial chang<:,
mOO<: concise.
Schiuclewolf (1940: 230) advocates a "neutral"
U!lt: of the tenn hom<:<>morphy to dMignate
"similariti"" in shape, ..... ulting from parallel Or
diverg<:nt COUTS<:S of evolution." (Tran,lation
from the Gennan.) In the ""me year Moret
(1940 : 27), after mt1Itionini: the l>OOAibil ity of
converg<:""" within ono and the ... me g"""'p,
gives the following <><ampl",,: " ..... inoi
Ammonites, d'oril(ine Ilf,1.tement differente, pen
""nt avoir nne COQuille lOUt 1 fait oemhlahie"
and CQIldudes: "on dit alors que"" IOnt de.
""pkes homb>morph .... "

or the various opinions jua' O<>Ud's
and Moret'. come cloecst to our own. To UI
h""""""orphy is the "",ult 01 a COD""rg<once
which has i"""'" far as to prod...:.: a similarit y
aife<;ting the wbole outer appearnnc<: "f the hon,.,.
omO<pho to "",h a degree that the one may he
mistaken for the other unl"", .,.".tain internal or
other cliaracte", requiring clo<oe examination are
otndied. Thu. conct"iwrl, homeornorphy "'pre'
"""to the decer>tive cliarnet ..... of all homoplastic
modification ... a. discu.-i above (p. 43), at ito
peak, """,,ittinl! Buckman (1925: 9) to speak 01
"hom"""morphic decer>tion." and Cloud (1941:
9(1) of "the !litfall. of homeom<>tpby. " Char
act.", whooe examination makes the distinction
of homeomorph. poMihie nfter all may be found,
e. g., in the earliest (innermost) volution.
(\VOrtenberger, 1880: 11)6-107) or in the .uture
Hnes of "the .... ;.. deceptively oimilar ammonite.
1921: 250; Haaa. 1942a: 6ol5).
The couo;.,ption of h"""""""'phy hen: pro-
cau ..... the boundary hetv.'eCn it and COn.
""r"me,", to be vague ince the forme. r>henom.
eru:>n is conoidered merely a . pecial ca .. of the
latter. Morro""r, the above of "inter.
changeability" "'qui .... ela.tic inler"",tation,
.. Sri" <'02\1) within .It<: ""Ioontoloticol
tunu .. lati"" to part.o 01 .... n ...... ooly, homoIor<ou od
.... ...-"" ... ..-- \he lau ... bdn, the """,Iy ...,...
--
OlTO HA .... S .... ,,[) GAYLORD SIMPSON
acoon:Iing to the ta.l<OOomic diltanoo of 1M ron-
v-oo,.; th".. in lbe lamoo. eumple
CaJcu/IJ. RidIiMf ... il might be
deemed oufficienl Ihal .. pcll'CYpod or a hrachi_
opod nlay be mi . la ken for a coral. in tbe
cue of intragencric .... in Wurten
beli"'. or in Nolan'. c,iDa.
quoled abo, ... (pp. 331. 3.13). only ouch lonna
at an: ino:!;'tinJuW!abIt excepl lor
oortain internal. not modily ., chanel.,..
may jualy "" callcd hon.oomooph&. ThtlS t he
Ihlft exam"""" of cilM 0.100, ... may
te<Ve j ua ... "..,11 ao6A/tlple. lor
bul of COOl .... th;" doc.-a nol 10 a ll cucti 01
IO""'1l:"nee. For ucell,ml ,)l ullrations 01 bo-
"'H1lO,phy ",Ie,nu il made to
Dacqu!'1 book 0/ 1112 1.
"The redefinition here attempled ;. bound 10
il>COltlple'" ao Ioog RI the
elcm(1t t is DOt tak(1t ;010 accounl. i-"., .M
q ucaion ... 1>0 may be Cipt."C\ed to miMakc or DOt
to m;""'ke the one thing for the OIMr. II..., ....
Wt in t h;" iCs;.,d the layman.
who can ra",ly. il c\w. di .. I brachiopod
from a pelecypod, should be excluded as well all
lhe apeci3.lisl . ";nee. c. I . a ,,ho
h,.. dOll work on rudistid. ,,ill
ha,dlr be unable 10 0.1 fiB!
"'lIh. , 3 homeomorplo.oul coral or brachiopOd ao
_thina: diff"","I. Ral loer ohoukl the im
prtllion and jlKlgrncnt of the trained natural .. t
be con.i<Jen>d dote .. "".
From thil angle ctain in which the
61"" I- ioo too
to mind ought 10 be u.cllKkd. at ", ... 1 from 1M
muning of thi. 1C1m. ]f . e. II-. an
uncoi le<.! . Ihn<lgh C .... taceou. amfJK)flile
from Angola (A .... o<"'" J<I...,' k .... U ... (pic""1,
var. ''';''0''' Haas. l!Ulb: XLVI. fill 2"J
Itri kinl:ly .--."blco ..,,,,,, ""terpillarl. thi ...,.
.... il1 nol fool trainld na.urali.t
"'ho k" ...... .....,lIlhat il ;'lhe.ne11 of IMammOll.
ill which btw""," and I""t catrPilIaB
l""k an oo\C1" lnlrJUn ..... 1 capable: of bring 1_
oilizoed in the way. M anmher """" io
point, thn<lCh perhapo a boodtr line caso:.
be cill:d the oimilarity of a coI<>n y of F_.u.. a
Palfowic tahula tt coni. and a hoocy<omb
which nnt only again and aa:ain miolead. Ihe
lay"",n but even account. for thl gl nric name
- honeycOmb). Here both otruau","
a .... buill by in""rtcbralH '""longing to grOUpo
lar diot.anl lrom 0\""" 10 I>ou .. the individu
al. of the colon}' in one cue and ic. oRaprin, in
the W.,..., it not 100'" t he lact lhal the
ie .. bjecl to foooili u tioo in tl\(: lormer
ca!ie bul not in 1M latlt<. the term 1I0moolllOO'"-
ph)' be a ppti.ca.b'" to Ihi, <umple CYen
un<kr our preceding ,""tri<.tion .
Thu . we arri "" a l lbe of hom ....
morpll)' a. a similarity 011"'0 00'" nlOU orpnioml
or IIrt1(:IU""''' alleelinll" tbeir ... 1toIe ooteo" ap-
platanee and ,,,,,hin, ouch a '''''' ''' lhal. unl_
CCtt;o.in inl nal or ob.:u..., eha""'t"" ;on, ex
amintd. lilt one may "" r ... the 0"",,"
... "CO> b}' a trained na".rallot. The similar
ors.caniem. or ItructUres art' <11.11"" horncomorpb.
in Il'btion to e..-h o.hc.-.
IIO"EO"ORYHY n. ",,,,en
Halh Iluckman (1901: 232) and Cloud (1941 :
899) lmpha.oi.., ,lie connrclion bet ... ... " l hese
t .... o pb.nomena. The l<>nner ronoiderlo mimicry
lie "one' f'hao<' of hornotomorphy", t he latter
u'iH "the "'idcr """II" of !be oecood t"'"" in
ptc/crrllO! to .be commonly m"","", word
. .. , ..
m,mocr)'.
It ie true that <:ertain cu<s of mimicry in"",,,,,
honw-oInorphy, 10< l he (predominantly
female) buttcrflie.. quite rently d;..:uoocd by
Goldechnl,dl (1945). which topy otloer ope<:ieo,
distaltclul or othe"T",i.., .... [lul. iv. to predat ......
(>T inocclO .. hieh imi tate in their ap_
peo.rance onn belonginil: oon>etinlCll ' o
uxonomicolly dislant I[1"OUI'" (..., Dendy.
19lJ : Jig. 111). """""''t"r. mimicry in_
clU<Je. onanr c.a ... ,,hich can by no_attI
be conlidcn:<l homtomorphic ao the is hen:
undtntood. T IM: gr:J.I8hopptr or butlt<8y ropr'
ing a "'a( . he Aicb" imitating
flit",. or. Ic.ul 01 all . lhe mimicking do-ad
Ie;o. ...... u vividly dcecribed hy Willey (19t1,
64- (05). certain!y do not come under our " bow
d(:finilioo of homeomorphy. O .... ,e .. parate<l
I,oon il' envirOllmenl . l he mimic could hardlr
r_ be m;'taken lor i", TDD<kI. nor lhe latter lor
the lormer. Furthcrmon:. nl;micry. aa de6ned
in IllOO.Itm biology. requira Ihat l he """mblan",
be advantageou. to tho- mimic. ,, ..... ou home-
no n __ ... _.. P'" """"pIIto . bo, __ 1
""'_01..-.:6 ...... ''''' _100M oad IM"';to. Piau-
........ on .... ""-. ,..-001 .... \)epffl!< (1'101, lll).
",,,h' _ .. . ...,.:1 ... m .... 01 ...... 1 .. ". pIoy ""'-
oinllo ",,1<""'""'.
Sl""" W;JI<,' , """ ........... Inc"' ..... ' '''''''J -u.:'
...... ,1\0 II<R <>1i00i """-.o: <>!'r. h ..
."_1 t, l-6) ,100, n;., \too oI:w\ou, ..... II .. 01 ....
.. "''''''\1''' _ ......... <On.--""" 1.- ... ",,,,".
ay ., I, " '" mo. .. "' .... ;, I..".. hoi >'oIIy .... kI.1oo be
.... lId ..... , h" _ ..... _. f.W ... _ tv. I'll- 51-
.. ) ... ...... """ H<z ....... ,.-
ANALYSIS OF SOME PHYLOGENETIC TERMS
01,,,,, ploy, as a pu..,1y morphoIoeic oorocq>t, doeo
not imply any such clement, nor any distinction
bet_to mimic and .....del, putting, as it doe.,
both (or all) homeomorph. on an level.
Thu. n,imicey i. nOl, AI hal -..etimes betro
.... mI (Buckman, 1901: 231), a "pll_" or a
lpeCial case of homcornorphy. Can, on
hand, Iauer pcm.po be in.dud<d in
mimicryl Certainly not , oince mimic a nd
nlO(\el canno. hu. liw loge."'" in li ..... ama. and
en";ronmenl, a <fquiremen. which will rome
tNo: lor only a V1.'I)' orn.a.Ill"'OIIOI"lion of an.....,.
01 hoo ll<OlUorp!>y , H .. de<! and oIJ.crvabic;n the
IO.kJ of p.ok-ontology. Wt Ihul alTi"" at the
cond uoion lhal the CWn con('Crue oV<!l"lap.
Duckman (IDe. clI.) utle ... Ihe opinion t hat
" Mimier)' may be ollggef, ed in rtgard to I>omoe-
omorphoo. spK ..... , .. giving am""", iIN of the
genera Dtl_1Uri4 IlAd GM .. ail an "".
ample. bul he add. immediately : "It ..-oukJ be.
pnh:l ... 10 mimicry in Iqard
10 ......... E' ..... whme in the olrm>eiy ...,.
Itricted Mki 01 aline' hon>eomorpho.. " 'hieh
liYKI not b ... also in the
...... aru alKl cn\lifOrlmt'tl t , should be oorne
realOll lor .Muming mimicey, t he..., will indet<J
be no " .. y of proving i t and. I .... 80. 01 cor
rectly disl,ilnll;ng tIM:: """ or mi mic and modl:l
bct,,--. lhe homeomorph.. Therelore. pale-
onloloa will hardly ever be ",,,,",",,fully 10
Ute the lern> mimicry.
' ''''01'(\' ''' : .... Ol<I>lICM I V "1I1t.Al TIl ... S
AI n ..... t.ioned a ........ (p. 329) . ..""., a .. than
(e. I .. FIawn', 1/)41 : 11, 49) uOC It... term ";';"
morplo y" inOlead 01 h __ pIoy: howeWI',
lor l he "'...,... previously .u.ted, il ohoold t...
...,jwd as a synonym 01 hOlMOmorphy aa
at of hornoplaor
T1Ie pl>onel icallr similar tcrnll "homomor.
phy (homomorphic):' "homon_ph,
morpho," and "homomorphou." haw """" d ....
ab<>ve (po 329).
By ...,..y of append", 10 Ih;, heading ,,'" may
brio,8y d.",. two terms t ha i h., ,," IK>I }"'I hem
w ll with in !he ... t.1OI fludy; 1/ .. ",,,,,,,- was
ori,inally 1""'" -d hy Bal...,., (1894: 85) ,.. a
oubllilutc for lhe "",Iier lerm """'tamorpby,"
to indicate "that """"' thing It.u been in
Ihe liko:n .... of oomething Bridges and
Dnbzhanoky (I9J.J: 515) defined " heralitaey
homnIIo.i." ... the "radical alteration of the
morpbolocr of an organ (owing 10 a mutalion.
or) ... di ... ppeara""" of Ihe organ and the
dtwlopment 01 another orga n in il3 place."
/INr_pIuI ... , is considered a Iynonym 01
hen ...... by GokIo<: hmi' h (1904 : .126) and Villoe
(19Uh). whereas Boytkn {I<)U : U T) malu>o the
1"'1ewin8 dittinctinn: " Uolhoeoeil ... 10 the
ouboti ... !.ion 01 itomol )'ptS".. IIt1iaI
I.". other, ... hom;u hetc""."" phooi:o ...... ""
d .. "'11lace"'<"n1 01 ..... '*Pn by any other
,,!tieh nd not he homol ypic." Dot I llat AI i l
may. both .......u are primarily l erml 01 """,Iics,
but Ih. underlying may giw rUe to
pI.ylqjometk changeo, aa d"" .. /.
p. llS. foot no,," 48).
HETEROCH RONOlIS AND ISOCHRONOUS
MOOII' ICATIQ .... S
Maoro ab<>ve (p. 326) .... i, her homoplasy
nor ilS I{lecial cues panlltlilm a nd con ....... _
amce" as IlUCh imply anythint" iU 10 (On",,,,...,...
of lhe similar cltanK"O in lhe two or more
li ... agu con"""""' . and thi. hold. I,""", for home_
omorphy iU .. ..,11, ... hieh is to u, t ..... ",,,,,II
or a ver y high del(l"t"O" of TheJ.:-
fore, the lime demen! has uORally b<!o.n "" 1 out
01 accounl in the _lion . I n the
Ihrte following howev.. .. , il -.meo
importanrt'. n.e lenn. "he",mch"",.
ons" and "'iIochronowo,"' I by Mnjoi ....
o.icI (189.1: S) and adoptM by lIuckm"" ( 1901 :
231-23.1) 10 dislinguish """"""''''''' po<aneou.
lIomotO,,"""" from more".. ... contcmponone-
onlonct. a"" then:f""', b<>s, introdUCftl at Ih"
point of OUr in....,.ligalion. They ""'y "'Iually
be applil to all kind. 01 phylOJ(!netic modifica
lion, " udied und.". Ibt Ia. t th"" lIcadings.
lSOClt RONOliS COSVERGENCE (OR HOME-
AND POl.yMtVLETfC ORIGIN
iU.,...rty ... Ln 1891 (p. JII! ) Scott .. rotc the
loo1iowing IJ'lr.lllraph ,
, .. ;t th ..,. ....... 'PO'" oj tIoo '",,"'nl
... ""y t!o.e _ <h:Inel ... ;ndepotldutly
oj _h 0"'" ( ... ,.11<:1;"",), .. if ,.,. -'""' 01 w;<I<ly
dir .... 1 ......... ""y ,tOdllOlty ... "me a "''''1tIOn
.. U" ... ly "1"0IIII .. 144 ) .......... _
. Ppo. .... 'ly ;n.pirN by hi, .. od.o'er ' 0 oeeu,. -...
*poco" I ... hio in <l<finod .. ultt ....... bilduOCOn .. _ to
_lrie, _' ''' .. 10 . .... oImilo,I,;". It
will be pojel<od oc, boloo. W( ........... , ......... r ... .
""'"!"' ' ..... 6", ....... y be ...... ood ... til;" _. _y 1M:
_ I '" _ ' ''. ot .. __ .. '"
--
,.,
0Tr0 HAAS AND GEORGE GAYLORD
like" ... (con'-"n" ). ,hen it i. plain that ..,ch
IUU. i, ... __ mbl'g. of I.,..",. of poIy_
pbyl<tic oriJio. Tbot _h paralleliun 0/ devoiop-
"",nt ck... oa:ur, "'-. ,h.lI ..., i. the <>ell' oecti"", .o.d
,b_.", fo< bdie-';"/I "' .. """' ..... ... _
;. .... oo'*"'. phttKI ......... as i. jOlo_ally __
to be l rom .. -lIkh it follow ... !['ft'
,hat "'.)' ",..eric 8 ..... PO.,.. not ap...,;o,," 0/
<datio<>ohip bu .... 15 ... , . _mblarn 01 oImilar
'or"",.
In writing tenten"" . be may have had
in mind \VlIrttn!x'R"'" and Hyatt'. c ..... of
in ammonitClieadinl( to
tic 0< ewn tpt<:" 138(1:
1(6). If ... e _ back to Wiir1aIbeip" , (aOoO
of tbe"";gin of the .4 ..... "",:,,,, "'Mlobilli rmup,
n:pcatedly di.cuooed .bow, may weD raioe
It.. question ... hat happm. in ouch a cate if
lin ... gu of dill''''''"1 origin can no lon8<'1 be
diounguiohed in thf:ir offoprinlir. Wonl'tltx.rger
himoelf (ibid .. 106-]07) wultill in a pooitioo to
tbem by eKaRtinini tbe in""", YOIo-
tiont. Howevoer. if tnon .. a 'rn>d to puob the
ancntral """ ... rant- and farther back ill the
ont<lgeny, it may ....,11 be upocted thai SU<.h ,.
du., inction miQ:ht no Iongn be fuoible at a_
wlut later .... oIutionary . tage.
Tal[t.)Tlomkall y. the '1l1Muon it .... ther
usily .., .... ....-ed : u-.,..., ;. no nlOl"e po.sibil_
ity of opIit tinr I1P a ifOUP 1(1 bt CtlnI-
P' 101 not tt"Ul y relatrd lin""''" into ie. com-
pr>ncnt. R . Novak. _ able 10 tplit up the
"form ... " &4,ItiI .. ; _ aboY<! . p- .JJ3). thrn
"'" ha ...... .unply to ","",pt ouch ktOI'.- at ta>:-
uniu ouch at opeclet. g''''''''. or
tven high.,.. grouP'." according to thdr hieran:hir.
rank. Thi. ie. by the ..... y. thf, point wbe", II..
t.emI. "ciI"culul" or "geologic gen ..... " poop, :d
for oucb polypbylellc by Bathe<- and
R...&:manu ... z]U>:tively (_ Grlobau. 1913:
PIa', ( 1921) in .. rest;ng C(WIIidera.
I thi. tak .. can: 01 the t.a><onomic
porieo """II aa the hiaher
taxonomic I1nil' bein;. a />I'krl. menta! concepu
(Simpeon. 19010: 414). The much m"", intria:u.
in; problem m;ght arioe whether or not ioochron.
..... t1)n'>'fozg'''tt. ;n &<>inr beyond ;e. hi;h .. t
d< .. ,....,. homeomorphy. micht be able 10 prodUOl!,
by facilitatilEi hybridizal ion (aa far ;tlZ t!nlz'lly
prnducinr ooxanioma an: ""tun!
polyphyle tic co.p;o.bie 01 poupaption.
HO .. CWf ...... "'r .... in from d;"'ullinK thi' prnb!f.m
wi t hin the ICOpe of t l.. prcoooni paper.
TIME SICS"TUMES
Thio io probably the 01 all u.., term.
deal. " ;Ih in the ... _ut ,rudy. 1M Irmi
"Zeitcharaktrre" (_ time bad bot"
u...! by Bnzrl<n (1929: 5(l), Dacqu4i dtvoud ""
cntire IIOCtion 01 his book 01 1935 (pp. 226-251)
In "7",;t/ormetlbiklun;. Zeitbaull ile. Zeiui$:na.
t ur ..... ( .. format ion of time character aTehi,
ltIP"1ll Ityleo of lhe period tin, .q;nuu_l.
uoin; all th ..... oIlheoe term. tnd;""';mi"" .. ly.
He attributn the 01 "Zeid ormen"
10 ..... predilection, a l otrtau. period. ol ' M
earth'. binary. lor definite "arthittural style.."
ailical .""",ination 01
.. 'Ofdr-nd moody <;on thi.
sub;ecl ........... 1. that b;' "Zt;tIO<Tne1\bildl1ng."
in lbe wide ocope given il by him. i. merdy an,
other flIjA z"m ol the JorP:aI n" ily for
nodUlion ID tau platt th ..... a" ti"",,"" 0( the
not ........ k_... f",,' II .. , e.....-y geoiop;
period Nd living bein[!l of Ih;'
VH)' I"'riod, Thm: io OO'tainly no n-' for
"IlK;"1 term. IOflIpn:II ouch commonplactS.
Only in" very rewic.ted .... .., might
roncept be "",,",pted 10 de"gtla" the repeatedly
ob" ... ""d oimultaneou. 01 ...naill.
....... tin .... IlIpemcial, ill difftrent,
moRIy ... ' .... MI n:lated a........ n...e_,
of rou .... nolhinl bu, C&II'I ol isoch .....
.,.... parallellom or WIIuZa<nQi!, :ao 1M may
bt. although lhen: ifl no orriou. objecl ion to
livinl Ihil phenomcn<;on a name 01 own .
T h", Pu'lJ<>'<' might belt be oerved by u';n& the
last 01 Dacqu4i'. th...., na.".,.. time ';ptur ... al
dN:riptiv ... not ",pla""looy lerm. ",jectinr
lhe ""'0 Olllen which..ern 10 be IOU IJf'I'lCfItiuuo.
SclUrIewolf (1936 : 10, I I ) alao __ 10 fa....,..
a ratht1 ..-rio:ted 01 theee urm. ol
and to re;c.ct the latt<r'.
inf ..... n<ft. He a dlatinction he' .. ...,n
time IIrna1ureo O<o.:urring .... it hln Ii .... agco rather
d o.ely relatod witb each other or deocendina
from comonun _ten. and thoooo occurrinK in
ANALYSIS OF SOME PHYLOGENETIC TERMS
'"
groupo not cIoscly related wit h, 0< far re-
mote from, each other. The former &hinde-
wolf interp<ets a. due to oorne common inherit
ance, but thit con,""ption make. it doubtful if
the designation time signatures, as meant by
and"" heTe under1tood, w""ld 1Mn still
he Ti_ of tm, ""O)nd
lcind helie-.! by Schind-olf to he """"Iy
accidental and attributed to "pure converge"""
or hon>eomorphy." Si""" time signatureo are
de6ned above "" .,. ... of ioo<;hronou. par.dlelism
and oonWTg<'n",", both of which m.a)' occur he-
twren related ao well "" unrelated lineage., we
cannot agree with Schiodewolf'. d;"tinction.
Good exarnple3 of time signatures are found in
the in""rtebrat .. and, among t hem, bore again
mostly in t he ""phalopotJ.. &hindewolf (1936:
44) cit .. tbo " .... peated and apparently indepen.
dent" {translation from
of triangular ronchs in various oet"ie$ of Oymeni.
ids of the Upper IHvonian. Brinkmann (1929:
2H) notice ..... uffllllige Trachteamoden" ( _
. trildnR d ....... fashion.) in the CO$lI1""",.,.tid. of
the Callovian. Haa. (I114Zb: 142) that
the spiral ornamentation ""perim.-I on the
primary radial one in .. "",.,.1 genera of Dipolo-
""ratid. of the Albian, a. previouoly notod by
Spath (1922: IOJ, 146) and Mkin. (1928; 234),
which reaches its peak'in tM genu. Elobic"'4S,
might be a ca .. in poin!.
Thomas (I'WI: 52. 5J) cit .. th. unifonn
change, in the eariy Silurian, of all
families to tm, morlOgTaptid type u another
instance. amOllK invertebrates, "where, instead
of there being later repetition ... as it were, of one
of Nature'. fashion. Ie!. the following l>ea<iing],
similar form. an: independently p<O<luced alm""t
simultaneously." Scott (1891: 3(5) ga"" a
ditocussion of amon);:" Tertiary mammal
sku! .. that might now bt interpreted ao exempli.
fyin);:" time signatu""" hut in the Ughl of pr<'ient
greatly increaoed knowlooge h;" Il""eraiization;"
qu .. tionable. We ha..., found no clear example
oltime oignatu,,,,, among vertebrates.
!lOMEOMORPHIC RCURRENCE, ITERAll0N
In a non.l"<:hnical way the term recurrence it
applied 10 ""I)' different phenomena, e. r., in
the 6eld. of Koology and paleontology to the
return, on,"" or .. veral time., of "cutain lithol<>-
Kic fades" 19U), rontinental facieo
(Willard, 1940), or certain fauna. (Williams,
191J) to the same area.. With a more technical
phylOgenetic meaning Hyatt (1889 : VIII) ap
plied thi. term wlten he noticed "in lhe di!fm:nl
genetic ""ri"" oi a Iype derived from "n, ancestral
.lOck . . . a P'Tpetual =:urrence of ';milar
form. in .imilar """", .. ion, which"'" usually
called repte .. nt.o.ti...., and oiten falsely classi6ed
together, though tltey really t..IooK to di""Tgl'nt.
i""etic .erie . " 00110 (1922: 216), on the oth.r
hand, in"ts that recurren"", ne""r happen in
paleontological oerie..
Mojsioovia!. (1893: 5) ""neotod tm, itrm
"heterochronous con...,ryence" fqr the similarity
he ob ... ved bttwtlth unrelated,
aneou. ammonites. Bockman (1895: ( 57), how
""", prel"", to call this pbcnomenon "hewro-
chranon. homotOmOrphy."'
Ha"" (1942< applied the term re<:u ....... """ to
the. reappearance of morpholOgic t ype. and
evolutional)' cycleo oi ammoni",," of
an tarlier geological period in a later one, equally
una"""" of the earlier more general state"",nts,
quoted abo,"", and of a prevjou. U .. of the same
tenn by o.,p&et (1907: 235) with reopect to the
same phenomenon."
In the ca",," ",Ierred to in the two preceding
pangr.>ph. and in similar oneo" phylogenetic
significance it denied to the ' .... mbl,..,.,.. be-
"""""n the hetemchranon. ammonite Iypes.
They may be con.id<:red typical examplea of
homeomorphy, ..... a rule, only ""tural differ_
enceo warrant their distinction. Soch .".,...
might, therefore, wen bt called """'-ph;';
..aurence,
.... oomtwhat related, though, at least theore-
tiC'llly, different phenomenon ;" that called
"'iterat''''' Artenbilduni' fonnation of
species), deocribed, "" follow. by Koken (1897:
40):
A certain type penio .. throu&h lon& pen<><lo .t.
m",,' "nch .. ,ed, hut """'meo ...--tedly .1><: .... ,...
inl !><>inl for formation cl >pea I",,"nandy
rad;"tinf in all dkectionL Theo<: hOi .. cl vatioti ..
and . peci .. Ii. on. abov. tho othor, like tl><: Il00 .. of a
buildini", apparently "'ithout .ny diro<:! jI"Onotic
connection. S;mUar forms recur, .11
from. con ........... ti ... p" .. nt otodi:, but nOt ,ivin, ri ..
the Ono to ,I>< Oiller,
A hriefer del!cription it gi...,n by Koken in
1902 (p. IJ): "A persi!.tent opec ... puts forth,
from time to li_, varitli ... which appear in
hosts, a. it wen:, be"""""n more 01' lest long
pau ..... " (T .... ndation. fron> the German.)
"_ til< c..- """'ition
"Rolt ........... , qui,. m .... _ly r ... rrtrO\I..,..;"" ....... tion
(_ im.r'" 1!l9l!: Jf, r).
"s..:, 0.1., 1911: 1-16, " . N.
OJ 10 HAAS AND G.\YLOIW
The original.,..arnp!c for Ihit I,henornf:non ,,-ao
ll><Iiod by Koken in Tri:lMic po\rOp<><.ls of tbe
W .... gTOup; nlher Uk'& .... . tudied in
pcctinid pelecypod. of Ihe Vot.. Iyp< by Philippi
in aniid brachioPOd!! of Ihe Silurian by ,'On
Huene lr><. w.).
ikurl<n (1929; 51) ptOpOtCIlbe oborte<" 1-Cnn
.. it" .... tion for 1M pbtnorrIcnon hen: .. -ith.
"' ... cribi ng il U lbe "pe'iodoll)' """"",,nl
10nn.1llion of I ...... m .. or similar In"'. within 1 ....
sa",.. "'''''k ('St.:lmm}," prHUl"J1>(8Rl( "no im
ll\f<liat .. ...... .,,_ ('ZueammrnKcl>oeril
kcit'j, but indiT""'- ",la.tiontloip" and statinR
tM' "crrt.:Un r<>nn:ttiw trerId . in""""'t in """'"
I;>'ilt. group. manif""l from time to
time." lku.Icn' hortcr le.m i. here giv.,n
pn::fff<"nCt" 0"'" Kok.n' . 11K, """""" lin"" il i.
nol. :II il lhe lauer, ..... 1,.;,,100 10 lhe opeci.-.
lewl and. lherd""", l'I'"T'ill of "..,..., g<1Ieral
applicalion. Iku ....... nnpl ... in Ihe di/f"""",,"
t..1......", ileration and <'OR'dlb"'" {roueh1y cor
...-.ponding to lhe one btn- bcinr moo. brl Wttll
il<'Rtion and .... 'OCUrnfI"") .u "'-
Schindt:wolf (11140: 230)" in <IH<1ibina; ....... nli
ally lbe ""me pmnomtnon ". "i ler:lIivt: rel"'li'
lion of always the evolutionary rou,...,.."
!-Io .... e""r. cardul e.aminalion reveals thi.
di /fcun"", 100. 10 be ",Iali"" only. <lep,,,,ding- on
I .... luonomic 1e,..,1 of th .. in,,,,"ligalion. TM
l\.,nxh"""",oIy horr>eomorphic amruoniteo.
ffioefIlionnl above, f .. eumpie. ttr .... inly do not
do_, .... Ihe oroe from II>r Olher and are aJllJ..
l"1r...- unrelaled, "" Ionl '" we think in lerm" of
",nCia and lamilirs. Vcud. r.o...."...,.-, .. n"","
1M "..,..., f'-"'Icr.ol a.pecl oIlhe onIer
ali an entily. th_ fi.'CUi"i'C1>' m<wpho!op: tyl"'"
lum out 10 be JUI rcpr ....... tML .. "" of oomC of the
".uCttoli,,, wa,..,.'" (Spath, 192J: 33) of IICul ....
turN ncntic linea.g<'J which. t hrougbout the
M.,.,wie. "Rain and aKain branch off lrom lhe
omootI! pelaiic "coo .. rvati,,, 1100:1<00'" (" Kon
K'n'al iv.lhnme" Salleld) 01 lbe pllyllocernlido
and lyt"""""ti<\o, 10 I'" h,.pod"";'
by Spol h {l91J : 14. 31. 3J; 1917:
6.H\o4; 1933: J; 1942 : 6II1-6&J} and Salldd
(1924) and .h ... .,1. IhouCh in ..... , .. w .... t ob-
KUrt: w:>y, by HlICk"",n {1915: 9}. 110;'
.. A""h i. deft h <llt-n ........ hnlicbt C".<tII .... "M
auf .. h. "" ..... _ I>n,.ntkluopo,.I
.... &.'n .... 0.10 .. '1.10.;' l<e4"" ....... K<In_,
..... .iolmeh .;,.. h .... I .... W; litto
Ikicb ... En,-.;ct!"",tablhf.. Wi< , I .. b;.. .....
:. ... I!;:,,';':-:-
t to Gnoodloee ""0/&, ."
i>bmornenon i. on an inmmp;o.r .
ab/)' ""ale, '''''''II,,,,lI
y
Ih" ....... <os
Kokcn- Beurlen. i","' tion. 100. 1hI-
1a1, .... 1 off....,,,, are not immediately int ...... la'oo.
bUI all 01 Ihem may "" dcnWld h'om ""me com
mon an""'lral siock. Thuo, the differe""" be-
Iwn hn""""""'l>ic and ileration .
., clr .. at a 10'",....- taxOnOmic 10",,1. Iooeo """" of
ill$ll"'p;I<_, n..y. d,,-indloo a .. -ay lhe....,..,. l he
pcnpttli", ",-idfruo .ndt>r "'hich IhinltS ,.,..,
look...! at. Just ",-rol" 10 Ooobom
(..,. Oobom. 19070: 239): " E",ry liv;nK Ihing
.,... ... \1 propt"rtieo to t .... 1 i. 10 oay,
it. fund:om.'t\1:>1 I""OP'rtieo 10 a homO\l .... y
rommon 10 il .. itl, all other livine thin2'. and u
you run dO"'n group within mu.t be
a ml)l"l) lI(lO<:ialiR<l homOll'"" Y afflinR mcmben
of .... aller and .mall ....
SUN.\I.-\RV
Owm. (Iperla!) honoolocY oloould tM, rntriclfll
to limilanl;" bel"'C('I\ I'UU. orcan., .. Urvcl-
...... 01 d,ff"""'l organ;"". allribulabi<: 10 com-
mon al\CHlry; homogeny (L ... then be-
coo, .. a lynOflym of homology.
Homoplaay bring. aboul simi.
Inri l;"" bel.....,..., organioml D. Iheir 1"" 11, orian.
O' due nol t. comn>Ol, anoo:otry b .. t
to ind"P""dcnl u Quisilion of t he limilar char
",,1_. E,,,-,,, when: this intkpt'f><lrnl a.cq .... .
lion m..y he attnootfll to oomc or pot .... tial
ptWi ....... tion inheriled by balh .... all lin .. COQ.
crmoo from a common ancutor (Oobom.
b ll'l1l or pot .... tial Io.om<>logy) , the t.imilaritie5
be ('()I)sid<:n:d '-nopl;l.otK: but "'" homo_
Iosrou.. (COJ>I'), homon-oorphy (Pal.
i .. orn .... phiom or ioomorphy (Po..""r, Far_
brinl"' )' homcogen"';. (Eimer), and morpho-
IOIlkal ""lui .. ""'n,,, (Hya tt) are rolloid'-""'<l
")OOfl)mS. "n"logou. variation. (O" .... -in) and
ind...;.endcm (Hubb) an ronoide<ffi
oporial.,..... of homopI..:aoy.
Otr...- "Pt>Ci:lI """"" 01 hornopluy a..,. coo'''''',
gertcoe and parallclism. d'Slinclion ... bid!
thoold be ba-' "" ,,hel r...- or not similanly in-
with evoIUlion .... <her t .... n on <leg_ of
,..,l.3lionohip brl .. "fl:'n the Ii ..... invol....J.
may load to hornoomorphy.
Tho: laltc' IL'i"m ouRht to br .... , ,.;,,ted to . i ",i-
la.it ift al..:ting Ihe whole OUler of
the h<>rn<-omorpho and reACh'tll ouch a dq.-..",
Ihal tho:- """ may be mi",aken for 11>0:: "ther even
by a ,<ained natu",liot . .. n!eM ""r\.>.in int..-nat or
oboe .... .... """ Cx.3minood. 110me0m0r
"NALYSIS OF SOME PHYI..OGENETIC TERMS
'"
pby it not equivalenl 10. '- doet i, indudoe.
mimicry. but the 1 .. '0 cot>ttpU .,....,.-Iap.
Ncilt.er homopluy '- paralloolism oor (011 .
and homcomorphy imply ""yl hinl: ,..
to OOCIlemponneify of lbe ';mil.,. dlanrel;
t'- phmomena may' be hele'ochronoo. .
or itoehr...-o ..
l _hl'llliOUI CQIlv"'l!<nQl! and Ioom.eomorphy
may. IUOIi""''''''lly. J.ad 10 pol yplo rJ.ti<; n\OI"
pholof[ical *,,1\Ci'a and hig" er uni ....
Dacque. ICfm, "ZeilfornoenLildung" and
"Zcilooultile" ore rejected. hili .. ZCi ... ii>\atu ........
(li me oign..IUTCf) are admitted ao " d_riptive.
nOI t.planatory. I""", .0 designalt cntain <:UH
of iIochrorooout paralJ.li>rn or
01 onnrpboIogic Iypro may bo
,,,,,,,Iy homtomorpItior r..:utn"1>Cl< or a.tnbu!.>.b,,"
10 Koken', .. il ..... i"" Ntenbiildung" (i"""'li""
fonnation of tpOciH - Btun..n. i ...... Iion).
REFI>REl'CES
O. IQ. "-........ 0.:1 ['eo h-v. r ....
..... U . """,.-N<. Gu, ...... la ..... 1M, (m }-
(UO).
.. ...... S. 191" 11._ .. Tuoa. Cr. ..........
["';1 .. U. ;', T..-.lloiJ . 2lJa.
N. 1921. Evoluoioo 000-"., aod d ......
.... ' (",",oSontiol A,Id .... ), p.. 10, .... .
$<i. C,.,., 1_16.
A.ute. W, j , 19U1, Su.n""'" of d EoropNo Juno ..

.. ;_; ;,;:11011. G.oI. s.c. A _. ,"" I-J4.

...... IN. M ...... l> f...- .... . nody of voN, ...
, "0.'''"1')' . 1", tho
oil _ "'"' """".. of
; 1l3-lf1.
_ 1_ ...... 1*,. Qoioo1. J_.
G.oI. 50< ' .... U, UJ-4.\.!.
_ 19. T)' po .....-.0 ........ I.ooftdoon. .. "" a
w .....
c..o...... E. I'NI. -_ ..... 1 001
iIIu ..... _. A_. J_. $d. Uf , t9t-

... ... D. 1861. OO'I>< ...... of .... ,.. P, ...
AcU. Nol . S<>. PIoiI<o. :!G, 2U_JOO.
_ 1'71. Ropon. _ the .. tinc' ..... tebno .. 010 .. 1 ....
i. N." M.,oo" by ... of tho pod!'''''' of ISH,
RII. U. S. G..." . S ... , ,,,,, ofUo. -"I ..... -
I""""'tok,.-y. P'. II ,
- ISSI. Orir;n 01 ,he fi,..... lioMyo on ovoI.""".
N_ Yori<, "nh _ ,
- 'Tho po-imuy f"""" 01 _nic _., ......
CIoall<'o Opoo. Coo ...
DACQIlI. Foo". 1911. V .... ,0600 f .
___ r ........ n' ", ... T .... , 1Ioo-1io. Iloro.
,_.
Orpaioc"< Moo."".> .001 ... _"1,, .
Bottlo., Iloro."'il.
D .... " . cw .. , " 11160. 0. tho iitioPn of '.' .. . "'.
New AppI< ......
_ 111'1. 'Tho" ," ......... . 001.......,. ;" .-.10' ...
... -. [a ll. s.,.. Yori<. AppI<_.
- IQ. 'Tho ...... of .. _',Iall . N."V""' ....
1 I ... A"'_.
Dot_ ...... ' .. OL 0.., ....... ovoIu' ..... 'Y ", .
oIon-. ld od .. Coa. .. blo.
0."1lliT. Otn t.U. '9(11. La "".of.,......,,,,,,. do
...,...10 ....... 1. !'&no. Flo .. m&rioo
D, , . , C. eno_Or< <lor 1I"'"a,!o .. pIo".
I,c;po.ic.md \ " ..... n Ooutkk .
Duo."", Lout. . I". ""pIo.kv-lto _1+0 .'
I ........... "'"', de 1"<!voIu';"". 1Iij*. DM-I , U"_"
A"" ll ....... ). A- U ,4. J: : lU_lUo.
TM. 11">1. 0.. ",,10 r ,ito.oooI .... ''''''''-'
I iii I 1_ ...... T .. _
....... - .... '-- ""- ......... _ . .......... .."...
O li O HAAS AND GEORGE G,n-i.ORO SIMPSON
_ 11I00I. n,..".",olF_ ..... I .. . 1IioL s.; .. --.
MiouM. Co/l<d. " (I f II).
_ 1910. P.Iooo>......,..ne! ",,_y (I. , Th< l'a\<.
"""*'lie R<ronJ). $d. M,..uJy 77: 295-291.
_ 191l. PnDciplood .,,., ... pt,y. N ... y",k.So;'o ...
C .... ou, ,u"""Uf w .. M .. "" ... RUD. 1911.
M"ta'ion. ol S,;.v" ............. , (Abo .... Pr.-.
s-.r. I ...... Z .... c..,.. _, IR4 A ...... .
IHP'7n_76&.
':',:, :.:'
J_. hI- 16: '.
_ INU. no. \' ..... , .... "" .... 01 C_ .. (AI
bio_)'''''' i_"-""'J<IIo. BWl.A_.N ... N ...
HUt. II , 1_2U.
tI&><nI'-, J. F,...,... W. 1.1. HI"""'_. 1929.
dictloftary 01 _tiIi< ......... 2d ed.. Edinbwch
..:1 U>ndooo, 00_ anc! 8of<I,
1t .,...,G, O. Hanc!l>II<tI dot ...."._n .nd
.. pai""","U En.O' ;ckl.nplth,.. <lo .. Wi<bel,lere,
J ..... t'-...
ItO .... Co .... L 1_. C.......,.. 01 I>o<t>oIoiY """
... Iocr. A_. N ...... n: 1_.
It Ut<T, J ... u. IMl. s.. V .... CuI,
Hun, AuIt ... " 1154. r-n oepb&lopodo ......
101_ 01 c.-__ ..... ZooI<oaY. PHt. A_.
A" . .I . s.o... <I _ .... Jl)-J61.
_ I,". Cp' . ' oI .... Atiotbcloo. N .... W ... c..'.
Z .... 16
_ 1Il90l . !'II"""",, of . n _iN<! _wi ';c.
I'r ... A_. Plnw. s... U : ""-6011.
J. 1946. Die H""""' ait oJ. A .... n.ock pmbol
lid>o .. /Iau"u .. v"," Ty""". Boll. !i. FrtHtu(,
S. N".l7 (l04l_I944):
K<I&Il, E. 111\>7. Die c ... ,"""""" do< Tn.. . ... HaU
"'L U . ,... 11'",,"""".40 (1196): 31-116.
_ 1902. h'-'olOioP 00<1 J_,
F..,..,..
. ,
_ 111Ot. 0.. .. ..
. ,.
McKn, Eo .. 'w O. ' .... 3. Somt ..... , .... p/Ii< pri..clp!oo
il ....... 00<1 by 1'0.. * dtpooiu 01 _ ...... "",-.
A_.J_. S. Ul , 101 _lot.
N'V...u, Sr. 1110. 0. ... _ 01 .... _"'.
..... 1 .. ,.- A.,. . .- 11. ( . N ... HUt . . .. __ ,.'
llZ-II I.
M.".._oao _ 1tI"' ... .lJt, HDoou...... 111')3. Die
pII.oJoo><rI d'" H.ollotl .... Koike, II . AlkNl.t".
,..,. R""_.'" 1<1 IlaU.
M""'Jn', GA,"". __ Il. I .... Tho ... lation.ttipbet_
...... 1 ."" . p0c41 ."....;.: ",,"1oci,1<o.
A_. N ...... 71:
Moo..', I.to>t. 1'>40. M.n ... 1 do ... ."i"",lo.
r.,;"1.1.-..
NOLJ<, H. 1_ . .-.- _ lot 0 ......... d. IfWpe d.
cs , ..... I W. c. II. Solo .... ' '''_ , Jd
_ .tI, 11S-11IIi.
007M", It u n FA,&f]&U7. 1'100. Tho ..... ical"""
Eo ......... ,0.- of I 'a _ **' Amorica d ..... tIor
Too.;'" .... . , _ ... ....",. d ... w ,
"n _, ..... 77:$01
110.
_ 1110;10, _. Eo .. 01 Ia_, '" _""
............ ,. A_. N"", It, U'J-17I.
_ 11l1)n. Tho laO' d adapt;'" "",;.,,,,",. A_ .
N .... '. H: Jll-:l63.
_ 1\1'03, Th. Id$ nd ....... ol ,,,.,d,," phi1ooopt,1co1
.... """y. s.u-, ..... 27, 9''1-401.
_ 1901 . Evolutioo ol IOIm""'l .... 0I0I0 .. ... ,h '"
..... 1..- ,II< ......... l&I ' ype ole. .:.1. by W. K.
Ct .... y. N ..... Yod.
_ 190' " E ......... _I'.--"""Ioo po_toIorio,.
Sc>-'., .... , 36: 7 ..... 749.
- 19Ofo. .... ol_tioatt<-
SO-. .... , 17, 14&-130.
_ 1_. C ' .... , ....... tioot w-ah
-. 1'-. 749-::5l.
_ "'0, P>.loontokci<: .... : ... Mc 01 odop,i", .-..1; _
.Iiooo. I'.,..u.r "_,, July. 1910, 77-.11' .
_ 1912 . Tho """I.""" ...-lei. 01 _"In uni, <il ... .
""ton -..... by pole<Joo"*'li", A _. ". .... .
40, IU-:roti, 14'J- lla.
_ 1911 . T ..... o!a>y, II .. 10 ... 01 ,ho /"", 1 ... ".,.obI<
f."""" 01 ............. J_. A<M. N ... Sci. .
2d_ ..
Oo<u, 7I.OCIUOD. lilY. 1._.. ,lot _"",,<l..,
.. _y.oad pIo, - ..... 01 tho i __ .... UaaIa.
I -'"" Loo.c-, B..,....., c... ......
_ lt1l. Ropn .,. .1ot "",,"'I',," ...... "I: .....
.... . ..... 1>0 .., ok .. M. K.,. 11/.\ II......, II';';""
Af_. A_ So' .. III9-UO.
_ 1161. 0. ........ .-, 01 _ .......... I. F'.-
.. '" Rtptilt.r. 1.oa<Ioo, Looc-, Gr-. .....
Co<npony.
J.,\. IMf. Ow pol";,.
<It, C<ochloch-., .. 1001 1 ...,_. 1;1 .. ""'"""'"
q;iod>t Uo_h.".. I ... "...., Eo,.""",
l"''''l<Q\1otu:, L. 1907. """ T .........
5 I.... la: _ d. 1'aIf.lIIooIoI. TU ..... ..
I (Car .. c ..... ;.,_ do 10 T ...... ). P>.t .. "' ........ 1.
1'1". J. z.. Krim "'" Go" .... k ;,... Y ...
_ . -.---... Ga. 11-';'" 7'0:
-"., I'.a..t ..:. Illll!. t.. .-... jo ..........
., <rl_ "-I do C
s..u .... , II. 19zt. [);o _"'III. .... K' "'''v.
. ., __ f lt "" 5._ --'-"10 ... .o"WF'
.,..,m , ......
idooo. CntDdboti<a I .... di.tr Erlonclw.. .... "",.
wldd ......... .0_ dot Jon . Dd
Lolpok. .
s.:: .. !OC1I, lI u OT G. 1m. Iltvle .. 01 'W" po ..... by
Norman D. N ....I1 OIl 1.0 .. Po,.".";' .. Iooe,,:o>;Io.
1_ ... P' M _ . 17, 6 6JO.
Sc.,..-..., Om:; II. 1'36. P.""",.,.,.le, Eotwlok
I 'i I I.. '-" Gr.otlk. Kri,1I< .Dd s,.._.
fItrIUt ..... _,
_ IMO .
ANALYSIS OF SOME PHYLOOENETI C TERMS
,.,
. ek.H_r ..... ......
.. , .............. .
"'" _ ... 01 .... Liuoic
in "'" Bri,. M-. .. (1'1 ...
DoGuoIr. IMO. r...a. ..... f"h .
r ...... ........ &.r. u ..... 5<1 .$. f.,.. 19J9,
If 'Off-ocf
'ltf '9t," 'il' 'ltr 'Iltf "")1"1( ....
"'0fr."'''' "'iP1fo"J
6/)' 'trf .( ... .. /'1!."',,) '"!'-
ttf'liff'm '''''!' ......
m ' ......... ...,
flf _,.,...,. I'U ......
U[ "MOIl"!""
Iff' . ...... "'>!'""
IU "1"" ' "I"U(I
rtf 'Itr 'iff ' / ft '>I"'I-llf4lr><' 'fluo
ttl. ''''''"("'' Inbo " od-oQ
litf '..,....,."''''''' ,.O!I"O
rtr ,.a..,o"
Hf 'IIO! ......
111' iff ...., ... 1'11
ttf 'lff
' Off ' ttlf '" , .... A!.bo \>' ......... l
af ';CIuo["I_ !901." ;
>11:-t:tr >! ""''''''!\l
,rr
'Mf .(, "."'I''l'4 ...... )
Itt ' Qtf 'PP"IX
5tf ',tf 'Otf 'fff 'A<>!"'I I'
,If 'otf ' '''''!\'
I lf
rtf 'lIT 1 'nr ""'I
9I'f ""'>amp , ..... .,
iff Wf- Iff .... ' " .,. -
"1"1'"'' (---< .. , ... , .. :IJ-....... X
ft, hiI ...... X
.,. "' ..... . " ''''''-x
'.
.If 'tt,-',
'(' '"''!II"'"' 0'""1)
ttf - .. .....--.v .


Itr itr ' '._,. ""'!'"""'I
Itr 'Itr ... r '"._1
IIf-ttt' 'I'ff .""'!'_(
'If 'ttlf "If " Ijd_1
ttl' 'ttlf
'"
'rtl'- Itf '6ff 'tH 'iN: "_>pOo1
ttl' 'Off 'af "'"!""'M'I(
'If ' , tf 'lIT '9i'f 'm'O('I'
-ilf '("'].lOU") (_)
.... " ,. ""'fI!'!fIb>o ,n ..... .,..,
Iff " ff '.o.L''''''"'H
Itf '''!'''''''''''''H
Ilf 'Ilf .. ' .. """-.<In
,tf '11i' ' 111' 'Otf 'liff
'lff ',IT-t:rr 'Ofr-r ''-'''''''''''H
fft
tf '61T "I.'r '9ff 'Iit
'(1(1' wr Wf 'nr 'tlf ' >!>
I'l ... .. " I" Il
I ff 'Uf '''''11 "J!
af '11lr ' Iff .I t! .. .'11
' Ii' 'iff 'U f .,.,...
,.-
.If ."' .... 'fdoo>'"
"r .... ,.
'11f 'toN 'hr 'Off
ltf ....... l)"""",)
rli' '",- ( ..
nf - .... 1 'f"""J)
l IT
. tf 'IU '''',.'''''"1
rtf ',ooaof""O'(
I ii' 'rtf ' .... I,..,j '00)"""" ""!' .... "',.
l tr .............. ,.
fK ..... n ... " "_j
'ff . .,.. , - " .", '," _ "'"3
fK '>Opt> "'"""',..,..."
rtf .... , .... W
Itf '6fr 'rn: 'P!I'-
rrf '(1(1' '...a....IP
'fI' ',.-..,."
l Zr
'Ii' '/fr 'O!'i' 'ur '(,,!Ofd
_) l"'t""'''' ""I .. ' 'N(tA!"Ir,.
6lf
..rr ''''!.,..... .. II
fff 'orr 1'o .. lQ
nT 'tIT 'Off "" .... ,10
It. __ "!'to' "!"'I"O
n f '&>(;>\;)
iff 'flT- lff 'N>' " '''0.
""'><07 ( ....... '_X)_ hil ..... J
fiT ._' ......
fiT 1I"",,""J
Iff '., _E' ..

IU ' lO"!p..oD
kpo ......
rr . 1"""!'' Jl"'OO
lit
'"
-.!If 'rlf 'llf 'ol r ';<><"''''''''J
tIT ....
tt1.. 'q><IlO .. !' ...... "'O
lli' ....... "'J

'1ft " "'Ii ..... l>Uh()
'ftf ' .. y ......
rff ...... ,,_, .",. , ..... y
Iff '\lff ... ,nw\, "'110["'",
Ilf ' 01:[ '" ..""."\.
ttl 'Iff 'IIlf "11U '" .""!'''!'"-'
ltf ..., ......
.t"f'tlr_ .... ....
'1ft 'rrr ' t tr .. ....." ... ...
fff _,"!4-' 1""!Iot ... ...
Itf ' -" IIV
Itf ''''*''''''' H .... -.llll'
fff o"!"'''I'V
li:f
Iff 'ttf .,,,,., ..,,
m . ..".. ...... , .
,ft ,,,,,,,.,..,,,,,
ttf ' 11'1' 'ftT '9ff'm 'orr
-t:" '(_,_) .... >eo..., ( ___ )
.'1..... ,. 'm : '"..- ..... "1 ......
Iff ._ ..... 110> ... _ :. ' "<'
(13sn SI'IIII l.L ;)1J.3.'I:i:J01AUd "0
ON01.\,' 9 0\' " SVVH nu.O
'"
,I.NALYSIS OF SOME PHYL(x;ENETIC TERMS
'"
1'uoJ ..... JJO
P,--,",<, 4 '-"'_. J1I
Ph<nO'ypic .. .,...,... .... lJIi
Phylupno'" ........ l26
_Iotonrun.,""' .. 1J(i
""""""'. J.U
Pia .... ll6
PoI""'y\rtic l19. 331.
lJ&, J41. J41
",igin. 331. 33' .H t .Hl
P"",.tiol t.omoIoc,. Ju. Jl1. lJ(i,.l
phyl_,ie, 3J(i
Rao;fi.ti<xt. wp,; .... 3J3
Rocapi.uIa,;';". 330. 3JI. 331
Rocti,:,..J.'ion, , .. .. l11
&.010........,. (Rd."' ...... l. ,129. 330, .Hl.
M,
homoomorplli<.lJO.
Repo';tioo. _rogo:n<tK;,
i' .... 'i ..... j15. j.[J
Ikpo..."",tioo. l19
R<"",_;v< J.U
K ............ , phyior<oeti<, J.U
SeIoc'""" ... ,.,.1. 116
s..lal homolon. 321, Jll, J14
... J41
s.n... panl'" (P",./IW"/wol. lll,
,W
.... ,1_. J..II-34J . .145
S;";lari,y. J14
SpoH" no. lll. llJ. 324.
"'
SpooW>outl. v ..... ,""'. 3.15
Stock .. """ ....... ,; ... .l
St"""" .... lZJ, JU
3:/8
T .. ,.p!aoy.321
Ti .... J41
"''''.'''' 3.,- lB, loU
T_"''''_, J4j
Type<, """pholooP<, J.iJ-341
\,ori.otioo, , . .. lIt\. ll3.
311 .l
,pooUneo<Il , 3.15
v", .. .n. q-;c.I. lUi
<!<fin; ... 317
ll1
. . . " . lll. lH
LriI<W.lW . .. ... .HI
...... .142
4iI4 ... ". J..II .141 , loU
Z<i< .. ".... , . . ,,".}oil