• TABLE OF CONTENTS
HIDE
 Title Page
 Synopsis
 Introduction
 Materials and methods
 Taxonomy
 Variation and predation
 Pelage patterns
 Color variation
 Size variation
 Origin of pattern traits
 Evolutionary sequences
 Incipient speciation
 Acknowledgements, Specimens...
 Literature cited
 Back Matter
 Literature cited







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Title: Variation and evolution of Gulf Coast populations of beach mice, Peromyscus polionotus URL: http://ufdc.ufl. edu//UF00001532/00001 Site: University of Florida Digital CollectionsVariation and evolution of Gulf Coast populations of beach mice, Peromys...http ://ufdc.ufl.edu/l/UF00001532/00001/print?options=1JJ 1 of 95 8/1/2011 8:17 A M

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Variation and evolution of Gulf Coast populations of beach mice, Peromyscus polionotus
CITATION SEARCH THUMBNAILS PDF VIEWER PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00001532/00001
 Material Information
Title: Variation and evolution of Gulf Coast populations of beach mice, Peromyscus polionotus
Series Title: Bulletin of the Florida State Museum
Physical Description: 91 p. : illus., maps. ; 23 cm.
Language: English
Creator: Bowen, W. Wedgwood ( Wilfrid Wedgwood ), 1899-
Publisher: University of Florida
Place of Publication: Gainesville
Publication Date: 1968
 Subjects
Subjects / Keywords: Oldfield mouse -- Florida -- Gulf Coast   ( lcsh )
Zoology -- Variation   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: p. 89-91.
General Note: Cover title.
Statement of Responsibility: by W. Wedgwood Bowen.
 Record Information
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0842
notis - ACK3660
alephbibnum - 000442999
oclc - 00468973
lccn - a 68007801
System ID: UF00001532:00001

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Table of Contents
    Title Page
        Page i
        Page ii
    Synopsis
        Page 1
    Introduction
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
    Materials and methods
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
    Taxonomy
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
    Variation and predation
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
    Pelage patterns
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 42
        Page 43
        Page 44
        Page 45
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
        Page 54
        Page 55
        Page 56
    Color variation
        Page 57
        Page 58
        Page 59
    Size variation
        Page 60
        Page 61
        Page 62
        Page 63
        Page 64
        Page 65
        Page 66
        Page 67
    Origin of pattern traits
        Page 68
        Page 69
        Page 70
    Evolutionary sequences
        Page 71
        Page 72
        Page 73
        Page 74
        Page 75
        Page 76
        Page 77
        Page 78
        Page 79
        Page 80
        Page 81
    Incipient speciation
        Page 82
        Page 83
        Page 84
        Page 85
    Acknowledgements, Specimens examined
        Page 86
        Page 87
        Page 88
    Literature cited
        Page 89
        Page 90
        Page 91
    Back Matter
        Page 92
    Literature cited
        Page 93
Full Text




BUL LE T IN

OF THE


FLORIDA STATE MUSEUM


H 0 OGICA I


SCI NCES


Volume 12


Number I


VARIATION AND EVOLUTION OF GULF COAST
POPULAIlONS 01 BEACH MICE.
PEROMYSCLIS POLIONOTUS


W. Wedgwood Bowen


A
Mr -


!

v'o.I


UNIVERSITY


01- FLORIDA


G a i nesv I I I e


1968


k .A P I












U N I V E
OF F .C


R SITY
) R I D A


.LIBRARIES




















VARIATION A NI) EVOLUTION OF' Cu.LF. COAST
POPULATIONS OF BFACH \ICE. PFROMYSCT7S POT.IONOTUS,


W. Vi- --y:w0D BowEN"



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1968 BOWEN: GI..F COAT BE.4ACH MICE 3

the coast where the dunes have been bulldozed away; ian inland
where the land has been cleared for reforcslat int, these u cc arc ir ow
extinct or rapidly bcomiiw so.
The palest form, inhabiting the almost pure white dunes of Santa
Rosa Island on the, northern CGuf Coast, was first described by A. H.
Howell (1920) arid named PermiqsrtsusF leurocephalus. The greater
degree of crptiHc palenr-.ss in this form (in contrast to the somewhat
darker mice living on other beaches of enalilv white dunes he con
jectured. offered "a strikhin ilJlstratioi of the cflect of isolation in
perpetjuatin g color variations, aInd in the case of the Ablbalnii repre-
sentativ'c of albifrrmis tiu- iiitadiquacv yf a p,.cH iai er-ivion rerit 3pa.rt
frnm iolntiron to produlrce similar changes."
Sumnner ( 192W), who was then -eil.g L'd in pion.er stadics to de-
tenrnine whether or not georaphic color vari tion w;.s sgpnetially
controlled, became interested in this case. His findings (1929, 1930)
have been widely discussed and quoted in evoluitioniry literature
(e.g, Mavr. 1942., J954, 196%3, HuRley 1943,; aildane. 1948, Ford,
195-1, I960, 1.964). In Smner\ tim's e the .,vstenatics of the species,
epccildly along the (-lf (.:st, hid nt been a.id', telv, studied and
Si.imncr's conclusions were conTsequntltriy based in large- measure on
misinterpretations. The first major purpose of the present paper is
thus to clarify the taxonomy of the Gulf Coast subspecies and so to
lay a foundation for a thorough re-examination of the course of dif-
ferentiatiou in these mice.
After collecting population samples on Santa Rosa Island and the
adjacent beaches tr west and east. Sumner (1929) xxt ran a line
of trapping.p stations inland at approximately '20 mile intervals, and
found what appeiarf"d to be a cliner terminatinLg in an abrupt change
from pale to dark som n 40 rmil.-es inland. Tinis he interpreted as the
result of gene flow extendi( inland from thle beach populations.
Evidently he was not LawIarc of th<. fact that be.twteni his coastal sta-
tions and his first il'and trapping si nation at ('rvstal I ake an extensive
swwamnp (the Devii's Swamp poses an effective Latrrier between the
two populations. Furthermore, for the past hundred years ( 10,4(00)
mouse generat!uns aik: )tle-t bArfrit-r tu ogeI-JN flow has exisvted, crcrated
L -
by the intraronai:1l hip cSinal I vn i( heI dseovprv -Of thesP biTarriPr
that first le-d to the present investigiatiun. Later it was found that the
line of abrupt change from pale to dark mice coincided F with the
highest (Okeft-nokcee) Pleisto'ene shorline (:' Mac'Nc'il, 19.50).
Four extensive field trips have bcen made to collec('t specimens and
acquire first hand kno.l, cdi-e of the topography and ecology of the













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BOWEN. GUL.LF (.) AST BEACH MICK


along the Atlanti c oast. On the third trip 32 days were devoted
to further studies of the nrthirn (;Cultf st .'rom Bald win Co., Ala-
:arn, .-st to the Ap;ilacihe Bav region aF d orth i o Crest i-'I
Marianna, and qa i inc;y tihe rt-e ainii 15 days wvrr then de-v ttid
to cornp;arial ive st udiec of tCoporaphical and ecol( gic-al conditions
in penstiisil -ar L"o r (UL.t i3c(!idi uli (tit- r;ang I :,s If ;ill thrve -1tl; rIt-
A.s.t sbhspeciis Fi.:idquarters for the fourth trip was t "e Florida
Slat:-' M tieue1i I. toi ll i \ Iti ri .11 ijb\ t'r L~I Ic ii, il! dirtctinits t,) cl} .k
jald arniplit t1 i' ir itf.rifldfpiriaf ;itaif.d c(clichlisio ns( draw i di-.riig ti. 1 ihrfe:'
prcvioul trip A t",ta]I of 11 davs vwas spt, inc th-e iiiIrt}ih.rn (G.ilf Co. st
region, and 9 days On the Atlhnti-" C .)ist, the ir tmainn 11 period \\'wa
devot--d to visiti;n various inland localities in peniFular Florida and
to mruseuirti studje'.: .it (:ainesvillh.
On all four trips ciflere t -i. rouite.s were ak'::n t through. (;l 'or.'ia. :Iarid
Somh aroia in urdter to s.te as much as possile- oi h af habitats
(if thle Hil.-id fi f ns 11nti some tri.pping was done .'[n r(Ilout Pirts
at IfJst oA i:}e r; p g('oS o if all t If 1 ;u}')spJ -c'ies w. I1re ti's visited d and .,(
Randd total of 92 days was spent in the area- of chief concernt- to the
pres.'i.nt study. Convestio na I .i 11 111 tu :.specimens vV('- n ('.cllcted dur-
ing the 19.50-51 trips, 1on th" 1961 trip the specinmns wert frozen in
a p,'rta~~d refri.-.ator- an d taken hack tob Hanover, N. H- for pre-para-
tlOil.
__ 10 11.
Oni this trip the, first live brcedin g stocks were colloctIi-td an d shipped
hack by air express All spein I!is ,I-tch'd in I 2 were shipped
back alive. For cuInsultatiun, in the fi-LId on all these trips, iaps of
soils, vegCtation. Piecstccen et sh)re line's.. geological leatnres, nd
coastal topograplhy we-re Ipr\ icsiv prepared in the map library of
Dartmouth college e iand plotted .ilt itrge sIa.Llc (o .nfltv Pmaps issued
bv the Florida State Road 1Departn,-'it. U.S Geo',loIic.. Survey' t'opo-
g:raphic' in:ps 1 :250,.000 sCaL'. a wevll as a rnunTr of th,. mIor
dletai. ed e ._I ), and the I- k-n8 intIte series 'r w r, also ised.
Museum studies A total of 12.54 specirmens was examirnid in the-
course of the syst4IIIatcLL par-t of this project. Most of this xzaterial
was obtained through loans I (see appendix ,i Two visits wrtn- madr
to- study the grc-at amIount of material in thi U.S. NatIOnal Mus-um,
X-Vashiington, 1. C. Standard taxonomic procedures haVe.- h IDten fol-
lowed in respect to size variation, in which rellin e must he placed
on me-asurement., ta-ken bvy collectors at the tincf of preparation.
Head-body measurelnents were obtained bv subtractm. tail length
from total length. Only ear length measurnen'its that were clearly
taken from the tip to the notch h.ve been used. Sku[l [easureient-s,


1968












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Special i.is were made to hol. d flat skins ini a manner ,r pern itt in_ a
pin-p iniifing ,f the c pt to bn measured, as well as to *L-xcltde external
light. Conventimonail iAh.-d muise ur s akins are not "eakiyv mea:su.'mred lby
this instrument. and no -la t skins of pe nrm dnri-;,s were availabi'. Two
Sets of met .ivreriin its w1enr take:l for ea.h spf-'cir'leI. -i one alon-ig the
mid-dorsa, stripe in the mid--hod'- region, the other lateral to it an.d
midway between the dorsal ripe aiid the outer marin of the Ior so-
latt-er'Il pi.,merit are a. Ii each 1a1S 10 in )dividual re.i diiJg.s were takenri
at cifFUr ,iit w- iv J I-jii4-gthiis an< i"iiI d(licE' !-iv rit) (.1 I, f 4, spe4 cial richro-
matic coeffici'nt ('mrpu tin form for ill 1ni nanit (" ( pro-vi ed with
1the isi 7 d n:F'n1 f I etermi in the l hree e r-ofipoliBE.tt ';aius-s, x, y and Y.
The Y value provides im index if refieetance_- at all wave lengcths, and
i.s therefore an indicator of rt.lative p.-i:tlT-i's. r.cLgfrdleSs 0f coilori. [The
x and k va] IL-I mi nst be p .! teoi on a c T.r )omiiti .ilv ia ialn to determine
the drm-iinxiant w;ave length ."iand the punitv of col.ir.
Tiw. prc'ess is .t l,-ingthy nri. wh icv'i hI-Is no t ;'et ,,erII cMOTmlp1Ited fr

all the 25(A) spi'ci,-ime.is, captured and IlaIboratory bred. Thf- vail1e Y
(refectance). however, i- L' I OW boni dter-mninIeCd for :t-l the speci-
mens and, for thelt pir-psr- or this report, about '20 specimens of each
wild-caught sulbsp eCies were picked ait rndom, their individual wave-
iiit.' i r ,iadn.,s ;i\er...a I_.d. ;id a vi ra, ,,'a it .. s thuis c.flt"rm irl.d for
dominant wvav:e l&i IIhl and punty (Fi. 25 ,
A de.sriptioIn of this i nftr~ment and its application to vs ter-ntih
studies h:Ias been pub.lislhd by S ;tlandcir 't al 1964 .
Breedinig .x~~ rime nt.: Slih-tly miore Ihan 2500 i were raised
in captivity. cbi.lIv to find possible traits that miafhi frin- ish clues to
relationships between the vLariUousi> 11bsplcit's ,. rnd so aid and supple-
ment the con.clusionls drawn frrr I xiot nic studies. Bre-din stocks
were obtained froTn the followingF localities:
1. Vicinilt ,f Pt'nrs.':o:,, F!.ri.i 1 Fig,. :; ..A .
2. GainrCesv]il-, .an 3. Vici_ nty of L[ -,' iai-N. Sp nTj s. Fi- ],r1i, Id Fig. 3, C .
4. Beachip.s of BaLdi.i.[i Co Ala-, n.,, iI:.:-l ! On i'[lnI .' Fi.. 3, E
5. B,'ach ,e'cti ,n lh t i ., P' -rdi,.o ,,nd P,",i,"S.-.i.,.. '-.' t"E ,.. I F
fR. *Soi. ta Ro, IB'l.-ii, ,. -'A r. Qid .lin-1 :'r iT l ] It .if.n i a.r t ,. '...i.V*,,'3rt t'Frid
SFig. ., C ).
7. Beach; sec'ti 'ri 11 h viw .'n C h,' -'..vla't t'ii'-c id St, A i.dre.w IB., i Fig ; Ti .
8 Vin.. itv of th.. Archbt i:i i.starr.l L bur.ti. v at Hi....rina, F ,,rid.. i F:' 3.
K2'.
Efforts to collect live speci.men.s fin, areas D, H, and J Fig. 3)
were unsuccessful.
Crosses were -made with more or ]e s si -C.( ss, between all these


1969














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of liedcI patt,. 'TI traits ot ''wlh.1it Nsiumilar undt! 1vol ving seven l-ci
has be-en c'onstru iced as -..VI koi kri h vp -thesis.
1Tlic mIll)p pitttfcm' trits 1h vt' hiL:-C'T pa.rtic'.iiltary' helph]l in tracing
gene fiovw anLd furishing clues go s'Lb specific reatiorishipS:. Were it
not' for the tract thai wo, (, diic'rmhit L-rifVtt1c's prrold(te) plit'v Ct vpEs
that are V.isuI 1i V i tin ui .Ii i -1 it Iii t I' p ..i l r to discuss
the tV"O 1. I uioar tlV '.L.s p:c l ere vi lh:u t'coui rs To i h.'se pr'w lip -;in; try
',il'tic rrsu tsi 1B it .ts ,th, Tyip )ft si.7l' sw st -ti peni ]T tCli C1.'O, 'iO'al-
irei it of tr;Mits rd ii s iti di.,S-.'v er,\ of thei presnc ,t- of tile factor for
"tapered nn.ip" inv ; 1in land popu Ixtii is c'irucii:ti1 to the 4volu ti Ti;Irv
disclIS.SiorI., t ils c rnpro T ii'(1L proc dt.111c is I :i'.(-ss.ir .

TA(X NOM1Y
Prior to Oscotod', : 1900 rt 'visi o of the t enis JX'rtmnii.ivmis. '.
po0iortiu, T .1 : 1n 'ie'".'Or(,IA fromrn A.:!t;i1rn a ilid n-orthw1 est lI'nirtda. Os-
goo d described 13 spEciriuens \V. E. Clvde Todd collected at Whit-
field. Walton C(7., Florida in 19(13 i.s a new si. i-.p ci's, P.. alifrons.
and 6 additional sp1c,'iivn.s A. H HI-. v- el collected aLt Hon Secour,
Baldwini- (no., Alabla, ma. i 190S aH ilnw.:.d hi iii t deftine the ringe of
this niiV' frrn ;sN "'.t.si of w ..S4 t'rn Florida Al; ba;i. .-" O() sg?(i dl'
m~ n1.IsC.-ript waS vid iiitl in pt-i.s. s W l 'ii le' C.'t.ivd l th ttr-i sprci..
Il is i .'vid hi.., foitnnie), arid cc nsid ra le i..~tl- now 'a\ exists .;s to
whetL er tit' tw': ,auiip ~ 'S 1(-Tl( ;.dv- f( .h-q .]iv omipar'd.
Dining the present study six M of theA original toIpot. pl series were
compa red w.itii : i i nhr .,f ,sperni ns frorn ti l.la hami.nia m'st. in-
c!(udinr 11 ovwel 4 ; I. S .')t r i', ii 111 i". TI li tW' pr)pui tilloS are
clearly not the sani. The Alk I.i,tin"' population lick t.he (diaun)s.ti
miediani ni os strict i.td is lism"!r1 .d ier no,.s -,riv":.r t t it topo-
typ'"L i albifr'ou t, It aklo -has ..i signific-nJitly I arg-r iind foot. Oth-er
populatioitton s hEic-rtofore inrlcld-d in the ian .g of ,al/.JfrTois .;Sc'h.wvart.
1954; Hail and Kclsonr, 19.59 also differ in various wa.'VS.
The beach ponpilatioin aTel.wEet.I P:i( lido B;a ,itid Pcrisac'ioula Bay,.
for instance, differs mark,.tdlv iii pehai e p.att.rn as w\ll aIs being p[lE'r.
lThe: popula.itions east of the (l'hoc,..\I\-hth.chet-e ti'-.r, which were
stuidie int.-'i-iv<.ly ;,~.nd r. Pist k. ',I]v idcti.ii t-'c.d a, 1,i if /r vn- }iv Sui n.-er
(1929, 1930 ), Blair ( 1944 ', ;nd Hivn- (I 1950I) (ifT'r in mcav rn-
speets from true llhifrons Incd,-d. ;.is wil! 1)', shown later., Ith'r
appears to bec a partial re'pr-od icti.- ba rtr- I 'tV tl -.'s and thd
true alb iftrm.n population. I.c-examination of the E-'aXC'inomy o-f all
the. (uIf (.COt Orm'. is liiis a necessary preiinTlinr to further 'stuc
of evolutionary changes withlin (he species.


196S Bc):WE N.












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!9, B)1E\,. C(.1. 4.! )OAST BEA<.:H \'11CI 17

Bov> ,1n SkullE an;: invws in ,Good condition, exc'Teptigi.L. a s'mail gap in
h.:e let t zyvgr-n.ati< Ir.ih.
uDia,_nishi: Pal(r and r..,sIr than 1 aifrlinm; :d ris&br.acatnu,
venter purr- white: hind foot faverflinflY f:.rger than in ,i:riseobracatus
and sirifi.ca ,ll tiid lar er than in 11 J r.-'J.i,; c',tT in tenr :ed1iate m in 'l t h
between th'l, tv ; I-'i<. 27:. Dicr:rphic i- rn'.pct to he.id pattern Fig.
2.0, B and C .and rP1.mp F'i. 5. P' aInd S)' Iisa-l-biir pattern as i.
i'111 5 .10 and 1 1 4. S it nli] 1 i no" o i -t u i.'cjphait.o, but ir tucl
tLrk.r a S)(m. iu-miil wav: i-'Onplt :1 up t o 1 c h ii th; m id- ,o rsa sItripe indisti;it Sin ti-
in .si/z to al:ifrons and ricorac cnius., but tail avera ging longer Iio




'1vpe T. in Spe mI i io"s,
,.ni' [ i s'1.., ] i
( i-pit b-n.i.- 22 21 .0- 2 '-4 2 30
B til.- r 17 4 1'7. 18.1 17 .'i
} .:1 pl i1 .2 7 ..1 ,' .1 ,.ti
Ew ii\ .' ',ira:iri 4 6 4...5- ,o 2 72
M ;I, tr" r 0..-T, v 1 2.7 *- .5 .11(
jil ~tR. bridih i ) ]I.f 1C24 11 B-
ir.i hrire, h 1 1.,3 l .2 -il 1'' ll ...1
I'ri r-, -,r;rb ai] ltcal s f .i.h, 4I u -1 '
(.: :<;ial iu t S ; ,.S.1 9.] ,.-51
Di.tri/ui.n: Cm0)afstaI] diurnes b.'twe.ccn MNobiler Baiy .nrid PTrdiid. B.ay;
a] sr On,. Islml, ;ifi the tni otlI of -"tr1d.ldo BIta,. Alaiham ma Fi"i. 3,, E .
'ftU/w o .f:f' .Alit'.n' ( I s;ind. b. t._,s ont' that ha rU.ts.

'. pflio (.->tu, fri'yl!f.'.i, new subspecies

iohltupec: UF 5521: adult, said bar east of Ptrdido Inlet i l'lonrida
Poiit '. Baldwin Countyv, Al)arIa, 22 (C)toblr, 1961, \V. W. How n.
S .ull and law\'s ini g.(ood c')onditi.ni
ia now .v: Pal-er than amm.orate.s and difiering in pelage pattern
Eri alrt fli.r si.;s -pHr'is. Sin-iilir to) o'bifron.v in color but much paler
i ,!. 25 i -' h li(LL!.i r'Hw 't-tiicr' \- !.11. hilin wv hii the r;ani) e i f len-
C(rephaus Fi. 7 v'tra do iLmaiit W\av' ].1 th 576.5; purity
-ae 1 4.6: rtlectance 2.3.5. Di inrplhic for head pa;t-terns I and I
Fig. 2)1 i and ru-np patterns S and S +-T sup'rimipo.sd Figs. 5 and
24'. Basal-h.iir patreri as in, figure 10. W see also Fig. 12. T 21);









1BI.3.,ETIN FI.(H)IDA STATE MNl lEUiNiM


tail-stripe absent; mid-dorsal stripe indistinct; venter pure white. Simi-
lar in size to anmmvbutcs but averaging slightly smaller Fig. 26 and


CANP.IA L.f I FASU IF.M ENTs


Oc i pito-..nasal
0Ba2ilar
P''st -palit;ia]
luisive frtramtic.ui
M'axil. tooth-row
ZygT.ma tn.: bread 3
C:,rniaLl b'readthF
i nt--r-orb:ital b:readt i
Craniat lie'lght


1 vp-.

17.0

4.9
3.2

102
4.0
8.6


Ten Specimenris.
range
20.6 23.2
16.0 17.9-i
7 5 8 f
3.9 4 9

910. Il .4
9.(i 10,9
:;.. .0
8.2 8 9


Distribution: Coastal dunes bet Ween Pcrdido Bay and Pensacola
Bay. Alabama and Florida (Fig. 3, F).
Etrymology: Tri L) three, syllepsis (C) put together, in allusion to
its postulated trih\brid origin.


P. polionotus allophrys new subspecies

Holotype: .F 8522; .2; adult, coastal dunes near Morrison Lake,
Walton County. Florida ( about 10 miles east o 1 ).stin 4 October,
1.962. W. W. Bowen. Skull and trunk skeleton in good condition.
Representative of morph 4 of this polymorphic race. Captured alive
as a young adult and laboratory bred for some time before being
killed at age 15+ months.
Paraiypc 1: oF 8523- adult, laboratory-bred from wild-caught
parents taken at same t imr and place as the holotype. Killed 3 April,
age 6 months. Skull and trunk skeleton in good condition. Repre-
SelTitlii v of n.i1 'o pIh 3.
ParaI-pe F.2: UF 8S524; ? adult, laboratory-bred from wild-caught
parents taken at same time and place as the holitype. Killed 1.8 April,
1964. ae S.5 months. Skull and trunk skeleton in good condition.
Representative of morph 2.
Paratype 3: UF 8525; ,- adult, captured alive at the type locality
in trap adjacent to tlIt wlhirh sec'iured tih liholotype ( sanme locality and
date). Kt-pt in laboratory for 18 months (infertile). Skull and trunk
skeleton in rgood condition. Representative of rnorph 1.
Diagnnsi.s': Polyrmorphic in respect to head and rump pattens, width


II1&arl
22. I1
16.8.5
7 97
4.49
8.00
1 .52
1 .20
3.89
&.48


Vol. 12











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cause of the intense whiteness of the hu.ach dsit.s. But- thr h-whi}i
ormsl show a cornidcriI., ar.lo'irftltf of vri;ttw.-o1 il cryptie paleness that
c.:nT11 11lot bl *\pl in .tI n the Ic.:is oi)t ,-It. How and isolation. a's was
t iou ht by I { ll3 ;I1J( SIII.IlitT.
~ tli useful itu i rlTIOni 011 il tihe ecology, behavior and p'pi latiori
d 'llI TI [( S o this species IaiS been pui hli e .. '( i!. ;-llid K.ii'- 1} ;,


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BPiLL..-TIN FL.ORIDA STATE N .11S i :i 1.


Hayne. 936, Rand and flit. 1942: Blair, 19.1, and others ), but bPe-
vond listings of probable predators, virtualiv nothing ts known about
predation anld mortality rates in this species. A ferna.e is capable of
protducin:g SO or more \youLng under laboratory conditions;. and litters
arr I.rn r -gula rlvy :t approximately .26 day intervals, so the atciiti t
rate mut: be very high. Blair I 194S) records the average life-span t4o
lC less than 5 mon Irths in P. nuni'flatus. and only f.ew live the full
potential of .3 or mnor, e years.
Saso na] luctuattiOrl ini populitioin 5170 ;ippr;ir. to he .great, as is
i i(ieaCc.id by thIe fllw viY ILi.)t. fun ltied .t v ,y r1-s N. Lav.yne, thien at
Cornel l.University: "We trapped two nights [between 27 March and
4 April on Sant.a Rosa, at the west end near the eritranTce to Fort
Pickens State Park. In 2..2 trap nights with live traps and 100( traps
with museum j)pecials, we took 27 potionotus-. They were really
sw,,anniigl in the area. A.i. ti rst we had te iiipression that they were-
iTOst abundant on t liC' Ld111 just behiTnd rthf' leach. but we later
took the.rmn in almcpst all the habitat types in the open sand areas. They
were even in the rather dense grTissy plot-s that conidined iinwas..
cilippi .s and burruws of Si2?lodnf. One of our party went out at
night with a htIeadllarip and found it quite easy to 0i)bserv. the beach
min:.e in their .Ictnvities. SI-me' were se('nI within 50 feet of a hom)e.
Tracks were verywhere, and hnre werrewere aiist no sea oat l,.eads
left. This v.as; iHn gr a.. cot- it-ast to w.viat I found in January when I
care through t ils an.ri-.. A.t that time flirre was a tremendous crop
of sa .'; O;t. I tr i-pped on lv one) ni jht thlet. ab .10U. tw.o Ilii -1 tast of
Fort Walton Beach. I took only one plionotu.s in 25 live traps and
saw relatively little si. D)nring periods of food scarcity starvation must take a large toil,
but at other seasons 1 pre'datilon is thl- most pot'n) t factt or coi mtrollfi-g
population size. The crn.uci.al (estion is whether, and to what extentt
predate ,ion is selective. The feral hunise cat is becoming an increasing
factor in the reduction of beach populations. DIu.riing the present field
work ( 1950) 1.961.) this became so apparent that in the later years we
avoided trapping wherever we founId cat tracks.
11 one strO tch of beach cat tracks were, aluiindai t for nearly a miler -
an either side of a lonely closed c-ottage. Not a single mouse track
o.r hole was d isenvered within the 2 rile stretch w\hih11 had hecomr
a virtual b irier to ',ne flow within the alreadVy shrliking rinit.e of
this c.los-1d p,11pu1atin. Not all tleh instances oer'rT,-d wvere as d(ras-
tic as this., and in on0e place mouse tracks and cat track. were seen
to converge on the entrance to a hole in a rotten log stranded on the


V.-.I. 12











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BO.\ EN. GL 'L COAST BEACI MICE(


mouse blocked from a hole will run for the nearest vegetative
cover, which is seldom far away :I igzs. I and 2'. hiding thus at the
base of a t.uft of grass or other vegetation, they are exceedingly diffi-
cult to see, as I have noted ii.mlli hinllls w' ictI di,11.'Hig their holes.
Complete elimination of pigment would undoubtedly destroy this
cryptic effect by nullifying coulnter-shading and the disruptive caNIu-
flage of a bicolor pattern.
In heucr-.paltal.s., where (dorsaln body pigm-inrit is reduced to less than
50 per cent of ie birth eitUiter-sha dini g cannot he efleTetive, and this
Dmust he particular so ii the occa.siona examples where pigment is
eliminated except for the rnid dorsal stripe ( Fi.. 1,. niirber 205.
Here a dazzlin g etF.ect might be produced by the white dlorsum in
contrast to the shaded venter ind pignented dorsal stripe. Such a
condition mrxay well produce a disruptive pattvei inider strong light,
and the su.lden change as the mouse reaches t'e shade of a clump
of grass would aid Im its conicealtrKi. This could be comparable to
insects that display 'tonspicuoi.I.s marki-gs in flight and cryptic pat-
terns wlven at rest. Thc fat ih -at te'sVe mic.C are nocturnil and usuallyx'
inactive on moonlit niiights Tiighit s_.-rJ to ic ga.te this hypothesis, but
a mouse teiTg chased fro-r its. hole by a digging predator in the day-
time o oon a moonlit ni-ght Tnight produce such a dazzling effect.
The dune habitats of ih1, taci iiu'.ee ,.i arc' rtot uniform; numi.Ierois
microhabitat diiferences (..cur within the range of a giv-en population.
In photographs A and B i: Fig. ). for example, which were taken frm.
the same spot in nearly opposite directions, the grassv foreground of
B is part of the same area shown in the lower left in A. But the (tne
formations and their vegetation are dissimilar. The dunes in the blow-
out area, B, are he!li together by the matted ,-roots of mwrubby oaks
(mostly uxfr.S Qrcu'i rala Small which make it virtually impossible
to dig out the holes. The dense micro .ungle abov the ground also
provides abundant food ( at certain .seasons at least) and excellent
cover which would insure safety from predatory attack.
In contrast, the frontLd (Idu -es, A, are easily d( .i arnd provide only
sparse cover for concealment. Food is moreIT1 plentiful aind varied here,
which probably accounts For the ,reai-r inunber of holes. On this
particular beach the allophrys population is quadrimorphic and pos-
sibly the microlmbitat differcitees help maintain the poul.orphic bal-
ance, but the number of specimens captured in the two areas was not
large enough to detect a difference.
Inland in scrub areas that were formerly leis ocenIe beaches, select
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Po ted I P--,, p.:'in'.ted -- .i 10 matin s. ave Il pointed (P -)t
S ,iqu r i S-- I' .4id'?i-i . *- 1 1i l.Hstil f .. ,v: 17 .ill.i -, ; _
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indistinguishable from sq iiar-c I ppSStt..
iSqur"t' I ppSStt 'I x tapered i pps-TT" 1 1. ,tiinii t, 1 ,.ou'b',- IIituLna 1_ ippSS! .
S.uterl ( ppS .t i p r at (' t t ;; ti .- i\pl :t;' i t f all ihalf-p. iir tI
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The results tof ('rUos.sts where tle, post-ulatci:l ge.'T (tl is presenIt are
givTii in i-ablh I In some other crros-es the rc-sult doit.) not confrnn to
the model. TI 'xpmlanatio'n of thli is not fo-..rthioming Prlalips aii.tlICr
g(CR. Ilin].st h temporal flhi.tiitionis ll ,pigentic inter:tction during molt. is will be
diL:cs:sed later. These ,- iot n.nforming results .tre.' 1 fullmows-
pt inft.d .r Ltjl"'e 1.'" O,. lati .' i l-p-int,.._ .., 12 s u.r. 2 j.iLLk-
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to .15 d -nhIe irn-i,. r- -. T .tp rI .
Nottwvithit;tndridn these une-vpLL ined resu lts, whir.'lI 11y:. i idicate
that the proposed iodel should Ile modified. the results of t- hrc be'l-
i-. experiment[ dselemostrate the valuOe of such experi'mcnrs as a m'ails
of .i-Lt-'Ctci m hidden traits in the vainous popult1.ti.rs. T he tap 'retl-
ruInp p1) n.t.. ype, or -'x.t.Inpl-, is, fi:Ond 1)only IoIg tile prc. SIt coa..st-
line i: Fi. 5, Ti and the- systematic studio s S iugcstd that it ,a- ,,
probably of recent mutiatlion;i origin. Buf a mated pair of aib ijror,
from 1),:. FViniak Sprinis produced 2 double iimadge rimps in 12 .si,
and as the arguments against gene flow inland from the coast (sf .
a:..bo-c .are applicai-,e here We inlust CO.-FiClude thla- enet- T i's o i mni
older or in and is possibly present throu.ghcmut the species,. Ih pe-
-1ibar geoIfraph ic interpolation of a lop. hrq,s (pointt1d anrd s5quar t
1n1]ips. 1 ctw-enlI h'ucu ( p/ralu.s t [ap.ereld rn.-p d:and pt.ni?.,.lari.
Ssdqu.tare and tapered ps11111) is also eptlin il tI eX C t riX it-ts
1,v the high friequ icy of P in tHI allopthr!rs popIlation.


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    th' (-ise' with ih 1. hu liti-S iin thlt h.dy ,irc -. a'S well ( !ig. 15 iT,,ht .
    A fi t''r zo ei o v worth \ pOi ; 1t is t 1h .t i-t ,. lles (c r t ~1', 1 li.l' 7tWt C'

    ]- _ii]~ ( .. li li I I ".' ..(.-. . i.....r" .. ."i,- lr t '. .. . .. .. L P 1. ,T.f1 '
    bc.vonrI (he midriff / diL" i tti inti) itih li;: l ;i wl hf-r t h, fat ult 1 i.rW-s
    occIr. In thr ntrk z7one pattern \ ,cLppt:'. -.r a -t sI spric. t is .S; i-n h1 t
    i-ntri tile sapul.ar 7i onrr thi, riiL'ht side-l. X ier it ni 't-'t, tlic' V pitter]i
    lat .rlf t. ihe L rT .Lu-, f .ine, Tiir.s.
    The pr.'.sf.'rnc cif pattern \ in Ithis stock c iact Il.ib t t .li pt.'1i& f 1u%.( i l ,-i!v (! I <,t fvro ni tic. adrN-a \q,.1th1 w ,_i t ,,
    PFill., I :.:hI li' .r *:i. -.':, .rr-i )O ury 5il d hi- I.r I t- sd ed rlit';i !'ii : i t i-illift. s
    t ri'.s /1',cp.sis front ) the ii1.,ii nL l Ip I f- :ri1tu,. p ) pi1]:ithi n ,r Ii r.lv a -}i Pi b !
    1SS 2 ( see list of miap p. .:.1 ind it is inti'r-sting to find evi den- ce
    of the finr erirg effects of this gene -1vchange.
    "11he appearance J such overlnipIs Ai-t- ween) f.,r-i' ?in ;l-id ,1ot her
    ca.l t-f S rt- t miii r in u i)I thl'r .skiin.s, )iut de iiorist r, tirij it phfotogra.lphi-
    ca lyv .s inot .asy. ITI spe(intWi BPR 517 1Fi 14.1 the whiit'-fipped
    hairs w' ithtliil tle V patternL a rt weaikly develIp'ed i.e1 tl h- s'capi.lar and
    Ini lriff yonrs, .h.it hair-developrnent is (stroi g. i1 t h :tre of ( overlap
    and sjlio S up is hvo nrrtu (cistiritIrly wV'it'" p.t(t'ic o! ithl ('r sidce 4f
    the hair tip pi. giielnt patt.-rn.
    't'he a )otve 4- ife all rathileTr .\tir-cf < it' .:MiNt, tl w,,111: d pro1Utllv no1 t strvivi uiinder
    ullhih r;i (',OPl it Cl IJut1 i lhauiilt'S f la It lii]s iAnid it n lil itil pIi t ila g-'
    li.rTildiltics 'AT]ri c f1,ind c i \ti V l. iid c 1 'lh i ', -spec'ially in the areas
    wih.er l,." 1 -.1!il 's if .s't' JJ,.ri ';v ;iidl r r"0,iieli apprnite.h onif, another.
    Sp'j i' i ii 1.;i tei i t]iL (.-) cal N ati I t al i 1"tFOr t, fo5- eit, iliple, hre
    ,q iiuentlnv si ," fr:ii 1l] I Ic i, .IG. S 237 and in th- a"r'C':i to tilt so uth
    V, *,cre the ra':. '.s c .', Ii:,". rr /1?i t7 dd rh : coi'-;t h (S urt
    t'CVC t li r11 YC f iT'f I ieti)..
    In t l .ds aLirc'ai icn-. rt T ia .r I. p I .puia I.l i KL3 i Fi .3 int er.n rtf.'ds
    w ith f t f I Z if j .'f ,t tl:. f j rt I.() do' 11 u1 1llill:1u i t'utti''', .ir"T 'xPhii d.
    inste-,d olf :1 -1 i' d fl Ivpr of iittc r:idI;.itli ii. os js ,s. Nallt h 1. 'i-e t.-se
    where tets., ltl):.s f i -.. : l-hte. ih mie ie ller- lernd I 1 1- .:' -iher wvhite-
    l},! lie'd ,'r ,ra l:\ e!ii.d. aLLti t le' prciportions 1 0 I" ITILT (. fr Tin 11T irt ti So lluth.
    ',1 h.. .v S t ; ppc ;ir f?- a tore or I e's.s clear crll \-greg .. t "_., .. rid
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    BULLLTIN L lt'F i [l1).1\ ,TAT- \IRI-, F M ,


    correspo ids I to the -sim ilar i-v aIgd st. shown hv C( ; l"v <.'f al. H, r'ad
    rn1 '" Ult'nc'S I!Va v 1" t IeI- WLJ w' rked out ill t hut ii. I, i irIl t it
    t.'o 1110Kt nr4 ion) 0C'cijlt vand rnose: iindic.le d 10v C-lins I tlM')
    C'orresp.llnd to sn .ii t reas-i i ifdnied i tI he preset if study A tiitrd
    f4<-14' p ri i. Imid.Wlav l 't.v(.'c]T lrild 'Ii '4-hiv v.en"ltral to th- -'v.s a.-nd
    "-rs ( Fi'. 17. 1 ) , 3Sl rt-.'i.;iill7.e(-l. 'h .is a1pp .lirs1 o orre.sp onS d
    to an a.i rrl'w sheONwI- in fi u1 i re n (.'tliri- .. I.

    Dr ilo-AH.4 AT. I [A. T--TP FP i AG L. N'r S PilESSION
    JrIn oF'_el"tJioli :.ipt i; fully pigni-ntl In others this pi.ineit s'uppoi 'Y.,rn. is limil.ed to
    i" (.Of morr f o tf i th :) ', o "s [il. I g. I .- spc-,il 2'- 2. 2-95 mni
    169, a< dkl 1i2 (. 1.9 At fir.t this ;.ibs' kei oC f ,1 fli.r O i rlump phi t)p .
    wa.s thIutight to signify rc<'ssiveness. Mial- 20, vilt no drlteral
    hair-tip pi (T I '..t, W; fri irs mated to ft hill pigi t int,-d fem--ale .
    nd(i a 1 ith- of six x' aS OI tai!-led, twOV Of wiich had suppr.fi .d p Tiil.nt
    JIN tii posterior Iodv 2iC. i ,29 Thi' rn.i wA's le' litit'd t1~
    a feliale-i ,Tbifr.iS (Fif. I p cimen S) ith pointed rLuINp. -11 -_CI
    fi e' ~Ci.ir'rt-n Iiip I le. F W v crt, -i bt ineCd. 1bii; -. t he e.\p - cross O.f poillnt-d N t.-pere. arid had th ti ial i t Cen recessVi'e all tl.-
    Fi shoii hive had po ii 1ted runIp. ()ne of the Fl was tir i ticl-
    r s.-ts.d o h 'r t, lia r. t"il vi litter'. wcre- .orlt, ilit C'11" oi 4' iof foIT r .1
    1rvi\ .-:., ',,. It i.d .1 a lf-p-imiled! i-uInp 1 ,371 ), tvwo were sq uare
    rili-prd ;in.i one was ,,'..i. 1 < il>!- iliin.. p li', Wotitrp i. 1 )80 )
    A. 110 O ti] Slic': lis t rill 1ing-. \vere act'- i."h \'.d. [ict coIli 'is;151 1 1. i..E
    I. dJ aTir. 1-Iow-v--\T th v\,steliC-, of t i e i11idividuiald Vith r.diIcc-'d
    pi-ilif'I1 Is 1 \% l 5 Lr t 0.1 .S .il t.d tr"( r(Mii",it"i ,, t I 1-1 i\ a' i 'p SOfj'C
    survi V..A I dvO miL i t, 4.'. iT i'n;ltio r d o h !- Li- profiles ;.?'ii ft--ded si kins
    1Fig. 19, A i shows that hah- lip pgin iiilt i si upprg ss ICl or diilute1d
    latlht'I. th;ii 1 3 iri. \V t1l liiillatcd 1 i i lt'st' ('as Ps.

    1tA1H-TIP H1-r.D, P:AYnTiH

    The Iahiral hlaid pat-cierI-s of tie various st1_1,IpJc- .s :.. Figs. 20 ITi1d
    1 tir1 rioi nii characters, huit are c')inp owr Tf v-1ri.,i, ('4 'ii, i1na~i otc
    i.d in tcac. iln" i i.I. s_'vlnra] iit pattl ern ic -.nts Tthis c,' be c i'dr!
    ,c. t: i i 6 cGiie" 21. wrl-'r" ItI coii'bities C aiid (; I cc trinfi es (C7 F -and
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    ("l^I(ert,, Wh .1ic] 1-' S'JS Ih ) iin Fig. 22. T 1h v rn)i,\ _' l.tii,,is ..{
    troni t1 he- (i.iir.al patlternis Lv tlith- l'ttering AE. BE. CE, e-tc. A and AE
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    width ([ig. 7.. (he palest population ( b-'ucfcephaiu : having als the
    narrowest pigment. There are also some indi..aItion.s (see aboveJr that
    dofr.a pi.r'Ti-rlt width rnaxV 1b' -D ,. .prtss1"ioNi of tbe an ui es cn.S, dliolt
    the three Trmp pattern traits, and,1 so' reflectane I(p-ens Cay Aso
    1be r-.dwer l.hte -n]Li e uen.r" control
    It, the Cas of allophrb t.i. where- pi1m. nt width :n11 id rc llect.ic'c
    measCrer1mnts dTire ^iVIi jl s(pIrtitIly for Ihii f'our ?ii.qrplis. ci possibl corrC1latxoni with crtaitl hea.id p.l .lern elements k ..aso seen. Norpih 1,
    with thc '. W it .st ;trI dtrk l 1 .t , pi)i.,lle .it. pattern B (Fig. 2().. wher0 as norph 4 las I' ,,d pattrrn F. Morlphs 2
    arid I are (-haJ.iracterized by' h..-d pattern s 1) iand E respect ivt v. bit ias
    tr.Iac:s of leil it ILE Fig. .22 i re oaen frouInd in t hese f.orphswi. their
    vi.terrnmediacy if. 1 p liiIi it wvidt ialic palei.St-.-. r 1-. dua.- to ti us.
    1r7t1e r 1 1' to .'fO .ll 'C IrCl aticAA 1v it01 tCII l i lf.lt s DE Itnd 1F1.
    The appa.n rlt tint' Siiri.le--r :192 91 rec.-ord.s in irm..-ru .sii !ilt,: lit,\
    iiid 'xt-. itel ity r i )f L '~it i .riiid flaii itr: t 'i -.t i 1,.,. i ),lt" _} Hi 'L rC-_I'd-1
    fcpr two rT',-n1 ll. I Bns ( ll- clt1iil 'I at iODns i i-lid(, thr'< .siu})>pecie'.,
    al ipirs usI ( shore), .i n' e. ri. ( CrY staI I.ake f, arid Ili ,in ctu.. i (7 hiplv l.
    'he B found Lake stat ion is within the art-.a of i terf rada t1 ion according r
    to n.:atenar studied 'ere.', aiid S1 SinnrPT had .oilv o) It sanfiple Sot c Ich
    sui. -S;pecit 2 U thI rlativ-'e proportions of the nin. morphs witli1n e.Ich
    sam jpl4 is; Hiii l v1 a.in i l (ind.']3 di; fi e"I i. e. 'si C 1.id ".s v aIv f t c _" i"'
    ret; lts.
    H;t\rTl ( 195; ,idtdid atot lir colltctilg stlat ion ( Sr'moniri 'l Hil,
    in his r"-.stuidy of the slibject. but his sample from thero w.s -1l-',r
    in color to that from Round LTake than to there Crvstal Lake s,.amprII-.
    ''he ri-ason for this was; probably due to differeiice- in the conmpos.iton
    of 1ne IWt o Sd riples. N1 or-ph .3 of .vl/ir l.nr'ii has ;at A.v r e purity vali.n'
    of little. oIvtr 15 ,I,* a ul st 20..5 f or molrph I in the- i- s- lCr iem its
    taken her.' ( Fig. 2S .. A larger pr portion of one or thf:e otither i'morph in
    oil-hr sarnpir- cn-utI this eaiiv effect tihe readiutgi for red, orcc.r, -rId
    L:ldh -\violet ill li i r-.si.ilts
    No (1 colr ai- lvsc. hi< -, t bfl .. I d i -. ietl er thai;'n the ob.> i-tnifin- of
    ave-ractiT-.e r 1-',i1 sampit.- r",adinlNs a sh own iLn fi1f urIE 25-, In one) or
    two inlst.l it' 'u'hert itnice of strikingly (ii fIler lcnit cr o,; to tlan (yeV )
    wOre mairl-, Hte _',suits ha ve b.cii exa';T.m ned cas1aiIv. Thes- appear to
    relatIe o di erten.cs i-n purity rather t ia.'i in dCorilrnii.t _lvvw' lon(th.
    lHure thle qieust.ion m1:i I.'. e r)C.i',sedi as to 'Iwhlether the punriLt- 'arlintgs
    represent actual diL' ir.nc'-' in 1( color saturation or are dIu,-, to the
    relative number of bllak-tipped hairs dispersed within thi ;- are of
    co'lor-it pper.d inairs. W do in. 1t yet know wlhet-her these Iblac'k-tipped


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    BOW\EN. GULF COASTT BEACH MICE


    19bS


    The surprisingly high frqiiu-cT'y of mIlorph 2, in the .sumneri popula-
    Lion ,( 6()"- -. rn'iv th-S be the resnl! of selec'ti v f-tv 1onrmg, a1< it.s m1an
    coCIicides.. with the prT'sTirned averiaget' substrate color, whereas os iv
    the 1 lower pa.rt of the ran-e of the morph I group i.s thus f vored. Tihe
    alirr-ow anld ri'laliv.iv iCOnpICtIO-u.s white nose stripe ,_if m-orirph 2 ',Ti
    h.irdiv lh of much selective disacdi vanLtge In norph .3 selection shou ld
    (avor [ {t' iJlor vr 'i.l y o -)_.2.- iTKI \ 1i4vi Ias, )but h.re t' _' Wi it t ilose .;I1(:
    m1O" re t ric ted hodI pi Imnit It In a-lso it disadvaiUaiIt _us. lThe
    ('o.):r ratn he 1f morph 1 prLSun1 madv falls below ih S!cti- rant. ;
    in the ii a iad population arid. in alT'v stric ted body pigni:ent would act adversely Thus I-,ay )e explained
    the, als.C C(ce ? moE)rph h4 f 1 '-ad it b.i 'e'it prtS_- sent (,nci i ly II ; ,tI Id. tIh- low
    frequency of riorph] 3 in .u/l ct 1ri.
    In p ririty o Co0o i 411 r phs 2 antil 3 f ,Au.unicii ,'Tr,.v d Iot'r i:ead-
    in-s than t!h' i uf alfc! iry Thib s .u ,t. p.'riI;ps.. th;'i0 slc.ti1
    acts in f.vor ,Ti lo puritin' it t I drth arker iiiti ;td s ,Ii tl .itrt, bit ill tI}
    c .LIS; f r'iOrph I the 1 siltiirn is. r.vIV"'ers0'd. PerhaIps .-orTff c>orrte'i.tiOn
    ex itS et)ween do11 iniiia t v< -ieti:g Ii and p1r11it', il which case it
    ,i t il. :O-tsirn ti i, LL t h 1- t e 'tw ir ,- in.st tl it. InI re I'r.(I ,v-- :1i1it..Q
    in ivd u I i f m -.rp 1 1 atit,.iial: I d.111[ i ..niinat Li. th.'JosCe w'itih hi cher
    pi ri(t '.
    The,- approxi tia- _i. 11 fr ij;e1i"p ui-- s of ti -1 inur iorphls Iii he '. k' -
    phjrqf.s ill .'-e i pr ip !. ti 'i, i V, riv s urp i '-,li in \'.i<"\ ("f t t,.- r titl ,r ,I.i ,-tI
    : t t .errl cl i e re i 0'_. 1 >)etvt-e:Ii theiii. F '1i. Vli.itt 0_ f 1 -.)rp4 i 4 a i it !- i te
    white nV,. iS' o)f rY1or phi .3 ms m 51t rely I O c\'pt adit. i Y: ;is 'l-'-
    Jgestcd earlier. AlSo .iC I." l:.trrA-,w r dlo0US[L1 )od p ig ent. i i1 7. ;rtd
    ..lui rt. 1 ir-ip.s I f tli("iC' t,'-t 1114 1'rplis si, i]dt 'i 1 ." I '' It I .- fIV a Ori] tL, but
    the itwo 0Iie -xteiisively pit.il-nt.cd i i[)rp}is pre. -'dontLiiatted I y 5 per
    C('o lt. Th 'rl e fif 1it-r Nn-t. 111:lit v 11 if ilperi r fihIu'ss t lr.t ,cirit,'ra"tI thImir
    CTr',',i) l is d\ l i( t.4 a -, is ,i.ist .1 t LS < (;.l..i Iftd -W1it i. Li f-'se irc (.'xttr si' v '

    The absence o the L, pti-ed frumai cliaacter i, the s,,llJphry,'S popu-
    L :tii dl its prh'sen<":"' in the b -' 'l popiIons n 1 ither side c an
    be cxplii-e'd b'y [the iresejne of til'- -enl-' :u po-itc'Cd runmp in that
    racE'. ThCe thtt f(-r pu)..iile.d ruip .o ive a square rump plhe)notype. The prc.t.n-'
    of the tapered rtiLp i) g.One i al1l)p.r.1s has h )t.i'n dt.'t 'ctt d i ti ,' labtiur;i-
    torn' v xpervitn.lk IIn tvit_ pe 'in.SU.llri. population ,0vc ,\e tlwo isorj v hs,
    one charac terized rlc tapered rumnp and white head as in Iecruepha-
    lu.s, the- othe-'r square rump.'ipd and with head pattern J ( Fig. 21). No
    indic.atimonNs oi interaction of the tw.Vo nimp pattern 'TnTes to give the







    BULLETIN FTE.-I1.\ STA'IE \1i.>KIM


    "dol, t e intu "t patr. ,, a1s in trS it p.s-iS have lie fL f 111un,1 (.11--
    fortu- i t' IV t -ffori'; to se'r-'i.r :1r b ed1iingi sIoc k._ of this su1! fsp unsl C'sf. l. d

    O.EN] OF PA'rTr-N T'I"RAITS

    Sukninr ( 192.S) dri--sri ]ed a niu tai t form of P. 7mnaicr-ihdtu. in
    whi ch 1 a "gri7.7cd" and occasional a nmost w hiFt, pelage appears
    usu.11.v aVf t-r fi.: : i e Co 6 mnI(rnths or more, As yvoUmn adults th,-sc' mieC
    .are nl.Im-a] in color a;IdI pttern and in some eases the ;ri77lin',i i tpappr ar';
    ais iealI as age 2 I uoiIths. He' classified them into right alTritrary,
    grades and remarkt-d t hat "th e condit ioi .i t le nll rt p f utinLOd o-f
    th .sr .rizzled liiC't-, iI!f thi.v P 1tlitirul'shl a seri: c 's,, beais T u 1(.lsidf.i .b111 C
    rcI-'Sni-a]irF-e tO- I L ef O(rtain. 1niiX( -.li]d; of a F lorida ace., P. plion".0-
    tu.'y albifrom.'s in whicni the dorsal surface of the snout lind thi top :of
    the he-ad, as far back as th1 eves are whitete' He aS r:-'ring., .:f
    course, to aflophr .?T 1 morph 4. not tl/hifro-.., but tht, sihliilaritv does
    'xist.
    'l'}r' possil.ility. of hion..Tl0 y i -btw,"en the two wa.'; intlil xiit bv-y
    the d(ieIav-d .] pp-a..irnr of. the riz7d] traitss, wlir hiv ar i-n:ji:o .fst
    from the ? ine thlat hair starts to gro'v in F. pj:.iifjtu.i aiU. plophr./. This
    c'oncli1on v, WN acLcept-:'d when the pr'-,t,! sti dyv was bciin,, started,
    ar]d Sumnr,-.'s ri Liz'd n-,Ct. t was piit .asidc' as irrTi.'V.v.rit. I[aetr Wv1en
    ,1n et' n it:'i -'v p.:tk- rllit:aL t fori n FI,,. 24, C) :ppe'arud. its imI]ilar.iy to
    Snirncer's grade .S \X; i -i t; r lh'd. PB, tlu.s; thri tO vc ,' v'rIl tt'ad pattern
    iil ci e*ilt'it in thle( I I <"ah mice had be:'tn S tdi,"d ifn SOI t d(et"'il. Ot
    rc-r calinig, SuiiktiAer's p-aper thie U-xtraordin :iLry sijilaritit.s iit dil.tails of
    hliat p,-itriIi. as shown in his} photograph. 'u11.'LestV::d muchl mTor{ tl.an
    ; si lpcrhcTia' r,:-.vlitlbbinl,- iC',tC is.ld i A' 1 Lg.nt ralize, typc otf pi.ri.nl,,t
    re strict on
    .S i*mn'i *I T grad, r .'. 2r' l,' rl.'rs . f',xtret'r diltnr- that ti- O (w, r-.in he' diit in. ish-d only by hv si and
    bodv proportions, but rranv of thet herd pattri unIf.it e(ITlrments found
    ijl P. p,:'Oionatu, cvan bhe distinanished in his photograph of the grizzled
    1w .'U I :ihtu f- s track. RWeC-'ipt of a skin of :a mrwnn icuats mutant r.sv n-
    l) lig Su ini .r'rs "grizzlt'd" hn F R. H1-1 'stis. I i. ersit v o. ( r< t.n,
    LIs furtnr stir-iegth ,n"'1d tl. grv.Winl 'cinvi.'tio n that l.],. ,LDil dg1i\'s
    i id.'od i'.xist. de';pit" thte d ,lav.'d ;appe'airane ,f ti let. 1. piJ)tttrl
    p i'not.tp- 's in ,.otn Sinn-iiir's a.rid TIi stis',,\ lItu aa ts. ',i? apparel tIlv
    rcc.ssiv-'c1t- V I' for the extreme dilute form appearin int mn stock may-
    hIe widespread .iI the p.irmnaf.tNs species, as is suIi- te'd v l.Di.' s


    V\ ,i 1:.








    ] ,.f;4,


    ( 1934.y rc,)o.Lld of 1a 3I:aiLLkev 'd whliti .p. polki t.' us frorin ( -,. .r-ia.
    His de5s.ription fits. rxct-pt that it makes no mention 'f gr.i" 't isal
    Ilir. r b t this is o4fit n 1s dIiiltC- .< to }-- aisilyv ov r.look.d. Alsso the
    fact thit sp 1 cielli is prN:'scTrv'Id III, alohIOI may have1 Jude it hard
    to) .'c .
    LExarmird iu' nd.r ;i l'-n s sp:'--i .m i::n ii ii tLer 2 in Stirnri '-r' ph oto.- i-ph
    sThow clearly a mn saic i11 Itertici( o;f tlle thI, .lrN f or 1iit .ea. n iits BE
    . I.ni (E. not I.nlike ev.iimphes to b.e fiounr.ln in p lri otu ts speIIt llS ,;
    ( l" i~.2.2., to(p row ). \'TI [1,in1 -tri" pirgn i'] t it l..'w t}'ic e ves is
    sh1ioT.. Spec-fin:-l rILinbt-er .3 in Sumntr's photograph r"s"..miIl-s .-ry
    Closely thel he:d patterns of A I, 1(04 aln A I J. 1924 : Fii. 2'3 Thr
    r'pr:eient the interactionts ,ft uni[ elthriiInts DE. FE and FE, )but the
    la th (ulatu.' sp c.i:in-rn .. kL k 't t '-h wit- Slp.)t.b :.Ltroiu 1d the e. ar .C possibly
    bi-CaIUS' oIf Ie prfiese 1ce cd AE.. S111.iflicr's specimen numri h-r suil ge sts
    th-i prs1 nc11C ff the g)i.'le for s u r Ni- 3n N1111p, it aso 1s t-'rs 3t <.ron :'-

    fit' pi) turiod r 4)B,.dv r.cgiui oi some ) l ;' Cfph f .\ sp,.'Cin ts 1 4'11 I. "!- ..
    niimbe r 2.9. In the-' lt ttr case p' -rhap the square riinp rn.iy v 'in'
    to tht- )bIlaX -Ilair pIltlteil ,\"U Wti.i t ;I aih r thai4 tO the gen. tor hair-
    hip StJUar-F r ilip q'i r -Yth w 1 t .S-t-S ] t ClerIC S i--i'I1. ?.iLri' 11 11 ip-
    p -r...S l N' I a I ] hit i n 101 *1 'hr
    t -it these r!e-i blalr e SiIlit-: s :r- vit .tled, E? v Ic I a.e exi lil i;.Mil rio
    Ii ; TillsIlln h s-riC-S of /"F 'ri'f/.'CU. sti (dv skins sP owvd sIeIerai in?.st an r 's
    of li;.-ad pauttetr trtlist ci taractel istic of i ttin ttus lY ii nni-t i -n lt'
    I ,Hil 'f i U T N.s ',t'Sris. .liiit not i 11 l tr sJi 'CI s i F i. 29 A rhitr
    fIn rt-la, :l ,p.t ."or i,,/.. p rli;i -I ; Io.IIp:artbl, with the \v ite foreloc-k
    in man, Vwas also found in }'ot h .mak),Ci 'ilatus aid iti ccotnS, tnd this hias
    inow appt.:p't 'd in t]hej p.-. i of ti s i;.al-.rittorx s ock. T uis mnai w, if liot
    all the pattie'rr gen: char.n 'ctEri 'ing1 the" b"eac'h mice may', have been
    alre.1.rJy pres.'si.t in the P. nw O. ir lats5 stock forn which 1. poli ro ttiu
    t \'V' [1(d. Bilt l fIw lit tf is th.' h -ltk .d.l appLiif'LIt o' tihe ri,!,/lt*t
    co'ldifl^l to N4 p'"-x i'
    Su n Fml r "LO aird, tIhi iC iti on1 As li .in t the r lt 'f ~t" rs tl.t
    prd'ld.i t hr ,I '.i,.. ,f'tf t t it, ii latr lift E.- If wv. e I- th1 e I -Ce eq
    for hiVs S(''C Lril-d .ed ,ades ,a IiI act the s',- i patten4 "Iienes tas
    iII t b.acl.h'I. rnilC'.'. ti '.1 tl L t i i., Ii )F ( prjI t lnit Eric S eC in the ;i r ly
    sft;t 'S: (if i '. Thu,;s P mn i.ri 'u, at.f. tfs t 1 pftM sc.s- :sn ,:' Ian r p .rl, ps
    Ititi1.a oli t. tlat af r t pti ln.'It Wj1r c.l ti).'.m il el )c.'e .rid ing liolt a ci
    pre 'cv fit, li-It.. pa-lt-.rrt ...'-els app,_'arnT ill t :arv lif<. A .ll so s ofF v'mi's
    SI nI i tI Us bN 1iidd, '1 aL p4 U [ asd 011 wit] .0.?it r'v, 'c 1in-r their i Ic t


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    P-\'s B(.)\ t[. '. < .\.T E.\(H \11[. F 71

    Tiflespan to trst o t ioAn cC ]ha;flir nvioiiunT rier is ]'- .1Tl-v,:r hiddi ji l.i.-s
    tiev rliv hi pp1 to ( iE 1'0 1I"T'V. woi ., be eli -in. ite. d. of c i rse, buti
    iI n 11l I I,.i I 'ir ft I t I t.f't4. t A. .1 l 1 J. 11."W o d .. sti l '. i (o itLCLt w ith!
    their hab-itt. somIe' i,, it Nreveal traits beneficial to an Iv.;asioni of
    the ne"w '1nviri'Ohinrcnitt.
    As tir. i hidden traits MVppt7ar ilh(vwi re f ro1) 2 6o 13 i ntlhs or 0Miore-
    after birdl, MC:'II individuals v.ou Lld. till Ib capal.-ic: of rimuch reproduc-
    tioIn ;and their offspriig Xv,.OI3i i.have selective favor. Also sel',Ltionm
    ,w ntu(l f .aor the earlie-st revcl.ctici of the hidd I .n benef-i.'r il raits.
    Eren tWa l1 this couIld i ad to :liiiiiulaticin of tho delayi i factor, as
    a.pp :rii-., to he the e,:.St in the Ip-rs knt-day ba:-;h mice. Instead of
    l;a"iIl i t'o p1 st)i,;-it :1 series Of IIUtLUdi Sl. occurring .t di t-if'rnt ltin-.S
    on diier--nt Iees, v. ( .Y \bi,:-1FV t-Ci pr.t- .- s t- s o l b c h I 'ln Se s )
    SpI'iit inT1. uS the 1 .r niJt 'iI ccli iitiult dtioit of VJic i iNcs 011ii -)21 tl trills If
    Fpr'vio 1 hi .d I tr i.its t aI ( diff ren t r ic ,clies, issola ted yv tim.) ;1u(
    spAcr Some of thfc t in4it I- w t.'1eficial. others less so, Jand stil
    others .imful. S election ouId di-t LrT i th eir fi t- iiid the I 4of-iirse
    of fu rt her diffter utiation.


    ]V( I I. r'TI J.) NA HY SS E '- N rC'ES
    Thrl pro a ,illit rit: i? F jli..rt ,u.s evoy \-ed nut of a P mlnii ltni
    stock is hardly to be qucstionEid. (Os).gid I, 99', pointed out the.
    close similaritv bet-ween P.p. policatfi lnuS, ad P. wnanictL2atu a palIe c rcns
    of T'ixas. and partly -ecae of this B:ir ( 1950) pCos late :a orri inr
    Pof access along th]-e Guii C oas. "Ai ec.logicail .rrier now stp.;-irates
    dtic tw ('o: eo .- Ti .s l :-~l iT l nds nlo-W o lp inig fil eh -O,;istaY plain (.t.;ti rd nrom
    eatrn .erx ," Ari' av,,id(. d l. V bo1th s peci.t-s." Cranting tl it this IIea.
    be so- 111 Te11 s d 11 1 A l ,-isi:ina, the co ITip 'le a.1bsence of r-*ords of
    oCCTIrri It' o P. io ionv ftu uiA II Ih- appIr1 t id I (pI utt *ili itS parvt of
    Ala m., a (. west of \I )ile I:.\b' and the .Al, i; a ma jiv r (' t uc a t w t h cx-
    plaini.,d I:,v thiis hvpotlr.ess.
    The ."sieri limit" of the ra g. of P. polionof US s c 1 ear, v dcfiz d iC
    t-le. h.;.irrihr fortied Iy 'V .lr h-sii aid poo)rlv diraiiricd soils ,ordering
    the A lPabai.i liver Fig. :', and teiwsc extnd northward t. j, in, the
    .stik-v soils of the "i:tL-k: itIt" of central Ala hama,. IlLad the species
    0.0 olv .d fr'11 an1, cst wrd x.tens ita of a.iunitfit is sto iOf TePs, the
    siitibiue f1 .Iitats in wt 'str' Alb,.-ma ,would si.i' Iv ht e ccI 'clupied to dat .
    Aa not her poiit of' iMnuigretion T-anst therefore .v soi'h, lor iralmv from
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    t t,-' rI.C- f 7( hini, di;s fi eirt-id I t I\'.,Ti hr i \l tIh,- 1 ','i] Itf t,. O k -frno-
    kece Ohl.rc- .ine., wh rerta' ciell'rle v it. rnil I.-' ) T ,'i" d to thi t hii is
    <.i)r iiVol tve h t iie ii'

    l- eI ii s O the 1icni n 'io't: h ]H :' wi S1om'e 50 leet ldf-l'w Il ie
    O~l~h-nk!' .shore( liii> ibe sira"tndvd Ve :itit rifler of the titririT I N'st

    1 ( o f*lati ]s (" l ( cllf.i 1i t It)\ I -:.1us 1. t I.. 'rriti.O \W A' i :-,S
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    I- i '- SilI-LhiL] tIa t .-2 V)i1. .i l..) I) Ci iAV I jllit FO'di3l''o d k b -II I i

    gIt.- for- e-2 't.r fi._;l cii ..i .\t tb,: .-tai e" tm,, part 4f tliis
    pp ula,. -i 1Fn W ti ar d rphic. \t'd ri ad Ue(ie for h] ctd-. pat:' II cicEtwc t ILL I 2' l. I" t iI < ..tccnti ia,,t-,
    r 'iii h r'ise to )1 ti 'lirnrip.i.. i. "k1'i r .i- ,,i'.n r 'i .\ n.i '"'i t.i( lO

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    ,ill tlft,'- t:1 i>1i1i4iS an4 v.'s s1il;.,,'ii,'in lv 1 fn ri-% in ,i., i' ,ir !in '
    p 111:a! i ns.
    TIM"- hrn-ak in the sandv corlridor wVh't:-r the Devil'-% ,"nip Fi. :'32.
    inse1E flnup. i' i iov\ is~ ate m m ri fto ]m ,, ib. n/,I, m: av be(' O r( iif '
    reco.nt dIj'.t. Possibly the hIurricai ne .f .4 13, w'ilh wri'. Si- guJt g.ar t
    h.AV'c, .111d h111P.t,-'d itil :.' K (f tl i,:.;ist;t] t fop irLph\ t'is n01 nr rn
    Gulf Coast rr-.T i n mri hi.X I ,; I th .ILUc4l It 'l.104 idw .tterts c thi-
    ( tl'-' .',i, li 'lt .ie I 'c.r _ci .', cii ha' b',rokcti- r t Yhrniv i th'i" "- i .liat Id tnIft
    tfr'tive r it"l lii (i1. -ik r I n -" p rcf i ils p 'I -i..l i i s-t r ll i.

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    of tal ir, t- e N II I', U N i l' i /-P a f- .! J,. 1 1 ; I I
    i ,'st,,wa~ii sp~ cpn-d ("f x.:r. 1." .' rw 'at' i S1'., 111 i, I t ,Id i 'a LiE. :i's." n1., (l1h

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    figrnes itII ri..id '!' < ., .1 its pa1. r hi < ior.:hLit fI '; .aik' 1t F i.- } A.]ind '

    Jir i rc en,,t LtI ,c !. a ;i d1 so th '?,l,. / I',t,"iltS,.'v i r4 i'S n-[i .,sl t .\ f [vein.-
    a i,.ii n-l I1' ftr -rJ. ;n 1 -/c' I 'ul .- 177-. k Lit. SNb ._ ,-it> J -/ .';cc / S iT .ti s
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    Tii0 C*iSlli" i Il 1 iii p ia l. i.-io-if u.Kicn ;j !:ift1-i' ; r i fliii t lie spccirs
    Was WI l(.V ll LU I t it ipate- d c at tI'k tin'n <: f the lal Ir tt r:It,' urt.,red'.Lii l t.esl..
    1'i i !'xp1 thrI-'I- I- IhI rl x r"TI. I. -II I ti -'d V it i. t.' i. fi l 1 111
    T Ii r init dI ( I d af v iil; t- 1i, ;i ili'i, ti to ot1h r i ti <'t, T1!i f 'ii j :t iiiII -
    i'n c,,'is rtp u r iu<-P C .','_ 't. l -; T l t. 'I '. ; \'i r I I : I I i i
    *h. u 'rf.i pbaI s.s iJiI!.1A~i Sirif it r f.i l r -r its n I i1 t cast um t
    I.6iC'/rE) 'l. I t Iiiiitn O (j 'n ri 4\ F.i 2.r lit t'I.-lu hiirIiil
    k I 1 F I '- N 1*f -t.
    " 'tio, l id i il 1 2 en 1 ,ln j. !, 'riri d r cs-. 'i 'c i .T ( fl m.t" ; T .' .il
    If.Eu or pc ai 1 .1 :1 1", I v I ti S If I I i '. I th 1" 4 i '" t l >!i / N
    1C oiCl.lcj p/hulu. nIiI E 1 4 p it' \' \flP':< V ."A .Li' n d A111 ptI!r i iC0I L ci-Att h 'C'-
    C'i't.rr ir: 1. p.i rillrs T i.' T'sn I ol ;i t tmptc t 'd m ,TtiiiL;s .1f rhoad.si. aJid
    (Up ri11, to th lic stock irc a f' 4)11ows:


    V .1 1 .








    I-'S HOWEN\. (;li.F :O A.SIT BEKA( H MICE

    (i. ,1 -.*.rI iI'..iti ]- uiri[i,1:. ,tainy:' 1i d0 .l.'. ,f i. t _'r :lf,, .. rt lity





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    d S I :,i 2 1. .. f i I- 0.0
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    . ,:; ., t "7 2 02 *I i.- .2 02

    /u-?.e ;, 11 2 1...
    *ar *jri.cnt min !rto Liitv Cti rre..

    ReduceiL ft rtilityv is mI ndic a ted I Itwe "ei rh I oad.yi and subtrivufis, ar!o-
    phi .r ciTd ?Jo i l ,11C. 1, a t ph.. ,. tri d s .i i
    albifrons, in ad(ditioin t t that with lfuroce'phaltis. As bas is for cotri-
    paui 5o0, the proportion of suIceC.ssftul i.ittingi of other subspecies is AS
    folow w.s-

    o iIhspf eC--s Nk r!in"r :f S2i s:if
    ,IIrI :;,''s 11, .4 i ';
    I.h S o.x lt' w? .7 13 I .
    Ub-r'.L.% , s',b. ri'.t u: 7 4 i 72

    ,ctrr. L,,-i;' ,du x Icu,1 ..-, phj. a I .1 )10 : ':'.
    ll.phr7 s d h1oph ix 13.5 21 15.7

    The first three of the- abov, listints suggest that he- expe'td
    norniaiI nu.mbPr of sucss, i fl is ,iti.1s, tindc. r lahbora ttor. r nditio LS, is
    lightly under .50.. The higher percentage Dawson (1965) obtained
    is probably u be aL'c(Co:urhit"d for on the basis cif food ( s-ee materials
    and methods In .ucocCphalus a sormewhlat lower fertility rate is
    noted, aind in allophbrys it is very low This was noted (-arly ini the
    laboratory txperinieunts. and is t Iae chie-f 1 -ason why so few attem pts
    were mad n.ie ttotiit-t thias riitC-, This fact must a so be tal n i) ito
    consideration in the interpre-tation o f the r.Lsults of slch outcrossCs ;is
    were attempted. The explan.tionr of thlis low fertility is obscure, but
    it is thought po.ssiblv the result of genetic di.shrmonirs (Iusd by the
    infilix of sumnfer2 onto this beach which was previously populated by
    a peifmnsuaris-like .stock, as su tested above.
    James N. Layne has suggested in. litt.) that "the lower fertility of



























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