Spatial and Temporal Movements of a Population of the Painted Turtle, Chrysemys picta

marginata (Agassiz)
Author(s): Owen J. Sexton
Source: Ecological Monographs, Vol. 29, No. 2 (Apr., 1959), pp. 113-140
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1942200 .
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 SPATIAL ANI) TEMIIO1RA MOVEMNTS (O)F A PI'(O)PULATION OF THE
PAINTED) T RTL, H(JIYHRSEJIYS PI(CTA MALRI NATA (AGASSIZ)
OWEN J. SEXTON
Dcpart nint of Zoology, IVash intool l7nill'(sity, St. Lollis, Missouri

TABLE OF CONTENTS

PAGE PAGE

INTROD (lrITON . . . . . . .. . ... . I13 Activities of the Ch ryso(myS reInining

within (Cr-me Pond . ................... 1 23
lJo(CAT^rIONAND IF
,)ESU(RIi-TION OF TilEF AREA ........ 1 13 The Aestivnl Seson1. 126
Spatialt reorganizaition of the p)opl0ation1. .1 26
MNETrHODS OF ISTruDY ........................... l15 lResponise of the populaitionI to vegetation
(cla liges. 1-27
A IB)RIEF 1)ErRuHIUI OTII)Y OF THE
(espoilIse of iindividuIuIh turtles to vegeta-
POPI lATrION ............ .......... ...... 1
tiolI ('1n:1 lges .1 29
F'eattiloes ('0111111011 to 1)18 iits utilized 1)by
Tii: ANNUAL (Y1CE OF' ('111!J1scings A(CTrIVITYr.. 117
A Brief ,;c'. ............................
(7 h1rYs 131
Synopsis of tlie Aniuli (y-#e ... . 117
Advanlit:ges of the T-stintettnie ........... 132
The Pievei IiI Sn 5011 .................... 118
1, Sensoi .......
The VXeiii D)istribuitioni withinii 1)1 llt eds..... ....33
1 19
The verIlIl I (ci igln tiOll ,l1n1
Activities of ChryScoys ollnIatS of floa tiing
Oil
the
vegetationll .13. I
ellign llts ...........b.i.l.. ........ 12\
Eln vi iOnlinleIntl fen tillures of (rn ne Pond
U(tillu1( COhrscoinys to activity
of arleas of
forI er ven ..I,S
s. . . . . . 134
,11141 Slllrrotlldluing ,1en durS11ling thle \'('1 11,11
The Antitimiiah SeaIsonl. 134
emiiigrntiol. .. 12
Tlhe 1thinluenice of Nvnter telileln-tUle-( 101i 1I)uSCUSSIO()N
.. . .. .. . .. . 13)6
('Illel-(t Oil ('liligl-nltioll .13. 1 SU M1\1Al ARY .. . ... . . .. .... ... . . . 139
The extelit of the
llispersn I lilovemelilelits . 124 LITERA1T RE (TEPI)D .............................. 139

I N\TRO
1)OD CTO-1ON I Ialipt()n (Carson gave imallny valluable sullggestiOnls.
There has been ofe an inerea sing a mount informa- The laIte Professor J. Speed Rogers and 1P1rofessor
tionI aitulalll1llbltill over the l)ast two or thiee dleeables T. II. hlubbell offered mlc the use of the E(dIwini S.
which dlea(ls with the territolries, a-letivitv' Irangles,
sland (G'eorge Reserve. The eurator of the Reserve, I)r.
home angesof nn111an imls, both vertebra ite and II Xill , Cantr all, and the custodian, Mr. Lawrence
invertebra ite, aquaitie aInd terrestrial. That this in- Caniburn, helped mne considerably wvhile I worked
ter-est has beein aretively slhare(l liy investigators work- there. I)r. F. K. Sparrow and Miss L. IIillis kindly
ing w ith turtles is shlmvnl by the pa1pers of Pearse idelntitied the plants. Mr. Laurence Jobiiie dlrew soiiie
(1923), Catg1le ( 1944, 195(0), Williamls (1952) an of the fig-ures. Financial aid was provided by the
Bre( kenr1lidge (1955) ()ii fresh watter turtles, b, thIose N'atioia 11 Science Foundation, the (e orge Reselve
of (Ca-ir &- Caidwell (1956) aiidl Ca17rr & Giovannoll Research Fund, an(ld the Summer Research Fund of
( 1 957 ) ()on imairile turtles, anruidb1y those of Br eder the Graduate School of Washington Lniversitv. My
(1 927) , Niehols (1939), Woodburv & 1tIrdv (1948), wife, Mildred Bloomshurg Sexton, was a constant
in(l Stiekel ( 1950)) onl terrestrial speeies. These inI-
vestiontiois, ais wvell ais mo((st others deafling with the
spatial moveui enlts of assorted anmails, Werle( ( orleoeelrled LOCATION AND DESCRIPTION OF TIlE AREA
w ith estaiblishing whether or niot the amnillsls studied
The population of turtles most intensively stu(lied
restrictecl their -activities to limited alres or, if sueh inhabitec( five bodies of water on the Edwin S. George
<areas were foun(l to exist, with their size, shal!e, aun(
Reserve, located in Putnam Township, Livingstoii
perluancnee. The lpresent inivestigaio 0 wa ls unldelr- County, Mlichigan (Fig. 1). These five are Southwest
taken with the follovinv , enls in view: (1) to verify Swamp, Fishhook Marsh, Crane Pond, the I)itch, and
Or refute the g e ncral belief that the painited turtle,
Cattatill Marsh. During periods of great pIrecipita-
ch r,,secals pichta, restricts its movellients to certa.inI tion, late winter, spring, and sometimes (luring the
small areas, (2) to identify axny physical or fall, some water flows southward from Fishhook
factor associated either with changes iii the size in(l Marsh into Southwest Swamip, while the remimainder
location of an activity ranige or with the spatial anid flows eastwar(l fromt the former iilto Cranie Pond.
templ)oral stabilization of those saiiie atreas, (3) to As the flow continues into Crane Pond fromt Fishhook
ascertain the seasonal vairiation inI at(itities (of a. Marsh, the water level in the latter drops below that
population of turtles over a period of several years. of Southwest Swaamp so that the water fromlll South-
A (lebt of g'ratitu(le is (due Professor Frederick xvest Swamp reverses its flow, passing hack into Fish-
HI. Test for his ail. Dirs. Pierre I)aDsereclu, Francis hook Marsh and then into Crane Pond. The wvater
v'ans, Nornian Iartweg, Herbert Wagnc e lr t ad exits froiii Crane Pond through an outlet to the east
1 14 ()x W en J. IN-1((logic I
I loiiogras)hs
Vol. 29), No. 2

45
H K/ ~~CONTOURINTERVAL-SIM,"
S~~~~~~~~~~~~~~~~~
| 5
5~ CRANER PON
SOUT..EST t O

FIG. I. (raiie Pondl a(1n Ileighib)oring bo(lies of water Fio'I. 2. Map of Crane Po01d(, Edwin S. George Re-
StLdied(l (lulilig t11e (COll^SC Of t1e presenlt investigattioII. serve, Livingston Co., Alichigani. The ntumnbers around
Sy1l Lake, not ini(licated on this map, is located about the I)eriplhery rep)resellt certain of the fifty-seven stakes
400 iii south west of Southwest Swv.;mii). Direction of arounidl the p)o01(1 itiargin. The inlet (IN) and the outlet
wa ter flowv is illflicatedl Ily a1 r1rows. (OITT) are shown.

and flows eastward through a ditelh until it loins a Since water ceases to flow out of the t)itch into
lrmanch of Honey Creek, a tributary of the Huron Rloneh Creek during early summer and resumes flow-
River dlraina-e svstemu in southeastern Mic'higan. ing in early winter or later, the five areas in which
Southwest Swnivp, about 1.20 in southwest of. collecting was (lone are frequently isolate( f rommithe,
Crane Pond, illsa;,n area of about 15 acres, of win cli other parts of the same drainage system. This isola-
S are open dulring periods of high water. Inl 1946 tion must have been even more comiiplete before 1950
a seni-circular moat was d(ug', ai(l this always con- when the Ditch was dug. For these reasons the area
tains water except during periods of extreimie drought. under stuclv may be considered as a partially separate
Frequently the more shallow areas are dry fromi midn- Unit.
suiimner until late fall. Individuals of COmrisemyis picto inhabiting four
Fishhook Marsh, about 2'0 Ii west of Crane Pond, other areas were collected (luring the summitierof 1957.
has an overall area of 10 acres, but most of the area One area, Sayl Lake, is a shallow inarl lake with an
favorable for turtles is concentrated in an acre ponid area of approximately 40 acres. This lake is 300 mti
situate(l just west of the inlet of Crane Pond. Fish- southwest of Southwest Swamp, but there is no (i-
hook Marsh becomes (li\ (lurin- the sumiiiei. rect connection between the two. However, prior to
Crane Pond (Fig. 2) is a permanent pond (lug' by 1872, Southwest Swamp drained southward, either
d(rag-line with an area varvin-
in 1946, froiii 4.5 to into or close to Sayl Lake. Hidden Lake is a senes-
5 acres., to its construction,
Previous a seasoiial 1)on(d cent boo lake with an area of about 2 acres. Al-
was present in the sallie general area after 1938, when thoug1h it is in the samne drainage system as Crane
the tile dlraining the area was destroyed. At preseiit Pond and related areas, Hidden Lake is separated
the maximum depth varies froiii 0.7 inll during ex- f rom thein by a tamarack anld re(d maple swamp, and
treiimeiv dIr autumns to 1.3 in during the spr ingt,. there is l(O direet passa-e through this swanip. Hid-
The water is clear except during heavy ra ins. The den Lake contains water throughout the year. Neither
substrate is clay, sand, and organic iiiuck. The ele- Island Pond nor Burt Pond, both artifieilal bodies,
vation (If the pond is 274 Ill above mean sea level, and are (lireetly connected with Crane Pond or related
to the north all1(1 south a 11 bills pa-irallelilng,the south- areas. Burt Pond has an area of about 1. acre, anl
west-noitheast orientation of the pond. These hills Island Podl, an area of about 0.5 acre.
attaiin n elevation of 297 i and clhannel the }pre- TIhllemiost llabundant aquapatic lllats tounol within
va ili weN''St(erly wvillnds over the p1)(1 surface. Crlame Pond, the aie ra most intesnsely studied, are the
The I)iteh, 170 in east of Crane Pond, was (lug inl followinig (1) emergent plalnts-T Iyph1m litit~oli,
1950 and consists of three linearkl arrange(l poildls A I is 0)( triiUle, A(igittO rwi latifolio, leersia or1zoides,
varying in depth from 0.3 to 1.3 Ill with occ'aslional a1(1l h'Ieoeh 0liS ocolotri s; ( ) rooted l)plmnts with
(leeper holes. Thes;es three pond(s ontalin water floating leaves-Potloinoq(ton i1iipl/'OolO/s' J
P. 100tons
throu-hout the year although the flow of water iiito all(ldNifpl)(r odrenai (3) roote(l plants with sill)-
<anr(lout of the Ditch is seasoiiul. Cattail Marsh is ierged le.ltves-.Potamog eton pe(( tinotus, P. folioslos,
iiot (directly connected to the Ditch, but it is withiii 10. inplflifols, NaOj(s fle(ills, EleocPl(iris O(Oic -irs,
15 in. of it. The area of the mautrshis about 6 acres amid hnon imliciflllS longirostris; (4) anchored plants with
although the amount of open water is variable. All submerged leaves; or thalli (hara sp. and (Certophlll-
except a small portion of the marshl becomes (lix loim demoersooinm;(5) anchore(l plants with floating
durling the summer. thalli-varlious filamentous algae.
 April. 1959 l'OPULATION OF TIlE. PAINTED TU RTEI I'1t)
Other a(luatic or* seilfi-aquatic plaiits ot lesser iml- this fence could also be easily accomp)lishle(l if the
pollrtiilce are C(qp)ersl.strs igosS, Elcochari8 sp., collector was iiot at hand.
A very efficient trap, which took advantage of thei
(/ mhphibrioum, 1'. h !dropiI)eroides, Ran1.1cUdlls scelera- seasonlal movements of turtles, was construete(l in the
tifs, S cIte/lla ri eJviloliifolia, 3e1nth ( (arvlensis, ailld ile1t of' Cranle Po1n(1of 1 in chickenl wire. A ciruliiar
Bideus sp. Epizoic algae are Basicludia cholOnliil1 area of the inlet channel was surroun(led by a feiice
ailld B. cro.ssa. of this itaterial through which a funnel was inlseite(d
The more iuiporbtait vertebrates associated wvith oil the downstream side. The fence was 60 cmii highl
Crane Pond inelude: aind was supported by wooden stakes driven intO the
AlIuIIIII11aha: raccoon, IProc!/lo lator; miuskrat, Onl- substrate. From each side of the funnel a lead fence
dat ll it)bethica. extended downstream at an oblique angle until it
Aves: pe(l-hilledl grebe, Podimlnihs p)odlicels; passed onto shore. One lead fence joined the retaini-
g-reait hlne heron, A rdea herodias; green heron, BIlto- illg fence described above, and the other exten(led
rides riresceenks; American bittern, Botamlirtis len tigi- onto shore for about 5 ia. Thus, any turtle mmmov-
Ilsls; spotte(l sandlpil)er, Actitis mncl darial; black ing up the inlet was directed into the trap aild re-
tern, ("hdidonils nigra; belted kinigfisher, Meguaeryle
taiiied there until the collector appeared. Since the
curreilt somiietimiies washed an opeiling beneath the
Reptilia Blanding's turtle, Emns, blan(linyi;
fence, some escapes undoubtedly occurred. After the
pIainted turtle, Chrysemijys picta.; common water snake,
Na trixi Sipe(lonl; ribbon snake, Thu mo/)lh
ii
use of a simpler model in the spring of 195:3, this
is SlO iriths;
eomiiuiioin garter snake, T'. sirtalis. trap was in operatioll continuously from early sprinlg,
Aiphibila c ricket frog, Acris in 1954 until June 1955. Similar traps were used iII
yr/i/ils.; bullfrogt
R(aa (atesbheila ; leopard
frbog, Rt. pipiens. tile outlet.
Osteielhthves re(lbellv dace, (Ih r oi0llis oos; goI- A separate identification iiiark was given to eaclh
lnl slliller, Noteiii iqon us crqs oleeau s; mud millnow, individual turtle when it was first captured. Each
Umbra limii. miar-inal lamilla was assigne(l a number, aiid anm'
combination of nunlbers could l)e given for idlenltifica-
METHODS OF STUDY tioll merely by cutting a V-shaped notch into the free
edge of the appropriate lamina with a penknife.
Aln attempt was masile to collect every turtle in
The data recorded at the timee of capture iilclude(l
the population by employing the following Methods:
the iclentification number of the turtle, sex, date, timlle
balited hoopnet traps, sunning traps, funnel tras)s in
the inlet arnl outlet of Crane Pond, a retainillng fetice of day, location of capture in relation to numbered
on land, inoodling or mnuddling froin a boat and oi1 stakes Pulace(l arouild the periphery of the pond (Fig,.
foot, use of a hand net from shore aInd from a boat, 2), association wvith, and relation to, other aililmlals
t11(l selning. aiid )lants.
The use of a dip net from a boat and front shore All turtles were released as soon as possible. This
wvas very effeetive. An attempt was mnade to net generally varie(l from 53 minutes to 1 hour. 1)ur]imig
throughout the entire pond within three or four periods of heavy migration, it became necessary to
collecting days, and the turtles( did not appear to be retain captives 12-18 hours. Turtles were always re-
-reatly disturbed by collecting. leased in the immediate vicinity of capture.
At times of low water, muddling, or feeling about Most temiiperatures were recorded with staiidar d
ill the water with the hands, was very effective from chemical thermometers anld Taylor moaximuma-mumini-
a boat. It was less desirable when depth necessitated m1um thermomlieters. Water temperatures were taken
wafding-, since this disturbed the rooted aquatic veg(e- at a (lepth of :30 cmii,but the thermometer in the inlet
tation. was in shallower water. Wind direction wvas nioted
Three sunning tralps were used in 1953. Each an1(d the velocity measured with a Birami anemometer.
consisted o(f four 2 X 4 inch boars(l nailed together Thiis reading wvas taken just above the surface of the
ill the forni of a square, 90 cm )11 at side. Beneath
water at a location which offered the least resistance
this tr anl1le Was suspenlded a basket of I ill iiesh
to tIIe wN ind.
cliickeii wi me into whiech the turtles fell wheml they
Silnce the distributiol of CliilJlsemys thiomghlout
(lived off the 1)oards. The trapls were plctell olly ill
(mCane P0o1( was associatedl very strikingly wVith that
(aIr(eIs use(l as nlatural SUn11ning' sites or. ill other regions
of coemientrati(n. Turtles (1Joul easilyv e(scaie if: the
of vairious )iilts, thme locatiolls
of the beds of aqiu-Iitic
collector was not at hanml, thus activity was not inter- l)lants were mialpl)ed. Each timiie a turtle was cap-
tlmed(l in Crane Iond, the types of vegetation iII time
rupte(l by a long captivity.
A rettaiiiinig fel(es 10 in hill and iniade of chiekeni imiiiiiediate viciimity Nvas recorle(l, mfmil iii 1954 an1d(1
wire with a mesh of 1 in wvas erected onl laod to the .19571 llial)s of the aquatic flowvering planm mts m11mdthe
\vest of Crane I'Pol, an(l it extended for al)lout 100 niomm-lalanmktonic algae were iosade at intervatils thtimougi-
to south of the inlet parallel to the Nvest shore of' the ouit th)e veaim These two miietlods eaiable(l oII( to fol-
lpomid. Its puurpose wvas to detain turtles tenmporarilv low the, changes in location of the plant beds for two
ais they moved overland at that poinit. Escape fromo yI ea r'S.
 1OWEN J. SEXTON E(ologicallMoniographs
V'ol1.2 9, N o. 2

A BRIEF DEMOGRAPH-1IC DESC-RIPTI O)N the iiilet andl outlet traps were responsible for the
OF THiE POPULATION ecpture of most of the animals. These traps were
evi(lentlv not associate(d with eapture by the turtles
Diuring the period of studly, froim Mlarch 115,19153,
aind the niarked and unniarked animails di'd not re-
uiitil Septemnher I5,, 19157 878 indlividual C1hryse4mys
spondl (lifferentialily to them. However, (lul ing the
Jpic1o were eollectedl nimarkedaii(l releasedl. Most of suimniel' when most of the aninials were collected by
thle aictive ciollecting wals (oiile IIIOI01 aro01ii1d CraIne
netting froi a boat, the inajority of the turtles whieh
Poinl, hut soiie collectiiig, Was also (lone 'in Southwest
remained at the surfaee as the colleetor approlached
Swav,nipiII, Fishhook Marsh, the Diteh, and Cattail
to within three or four feet prove(l to he uniiiarked
MarCishi.The Craiie Pond( section of the p)opulatioL1
ninialIs while those which (love beneath the surface
was ohservedl aliiost (oiitiiiaill-v froiii March 15, 19153 an(lswainl wayv while the netter was still 10-15 ft
to April 30, 19515. Less Hitensive studies were ear This (IiffeellrenC
aiwayvwere (chieflv marked inlivi(luals.
iied out trout May 1, 1-955 to -Aug-ust 20, 191515 ; March in the behalviVorof the nlarked and unnIarked turtles
23, 1-956 to April 1, 1956; April 1-3, .1957 to April illnreasedl as the studly p)rogresse(1. The lower pro-
21. 1957. Another intenisive study was iladle from the
portion of imairked aninials during suinnier was
U11CI, 1957 to Septemher 12., 1957. InI addition to
.Jui also (dec to the ftat that the young of the yeari ( de-
thle 878 livinig aniiials calpturedl at least once (luringn
posite(l as eggs (luring the previolls y(ear) eilierge
this'- eri-odl thle following iiibers of (lead, nakd froimi the nest ill the sprinog and 11 a ficst axvailable
turtles wveic (ollectedl iii the area: :1953,103 ; 1954, 14; for cli j)tulle (lulring the sulilIiler. The grea test pro-
1955), 1; 1-956, 21 ; 1957, 2 ; t )ta 1 141. A few Ch rq-
portiomn of Ina rked turtles collecte(l ( 32 of 34) was
sum pluo wee ailso co(llected(lat thle lollowil1g loca- from Aln ay 19 to A\lav 29, 19 515; 1miost of' these aiiinials
tioiis (llriiig the suimmerc of -1957: 11idden Lake, 17- were lirge f'eiiia
lelles iilvillg a1bout oii Ia1l(1 Ilringll the
Buit :and Islad Ponds, 18; SalLake, 34.
II est iII0' S(elas(o.
The prop)ortioni of marked -a11( unmarked a iii nals ()I thle hansis of the ab)ov e Statemiients it wo muld
(a iitiredl thrmoughout the course of the Stli(Iy variedl appear thiait al miniliui of' (9-S4'` of the aniiialn s ill
regula1-k ulv fter a ceita iii1 point waV.s realChiel JTahle -1).
the poplillhitioi llan been mari-ked, amid a value ((1t
.'a.()'/, iv be ta kei as averag-q(1,-e.There werIe at leclst
TxnuEi 1. '1 lie percentaige o)f nia rked anim.iflas ca.-ptured 716i (Chril/se s (i'(t aliIve Ill 19513; 5(0 (hilitioiia
lutring the varions timues of the yea i. The decrease in iil(flii(lullls of ulncertafiii(W'a w(ere Ponba bl1aso 1iieni-
thii eeci(iit~ige o)f imarked amniua is froiu the spring a iid bers of this ye(allls pop)ullltion. tIsing these ligllres,
fall1 to the Siummiier Is the resuilt of the young aljlimal's it c11ibe a ssuil ed1 thalt there were ughllv 1 0(000
eiiteriiig the o('pulatioii a iii of the (liticreulltu~l1response of west Swnll lp Fish-
pll inltc(l turtles iiihacb1itilng Southl\0
theima arkdclanil ttimiu ked1 an iii as to the (01 lector Iduring
l1ook Mli sl (illCine Pond, the I)itelhi a 11(1 (-tttliil
AIl rIsh.
'Tlese 1000 ti ltles were (listi'ib)lute(l ONver.lIl aii't'a
Total P~ercenit-
xNhlliel vnIlIe(l froiiiI .1bout Ii li'ies (1r11'illo 1(l0(o(I ofI
iumbher age of
Period Activities of the of marked ext1(11e a (1rlloglit to about 25 11tes (lurinig periolls of
anuimials anjinals animials IlighI wnter, (a1tll-1thog ill neitherl.l nse wXIIs thle tOtIl1

ea
1(11limli'olrilX'1! suitalble for the turtles. Thu1Is, tlle
M~ar.25-Mlay 1, 1954... Emigrating 372 53.6
Oct. 3-Nov. 19), 1954. ... 225 84.0 eCOlllllI il (lemisitV varied froill 166 turtles per (1Iler
Emigrating
Mar. 25-Mlay 1, 1955 ... Emigrating 103 76.7 to 40 turtles per lore. The highest (lelnsitv oll)serelel
June 1-Aug. 20, 1955 ... Various 44 45.5 w\1II aillg giegil tion ot(f at lelAst 422 nllniilals ill a bed(
Mar. 31-Apr. 26, 19)56. .Non-emigrating 01) 65.2
turtles pills of Poto(11(1oy( tonl JfoliOSll5 illeasurilg 15 X 5 ill ( 807
dead ones sq ft) a (ei lsity of 2265 turtles per (acre.
Apr. 15-Apr. 21, 1957. .Emigrating 68 69. 1 The )ge 1111i seIx (listlibutioii of the 75fi turtles
turtles plus known to be alive ill 1953 is shown In Fig'i. 3. The
dead ones
Junme11-Sept. 15, 1957.. Various 121 54.4 iiethtod bv which the age (f the turtles wsx .l estilllnltc(l
will he (lescribewd in (411ilil elsewhere ( Se'xtonI, 111ivu-
scrlipt). Briefly, it a.tteiiipts to cstiniiiate the agOe o f
During- 1954 the numuher of turtles (captured (lurlig10 inllividllllll nIninlals by collip)aring the miedlil length
the spring10 em11igration waIs still rela-tively 0 w, hut dur- (If the growth aniiuli visible oin the right .abdomilnal1
ing) any eiiiiratioii thereafter, the perceniitage of l~l illn~l to exp)ectel vaflllesxalculate(l froiii the enitire
marked amnimailsea1)pturedl varied from 69.1 to 84.0. :opilll.4till.
Af ter thle sp)ring eniigration of turtles from Crane Fenili-es evi(lenltly olutnuillber 1111('s ill this lolpull-
Pond, few returned to the area until the mntlvi-iig tion ; the Iletusll sex rnltio is 1.49 Y Y / 1.0 ( for- the
hodies (oh water hega-n to dry ulp. The p~ercentage of (G04 nlililllwlls whisbe sex w5s known. Since soiiie of the
marked animals among those ea lptuiedl in Cranle Pond( juveniles were not c.ipturedl tifter they had(1 beconle
during- the suimmimerva-ried from 45.5 to 54.4. This .
Sexu'lll llture, theil sex couldl not be detelri ilied,
i .l

drop in the perecnittmge of marked aniinials, from thle .111d they were cllssed(l .Is juXeililes. Al ll(es Illlture
spring to the suntmier wa,-s almost certainly due ta the en rlier tharn feilllm1es; the sex ia till of thle II iiiiiils
differenit c-ollecting techniques usedl. During the spring- inrge eilnougli to he sexillydv imlltulre is 0.76 Y Y / 1.) .
April. 1959 POPVILArION OF TIlE PAINTED TrlRTL]E 117

OVER /40 Oft

13 BEFORE
FTTI
13- 141

12- -'42

I1E .-'43 z

0 - FEMALES ad/MALES -'4

z 10 446
0

? a8 //////// \\ \ 46

?7 -47

6 48
wd 0
5- _ '49c///,////////
4

4- *50

3- -1

2- .-'52

I_1 _ _-'53

no 40 30 to 10 0 la t0 30 40 50

NOS. OF INDIVIDUALS
HL 3. The ige 111(se Sx diistribuitioni of756 iin(l dllvldd Cli/.sm i/s picta uuilimibiting tue (1811e Pond a rca
11llrilng the growvlng sel0so11 of 1]93. Tue solid lhorizoiita' 1)818s relpreselt youlig turtles whliose sex wvas niever lefi-
iiitelv d(Iterhile(l.

About 210 ()t the 756 annals were ol0( enough to a1)le to cmjilp1are the aver'gie nata lity ligure for 1953
tliltedlte the ((nsti-tictioin of neIond
CraneI in 1946. a 11(1 1954, 58 young per sea-ison, with the overall mor-
Thiese ()1(1(1 turtles uay wenl1I1ive b)eeInresident in the talitv rate for 1.953-1957, 44 per year. Although
teilln 1)01y pmids an(11dsw8i1lll)s \vhieh existedl between there aIre still ialiiv sources of erilol ill these esti-
I938 a nd 1.946. At present there is aI simnuilarsitun- 11a.ates, the rough laIta. x(,o111(lseeni to indicate that
tiOl ill :iotlthier p'i)t of the G(mcl-o Reserve. Two the ntlinler ot turtles in the population is- relatively
small, tepllpmoiay 1)ponds support about 50 turtles in stahible.
their two 'lwres olu'ing0 S0111oieyears. These ponds be-
couiie liv' (lulring uuui(l-sunmlmer and remain so until TIIE ANNUAL CYCLE OF
lite fall oi earlyI i IIt er. (CHRY/SEWM S ACTIVITY
The renlminilln 546 turtles began their season (o
glo(Wth (l11ridini 1946 or-hlater. There has 1)eenl a notable A BRI EvF 8Y NOPSIS OP THEri ANN A
1L-AI CY
(CLE,

(leeline in the nuimbers of young in the two most A short review of the annual cycle of the l)ainted
recent agoe classes, but, otherwise, there seems to have turtles inhahiting the Crane Pond area is plresenIlte(l
been a stea(ld population increase since the formation here so that the subsequent detailed study of the
of the l)permanent bodIies of water. The cause of the spatial and temporal chanoges in the location of the
decline in the first and(] secondl classes has not been population may he seenmas parts of the entire annual
i(lentifie(l. cycle.
During the entire period of study the following At least a few Chrysem fs are atctive as soon as
minimial numbers of young-of-the-year turtles were the ice coVer hegins to thaw, uisually between mid-
a(lded to the po)ulaltion: 1953, 63; 1954, 54; 1955, February and mid-Mareh. During this initial cool
21; 1956, 17; 1957, 1'); total 167. Corresponding period the movements of the animals are limited to
mlortaility figures are: 1953, 110; 1954, 37; 1955, 7; ptassing back and forth fronm the (leeper areas of thme
1956, 61; 1957, 7; total )2)2. These( data seemingly ponId to the more shallow areas or to the shore. Mor-
indi(iate that the p)opulation is in a slight decline, tality is very high at this tinle of the year.
but while most remiaiins of dead turtles (except those As the water temperature rises, the number of
of vervy youngo ones) (1o not disalpl)ear for manmy active animals increases, an(l when the water attains
months a d tcainbe found relatively easily, not enou1gh 80C, usually in late March or early April, the animals
time ws ,s1)ent in collecting, the young-of-the-yea-ir emlirClte, leaving Crane Pond. Most travel westwar(l
from 1955-57 to make sure that most of these small to Southwest Swamp, but others move eastward to
aninnals were obtained. It is probably more lerason;- the Ditch. A few renmain in Crane Pond. Although
118
I Ill-~~~~~~~~~ OWEN J. SEXTON
iilost of the e(xo(lus is over within aI4-wk period, a few
stlraggler-s (1o not leave Cratie PXond until mid-June.
A va8rietv of activities assoeiated with reproduction
takes place during, the s)riLng. Breeding, although it
may tfatke p)lace at other setasons, rea'ches a, highl point
during the s~pring. It is first evi(dent about the time
that emigration begins. The turtle hatchlings, from
eggs (letposited the previous year, emelllrge fromn the
il est cavity in lnid-spring or later (Sexton 1957), hut
thev' (l() not emigrate. The dates on which the first
hatc1ht>ling0s were observed for any palrticular year are:
May 6. 1953; April 20, 1954; April 19, 1955. Nest-
in occurs1 (luring late M1,ayor early June, and, judg-
ing ftromi amy records, most of the females deposit
their eggs within a relatively short period. Positive
lnestin)g recordIs are as follows: ,June 7-June 11, 1953;
Mav' 21-May 24, 1955; June 8-June 11, 1957. These
pemrio(ls were usually associated with rain. Other
workers have indlicated that the nesting period of
(Chrsetiois may be much long er. Legler (1954), onl
the basis of his own observations, states that nestilng
took p)lace froni Jtine 16 to ,Julv 14, 1953 at Sand
Lalke, Nicollet Co., Minn.
I)nrin1g late spring and early summer most of the
turtles arne distril)uted
between the bodies of wa ter
leril)her'al to Crane Pond: Southwest Swamp, Fish-
hlomk Marsh, the Ditch, anld Clattail Mlarsh. As these
areas begin to dry up, the animals gradually move
back into Crane IPond,
(aid bY fall mimostof them are
re-locatedl there. This imm111igrationis not as distinet
as the spring emigration.
Growth of the animals and she(lding- of the lami-
nac also occur during the summer. Recogniizably new
in HIem1entson the abdoniminal lanminae are first visible
betweeni id-iMay and early June. Growth ceases by
the end of August.
The activities of the painted turtles nmay follow
o0ie of two patteriis (luring the fall. If' the water
level remains low so that Crane Pond is the principal
ref ugiumn 'in the entire -area, the animnals will reimmain
here. As (0ol(1weather approaches they will move into
the (deeper sectionls of tile pom(i. At least some of
themmiwill be active until the perlmnalellt ice cover
fi'omns in late Novemmiberor early D)ecemuber. If, on(
the other hIamid, there is sufficient l)recil)itationl to
lraitse the walter levels of the adjacent )onds and
IIIarIsmes, aI fall emiraltioln will occur, and tile emii-
gramm1ts xx'ill hibernate in the outlyvin' areas. Those
turtles wNlhich meimmainin Crlane P)ond ummder such con-
(litimos will emlmgraitethe succeedling sp)rillng.
A s im(liicate(l pireviously, the po)ulmlation of l)ainte(d
tUIrtles living'1 in thle Cramlle 1)()11(1area is relativelv
stalble in mimummbers, but it i ia.1bits aim environmiment
wvhieh ulm ll(Idergoes imma rked seasonal fluctuatiols, e's-
pec^iillx iii temmml)erature, water (delithi, surface area of
tIme waIter', a111(1iii the 1imatllreof tIhe a(quatic vegeta-
tiomm. rThe aIctivities of thme tilltles w eresetronglv
aiffecte(l by these chmaniges, amil it was the pumpose of
thias imi1vestiga.1tiomi to stll(lv tle effects of tile envilmomi-
mielmitalfluctutationls upon the (listributioll of the tur-
tles. Tlme behavioral patteirn of COhisoeiys aimd the
Ecolo-ical Monographs
IVol1. 29.) N o.2
phenology of' Crane Pond are sufficiently (cistilict so
that five seasoiis may be recogiZn(l: the prevern'l,
the vernal, the aestival, the autumnal, and the hiemiml.
In this paper the seasonal. activities of the pJaillted
turtles have been descrihed ont the basis of the first
four of these season-is.
TmHElREVERNAL SEASONT
The plrevernial season of the Chrilsem:ms annual
eycle begins with the fin.-al ilnitingb of the winter ice
cover. lii 1-953 the ice (over hegan to melt duriiig
the first week of March an(l had (ldisappearedl by
March 11 when this investigation began. In 19.54
the ict cover started to disintegrate on February 16
during an unusual warm spell, and a third of the
Crane Pond surface was open water by February 25.
however, the waters again becanie completely coverel
with iee subsequent to February '25. The final imelting
of the ice starte(l on March 13 aiid was conlleted hv
March 20. In 1955 inelting began on M1arch 11 and
was finished by March 13. In 1956 the iee begaii
melting at a very late (date, March 31. In 1957 the
exact (late when Inltting was initiated was unobserved,
but it was betweeii March 2' and March 18. The fif-
teenth of March represents an approxiniate averaIge
for the (late when ice cover is completely mieltedl.
The end of the plrevernal seaisoni is heralde(l l)y a
mt'ass emi-ration of Ch rlsempqs out of Crane l'onld.
IIn the spring of 195.3 the first emigrations were no-
tice(d on Marchi 21. Eniigration was well under way
by March '25, 1954 and March 31, 1955. 11owever,
the data for 1955 are scanty, for most of the turtles
had left the pond durinig the previous fall. The date
for 1956 is uncertain, but emigration evi(ldeitly oc-
curred about April 7. SomIe COhrqlscms were movingl
out of Crane Pond about March 18, 1957, but an un-
seasonial cold spell interru)te(l this m1ovellelnt, and
emi-ration was not resumiie(l until about April 18.
March 25 is an average date for the termination of
the prevernal season.
Environmmiental condlitions during the p)revernial
searson are such that the activities of the painted
turtles are severely curtailed. A niaxiniumo-mininiun
thermnomimeterlocated at the western end of Crane
Po01(1 registered air temiperatures which range(l froij
-9.4?C to 150C in both 1954 and 1955. Wcater tein-
l)ematures (luring the saime yaIrs
, varie(l from I C to
1)CC. The observed lmlaxilllUIi1 velocity attainie(d by
the wind 1roaring over the unmobstructed1)0o1(1 surface
was 380 ft/nunin. It is these two factors, teimuperature
l1 windl, which influence the aictivities of the turtles
verv strly-mi (-INliiii this tillie of the yea in, but o)ther
such as redu(lice(h oxv'e tesllsiomi i the
faictorsi wa'.terl
a'Ie also iniporta't1t.
I)uming thte late hietmall (wintertl) or pIevernal
periods miost turtles are observed only ini those p)laIicS
which aire p)rOtecte( froII the wind. Such 1)m'otected
)lI(''es include i(T-coMvere(Id wa1ter, the lee si(le of shmre
pojeections, alnd areas protected bv veg-etation, such
as clunmps of Typlhu. Tie few (Curyscouqs observed
in situations where they were eXp)Ose(l to theft orce of
the wim(ls durinig these cool pmeriods were alimuosthelp-
April. 1959 EVIDENCE OF PHOTOPERIODICECOTYPES IN TREES 119
less. On several occasions Chrysemrys were found 2. Cloacal temperatures
TABLE of Chrysewnys active
entangled in floating vegetation which had been cast (lurilig the prevernal anid early vernal season compared
with the temperature of the miedium in whieh each inidi-
up O1n shore or rolled up onto the shallows. Such
viduall was captureml. The temperature of the imedium
animals were unable to escape from the debris. Tur-
was measured twvice, once with a black cloth wrapped
tles observed in open water were carried in the diirec- arounl(l the thermometer bull) to simulate the dark cara-
tion of the prevailing winds and coul(l not swilm pace amid ommeewithout it.
a-ainst this force. One aninmal, seen March 16, 1955
was in shallow water and was being transported in an TEMPERATURE IN ?(
elliptical path perpendicular to the shore by the
waves. The cloacal temperature of this animal was Medium
7.SC, and the velocity of the wind was 380 ft/min.
Date Activity Cloaca Bulb not Bull)
Turtles collected in protected areas friequently had blackened blackened
lower cloacal temperatures, sometimes as low as 1.4?C,
March, 1955
but they were able to move slowly away from the in- 1. Sunning 1)o shore 10.4 12.8 ....
vestigator, indlicating that they were still inl control of 11............. Just on1 shore 8.6 9.8
their m11ovellmelnt.
13.Sunning o01 shore 19.8 9.8 ....
The Chrqsemojs observed swimmingl beneath the
Resting in water 9. 2 8.0 ...
ice appeared to be trying to reach the air. One of Swimming 8.0 9.4 ....
the nine turtles observed swimming beneath the ice Sunning on shore 15.0 15.4 ....
sheet on FebUarY '23, 1954 was Iiiovinu, shoreward. Just crawled up shore 10.2 14.6
The water temper ature was 1.5?C, and the dissolved 14.Sunning on shore 27.0 10.2 ....
oXg enIl was 6.6 ppm. The Chrysemys alternately ex-
teIl(le(l and withdrew its head; the rostrum was 166.u.. . Suig o01 shore 10.0 10.6 17.4
against the lower surface of the ice. Occa- Im water 7.8 7.0 ....
piresse(
sionally the mouth was opened, possibly in pharyn- 199. . I ater under ice 6.0 4.0 5.6.
geal respiration. Other turtles were at the edge of the Sunning on shore 14.6 8.8 20.8
ice sheet, the head extendingli into the air while the
20 .Su.. . ing on shore 32.0 20.0 29.1
body was still beneath the ice. One iIl(lividual pro-
In water 8.0 6. 8 ....
tru(le(l his head through a small crack in the ice.
Similalr activities were observed (lurIing the late winter 21.Sunning on shore 8.0 7.4 12.2
aid(l early spring of three different years, indicating
28 .Sunning on sn)oW 8.4 4.3 16.4
that this type of behavioi is not uncommtion for Chry-
29 .Sunning on shore 24.0 15.9 24.2
D)imriiig the 'l)evei-ntal seasoii the turtles move into
30 .Sunning on shore 27.9 16.4 25.2
locations where their body te ml)eratures can1 be inI-
(1case(l, that is, iiito sh-allow waeter or up oil shore. March, 1956
Ill suclh situations the (Ialkene(l car-apace absorbs the 31 ................In water )11)derice 5.0 2.8 ....
loget-r wave lengths -11and the body t'eml)crature of the II) water )1))derice 1.6 1.6 ....
Im water under ice 1.4 1.2 .
trtlfe many he higher thanl that of the sullrroudl(ling
Il) water urlder ice 1.4 1.0 ....
mendiumn. Foi exampl)le, onl MAlrchl19, 1955 a Chrly-
N,*M'I/S (Ia pltu ed hellenath the ice had al Cloacal tem- April, 1956
1................. I water underice 2.9 1.4 .
l)(i'-tuic of 6.()C(0 while the wa-Iter tinepcrature, mneas-
Is water urlder ice 4.6 2.4 ....
ulred hy thie same tlhelrilmonletetl, wmasonly 40C. This
in(lic(aItes that evelnt turtlesXhelneth the ice are ab- April, 1957
SOrbI hgil enroughll of the longel wasve lenllgths to raises 15. SunIIiIlg on1 shore 13. 8 11.8 .

theli! teilhh)craltur e aibo-eA that of the water. Addli-

tiolial (Icses of diffelelles betweell the temhl)crature pertio(l of late winter 1a1(1 e(ally spring. In 195)4 five
of individna(ll tllrtles a lmd that of the sillromil(dillng medi-
Ch ry/S(slyo4 wX'(r captured two or immor'etillies befor el
umn a ic presente(I in Table 2. There ac-e four ilstaneces thmeadivenlt of enlligrationl; the llinilllml (lileet
(iitallnee
in xlhieih the temnlm)erlAtur-eof the animal is mtorethan between ftathest of lz-mtllturewitilill lCUIV OlIC
points
l C lowerl thian thit of tile surlrou1ndilln medium, but actetivity area va iied f room 3.4 to 124 mim, alVela gillg
ill thllree of these thle tur tles were ('1o1ining from loerl, 41 lot. D)ulin- the analog-ous in 1955 sevemn Oh roil fps i
(m0(1 (l1)thls iinto siallow watecr or up) oIn liI(. There NNere cal)ture(l two o0 11mor e tillies. The (distallces
alre sevell tllrtles with cloacal tenIl)erl-atires withill 1C b)etwVeen1 falrthest j)oimlts wvithiim at sing-le aeftivit tarea
Of the 1me(dIIu; (t thc( o al teilll)ratulre of the Ie- vallie(l fromim 9.2 to 92.3 oe 1111(1averaged 45 111.
HICIinmiing 14 Ch iS-q .mys is nmor1 thani 1 above that
0C"
of the suromun(lidming medium, iii some cases 100 olr THE VERNAL ',SEASON
mmc highelr. The grI-eatest (discrepalnlecs. ill tellnpera- The vernal season eXtendls 0111 MAarch26 to Alay
tulre ocur ill those an11iinmlals hlae sunlnling. onl the
vhicll .-31. Thle erite-iomi by wilich tile first (late was seleetel
shor~e. Iias been Imlenltionme(d p-eviously. The vernllal s(eas0o1
The movements of the )ainted( turtles withini Cramie is eo( elu(I( whemi Ceetil11 Sublnerge(l aIqUaItie pllIXltS.
Pond~ alppear to be relatively limited during this wimiell ale imimpor-tamt to (Io ry.semoys pict(i to to time
 4. Monographs
120
I 2 () OWEN e1. SEXTON
OWEN Ecological
~~~~~~~~~~~~~~~~
29, No.2

Wvater sflilwf. Jli 1953 Cluirai sp. was first ob- varie(d ftroit -3O to 30'(CC. There were few tinies
s(iV(l nit the suiifawe of Cra1Ie Poind on June 1; when the nir tetmperature fell below 0?C in 19.55,
u (
Rmm llU IU1U s lon ustris anldl soille otlhei species hadi the last time beiiig between MAylv10-14. The water
nttai(d(l the suitaict 1wb June 14, but they had prob- templeratulres (lduinig the saiiie year ralnge(l froin 6TC
a ly be1) 1 at the surface somiiewhlat earlier thiai this. to 260C in the verna'l season. The inaxiinnim Nvind
In 1954 (h Ui usp)., Vutoilmouyton foJliosmU, 1'. pecti- v(eh()eittvwvas 480 ft /mnin.
Jul.I(si (11 Rd 1llu(ld1Us iou ail-osti s18 beca e evid(lent at Throug-hout this stu(ly the terni mi-uIratioii will l)e
the Watel surface l)btweein Max )20-2. Iln 19'5., (Ch(,ura 11se(l to Sinifv ". a.i 11iio1v eiit whicll involves

S1p., Po(mlo(JctoIJ f)lwslfs, and ll/

Rmoll 111d111 Ili- a jo)Urney(\inlto .1 (defillite area, and a return j(lle to
rostris -1rew to the Surface b)etweeil laN 1:3-19. III the 'nin whence
A the arose
niovemIent (I lealp)e
1957) 1Potmnoyc((J(tons foeli.Slwms P. p)ec('nil{{tl..
fit'le8lun- 19321). An emi-igration will l)e a imovemiient away froiil
c ills iuuyii-ostr is ar11ius
ad111( specles f(f iilanelt'llls (Cane Polnd, aild aI i liil-ngrtion will be a IiiovenIlent
ait'1{ wxveieat the surface by eJuine 15. These varioii5 iiito Cr.Iie Poind.
(dites ilnlicate that iAI v 131 is about the tiiiie aqinati c T he Cc iiol O
i(] 0 tiolon 0(1 t[I CmCO iy ((tilts. The aiii Im'd
phiits iii (ia ic Pond reach thel surface of the water activityrccle of. the (pi rsciiofs io i the
J)0J) olotioii

acilidtllor exteiisive floatillg Iiiats oflleaves aull thah1li. ( i iil(e Pond area iicludes at least oiie migIratory
Ihllis lpeniilIl l rilliniily b'
is ('cli:arateriZed a.1111ss n(ovemluiemt allthouhall- aIninlulaIs (10 liot aIppearl. to
(1lilIiiltioil ot iilost of the (hrls(111/ ill (Irane Pond pa1rtici pate iII the 11iiiigatioii. Typica'
ly, the emlimgra1-
to sliriolllnlllhilng bodies( of water anli bv the localiza- tiOil takes plo ce (durilng the veiiial seasoil, the (e11-
tiol o t tllse tur tle s II;I ill
(i 1
C1rne l'ollonl to gra1tiing turtles hleadiiig eastwar(l 011(1 \vestar(l dawav
ceitail ii a ie in
iWhich there aie Stinniino sites. f rom I(riie
These oin l
(Tamnble 3). The hills rising 20-225
tWv) o1(1(uli (i5esi the eilli-ngtionl anild the localiZat(li60, in an 11(1 iiore above the 11(11(surtace to the north and.
-i ie (N-iliietlv il-l/itedl to tIe hi k 1)11 surface ve-(t a- sout l)l ielule a iiy extemisive iiioVeliienlts iii these (Ii -
tioll ini (raile lPondl at this tille oIf year. Other iectimi s. The aninias disperse emitlier hy tr xne'ling0
(evilts lii Illh Illiacterize this sea si0l11aie tlle initia- overla iid or biluOving throllugh the iiilet or onltlet.
t oillot fct(dini , niiating activities, aiil(l tl( eiergei(ice Those going0 Overlaiid (uld stw-ard froimi (Crae Pond
otl ti Iultcllliiig turtles fillm the nests of the iprcviOUis prl)i()lbYs aire soon ftuiieled into the outlet by the
va11. Nesting takes place (luring the eindl of the top)ogl~r.phy. 1 i1(lee(l, aIll turtles miiovill eastwanrd
xciria I ealson ii1 ai the beg-iiiiiiiig of the aestivzl one. tlllhoughl the outlet mlust first (c1rossanar111ow\ piece of
Tlie teililmperatule is very variable (urillg the vernal land, ahout 12 ini wide, hetore e iterimlig) the ()utlet
seasl Iil (v'ei it it is iCIlTeasedl over thlat of the pie- pro)per, for they will not pass thrlough the b)uried
veriia 1 sealsoi, an11dthe wiilll veloeity ('all e qIuiite high-l. 111lvert which dri mis the ponud. Turitles nuigratillng
I )uriuig 1954 the ai r telmllerature at the westerii eud of overland towa d(1sthe wvest iuiust travel across a 111i11i-
(ane Ploidl vailed froii -12'C to 32.5?(. Air Ilmanunof 120 in of gransslan11dhefore aItta.iningt, South-
teiiimperatures b)elolw (OT Wverl (lonllllllonly reco rdle(d west Swa mop. Thlose which emnigra-te through the ill-
tlrllnollulOlt the vernal scasllli o(t 1954, the last such let lmeiuulin In water for miost, of the jonrlley to Soith-
lo1W tlill)tlati're lre bliog r llrdlll be(tweenl Many 25-28. west Swamp, but they first cross a road which is 5
\'0~~~~ Wei~illlllllillli
Noliixiui-iiiiii therniaoiiiters
h.lllIltlSstre wr keeti
pt ill ]In wide separatiing C(lraie Ponid fromuuFishhook Marsh
the water (Iduinng 1 9;54. The wildls attaiilled velocities and then cross amnilumuni of 40 iii of 1land to reaclh
olt at lenst 740 ft/miii just over the Waiter surface. Southwest Swanimp. -It is po;sible to travel between
T11e venial Seasoni (I .19.955 was i1ul11 illmOie placidl the swapl) and the imaIrshwithout crossing such an ex-
tliaiil that lt the pvrecedlinlg vea.sl The air teiiiperatuies; tensive tlac( t of laid, but there is no (efinlite water

T~n.BE 83. NUIIIbelrs Of tull-tes uuCoving into aend out of (rnaie Pond during the various seasons. The seasons are
alblbreviate(l as follows: P, prevermal; V, vernal; Ac, aestival; Au, autumnal. The (lashes indicate periods when the
tralps were, not in contiiual ol)eration.

YEAR AND SEASON

1953 1954 1955 1957

P V Ae Au P V Ae Au P V V (part only)

Turtles Leaving Crane Pond by

Moving:
West through the Inlet ........ 16 37 0 0 2 277 12 175 12 61 35
West Overland ............... 2 3 0 0 1 20 1 5 0 0 20
East through the Outlet ....... - - 0 - - 9 6 6 0 15
East Overland .......... .. 0 0 0 0 0 4 1 0 0 0

Turtles Entering Crane Pond by

Moving:
East Overland ...........0..... 0 0 3 0 1 6 5 0 2
West through the Outlet ....... - - 0 - - 6 15 7 0 2
April. 1939 POPULATION OF TlE IAINTED) TUIRTLl 121
li1111(ll1 betth.eel the tNN-' aireais. The turtles dlo not a hdomlina1 hi la Vwhoselelltii is betweell 2.0 and 2.9
seeuu to t vol tIns ioute, for ione Ihtsveis beenl
eI (i Cll11(1 which (1o iiot have eioniigated (laws. Thev are
tiured(l osillg it. I telee, aIII Illngra nts (lOss Somue bind. 5exua li iiiii.atu Fe. Adlit feiiia les are those turtles
The 1lii! herl. of ( '1!/.11st,1 ios imveiielits througih the ill Nviliiel the length of the ahdoiiiia 1 hiiiiiial is gr1e1ter
illet dIin ellmilra tioii is shlowxnlinI Ta.ble 4. These tim21i 2.9 eiim 11(1
wnichi (10 iiot hita (e elonateil ela whs.
(hiti (10 iiot 1e(1)reseiit the total iiumiiibei of iiidividual. hlie 1ll:1jolitN of these ate11 Se xuatlyllv itiature. As i
tlrtle s a,.ssill tlllh()Ughl the illet, for Smiiie eliiiigrated 1)si toi eoiiomparison the 756 inidividuail (Chrlscm (/
tllolmli1 ocl(e, milv to returnl to Crane Poi(l and then knowii to hbe alive dIuring the .1953 growillg sea2son1
t ,vel n11pto the illet againI. lIi the vernl" seasoll of, (FAzig. .3) eoiisisted of 2.8% ,juvei lies, 20.3% slba(iuit
1 954, 18(5 r/5s(y00/ moved out o.t (ia iie I'oli(l twi(e, teiiiaides, 20.2)6`c teim-f1(s , anid 26.7C/,
eiiault d dult im-des
2 three tinie.s a1inl I 1(111 tiilies. The iiioveiieiit imtae oiista uit cheek was iiot nillait:it1(led over the emiiiiral-
diii-in- the tall of 1954 will be coiisi(lerel laIter. tioi duiriing 1953 aild 1957 so thait the data tfoi these
two yea(1rs ale incomill)ete. 1li the spr nigo o f 1954
T a~m:4. rThe mimliwrl (ff (C FlSccl1/S eiiiigratilig 19.3%, of the eliigtaiis were't juveniles, 21 .4 suill)-

wcstws.trd( tilroiugli tile inlet daring the spring of 1954 aiid

a iliiit teiiia21 les, ')29.(i%("aidiilt f les a iid 9.6
2 )(j a (hlt
19 55. (O1l tihose Ia tes a re inlldiea.te(l on whlic tartles iiiaies. InI 1955 11 of the emioii2graits wvere juveliiles,
W01 c,1
.Iturled( inl thle ilde(t. 2)1.9C`j, sub~~adult females 34.2(/)/' tIdult f-des 111d
32.9%/' atduilt i.iiles. It call, be seen that iewer iiUVe-
March 1954 1955 April 1954 1'955 iii (elligra te tihmii oiie woulld exp)ect oil the basis of
12 0) 1 2 15 7 0 ('hai111ee, l)llt iliiore aIllilt telles an:11d iial-es ellmiigrate.
14 0 2 17 12 I rl1lie pIiiie italeo of slilubidult Ftei ales ill the (iltil i(
15 0 5 1X 0 l
1S 2 0
1O )9 0 3 1953 plolllatioil aiiil ili the pl)piiiiationi of eiiiigaiits
20 0 1 21 15 1 is all)1bt the saiiie.
21 0 2 22 0 1 Thlie(dita fio Tab.1e 5 iniidieate that a1 .hii-rge o-
25 IO) 0 24 1 0
27 1 0 ti(oll of the si1iillerl turtles ieii iaiiis in ( a11ne Ponid
25 1 0
28 24 0 2 2 0 while the 1:Iagelr illflividulals of both 5seXes elio-inate.
29.) (0 l1 29 2 0 Sever'11 iollieetiis ill the 1)011 dur
W(e'e( 111a(1de (1111
30 0 ) 30) 0 1 of hea.1vy e(liiri'g1tioil
peliols to deteliiile the stinle-
8s1 0) 11
1\lav tiiie of the( residuia,1l J)i)f)lopitiioii. I)uriig A pi-il 9,
April 1 4 0 1953 tlhere \va s an11 elij-orati on of 27 Chrisc ( hlqs id
1 1 5 12 1 0 wliic 3.7(4 were juveniles, 14.8%6<subaldit fenaul1es,
2) ! <3 it) 14 1 0
:8 25.9).( a duitleslteiia , a1iid 5.6((/ a(dllt iiia es. ()hi
0 15 1 0
4 O ) 21 1 0 thle Sa ii(e (hiy 11 pi11 iitedl turtles w(erIe Iletted wNittlill
a tille pt ll, amil thlese cioisistetl of 72.7%j' uvelliles,
6 43 O) Jlne 9.1 % suba dult feiiilesx, andi .2% a(luitteamt ies.
d
b
(;) 0) 1 1 0)
9 3 6 5 2 I)uriiig A pril 6 and 7, 1 954, 1 1 ) turtles wer e eanlglit
O)
IO) 1 2 6 1 0 ill the iiilet a 1d(1 14 ill tillp loldll. The emlli-ra tOi-v
11 22 (O I1) 2 0 groN)up) comlsistedl oft 14.3% juveniles, 19.6%. subaldult
12 6 3 21 .3
l7 feiiales, .34.86%Vladlult fteiiales, 11(1
n31.3%1 allult
*( 111iides.
1: 4 22 1 0
14 23 (O The resi (lelits were c l)opsed(of 71 .4%f ,juvemiiles,
July- 7.1 subad(mult fe'males, and 21.5%/, a(lult feiales.
5 1 0 Even though the lnulmliers (If turtles ca1ptured withlii
the ponid1 is iiall, the results seeiii to (orrohorate the
belief that the younger turtles (d(o not tenld to elili-
The miunmbersof niovemnents iln the inigration rapid- grate .
ly reac.h aI peaik and theni (liminsh, but. the eniigra- There is also a tenldellen for iiiore females to enlli-
tionl 111ay1V
eonItillue ill (lesultorv fashion well into the g-rate than niales. Whereals the sex ratio of the adlult
aestival period. These stra.ggt lers mtay be moving aiiiimals in the 1953 population of 756 Chryserntys was
aibout ait raml(doIil within the lower areas of the Reserve 0.76 Y Y /1.0( ,that (of adults emigrating through the
so that it could be erroneous to consider then here, inlet during the spring of 1954 was 1.0) Y /1.0 S
hut they aire iilelu(de(d for completeness. a8(d that for the spring of 1955 is also 1.0 Y Y /1.0 .
The percentag-e of juveniles, adult niales, subadult hfence, it would appear that although prol)ortionlately
feiiiales, aid
11(1aIult females emigrrating through the in- more adlult (Chryseiails are among the emigrating,
let is preselited in Table 5. Juveiiiles are those ani- group than One wouldl expect, the two sexes responl(d
mnlls vhich have anl ablbdominal laminar length less dilfereitly. The (chnge in the sex ratio is evidently
thanl 2.0 (mo anid whose sex eainnot be readily deter- broug-ht about by the increased iiuiiiber of adult
lninedl. They are sexuailly mimnature. Adult mnales femiiales which eniigrate.
a re those Chry/sem, i/,S with an afbdominal laininar The relative abundance of each of these four
length of 2.0 em or moroe ahi(1 which have elongated classes, juveniles, subadult felnales, adult feminales;,
cehiws (onI the lorelimmibs (Sextoni, manusicript). Sub- and males, didmnot remain constant throughout the
adolt f1emales are those p)-iilted turtles having an eiiiigratorv period (Table 6). Iin 1954 the adult
129 OGWEN J. J.SEXTON Ecological Monographs
OWEN SEXTON y~~~~~~~~~~ol
29. No. 2
TABE 5. (CompositioII of the total iumber of painted turtles emingrating through the inlet.

Mar. 21-May 27 Mar. 18-May 31 Oct. 3-Nov. 19 Mar. 12-Apr. 30 Apr. 17-21
1953 1954 1954 1955 1957

No. No. e No. No. % No. //o

Juveniles. .. . 3 5.7 54 19.3 35 19.9 8 11.0 1 3.0

Males .. . 24 45.3 83 29.6 53 30.1 24 32.9 15 45.4
Subadult Females 8 15.1 60 21.4 43 24.4 16 21.9 4 12.1
Adult Females. 18 33.9 83 29.6 45 25.6 25 34.2 13 39.4

Total .. . 53 100.0 280 99.9 176 100.0 73 100.0 33 99.9

TABLE 6. (Coiposition of the emigrants nioving west- ing aereas datring the vernal emigrationt. There is
.ird through the inl1(t. little surface vegetation present in Crane Pond during

COMPO- ITION OF MIGRANTS IN TABILE 7. The composition of emigrants leaving

Date in Total (Cran
le Poll(d via various routes.
1954 Subad. No.
Juv. Q
West West West East
Mar. 25.. . 10.0 20.0 20.0 50.0 10 through West through West through through
the inlet overland the inlet overland the inlet the out-
Mar. 28 4.2 41.7 8.3 45.8 24 let
Apr. 6. . 7.0 44.2 11.6 37.2 43
1954 1954 1957 1957 1955 1955
Apr. 7 . 18.8 23.3 24.6 33.3 69
Apr. 11 . 9.1 18.2 45.4 27.3 22 Total number of turtles 280 21 33 20 73 15
Apr. 13 . 35.3 11.8 29.4 23.5 17 Percent Juveniles. 19.3 4.9 3.0 5.0 11.0 13.3
Apr. 14 . 26.1 21.7 30.4 21.8 23 Percent Subadult
Apr. 17 . 41.7 33.3 16.7 8.3 12 Females.......... 21.4 33.3 12.1 15.0 21.9 26.7
Apr. 21 66.7 13.3 13.3 6.7 15 Percent Adult Females 29.6 33.3 39.4 45.0 34.2 20.0
Oct. 12 . 25.4 27.0 14.3 33.3 63 Percent Adult Males.. 29.6 28.5 45.4 35.0 32.9 40.0
Oct. 13. 25.8 29.0 29.0 16.2 31
oct. 14 . 6.3 50.0 25.0 18.7 16
oct. 23 (0.0 9.0 54.6 36.4 11
Nov. 19. 27.3 18.1 27.3 27.3 11 the vernial seasoil, for Nuphar (aLdreiaais the olily
species which grows to the surface liuueh before the
enAdn(of the spring. This species was first aj )j)alrenlt
fenuuales were the fi rst to reach their peak of abun- onl the water about April 14, 1954 anld Marecl 9, 1955.
ulainee, relatively early ill the eiuigration. The males The majority of the plants reachetl the surlface and
attaiule(l their g'reatest relative abun(lanee shortly fille(l ill imost of the outline of each lily be(l l)v May
aifterwar(ds, n11(1the niuales were tollowed( by the sub- 6, 19.t54 aid(l April 222, 1.955. Thlis samie (colIditioll,
ad(1ult teiiiales which reached a uuia1iUIui on APril 11. lack of suiftace vegetatioil, dloes ilot prevail ill South-
rpe pereueitag(e of juveniles ainiolng the euinigrnits was west tSwam11p (Illg the Ver1l: seasoll. IHee the
erratie durii-g thie (elly Ilart of the einigr ation but area colltalilled withil the co0lifIles of the m1oont re-
iiCre 11asedl greatly towar(ls its e(nd(. It is interesting sen bles Ctma uie Ponl, but the shallow a1reats oft the
to note thlat tie seqjueince of these four classes ill endi- Swainm) supp)ort a relatively high corcenltmlatioll of
giatioii is the saniencs the (lescending order of aibun- surfa'ce vegetaItioll, including such philits as Lcc)s,
(lanec of thiese ftoir (lasses ill the overall make-ul) of Oei,.'o i(led.s, )p(irgaoa ila m ea Ci.jarpi)a _.1isi)i )a t i viale,
tle enill-(laInts. That is, the a(lult females are the 'A qittirioa ltijlia , Sp)irodela p)ol!/rl 'a anl Leia no
fitSt to eililgate in numibers, and relaltively more trisalc(a. Durling the verlial seaIsoit IllalilY of these
feniules elntigrate thian (1o ineiubers of any other class. 1h)11its, especially the gra(mminoids, (o llot project at1bove
The relitive abulndalnee ot these saine four classes the waIterl. Rlathler, tile listal .ttlds of the plaits., float
a nuoign the eluuigrailts leaving (C1ane Pond bv other 111l)01 tle suifa( cc, torm ing a imaIt. Sp1irod elp )OlI-
ro0ites is )Iesenltd(l ill Table 7. tnfortunately, these rb La hebcomes eitaligled( il this nauit ami1i l)ovitle s;
(lata alre tenlious sinlc( thie aisolute nuiubers passing addi tima I (cover.
out of Cre (ia Pond throughli thefise other egresses are A temperaiture giradienit exists withini Cra,me P'on(d.
low . Even so, it (a1ppel uIs lthat the uveniles d(1 Ilot Time thler llId va.1na1'tionls of the illel(t currellt w\vere (0omm-
leave thie 1)0on(1 by m)oving through the outlet or by 1)p1ittI to those re(o'(rded(l ait other statiolns throughout
going overlai(l, n111(1 it apl)nlrs that relatively high the Poinld. I)uringii the prevem'ial a lid vermiad1 sea sons
p)roportiOms of a(lult 1mafles nil - feles (,compared a mmI1liaxmmmllm-mnimiiinumn therilt)lleter was place(d Ill the
to the a niiuals a live in 19c53, Fig. 3) (lo utilize these waltel at e(1ach of the followillng statioms time ilnlet
othier p)antis. 'The relative Abun(lanee of the subadult stakes 57, 13, 32, and '26 (Fig. 2). 1)urnimg this
felIlales varies aI grieat deal vear to year ain(l exit to peli)(1O the i ildet reg-istere( the greatest (lifefmemce be-
exit. tweemll illmax1i mi111aimd ilii nltum temllp)eratures ill 15)
En r iron ln n tol1 e(ntn res (If C rnine Pod a(1 d11(1 sii rromtnd- of 24 timmies. Il. 5 of' the remnalinin- 9 cases, the dif-
April, 1959 POPULATION OF THE PAINTED TURTLE 12.3
terence obtained at the inlet was as great as that occurred most frequently between 12 noon and 3:00
obtained elsewhere so that there were only four days pmll.
in which the differenec between the two extreme Little or no emigration took place unless the water
temperatures at the inlet was less than that obtained temperature in the inlet rose above a certain critical
-at other points in the Pon(l. Evidently, the shallower level. This level was 7?C in 1954 an(l 6?C in 1955.
water in Fishhook MNlarshis heated imore quickly However, 8$C appears to be the lowest water teml)era-
(luring the day and cools more rapidly during the ture at which Chrysemys migrate in numbers (Table
night. As this water flows into Crane Pond a thermcal 9). The discrepancy between cloacal teltl)eratule
grladient is established which is reversible, for the
inlet current is wariner than the water in Crane Pond TABLE 9. Relationship l)etweelL the nuImi)er of enmi-
during parts of the (lay but cooler at night. granits moving through the inllet ai1(1 the teillp)erastu ie
In additioii, the difference between maxiimumn and of water at the inlet.
minimum values recorded at stake 57 was greater
than the difference for similar recordings at stake 13 Inlet Water Temperature in ?(

a total of 11 timues out of a possible 14. This indi- 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19| 20 and over
cates that at least the western third of the Pond is Number of Turtles
divided thermally. The inlet (current has the muost Spring 1954. 0 0 2 31 0 17 1 11 5 36 0 1 50 0 0 0
variable templlperature, and the water off stake 57 is
Fall 1954 ........... 2 o2 2 2 9 011 8 0 2 12 28 21 0 0
less variable than that in the inlet but more so than
that off stake 13. Spring 1955 ......... 0 2 2 4 5 2 1 12 2 11 ( 21 1 3
Fluorescein a1( lcrvstal violet added to the inlet
current in(licate(l that the water passing through the
mimainchannel of the inlet bifurcated into two branches and the teisiperature of the itiedium whichl was noticed
soon after leaving the SalixJ zone which delimnits the during the prevernal season (Table 2) was not as
mouth of the inlet. One branch continues eastward extreme in the emigrating Chrysernys (Table 10). -No
into Crane P0on(1or mOves parallel to the north shore. turtle captured in the inlet had a cloacal temperature
The (listance over which this water mnass will retain below 8.4?C. These data again indicate that these
its integrity within Crane Pond is not known, for the is a threshold, approximately 8?C, below which ena-
gration does not occur.
(lyes become dissipated too quickly. However, the
lve is evident for a( distance of 10 mnfroin the inlet
mouth. The seconid branch miade a sharp turn south- TABLE 1(0. Conitrasts ill temperature betweeni imidi-
ward and continued about 3 muoff the western shore vidulal emigrating Chrysemys al1id surroumIdisg medium.
for '20-25 m. Asterisks idl(licate animals which had beeim ini tie sir.
The influence of' wilter temnpj)erature and current on
emigration. (Chrilsemils is primarily a diurnlal ani- Water Temperature of the Cloacal Temperature of the
ial, and emigration is restricted to the daylight hours Inlet ?C Turtles ?C
(Table 8). The (lata in Table 8 indicate that the 6 8.7, 8.8
dlaily peak in emusigration (luring the vernal season, 7 8.9
measured in number of turtles emigrating per hr, 8 8.4, 8.6
8 8.5, 9.0
10 11.3
11 11.7
TABIJE8. The hourly variation, in miumber of Chry- 11 11.8
i
c/S1 msemigratingper hour, omi those days durinlg 1954 01 12 13.4
which 15 or moore Chrysemys nlove(1 westeward through 13 14.4
the Crane Pond inlet. The questiomi marks refer to data 13 13.2, 14.2, 14.3, 15.0,
for -which tile rate camimiot he accurately deterniied 15.2, 21.2*
;although somue emimigrationi occurre(d. 14 14.6
14 16.1
15 15.7
Mar. Apr. Apr. Apr. Apt. Apr. Apr. Oct. Oct. Oct. 15.5 15.8, 15.9, 15.9, 16.1
Time in hours 28 6 7 11 13 14 21 12 13 14
15.5 16.2, 16.7, 27.0*
6- 7 0.0 0.0 ? ? 15.5 16.1
7- S ....... ? 2.7 2.0 ? 3.0 2.3 15.5 16.0
8- 9......... ? 3.0 2.7 2.0 0.4 ? 3.0 2.3 18 18.7
9-10 ......... 1.3 3.0 ? 2.7 1.7 0.4 8.5 1.0 2.3 19 21.0
10-11 ......... 1.3 0.5 1.0 2.2 1.7 0.4 8.5 2.0 1.0 20.5 21.5
11-12......... 0.0 1.3 0.5 1.0 2.2 1.7 0.4 10.3 2.0 1.0
12-13......... 9.5 10.0 11.5 7.0 2.2 3.2 0.4 10.3 8.5 1.0
13-14.. . 9.5 10.0 11.5 7.0 2.2 3.2 2.0 10.3 8.5 1.0
Table 11 shows that certain telslperatures niust
14-15 .... 2.0 10.0 27.0 3.0 0.0 3.2 5.0 0.4 ? ?
15-16.. .. 1.5 7.0 3.0 0.0 3.2 5.0 0.4 be attained in both the inlet water mass and the water
16-1 ......... 1.5 0.3 3.0 0.3 0.0 0.4 within Crane 1Pond( before entigratiom. will occur.
17-18......... 0.0 0.3 0.0 0.3 0.4 During the short l)eriod in which emigration was
18-19.... ..... 0.3 0.3 0.4
19-20......... 0.0 0.0 0.0
studied in 1957 one maxiu-mn -ailminniuss thermsomuieter
was placed in the inlet ancl another just benseath the
 124
124 .r. SEXTON
( ) Wl<. N J.
~~~~~~~~~~OWE:N S ExrrON Ecological
IVol. Monograplis
29, No.2

sulrface of tlhe water wvithin Crane Pond at a point reverse(l, 11(1

1ln11d (1()
Idot exit before it is well es-
48 in east of the inilet month anll 15 in off stake 9. tbilishe(d.
Ihe water ;
'as (57 le
i deep at this position. Chrgq- WX hile the exodus of the (Cbrl/sC/i//s is corre'ited
.Pemi diot
li(1 eiiiigrate in numbers uniti.1 both the with (ertatill temllll)peatUre ('a ('teristics oft the ValliouS
Iiiiillil 111d(Iiiaxillial tenllwIratures weren bove 9-1()0C Wat'1tei' mssesT
l there are inl(Iicltionis thaif the crlIlelnt
ait bothphila e-s. itself ix (-I 1ositive force iniflueni1nc t(lli-i ItioB. N.
11188 (Islep tUt're of turtlest tromi the Po1 1was ever
T.uIIK II. The muiiiher of emigrants aiid the mininnal ()obsl-ere (xce})t during plios when wnter wv1asflow-
.11d11a 11xillal water telllmperatuies ill the ildet 1lld1ill (ilaiie iII- into the Pond tfrom0 thle test. Su1cvh occasliolls
Po011( hetweeii April 15 a1u1( April 20, 1957. A naxinxiuni- occiurtedl uriing the springs oft 195:3, 1954, 1955,
millimiiuiii tlieriiioilieter was pIlace(l ill the ililet for this .1957 (1lrin ill the fall of I1954. Ac t.l.lll!, thev Ull-
elitile p eriod, indl aiiotlher wa'S sAitnated. just leatilhtl the
(lOubted(ly take place eac(h sprlling n(1iiit those autuiiins
surf fwe of the wnatei at aI poi t iii ('ra iie Pond 48 ii
inll ii(h ate(lite tnters X(r lle Pomld. ID)lll il tleh It-
wvest of the inilet Iiioutit.
ter' pnrit
oft April, 19 55, the inllet (clltrelt was blocked(
to see if there w( S (a111 ('1a111ne ill tIe lIuIIIhes of
WATER TEMPERATURES 0C NUMBER OF
(71/F/'scml/s emigl-ratinog 01 if thci'e wnSal lay (lillli ilitiOii
EMIGRANTS
ill thea (tiVitv ot, the all111 ils a bout the inlt. Illemlotlh.
| Iilet Crane PoIid West West AMost (4 the tinlte ti (Ullrent eould not IW sto)pped,
Date -- through Over- but it x-aIs reduce(d to at least hnlf speed. Severall
Max. Miin. Max. Miii. Inlet land
ti iies theretv'ws no flow of Avatei throu11g the inlet
April 15. 14 : 3 11 2 0 0 It (a111. These experimllents Wtere vel irlyoiiliisi te
April I . 10 5 1(0 6 0 0 silnce most ot theallt liaxlIls had( alIeadv (Itt (rnle 1Polhd
April 17 1 1 8 10.5 8 0 0 so tfint too few
0 tllltles Wtr' (cla)tulred to iiake a
April 18.. 12 9 12 9 2
April 19. . 19 10 17 10 12 -13 (
generat' .lI iz(t)11t . lI oxtever, shor tly- a ter the rI-
April 20( 21 15 2 0 1_4 l_ mlov:1I of the (la I11on April 16, there WalS 8111 inCr(ease
ill the a iulliit (t aIctivitv about tle itoutli of the iII-
let aitiltOUil- 11104 of thie tulrtles s(Ieed(I to he seekillg
Fi<li ''e 4 illlusrates the reversal of the theriiial h.lskiill Sites. Trle eXperlimllelnt Iavs rep)e ted1 )ii April
i1(li'lilt wi thiini til westeiii a1 11t of C(Ial |ie Ponld. 21, -19571. rTh1is(IaV Il( Ibe } by two (Iays Oil
)ltbnpeceded
The ulilrXv tIl thil tepillpera turie 1o1 till' ilelet raittl 15(1s whlihll fa ilbtv (XtellsiVe elllira-Iti(m1 Ila (1doccl ied. Tlhe
111101t' steplyl thii that olf till' \\ltl o1ff staike 157 oi ('lille'llt ba ciked( at 8 :20 a 011 the 21st. Tilet'
til l
(o11er 11os05'1 the aittei hun iig till 111011111111.
d.Ia111 dhid IlOt inllttlfel(c ithl tile 1llOvemllelt 0ot the a1l-
ElIligiatioll is llaexie'st wiiell till' culive1 o1 till niet
11 1S illto the tir p. No (Ih/rye is wvere ill tile illlet;
teiii 1111r1tiil s ablove thiat otI the 1)1ond1 t pli1)tl iatUl.
tia p at thl, tillte; tile Awater teuupeatuve NNa8s 15 (C
e'lia s)s it iS thiis 1)11111111i i ll11 I
oI t 111111111 Ieve Isiliility
('eltajill 'V a (le'(j
I te for (illnigra tiOII. No turitlhs were
Whicli ii Ilelulnts for till 1)ank in the lill (vel oIfI lill-
found the tia p at 1 0 :35 nll ( Iwatei tetll peratuie
ill -
ia
Iti} 111 11111
(U riim ill the a ttellIooll, fo i flw turtle.s;
177 ) 11 :45 nill (Nwater tell)tlrtllle=
1101 at IC(1).
mlovt' Olit bef'ort' thet tt'lplrnt'lxirel po0h11 ityt }11s, bet't
Thre(l( Wele 4 Ch rposeiim is sulnnlling at the louth ol tie
17_ inlet a ld 6 ill the water oft the iliet aIt this laltter
tillt. rhe block was reuoved(l nt 12 :3() lil ( water
tetm)rlature = C19 ) anld rechelekei at 1 :00p)(I)), but
'5 -
there Wtlte no turtles ill the trap. One (71,ryisecmys wtis
(81)tul(1 tat 1J:130 pill (water t plleltatule = 20.5 C),
an EmyIJs bl(uidyi fit 2 :30 1)1i1 and a second 011( ry-
sennsa at 3 :45 pin (water telmlp(r.ittlrle - __ . The
N M N M N M N M N M N M wHOUR
(lataI tare XtlremlleIV illegl, rbut theyv at least arce in
w / 1M keeping- with the idea that the clurlenlt itself is fatactot I
.-s in (iriectihg the emi-atillg Chrlvisemls to IlnoVe west-
13 -
walrd.
The extent of the disper(w l mol(olmenlts. Tultless fTol
all parts of CraIe Pond enlti-late through the inlet,
:1ald the dIlrleellUlhllll (11re:t (listanlee a turtle Illust travel.
withiii the pond before reaching the inlet varies fromt
0. 111 if the anlllhil i's Cat the inlet Illouth to 315 1i1 if it
I M'GRANT 2 11 5
is At the etastrern (1 of (CraIne Pond. Ir 195-4 ?
I0 2 w
Ch ry/se/m/s wCeIre(capture(d a total of '27 tillln8 in Crane
Fim,. 4. R~elatioiishiip (If the water temperature of the Pond hefore they were again captured while cinigrtat-
iniiet (solidl line ) and that of Crane Poiid (d~ashed, line),
iIItg through the illet. The Illinill
n di icet l1(li) dis-
as11rlPlreseiltell hy the water off stake 57, to the num-
tanlce between the site (f original capture tand the
1hCr (If eiiigraints pIassinig up to the inlet. N is noon
a1-111dM~ is midlnighit. The danfti are froiii the spring of inlet mouth was 41 i11, the 1im-1?iXmum315) ill, the aver-
1955. age 15) ll. Trhe shortest time intervnl between the
April. 1959 POPULATION OF THEPAINTEDTURTLE 1

orlgitiotl ajptUre wvithiti C(atie Ponid and the capture Southwest Swanip. On May 1, 11 ChrYsemys were
ill the tilet wa^s 3 hours nil 15 iminutes (the tinilltullm coUlite(d in the Swamiip and only 1. in the Pond; Mayv
tIm-a) (distalice traVerse(l was 120 il1), the lotig'est spatn 6, S ill the SwaIIIp anrd 0 in the Pond; May 9, 3 in
was 4.3 (days (two cases: the (listaite tlaverse'( was the Swamiip and(l in the Pon(l; May 1-3, 22 ii the
65 III ill otie itistaitic anid 295 i1 ill the other), anid the Swa1111) -and 4 in the Pond.
avelage ititeival was 16.5) (lays. It became evi(lent that the (listribution of Chrq111-
Thelt is int) evideiiee thalt the plainitedlturtles closest S(?sc q ent aingin
l/. the Pn11(1 was correlateti with
to the iHilet iOUth artae the first to etitigrate. Three certaili envirotimentat-l factors. Onl April 22, 1955, aI
p)ailtted tllrtles were capitUrel two or titore tiIIties ill Iiiap of the surlfaIce vegetation of the pond wa-s (Ira n,
(Ciatie Ioidl prirl to their etlig-ratiomi. All three had(I a i(l onl April 23 the location of aIll of the Ohril yse qs
nioved awayv tfrot the tilet rather thean imoving di- observedl iII the p)o0d1( XliS superimlposed ()io this bac-k-

twesttl to thte itilet. Il (l(l

ditioll, xvhelt testedl for a ii v g,1r1oumid mai-
lj1 ( Fig. 5 ) .The lcatiomi ot pa imited
(otelatsiotil i
eteet\ tllheti the two va riables, niuniui turtles (captured or' observed dlrin' Mayx 1953 was
tia (Plitsta(te l)etweetll thel site of the orlginilal C1)tpuie
c also recoilded on tills nap. III comiiparii n, the (listribu-
witliji (talle Potllol aiol the illet touth an1ld the titte tioll of the anli tails (lulrinlg. these two vernal'l1seasomis,
ititrtvi I hetweetll the orlgi itialI (a.pture a llot the itilet it hecOities a1pp)al'.e1t thaIt there a.1re four aIrea1s. ()of
(aiptute, were showit to be itimlependlelit with r - 0.24 turtle conicenltra tion0 (conitoonolto the two utitaps. The
atio1l N -= , use of these tonui areas by (Ci 'ylesiolis aippears to be
Tiihe iautt se"as
a into which Ch 105s(e0 /s5 jourlltes relted to the fact that these arealrs Provide the onlly
firotii (i 1ltllt aIte tot the xvest a tith southwx-est a iold sunuting sites ait this tiutie of year. One sectionl, 1i-
tot thle ea.1stott (Otata Pottl. Southwest Swaitip sul)- ceate(dbv the fallen trees, haIlssshifted sllighltly over the
po its titatiV itihiolltadls ot, this species, toiitst if titetll two yea ll perl iod. Inl -195 3 there wN-ere only ai few
frtoti 0(t tie Pond. Thlis ltiecessitxtte's a jouittev ott at fallen liuiibs which xx'ler eXposed ellough1 to p)rovi(le
leaist 120 ii to iealih Sotuthwest \Sva npl) atidl a111toe- (areats. D)ulioig the spring of 1954 two large
sunu11inig0
t'oitit ( tatimothler 280 ito tot teach the titiost dista, ut oak trees topp)led into the waiter, and thiese f)m'ied(
,ollt ol thle Sva tip. Other Ohrittse(!,ys, p)teviouisly more de-sira)le sunllilng' places. A second area used
'
tor ke(l illt Crtatie Pontid, have beent reca iptnte ill the' by the Chi-'qsemoosis the Thipha bed. Tle ciuui(l)sJ
I)itelh, 1 ,0SO( ii(llowlistrea i tfrotm Cratie llPtiol a.1ild tin dea(i Typhi pr'ojeetioig ab(ove the water surface pro()-
a hillsi'de eaIst of Ca1ttail,I Marsh, about 1 000 II dowI- vided pl)atformiis suitalble for sunning. The dIeadl
stteat11i trott (tatie Pottiol. .Salil in the mouth of the inlet is the third allerl of
,
Tlie o1lispetsalofX Cit
P//seat/ s ftot i (ia ate Potld conceuitration. rthe, fourth arei, inI the xest sectiOli
seettsI to he lillited tot Soutlhlst S wait,)p, Fishhook of' the p)oid, is the bedIs of NAfooh0' (ad1(r(a whose
iNIalni4, thle D)itech, atiol Cattail MLarsIhs I )urilln the floatillg leaves plr)ovidle sunning stttions. In 1,955,
SUtIttie(t' of 1957 foulr b)o(lies otf water
peripieral to the turtles wN ere alsssoeiated with the mat;s of'
the (i tie lotlid arei, RIiddetl Laike, Islaln Ponld, filamtietitous algaie which gr e\x' alongside ai1(l within,
ButtlloPid1 atlld Sayl lake (Fig.. 1), were ilivestigateol thlei I)ltcles of Nolphor. It is noticeable that few
to see it aill\v ttarkeol a nimt-ils frotti the Cralne Poitoll C(hryscmlss were founl in those alerts lackilln, sUnrIng
a tea lot Ild ttoved itito thetti. Althiouglh the fi st threc sites. i)urin, the vernal season the banks X\'ere nlot
ata'11 are Withlill the saille (Itue lit"iagesvstetii as (t11Ittie used for sunning ais frequently as the' wx'ere dur1ing
Potld. tlere is nlo dlirect water h t li cl cotlillects the pl'eve'rdal sea soli.
tlietli to (tatiePotdl. D)uring periods of highlipie- Thie noiniuni (listalices between falrthest lpoints
(i i tatioti watet ftrot Hiddetn Lake anld Burt Pond of (ca-ipture were iiieasured for Chrj'lseols calptnlrel
flow.s itito the l)itcli after passing through extetisive two or itiore tiues withini Crane I'ond during the late
tatslhes ati(lndswattpl)s. Notie of the 17 COhr oilsettot vernail seasomis of 1953 and 1954. The limiting (lates
(altlutel ill IHiddenti Lake, 12 juveniles, 3 su.ad(lult were airbitratrily set at April 15 an(ld May 31 so that
tetlta1's, 1 aldult feltiale at(lld 1 adult male, were -1- tti tiovetentts; (lue to eiiiigration could be moore or less
mals whlic(h i lot )eet timarked l)ieviously. Nor were eliminmate(d. Nine painted turtles were caltured two
tillv of the 18 p)ainte(l turtles collected1 ill Burt ati(l or moore times during this 6-wks period in 1953; the
Ishlatd Potids from Crat-ie Pon(l. These 18 attnimals distances raug-ed from 0 Im to 301 ill aftnd aiveraoged
etisistedl of 6 juvetniles, 2 subadult females, 7 aldult 144 ill. Eight turtles were collectel dulring the same
feutiales, atd( 3 ad(lltlt mtiales. Savl Lake is 40() ll period in 19.54; the range of variation was about the
soUtthwest of Southwest SWallltp anld separatedl trotti samiae as in the previous year, I ill to 279 ill, but the
it hv a toad anh by (ldry gtound. The total dlistatice niverrlge was onlly 63 ill. Both (If these aiveraiges are
l)etweeii Crane Ponld anld Sayl Lake is ,about 800 tll. higher than those observed for the prevernal season.
Notie of the 34 Chrqlsenils captured iii S.-ayl Lake Chrysemiys begins to feed when the teniperature
were reca ptures from the cetitral population. of the water rises above' 1j50C. Anim,-als were ob-
A(tilities of the C(hrxsenoos remoining within Crane served eritimig in Crtrne P0n(1 at temiiperatnres of 16CC
IFo)I(. ()tldv a few turtles remail within Cratne PotId and(h17?C. Turtles froiti areas wNithin 1(0 kill of the
for the entilte veliatl seasoti. Counts of turtles were George Reserve were kept in captivity ill a large
titatle iii the spring of 1.954 to give some itndicatiomn 1)001 ul1(leil fairly miatural conditiomis. Fee(dinmg be-
of their relative abutd(lance itt Crane Potid in ( Otl- "nite eCxtrellely (desultorv as the falling' teniweratture
trast to their abundamice itt the northern one-fifth of rapl)Ioaehed 15?C.
126 OWEN J. SEXTON Ecological Monographs
Vol. 29, No. 2

A.

c23

FALLEN
FALLEN / TREES
TREES.

NUPHAROD2NPA

0 26
SCALE INM.
I
A. MAY, 1953 ALI B. APRIL, 1955 SA

FIG. 5. location (circles) of all individual Chrysemys observed in Crane Pond ill May, 1953 and April,
1955. Each circle represents a single individual except as indicated by the attached numbers.

THE AESTIVAI1SEASON in intensity with the emigration, for the animals

This period of the year extends from June 1 gradually filter baek into the pond from outlying
until August 31, although there may be a leeway of areas. If one assumes that the first aestival recapture
one or two weeks in either direction. As iiientioned in Crane Pond of a known vernal emigrant indicates
previously, most of the aquatic plants which are im- the latest date by whieh that individual returned to
portant to the painted turtles as loci of activity have Crane Pond, one ean obtain sonme evidence of the in-
reached the surface by June 1. The vegetation con- tensity and duration of the immigratorv movement.
tinues to increase inl weight as the season progresses The dates on whieh known emiigrants were reeap-
until the biomass of the air-dried non-planktonic vege- tured after returning to the pond indieate that immi-
tation averages at least 1300 kg per acre by amid-July. gration is a steady process whieh begins during the
The air and water temimperatures attain their annual emliglation period itself and extends into fall (Figure
imaximummiduring this season. In 1954 the air tein- 6). It is also indicated that the height of the inmi-
perature at stake 57 varied fromt a minimum of 20C gration probably occurred during the latter half of
to a mnaximumim of 35TC during this period. The July and miost of August in 1954. There is a three-
water temperatures are less extreme. In 1954 the week period in late August and early September for
minimum water temperature registered 13TC, the which there are no data.
maaximumii 29TC. In 1957 the temperature of the sur- The entire nmigratory cycle in ChriJsemjs., then,
face water (shaded) ranged from 17TC to 33TC while consists of a sharply peaked movement out of the
the temperature on the bottom of the pond in water pond during the spring (occasionally the fall) and a
53-65 eIm deep varied fromn18C to 31TC. Thus, the gradual movement back into the pond in the summer.
water temperature rarely falls below 15TC during the The peak in the former is very pronounced and
aestival season. This is the temperature at which occurs soon after the start of the exodus. There is
Chrysemys begin to feed. Renewed growth of the no distinct peak in numbers of returning animals.
animals is first noted during the first week or two of Table 3 shows the number of animals caught enter-
the aestival season and continues until late August. ing or leaving the pond during the course of the
Shedding of the lamiminae or scutes also occurs during study. It can easily be seen that most of the animals
this time of the year. caught in the spring are leaving the area, while a
The activities of the painted turtles during this higher proportion of those captured during the sum-
season are characterized by dispersal amovements from mimera-re returning. This is true not only for the
one body of water to another and by movements from inlet but also for the outlet. Actually the six ammimnals
areas of one vegetation type to another within a single miovirmgeastward out of Crane Pond in the aestival
pond as the water depth fluctuates and the nature of season of 1954 and ten of the twelve turtles leaving
the surface vegetation changes. via the inlet that samie season were stragglers frommi
Spatial reorganization of the population. There is the vernal emigration. These sixteen turtles left dur-
no mimassiliinioratiomi of Chrysernys which can comimpare ing the first three weeks of June. In the vernal
 AI)ril. 1959 POPULATION OF TIHE PAINTED TURTLE 127
NUMBER RECAPTURED EMIGRANTS areIas which had vegetation floating at the surface.
()II June 22, 1954, the ratio of the area containing
4/14-4/20 surface vegetation to the area with none was 3 :1,
while the ratio of numbers of turtles observed was
8 :1; the difference is statistically significant (Table
12).

5/26-6/1
TABLE 12. Chi-square tests for habitat preferences of
0~~~~~~~~~~~~~~~~~~~~~~ Ch rysc myis.

Per Cent
Contrastirng of Total Nurnber Number X2 P
" 6/30-7/6 Iate Situations Area Observed ExIpected

June Aquatic Vege-

22, tation Minlus
7/28-8/3 1954 Nuphar 75.5 63 53.6
6.81 P91 =6.63
Open Water 24.5 8 17.4

June Kanunculus 92. 7 39 54.7

22, 61.8 P99 =6.63
1954 Potamogenton
foliosus 7.3 20 4.3
9/22-9/28
Aug. hanunculus 88.4 134 127.3
FIG. 6. The return of vernal emigrants to Crane 13, 3.04 P95 =3, 84
Po0n(d (lurilig the summtuer of 1954. The ord(inate repre- 1954 Filanientous
snIts the Inuiiiber of emigrants algae 11.6 10 16.7
recapltured in CraIIe Poi1(1
after they l1l(l emigr-ited, 1ind the abscissa the week
June filansentous
(lurilig which aIIy emiiigranIt was first recapIture(l in the 15, algae 38.6 53 26
po01(. No collectiiig was done from August 25 until 1957 45.8 P95 =6. (3
Septemiiber 16. Potamogeton
folisosu 61.4 14 41

season of 1954 thle greater number of C(hri/sem/s

moviii- tlfiougli the outlet are headed eastward awav Chi-square tests were also imade to see if ChryI-
firoi thie pond. II the aestival perio(I the treLId denojs were distributed at ranldolil aiiion- several (
tlhrough1the outlet is reversed, for itiost of the painte(1 vegetated areas colmlpose(l of different species of plants;
turtles are now iliovinlg wetward into the pon(l. (Table 12). On June 22, 1954, itiore Chrylsemls were
Response of' tie p)oulatiotn to regetation (chan>ges. found in beds of Potaumogeton foliosus than in ones
I)urimn the afestival season of 1954 three censuses of IRanunculus lonyirostris than could he expected on
Nvere 11d(le the Chr/j semnqls in Crane Pond.
of There the basis of a randomi distribution of anillmals. How-
Were ')2) ill(lividual Ch ryjsemns (counte( on June 1 ever, on August 13, 1954, nO difference ecoultIhe shown
(l1 i~ltt the fi st celisus, 71 oII June 22 during the in the plroportionl of animals found in the areas of
Second cemsus, la(nd 152 oni August 13 (luring the final filamlerltous algae colltrasted to those in the areas of
(Celsus. This inCrTease in numbhers of turtles as the I(tonuneotl1otslonyirostris. No other tests eould be used
suIIIIIIer proglressed also ili(licates that turtles returi to( compare the distribution of ChryJsberros in different
to Cra iie I'(:l(l (luriLng the aestival period. are1.1s because the relmaillimig aireas were 'ither ra small
The census of Juiie 1 (lid iiot illclu(le iapping of irl 1 954 and did not contalitl miarly turtles. There
thie a(etual )ositi on of each ill(livi(lual, but 3 were were sigiitificaintly imiore painted turtlesI observed in
,suiiiiiii On exPose(l branches, v
were swi lliing in areas of filamlenlto(us allgle thaIrl in a1reas( of PotamoyFge-
open Water, and 17 were associated with ql(1ultie ton loliosY.s Oll Julie 15, 1 957. It is so(miewhat (liffi-
ve-ebttio((, iiiostl1 R(Jinfc ifllfls lonytirostris, Potamin o- cult to iaterplret the status o( filaimentous algae.
flo~toll (o istilssWs,andl~ I). ~trictil o lin s. Theie is aI temndemity for the 111-11 e to o(lclur isli ilower
r1iioI to tlhe Se(oIn an(Id thil(l (ensIUses in 1 954, water than (1(1 the (other plaiits. Hi owever, there' is
1h8PSp of Cr(1ie Pa1,(1 wvere 111a(e (depictinig the 1y1eas 11(1 si-iiifilant (lifferenlce in tile (listributi(Ill (If Ch rij-
of openi water a 1(1 of surface veogetattion. -In1aIddition, .se (l lS within beds of filaimlleltouus algae wx'hich aIre
(8(1 }lplant hed( W8s (described( by species so thllat :11i along shore 01' in the lli(l(lle( of the poiid.
)
(Icliate picture W.. ts flored( of tfle vegettation withii I )uring theSt1111111er (If 19.53 the surfae ( If(o CranIe
the pOnd(1:it the tiilie of each1 ensus. Tlhes position Pon1d was aI nll(sa;it (If various p)lllts and openll waIter.
o-f c(a-li turtle was superimip)osed (ol ttiese, base Iiiaps. TIl )lalInts reclle(l the surface ait (hiffterelt tiiles sO
A similzir imapwaxv,-s isimde onl June 15, 1957, anlaid 78 thalt aii idlidictioll oIf the preference by turtles for
(ChrySc(my /s were obse Ived. certa ill vege'tatio(l tyl-es coul(I be (leterllleide(l by ob-
Ini extai-iiriiiii the 1111)s, it becaitie ap)p.1C1rt tflitt servillng the miiovemiients (If turtles' i'itol 111(1 (lilt of
tlie turtles wvere not (listributeCl ait iandoimin ill reh l- (crt'al iriol r'egionsli as new l)lanlit beds (develope(l. By
tioin to tfie 1)ond1( surface but vere(clumlped1 ill those c(alipiilr'ilg- vege0tati(Il I)as with thle' distribution of
1 8) OWEN J. SEXTON E(OlOgi(alMIOngra1phs
Vo1 ,N(.2

Chrysemys captured by nietting and inuddhli, it was

possible to follow the shifts in population. centers
(FigS. 7, 8).
The vernal distribution of Chrysemys dluring 1953
0 was centered about four areas, and it is evi(lenit that
STAKE 50
T'Iypha latifol/ia and Nuphar advena were the only
aquatic vegetation utilized by Chrysemys (Fig. 5).
About June 1, 1953, Chara appeared at the surface
in the area off stakes 50-51, and Chryseoiys quickly
A moved into the bed (Figure 7, A). At the sinie timlle
the tuirtles abandoned the Naiphar which never again
was highly utilized that year except occasionally about
the periphery. The floating leaves had risen above the
water by June 19 so that the plants were emergent.
Painted turtles were not con1on. in other parts of
the pond at this timle.
RC New clumps of Chara., llixe(d with Ranca/luas,
STAKE 9 broke surface off stake 44 about June 14-16, and
STAKE44 turtles were evident in this area about June 20 and
' 0
x 0 '21 (Figure 7, B). Chrysenmys had also nmovezdintO
an area of lianunculas which had reached the surface
off stakes 8-9 (exact (late unknown), and they wexre
R,6-14 still present in anl extension of the Chara bed near
stake 50). lowever, the Chara in the area adjoining
stakes 50-51 never again attaine(l the number of turtles
B it hlad previously contained(.
Aln extensive belt of Cliara apl)eare(l at the Surface
fromt stakes 17-30 about June 20. Turtles wvere found
STAK R, C, 620 in this aIrea'I (Figg. 7 C) , but largely iln those pmortions
;0~
()f the (C/.ar whilch also eontailled a highI p)eleentage
otf Rana nca/as. Scattered patches of /ao a ii('01(s in
C %%
the a iea froni stakes 1-4 harbored soiiie turtles. 'I'Tle'
6-20 area aft Ranimcitlta and Ch(ara (Ramlnc/fas heeaiiie
C 6-2014 the iiior-(eabund(ant) off stake 44 still containedl (Irlr-
\ - X , ,tA KE
yscmij in Somilenuiiibers.
OSTAKE On July 19,IPot(an oycttoanfolioos wats lat the sur-
17
face off stakes 11-141, and ttliles so(11 appear1Tdlill this
a1'ea (igUlre 8, I)) which had Jpreviously been devoidI
of teiiti. Ainothei area o'f J* fo/iosl was at the
surlface by J ulv .12 par'allel to stakes . 52-5 3. Ch)'i-
s('IOI8 were ca ltght in this alrTca.
Ilie most inte(esting shift o(eurred in A ti-tst of
195a3. Thebelt of Clr(/ia iti the eastern porotion of
(8lalev P(id now exteanle(l froiti staikes 1.7-sl. (Fig. 8,
EJ) It had previously beell inhabited iln number s by
painted turtles only ill areas where Ranatm itlit.> o was
(o-(lOiilillant. (,n)et species of the aalg,(alI'a olily Rivu-
lalri-aReaC ( p(robably Ri ca/aria inin a (a, although
P RE 7 - ,)- R (Glocotric la niatanfls is also, found in Cr.a1ne IP1(l),
which lhad beell go'winglle' adnate to submllelged stemis
FIG. 7. D~istrib)ution of' eon1Petratio11s Of Cliryscwmys a11(1 (lebris, broke loose and floate(l to the surface.
ill rela-tionl to tile apmpearancee andl disappgearanee Of The )revafliing westerly windls swept these Jplaiits
ni, -ua--tie; vegetfintiol .lt the surface durinig .1953. Each
eastwar11(lwhere they becaelle entangle(l ill the CmI(o'a,
eire le reptl esenlts anl inldividual almimlal caught lby anly
metliodl e.xcept balitedl trapls. Solidl b)lack aI eas alre b)eds (speeially off stakes 29-31. This aeullaulationl of the
of' .Xtp~ltar advt ena; area.s encllosed<- by) dasiledl files tire rivlarliaceous 'algae in the Chaio beogan about Aug-ust
o)tlier plint beds(1. II1 e~ieli nlofiition tile letter indieti;tes 7. There ivas also al heavy influx of CrlS rsr ilso into
Hlev pblflt loenited inl tilat arleal ns f'ollown s: C8, C/lara~; P, this aretla which started no later than August 14 aln(d
Ramiancidlus longJiro~stris; P. Potanl0ogetov foliositsx; RI, (eoitinue(1 until the en(l of the month. The floatillng
riv-ulkrine(eous (i~g,-ie. Tle nlumb~ers inl enlel. nlotation refer
algae then d (Iie(l, and( the C rlyseiiils moved out of the
to tile jumoiti li:md dafy n 1),,rticulnr pbluiit lbed was first, ob)-
tile sul-ffle of tlev wffite~r. wnitli Figulre
area..
served'( a:t C:omlpare
X8. The situationl in 195)4 wsas ver y (lifferenzlt, for
 April, %159 P0PU'LATION OF TILE PAINTED TlURTIE 129

\, '4 0

\ \ ~~~~~~~~~~~~~~RI,C

C :0

/~.STAKE 82
14 '0'

1
\-0As~liO* 7,19 1 OR
I 'p SLIGHTLY EARLIER 0 %
100 %
II

0
ItI

D~~~~~~~~~~~~~
0 00
~~~~~~~~P,7-15

FIG. 8. 1)istribution of (coliceiltr-ations of Ch ryscoins in r-elaitioll to thle ai"i(earic 111ead disappearance of

a(luatie vegetltiOIl at the surface (lurilig 1953. Each ciicle repliesclilts aii ilidividual aninial c'auight b any
lie'tiO(l ex(epC t ba ite(l t-aps). Solid black areas a ie(e(lds of Aitphar (t(r1(1na0; areas eliclste )~ daslie(l lilies a ic
othert l)aint beds. InI each notation the letter iln(licates the plant located ini that area as follows: Cdashed
C, alR.
Ranitt1i(t11is lo0. firoNstris; P, IPotamooioton foliosl/;; RI, r^ivulariaccous alfl/a. The numbers in eacli 1otltioli
efer to the iiilot anid lay aI pairticiular plant bed was first ohser-ved alt the surface of the water. Coiiipaiye
with Figure 7.

(01(l( {Ihfs XvS the n l1ilt of the entiie inva ders of new masses of
pi)le(lo)lillinlt aquatic veget'Ition doliit-
1o)l(. HIeIlce, this Pl0(I'(lIle o(1 (0o1ill)arlsOl1 (OUl(l less IIttnt
are .t ie t I il,,ligralits
111
to the p)n1d(1, bIlt
not he emipl)l0Ve(l
(dil ing 1954. it is known ti alleacpture-reea 1)tule datac1tllhat soli e
Chryscti(l.s 'apipp vs to hanve 8 hilei i elly of plfetlr- (t' tile individualis wvhich licave previously h eeill resi-
(lice o01 ceit'aiill 1)1h1t t 'y)es 811( eoliditiolls wNithill (lhe t iHt ar s(lll1 lwat restrlieted a1l'e will ov11 et'i Iito
rane P11(1. Tih is is the or(der oft illerensill- )itreir- liewviv d(levclh(11d regiolIs. Indeed, if' ceit'a.ill tvl)es of
t'll(" (1l)(p waletr, floa titing N (I-(o (d((1co,1 C(hof Sp., l)iailits ale' p)rertlred over others hv (CV11qscoo,,s,this
R-((0 i(11/15 /onl Qirostris ( 81((lie or1 with (o11r/mo sp.), hellmlviOl \rXv (l hbe expected. The stiulinli lrespollsilie
tilhimllitolls alglox, h
rlXIla'iCe(' With (l/ol) Sp., 81(1 for i li t i Oa
tiigl li'eint out of olc carlea
a'c Id il[to
Pot(e 1lc()!to()1 f0i/osols. H0(il(('(1cof((s, filainentoliutos lI'xlgle a ill (the( (a.Ill he l)ositivet' 0I, Lnceative In (lh ia (tel, 01
111( th e Ii vuIhiii8ee (115OUS8lgo8 1ip8p)t1 Ir 1 8bout equally 1)0tl. Thu11Is,the animal ayleative ommea.le heao'1se
ta vo)'(( :11(1 8lre so eollsid(11r11) hi(11. V). folioNs)) is thiat airte I cks a necessa lv reqluirelenet, su(h cals
po ohatbly thie p ie1f'11r1( i
t pI ut ill this p mld, und(ier1 the (l(l((h miica'cl Siloplpot, a 11(1 may restrict the aletivities,
(Oliditiml s obs1ervd(l. to a Ilothelr arlea h-,icli supl)lies the Illissilng fteature.
0(cr alt Ijit1ic p) lilts whlih (11Sll to be alvoi(le(l, at A hed of Pot(imofyeton foliosifs realle(i the soil fcee
It st (our 'n the aestiv Il seasons Studied are P'ot( oa- (if the wvater off sta-kes 11-14 ahout MIay 26, 195,4.
y/-to?) mli(t((s, 1'. (mp 0/li/fifs, 7p1/1)1( 1(th(/ oli, Cf'roto- All ni(midentifie( agelicv destroved this vegeta aftt' tioll
/)1l/l/lI) (/(c'/("fsl(/i/, eiieigeOtiit ' 01)h(/r and E/eocoris tJulv 8 so that the area was open water hy Julyv .18.
s). The T1/I/h1 sectioii (f the ponid is heavily popu- Fios. 9A ain(d 1(C show the successive sites of eal)-
aIted(, but this aipea 1ils to he (lue to the nailts of lila- ture of' 2 Ch r(/sei,, wxvhichhad beenn established in
telietus 8l-nae extenlling out fiomii tiel 111)h(1. p Pot)(- the wvestern pt'lart of the p)on1( prior to Maay 26. InI
a1loyeton /)fftim(l tis is plrohahi I8 Prlelferel dpi lit, hut June hoth turtles m(oved into the '. f!olilosus and sllr-
the area lovereli hv tilis speele(-s was too sii,-ill to IoiIIlidillng arei(5l1. A third (Chrysem /ls (Fkio. 9,B) was
determine the plIee it oIluepiedl ill the hierarehy. also resident in the same area Which had l)eenl occu-
Ifespn)oses oIf inl/iridll) thirt/es to eget(otion (h((ayeCs. pied hy P. Ioliost.s. The turtles shown in Fi. 9
The turtles llovino, into reeenitlv suitable area is of Iovel into the western section of the pondl after the
Crane Pond (o111ul eonceivahlv (colife troiil two source's (lestruction (of the l'. oliosius and didl not return to
nilllals whlich enli-ratedl dlulrilng the spling and have the mlidlle imimit;the indhividual in Fig. 1(IC remalined
recentlv retlurnell to the pollnd, Il alliliiis; which have nearbh at least until the miiddle of July but then
heen establishe(d iii the 1)pon(dsolmle tiiiie p)rior to the took up residence along the shore fromii stakes 44-48.
appearance of the surface vegetationl. Many of the The shifts iii the honme ranges, or activity area, of
18() ()xvv:N ,J. SEXTON dIIOgic
al
.NEMoiio-,raphis
F.>
Vol. 29). No. 2

these aummials were coreilted( with the fo-rmation and endl of Agmust. By late June a painted turtle (Fig.
(lestructioi of certaini vegetation units. 101) wNasalready( established in an area of Ran aun-
c-ahis and( Chara( off stake 26, but it move(l into the
\SCALE IN M. region off stakes 29-31 some time in August although
nothinig, appeared to have change(d in the original
)@ R |TA~TKE_14 aiea. After the disappearance of the rivulariaecous
algae this imdividual moved westward away from the
8,8t}-0 3 6-28
former areas, going into the central section of the
pon(l before returning on or before october 3 to the
eastern part. It seems noteworthy that this turtle
(lidi not movet into the region off stake 30 before the
rivulariaceous algae were present even though the
Chara wv.ais-.Irea(lxr at the surface, aioll tilat it left
the area after the rivularitaceous algae (lisappearel
altlhough the Chara was still present. It is also
interesting to note that in the latter part of Septemn-
bler it remained in and near a be(l of l'. foliosus
P,PTR R FA
wIichl relachcd( the surface about August 21, but which
9,I0-12 6,8-6
I ,4-17;2,4-2 5, 7-16 (lisapp)eare(l about October 4. In October it returned
to the eastern part of the pon(l.
IN 4TH It is easy to imagine that animals just returning-
A. JUVENILE to Crane Pond encounter these v'arious areas by
SEASON chance as thev move in. Less clear is the method by
which residlent animals become aware of (lifferent
SCALE IN M.
areas in Crane Pond other thanl those in which they
\' 0 26 are locate(l. Presumably all established aninial is
6 -2 1 soMIewhat sedlenttarv as long-- as its reqluiremnents are
\ I tei-t5,
1(t lv its imnimeliate environiiciit. As examples, all
the turtles in Figs. 9 and(l 10 remained in the one
*' !O t4,6-9 g 14
,8-21 areal w'ithout moving ilntO a(ljacent a reas. When
\s --", \ /ffi~~12, 8-15 they (di(l leave, they sented to leave for good, an(l
\ ~ ~~~~ 15,99-25
exotlus was correlate(l with ain environimental chalge.
5/
<i~~~~~~~10,8 - 4 Iloweverl, mity (lata indicate that (iChi/scm!is somietimies
iakes extialiinital jOUrneys. 1)uring one 18S-dlay
6, 6 2 8; 8,7-16 I, Ierio(l in niid-suinmer, a nmale occupie(l a small area
7,7-16 a%
within the middle part of the pond, left it, movin-
eastward, an(l then returned. The shift was evi-
Z,5-16 (lentlv not aassociated with any known vegetational
chaiige. This type of movement could( indicate that
16,10-9 3,6-6
an occupied area lacks sone essential, or at least
1, 4-7 preferred, feature, an(l thy deficiemicy acts as a,
B. FEMALE IN POST-I5TH stimulus. The response is movement out of that
area. If a more favorable site, e.g., one containinrg
SEASON necessairy sul)port elements, is located, the turtle re-
FIG. 9. Loentions ((irclesn of two Chrysemys in
maillns hiere. If oillv less favorable ar eas are en-
relation to aquatic vegetation during the aestival season
of 1954. Sequences are, shown hy the first numnber ill coillitere(l, it is conceivable that the tur tle woull ie-
the notaition; the remniajing figures in the notation refer tullrn to thle original area or coiitinue wanlering until
to the month imld(la1,ys of aphture. The vegetation types a favorable ole was found. Ill the aestival periodl it
a11reill(licatte( by letter as follows: Chara; R, Raom.uii- ap)p)e'ais that the structum(e of the v-egeta tioii is the
colas ion yirost'is P, 1Potalloyctonl folio.is; ll, rivul- featiture of tlhe habitat iiiost iiiflueitial in (letermililln
irianceous algae. (lis~tib)ltioll

1nuinbem's
Th1e of Chr ilsebnqs which slift the site
A slightly iliffereiit case is preseate(l in Figure
of thleir activ ities within (CrallPeII(l(Illluing the
101). A strip) of Ch/wr sp. had reache(l the surfaces of
aItstival seasoln is iiot known Imr certaiin. Since ther-e
the wa-iter off stakes 29-31 before Aunoust, 1953.
About August 7, colonies of on(eAof the rivulariaccous is nlo iil1)i tial niethodl fIor judginl whether or nmot
alolac, which had been attached to submer-e(l stems ain alil mal's travels alre all w ith4ii the b)oul(ls; of a:1
throug-hout the 1)o01(l, floated to the surface 111(1 single activity area, (an a rlbitr-a stanl(lr(l was used
w ere arNied eastwa dl( )bv the wiryi. The rivulari- to g(aill a Ilrouh estimiate of thle number of aninlals
wh-11iel
Icestric;t tleilr activities to Ole l'fllo e thlroulglIolt
('e(oUs (algae heca(imie'ent.aimgle(l ill the eXp)ose(d portions
of the (Cmra off stalkes 29-31. and formed a, fairly tIle aestivxll seasomi. Tlleme wN-ere22 turtles which
fle'llse mal1t,there. This conditiomi remoaiime(l until the wi'('l capture(l 4 01r imore times (dii'-ing either the
ApI~ril.1959() ISPOPIULATIONOF TILE PAINTED fURTLE 131

SCALE IN M. succeeele(l Iy clatures within one locale; 3) shiftillg,

0 26 if the turtle had been sedentary within one area but
5,66-29 had later shifted the site of its aIctivities to another
locale. Using this arbitrary (hisstifitication, approxi-
7,7 -17
iatelv 20', of the aniiiials were restricted in their
6,6 -3 0 4,6- 21
moveitients, another 2()0s, had extralimital movements,
STA KE 4 anld about 50-60%/ of the Chrisevos had shifting
9,8 -20
9,8200
~ ~ iSTAKE 11
- Iiioveinents.
During the aestival season of 1953, 68 painted
turtles were captured 2 or niore times within Crane
P0o11d. The minimium direct distance between the most
8 ,723 3,5-2 3
distant points of (eapture for any one turtle ranged
fromii 3.5 ma to 300 iii and averaged 89.5 iii (the
2 ,5 -2 2 4 --s_- stan(lar(l error was 15.7 iii). Fifty Chrjse;m ys were
1,4-14 colltured two or iiiore times duriiig the 1954 aestival
selsorn. The niinirnumii direct distance between the
C. MALE IN 7TH SEASON liost distant 1)oimlts of capture varied froiii 6 in to
2)92 im and averaged 91.2 iii (the standard error was
2 ,I-1 STAKE26 16.7 iii).
1,6-27 Features common to plants ntilized by Chr.semqs.
A comaparison of features comnion to the several
3 ,8 -23
species of plants which are preferred by Chrysemys
RIC , many reveal factors to which painted turtles respond.
The llaits iiiost often selecte(l by turtles included
8
STAKE3I Potamoogeton foliosls, Ran ue11fwkul1us
lon girostris, and
filanientous algae of several species. A second gfroup,
so(iimewhat less )referre1 alnid consistinrg of Leersi(a
6,9-12 oryzolleides with Spirodela polyrhiza anid Chara sp.
with rivulariace(us algae, were conipared to this first
group. The plants of these two groups shared several
10,10 -3 (characteristics. A third lot of lplants consisted of
uiiused species anld species initially used but later
C abandoned, such as Chara, sp., Ceratophqllum demer-
8,9-28_ -_ \
9,928 ) stun, Nmphar a(idena and Typha latifolia. These
plhints were comiipared to the first group to establish
-
7 99 - 20 *P, ' \
outstaiidiiig differences.
Other authors have suggested that the structure
of the vegetatioii inhabited by animals is a feature
I (
which they select (Beecher 1942), and the plants in
Crane Po01d were examiine(1 with this in aiind. Struc-
'SCALE IN M. - -
ture in this l)aper follows the definition of Dansereau
0 26 (1951) : "Structure is the organization in space of
the individuals comiiposing a vegetation type or asso(ia-
tion."
D. JUVENILE IN IST One structural feature coimmon to both used and
abandoned plants is the mat of leaves or thollli at the
SEASON water surface. Turtles avoided areas of open water
Flni. IU. Loc.utioms (circles) of two Chrysemys in if otherwise suitable submertrged plants were ill those
rehi tioii to 8(1mi8tic vegetuitiom durhiig the ilestival sc.ison so111ie almoos, amid thev also aivoide(ld regions in which
of 1954. Seqiiences ;i re shownv by the first nuiiiber in
11 the 1)10mits l)oj'ecte(l beyonol the sufo ie so that no
tlie iotfiltioll; the lei illilig 111111 ic rs refer to the 1iioiith
nid (hiv of e .iture. The veget~ition typies are in dic.-ited
sut ce III,-tfolrmied(l.
bY letter Is follows: (C, Chal; R, Ralincll iulas 10u(1i- The preferred )lomits- possessed onl( e character
rostris; P, IPothamoo('tol foliosus,; Ifl, r'iVulv.'in.icet'oUs w Ii1ch was evideiit when the beds were observed in
ilgo'.c.
., 1t.9 vertic-al cross-section. These )lhint iiisses could be
cmisiilere(l to hotveo 'F-.stiuiitiie. The horizontol bar
1953 or the 1954 tiesti vii seuisoii. Anps sioilig the of the T -ejl)teseiits the dense iiat of terminal leaves
oiiits -at xwii i each iiilivi dunil wns (eaptilledl wver flootimig fromii the surface downwardl for about 10 ecii,
eXuilililid, (iil1 tiii siasoll's lilovelllits for each iliiii- while the verticail b)ar rel)reseits the isolated stetis in
viiiual Were (classified n-,s follows: 1) restrictei, if the the lthlerwise Ol()ei water heiieatli the miiat. Since
enphirtie polilts were all w-ithiii aI sing-le limoion'eIemisO the mtllia steiis branlch out into great miass'es of leaves
part of ( nine 1Pond; 2) exttnilil itnfi, if a sing-let (listailly, there is miuch greater milass near the surface.
("iaIptuLe oiitsiiie ofi ,-I restriited area wa5 )(((l
prei d a tid This arIa migeii-eiit is miioditied for tilhimmintous algae,
1 32 OWEN J. SEXTON
OWEN 4. Ecological Monographs
SEXTON ~~~~~~~~~Vol.
29, No. 2
for most of the filaments are at the surface. Some of (l)ansereau 1952). P. foliossus increased from 3
of the thalli are attached to submuerged plants and (small, wvell concentrated colonies; local coverage 41-
(lebris, and others are submerged in shallow water 60%) in the spring of 1954 to 5 (pure colonies; local
so that a T-structure is attained here also. Actually eoverage 81-100%) in the summer with otily small
this vegetation type is better compared to a table, amounts of Ranunculus present. The P. folios/Is de-
with the top the thick mat of surface vegetation, and elinued an(l was replaced by many other species, but
the legs the rooted steims. the area was never again inhabited by CI/rysemils in
Another common feature of the structure of the ally numbers. A typical Rainncultus longirostris
preferred species is the porous nature of the surface stan(l was scaled at 5 by late May, 1954. The IRonun-
niat. The finely dissected leaves of IRanunculus colus continued to be (lominant throughout the aestival
longirostris, the ribbon-like leaves of Potamogeton season, but some filamentous algae appeared in. the
Joliosuis, and the filaments of the algae have little areas of Ramnnculus by July 4 anrd Ceratophyl laia
in common individually, but when each species grows demersum by August 24. The areas of Routiaacullis
inl dense aggregations, the floating leaves become en- continued to be inhabited by Chrysemijs all summer-.
tangled to formi a mat strong enough to support Adleantuges of the T-structure. InI plants wit h3 ;a T-
turtles. However, this mat is composed of so many structure the mat of leaves floating at the surface pro-
sub-units that it retains a certain lace-like quality. vi(les a platform over which all but the largest Chryj-
Leersia ory4zoides, with Spirodela polyrhiza en- scmy/s can move. The turtles are not comipletelv out
tangled in it, forms a mnodifie(l T and is evidently of the water, but at least half of the shell is eXpoied
use(l by turtles during the vernal season. The gramni- .1and the head may be easily exten(led into the air.
noid is not emergent at this time, but floats on the At the 5same time the animal is in water (lep ellough
surface of the water forming a thin mat composed of so that when the head is extend(led straig-ht forward
linear leaves. Spirodela engages in these surface or (lepresse(l, it is beneath the surface. The turtles
leaves, increasing the cover. forage in this position and food may be swallowed
(7ha'ra, an abandoned lplant, has a cross section easilv.
which is radically (lifferent fromt the T-structure. Thle pOO(US and yielding nature of the mllat per-
The strandils of the algae fill miost of the space be- minits easy access to the surface fromnm benieathi .ai(1
tween the bottoml anld the water surface an(1 resist elasy egress fromii the surftace to the deeper water so
1m1ovellmietthrough thilel. Henlee, (hora niasses tend that an alar11d(1 (ir/semoys milly (live quickly throulgh
to he cuboi(lal an(l fairlv solidl. NAa1)har adl 'na, the malt anld swill awNayl1' ill thie (lealed regionll bellen.eth
another al)-iii(lone( l lant, has a T-structure when the the maudt.
leaves float at the sullface, hut each leaf is a sing-le, Tfen)mi//atures taken onl August 24, 1954, inlhieated
in ige, cirnla m'ele aent ich contrasts stronglv with thlat the sum1face te'lllplc'aturc of the w\ater' ill l)lait
the leaves of the pireferred )lalnts. beds was within 10C of the' .lir teml)elaltuile. I Io w-
(,., ,topllo le(}eisa11w, eigenerallyan1 aban)llen(e ever1,? the waterl just above the bottoiti wa 30C to(
)h mnt diii ing the perio(d of stu(lv, is coluana'- r1 inl 90 C cooler thanr the surfaced water. Tlis lr(''elrOVl
shalle when growing a ttache(l to the substrate as it of emol water could provid'e a shelter (dl mg i lperndlls
(hwes in (Crame Pond. At tin0es( this p)lant foills imoats. of ligh surface tlll)emeratules. Inll conltrIst, observa-
at the sntuace when the stemis are longer thanr the tiOmIs mitad(e()011 Septemtiber 24, 1954, iii a bel of fila -
aiter is deep). However, the sublmlerged portion of nemito;is algaze inlllicate tlmt this wa 11111 111)1i('1 ltyeil
the pla nts is still quite colnmnar . may also be advantaeaous. The temlll)eratll'e of tills
The h(Oh aa-rivularltiaceosalgae combination of sutrface water w1as 18?C. That of the suirfaewe watert
Augulst, 1 953, is miore difficult to interl)ret. Perha-ips 6 Ill distailmt amld ill whichl there was no suit a cc ve'g-
the rivuhlavia ceOUs aIlg'1ae on to1) of the Chll0 folrmis tatiOmi iegistere(l 14.50C(. Themte was ai (/)l/n1/sC i/s
ai miodie (l T-strnicture althotughI a 2-layered( unit feeli ig at the tinnc ill the fila inentous miHiat. It ((mild1
would a ppear to replresent the situation better, the easily be that the water surfacee of the IlIat nay w.arm
Upper layer' b)einlg cmposed(l of spheres an(l the lower Ul) quickly inOughi (luring c/ooil perio(ls to permiiit
layer oft cube of dense (Ihara strands which di(d not Ch/.sc//uils to feed there while nearby aireas 11/v be
Ireach the suimfa-ice. There is also the distinct p)ossi- too ((//. It was ind-licate(d (arlier that (/Ci/Se// *s
bilitv that the turtles were fee(ling0 upon the rivulalria- lrobably begins feeiling when the water temlpllerature
(eous planllts, but this waslmever observe(l. is aboiut 150C(, so that the importance of this xZamine'r
The plants in Crane IPond(Itended to grow in pure stlatumll would inl(crlease at this critical level.
colonies until there was a (dras,-tic change in soine en- Perlhaps, imlstea(l of inlhabiting certain plant beils
vironmnentll feature, g enerallyl water level. When- because of structural advantagq,,-es, painted turtles ill-
ever'l this changii-eoccurre(ld it was followe(d by a ra-ipi(l vcadle these areas to feed upon these plants in prefer-
increase iii the nunaber of l)lailt sl)ecies foumlI in that en(ce to others. Two exeellemit studies ofi tlhe fee(ding0
a rea. However, the areas of I(mananelus and IPota- habits of (hrly.isciiys (Raney & Lachner 1942; I agler
anogeton Jolioslts in which Clh'irseoyJs were found 1943) indicate that this animial is a very generalized
were quite hoiogeneousu()u n til August. From the latter feeder. The former paper showed that volumetrically
l)art of the vernal season until the ini(l(lle of the aes- the fto// wa1s about evenly (livi(led between lplant an(1
tival season in 1954 the plants found withi aill area am/il//al sources, while the latter autlhor foun/l a slight
were scadle(l for sociability oli the basis of covera-lge pieferemIce for plant mnaterial. III any of these
April. 19.59 POPULATION OF THlE PAINTE;D TURTIE 183:I
stu(lies it is difficult to separate the plant materi,al This would seem to indicate that although Chrysemtys
elatexi per se frao the plant material aecidlentally in- is clumped in areas of surface vegetation, and within
geste(l with animail f(1oo so that the, actual volume of ainy one plant bed, the individual. turtles exert little
i)Iant foo0( may he slightly less than indicated. At social influence upon each other. Thus, in(lividuals
any rate, neither paper shows that Chrlqsemys had a, ap~pear to be (listribute(l at random inI these sub-
strong predilection for any particular food, plant or clulilps.
animal. Plants found in Crane Pond an(l vicinity Ac tiities of Chrysenqls on mats of floating reyetation.
which were mentioned as food were various species ln(lividuals of Chrysemys were commonly observed
of filamentous algae, Nuiphor, IPot(ilaogeton, Najas, feeding as they inove(l over the mats of surface vege-
ASpirodela, Lemna, and Ceratophyllw1n demersiimi. In tation. Field observations indicated a general simi-
a(ldlition I have sight record,s of Chrysemys eating larity in their actioIs, aliel a (lescription of the ac-
Chara sp. and Ranuncubls longirostris. The wide tivities of several foraaging Chrysemys should suffice
variety of food eaten, consisting, of plants hoth fav- for a general account of their behavior. A Chr lsenmlYs
ore(l and unfavored, and many animals, seemns to in the first season of growth was observed 10 :05-
miegate the idea that Chry semnys seeks certain areas 1() :30 am August 21, 1954, feeling in a patch of
because the plants therein are preferred as food. The tilanIentous algae over sonic Raanflculus lon yirostris.
method of huinting(, to be described later, also pre- The path of this aniimmalwas about 11 in in length,
eludes this belief. ai(l it generally paralleled a I5 lii section of the shore-
I)istribution 'within plaint beds. It has been shown line so that the path turned back on itself. Most of
that Chryjsemys is inclined to inhabit certain areas the feedin- was (lone in the areas of filaimentou.s
more than others, so that their (listribution is clumped. a,4sIle, andv while the turtle appeared to eat the algae
It (cala also be shown that the distribution of painted per .se on onle occasionl, iiost of the effort was (di-
tur tles within ai society of one dominant plant species recte(l towards obtaining prey. Theeeile
ju iiioved
is (clui)edl anidl not m ldomilize(l throughout the entire over or' throlugh the upper layers of the nat with the
bed 'evin though the vegetation may appear honmo- ca1ra-1p(aCe just above the sur face. The, head was ex-
ge'net'OUS. (lark & Evans ( 19J-4) have presented a tenl(led forward or sligltly (del)i'esse(l so that it was
letlho(l utilizing- dIistatice hetweeii meiarest neighbors bene'lath the water wh1len hunting. Most of' the tillme
ais ai metaiesure of the sp)acing of indi-;d(luals in a popu- the jIlvelile nosedl ahout ill the Xvteoctiotni as, it iiioved
latioll. Fdllowing this technique the (listance l)e throlluh it, biting illto certainl)atcllhes. OI one o -
tweeii nearest ie-ihlbor"s within al Pea"rls of Baa ofwlus'111?l SionI
the turtle ('aught Sontc uni(ldentified(lpre which
WaNs teaIsuure(l atied the lR values (leterminlel for their
re(uire(l albout a iinute to (crush and swallow. Aln-
(ist i bution on .Junei 2'2 ad(1 August 13, 11951. Clark other ti iiie it dlisturbed sOIiti alinil 1 liwhc
h (larted off,
& Eva us state that the va:lue oft R will 1iatige frotii 0.() with thei turtle followinig for 5-1-0 (eniIbefore (ai l)tur-
foi comi)lete ing it. A thiid ('alptuie required( 1 5-20 secoiI(Is ot'
clumtiipti through 1.() rndoim (listri-
butioti to 2.1491 for uiiitoiti
chiewiI-11'llgTe Oh I7/(Si/s l)asse(l a Smaull stick which
(lispersioln. The values
)rloje( te(l almout .-3 ('ii hove the ot thle wa teml.
ohta i lled for Oh r(/seol/s ( 0.453(3 amd 0.42504() respe- a surfailce
As it Passed beneath this sti(k, the turtle struck per-
i
tiv'elv ) ilndica1te that th1e a nlilila ls were ( l1lped within
ftiictoII'tilv aIt it, ilissed, alI(l ('olltilliued o( . This
a11eals of Rmana l(alas, a11(l the average (listance be-
s(11ils to ellitolIlize the 1111hitill techili qne e'iiq)lo ed
tweeni iea i-est neiilihor \vas 71.(itii andil 3.9 III, r-espwe-
111)01l imiost (i('caliOlI0 : exp)loratorly strikes of thle he'a1d
ti v-el . Al sitilala teeliti(ile could not be c'iiiploved
iiito Vcgetatioii to (disturl) potelitia1l pl-ev siificieitly
fol other lla
plt beds, for thlelre were iot etioughl turtles
to tiake it move, theil active puisuit.
iii them. The two R values, 0.4536 anmid0.4250, ate s
Wh|en' aI (I -/sc 8l/s1 11m 1)1ev
XOII0 at (a lidst'iiii'e
not sinitficatitltv different, indicatinlg thlalt the popula-
e(very effort wais iiade to cai)ture it. A wjuviisile was
W(Is ohal)lv respond(hing siiiiila lv- at these two
tiI ( wa5 s(ell to tove raplly ov lIer thet Sntrace of the ve-l-ta-
(i fferenit titles. tiOII ill ii liusu('ccessful attempll)t to capjitiure aii odot-ite
A (lifticulty in atiV stuldy of sI)ae( illtg of wild alli- rIe'stilno onl the vegetationi. Onl June '22, 1955, al lairge
tals is that ot1i (1 dme itot ktiow hlowNiiiuch ar-ea aliI ma1le paiiited(l tuirtle was o1)sel'ved beneath the surfa c e
atiiall or populatioti is a(ecu)yitig. This is especiallyv ill a(nI aret1ca of failly opell water aS it Swa11ii tIIwai(ls
true if th'e allil at (liestri buted thlrollulgout only Hana
ai traiisforimaiiigtemSI)eiO
(0 floa tin OII the sur-
onte ))ti)t oat a iin a ppa etlitlv hoollogeieous allrca, asi fitce. Th'le(frog julped(l awav when the turtle wa\ is1
is the (c ase with (Ch1r/semy,.;withimi the areas of Rana'a- still 20-30 cia distant. The Ohry/S;cai/s l)e'esiste(l ill
co111a1s in CraIllie Pon1(1. Clark & Evxan1s (195,5) have -following it iiito a patch of Iot a)mo getOan folio.,ms,
1)l-(t'ided a iielithod fol detttoitlintig the distributioti of where it (attetlptetd to appioach troni hticatlth. Atte'r
atiiiiias which is iltidependet of area. This technique two miliire atteniipts at capture the tu'tle g1ave Up the(
utilizes the 1)ereenItage of' reflexive pairs of i ndi- (hase. Onl the saiiie day a haige teiidile h rqisemayp
viduals, that is, the iutohbelrof palirs ea'(ch inldivi(lual was obseived huntin- frogs ill aI i iilaI failshion. The
of whllch is icar1er to the othier than to ainy other turtle was ait the sui'face (ld evi(ldeiitlv saw a siiiahi
indlimhiitital. For randoImIhilv dhistributed( populations this 1?. ca tesbeiana about 30 ciii ahead ofi it in s0iiii Ch ara
ert'i
(tage is 62.15. The paopulatioii of turtles in the spl- The turtle swain beneath the suiface to within
beds at R( n1lllWis far the Jute atlid August census. 5-10 (cmnof' the frog, but the fiog leaped aiwax'K. The
(lata had a1n3 (1
d 60.5%/ reflexives, respectively. (h rlse1oy1s siirfac(ed, saw the fiog again, aiila ti'ied to
134
I' 4 OWEN J.
OWEN J.
ecatchl it in the same way. The ranid again eluded
the turtle alld disappeared.
The following is another fairly typical example
of fee(ling activities of Chrysemyqs that also describes
the hehavior of l)ainted turtles when they come to
ulldesirable areas. This turtle, a juvenile in the third
season of growth, was observed from 11:20 ama until
12) noon onI August 9, 1954. The animal was origi-
allv feeding in Rananclus, moving at, or just be-
neath, the surface with the carapace partly exposed
at tinies and covere(l at others. Its head remained
beneath the water except when the animal breathed
or watched inte. Movement through the vegetation
was fairly steady but interspersed with frequent short
rests, a minute being the longest time taken. At times
the juvenile appeared to eat only the Ranunculus,
especially ol0(, brownish pieces. Even when feeding
on this the turtle continue(d to move about. At other
tiimmesit seemlmedto be searching for, and biting at,
unidlentified prey with rapid thrusts of the head.
Initially the turtle was located in a bed of Ranunculus
longirostris, but it soon entered aii area of open water,
a space 1 ill inl dianleter. The Chrysemys swamn
through1 this area quickly, returning to the Ranian-
('8105. It continued moving through the Ranuncitlus
umitil it caine to the margin of a Nuphar bed. The
turtle starte(1 to enter the Nuphar but then returned
abruptly to the Ranancalas. Finally the aninmal
caimme to the open water at the edge of the Ranunculus,
then it turned back into the Ranunealus, in spite of
the fact that I was now in front of the animal. The
juvenile continued in may direction, moving through
the Ranaitnculas until it becaiie wary and entered
soimmenearbv Naphar. During the entire period of
observation tlme anmial. seeimmedto avoid areas which
-re less favored by Chrqsemqs, i.e., open water and
and(1Nuiphar.
Oin other occasions it was observed that a Chry-
senJ.S, mnovinig across open water fromn one patch of
vegetation to another, generally would swim well be-
neath the surface of thmeopen water and frequently
just above the submnerged vegetation. If the dis-
tance across these areas of open water was not too
realt, the annimmal(lid not coimmeup until it attained
surface vegetatiomi. Chrlsemnys were also seen swilim-
maiming just beneath the mnat of surface vegetation. At
times they would remain there after they had become
alarmed.
Return of ClhrqsewJs to activitq areas of former
u/eaNs. It has been shown that somtie Chrijsemqs move
from one area to another during the aestival season
of any one yea m and that other turtles will reimmain
within relatively restricted areas for the entire season.
There is also somlle evidlence that so58e turtles will re-
imlain withlill, or will return to, the stame general. area
iin successive aestixvl. seasons but that other Chrqlsemqs
will not itlimabit time sammegenmeral area in two succes-
siv e sulmlimiems. The extent of this voluntary return
to aireas formmierly oecul)ie(h was deterimminmed Oil the
hasis of tile l)l)ed aestival (cal)ture points of 33
mab
O1hrysemys collecte(d during the suimmmnersof 1953 and
1954.
SEXTON EcologicalMonographs
~~~~~~~~~~~~~~~~
29, No. 2
It was arbitrarily decided that for each animal the
activity ar-ea of one of the two seasons mtiust be repre-
sented by at least two captures while the area of the
other season must be indicated by one or iuiore cap-
tures. In those instances in which the animal was
caught umore than twice, the activity area was repre-
sented by a closed figure made by drawing a con-
tinuous line tlhrough the fewest number of points so
that all of the capture points were on the line or with-
in the closed figure. Where only two capture points
were recor ded, the area was represented by a circle
whose (liamueter was the distance between the two
points and whose center was the midpoint between
themum. Single points of capture remained as such.
If, for an individual turtle, any portion of one bound
figure intercepted the figure representing the area
occul)ied during the other summer, or if a bound
figure contained the single point of capture of the
other season, the two activity areas were considered
to be coincident.
O(n the basis of the above criteria, 43% of the
turtles returned, at least for a short period, to the
sam1neactivity area occupied ini 1953. The remuiaining
57%) did not return to the same area occupied the
prece(linlg year. Actually, only 13%f of the animnals
restricted their aestival activities to inore-or-less the
sammiearea during the two summummuers.
THE AUTUMNAL SEASON
The autummunalseason, extending froami September
1 until the formumationof the permianienit ice cover, is
one of thermal extremes. During 1953 the air
temmuperaturevaried frommia maximum of 33?C to a
minimiimummof -10?C. The air temperature dropped
below ()C before Septemmuber15, 195:3, aln(l themi flu,-
tuate(l considerably. It reached a second maximiiumi)
of 29?C before it dlropped to -10?C at th, en(d of
Novemumber. The water temperature ranged fromii a
high of 27?C to a low of 0?C durillg the fall of 1953.
The permanent ice cover formned on December 13,
1953 and on December 1, 1954. Somie painted turtles
were active until December 6, 1953 and Novemuber
20, 1954.
Somite of the activities of the turtles during the
first immonthof the fall differ fromii those of the remiiainl-
ing part of tlme autumnal season. I)uring mumost of
Septeamber the animiials continue to eat, but they (lo
not (lo so later iii the autumnal season. On Septellm-
ber 20, 19.53 a Chrysernys was observe(l feeding in a
l)atchm of filamnientous algae. Thmewater temiiperature
was 2'C. Oii
) the saimiie(lay anotlher 1)aimted turtle
capturedll(l lm(partly (levoured a Rane (atesbeimln 5.3
(lii in simout-venit length. The latest date on which a
Chrlusemilsiwxas observed huntin- for food was on
Septemiiber '24, 1954 when two turtles were acetmve in
a bed of aq(jualticvegetation. The temperature of the
sul'face water was 18C while thlart of tlme wtter be-
neathi the malt was 13?C. The insulation of the upper
stratumumof water fromii tlme lower, colder one, by the
vegoetation iumatprovi(les a volumite of water whieh cami
Jtc warmmme(l sufficiently (luring the (dayi to enable
(?hrysC1111Js to feed until late in the season. Since
April, 1959 POPULATION OF THE PAINTED
C(hrysemys feeds lethargically at temperatures of
15'C and below, the presence of a warm upper
stratum of water is undoubtedly an additional factor
in influencing the animals to move into or to remain
within areas of surface vegetation.
Although the painted turtles continue to eat dur-
ing the first three or four weeks of September, they
cease growing by late August.
The spatial distribution and movements of the
painted turtles during the autumnal season differ ac-
cording to the height of the water levels of the various
bodies of water durinog the fall. During a dry
autumnal season the turtles concentrate in Crane
Pond; during a wet season those animiials which have
immnigrate(l into Crane Pon01 will emigrate out to the
surrounding swanips and marshes.
During,+ 1953 only 26.1 in of precipitation were
recorded at the George Reserve weather station for
the entire year. Neither Southwest Swamp nor Fish-
hook Marsh retained standing water during that fall
although the substrate remained moist in areas. The
only active Ch'rysenqos observed were in Crane Pond.
The behavior of the animals within Crane Pond was
still strongly influenced by the location of certaiu
plants, but the effects of the wind blowing over the
water caine to play an increasingly important role
in influencing the distribution of the turtles. The
effect of the wind and of the concomiitant tempera-
ture changes was demiionstrated on September 27,
1954, when the distribution of the turtles visible in
two areas of the eastern part of Crane Pond was
correlated with variations in certain environmental
factors (Table 13). Light, water temperature on the
TABLE13. Grouping Of Chrysemys in two contiguous
ireas with cerftain comitrastimigenvironmental features.
East end of (CranmePo0d(1o01 Septeamber 27, 1954.
TURTLE 135
probably mixed the warm upper water with the
deeper, cooler water. The portion of B which was
least inhabited by Chrysemys at the time of this sur-
vey was the site of a dense concentration of Chry-
semris during the aestival season. This was also the
area most exposed to the wind on September 27.
Thus, the preference for A, or the avoidance of B,
is correlated with the relative exposure of these two
areas to the westerly winds and to the difference in
the temperature of the surface water.
The distances moved by the animals during the
autumnal season of 1953 were greater than the dis-
tances nioved during the aestival season. The iainimum
distance between the points of capture most distant
froni each other varied from 0 to 336 ineand averaged
130 in for the 54 Chrysemys captured 2 or more times
during the autumnal season. Since many turtles emii-
grated during the fall of 1954, comparable estimates
of the extent of movements were not obtained for that
year.
Field observations indicated a general movement
by Chrysemys resident in Crane Pond from regions
of shallow water into regions of deeper water as the
weather becamnecool, and animals were relatively more
abundant in the eastern deeper part of the pond
during the fall, prevernal and early vernal seasons
than in the middle and western sections. Evidently,
for hibernation, painted turtles move into areas which
are mtiorestable thermally. As a check upon these
field observations the pond was subdivided into 3
units, and the positions of successive captures of indi-
vidual turtles were followed to see if there was any
directional trend in movement from the aestival to
the autumnal season (Table 14). The 3 subunits con-
sisted of the eastern, middle and western parts of the
pond as defined by the nmajor natural constrictions
of the perimaeter.

Variants Sector A Sector B TABLE 14.

Changes inl location of Ch rysecisys within
(Cratise Po01(1 the aestival
fromn season to the fall. The
Percentage of Total Area ........... 44.7 55.3 subi-ponisis refer to the natural sectiosis of the pond which
are formed bsy the indtlenitations of the shoreline (Fig. 1).
Number of Chrysemys.............. 45 5
Wind Velocity in Ft/Min ........... Position of Chrysemys in the fall relative
130 430 to that in the aestival season

Light in Foot Candles.............. 6000 6000

Percentage Percentage Percentage
Location in aestival Number of showing no moving moving
Temperature of Surface Water in season individuals change westward eastward
Ranunculus in 'C ........ ....... 22.0 17.0
1953
Temperature of Water 30 cm below East sub-pond......... 19 73.7 26.3 ....
Surface in Ranunculus in C ..... 14.4 14.4 Middle sub-pond ...... 14 35.7 28.6 35.7
West sub-pond ........ 32 28.1 .... 71.9

1954
bottom beneath the niats of vegetation, relative East sub-pond ......... 7 100.0 0.0 ....
aRmounts of open water annd areas containing Ranun- Middle sub-pond ...... 8 38.0 50.0 12.0
West sub-pond ........ 9 77.8 .... 22.2
(ulus were similar in the two areas. Area A was
well protected frommthe westerly wind by a peninsula
which projected out into the water at stake 38 (Fig. I)uring the fall of 1953 the amaxinsumodepths in
2), but area B was subject to the full force (of the the eastern, mid(ldle,an1( western units were 24 inI
wind. The wind velocities of the two areas were 130 (61.0 cm), 12 in (30.5 ciii), and 18 in (45.7 cIii),
ft/main, and 430 ft/min, respectively. The tempera- respectively. However, most of the eastern section
ture of the surface water was higher in area A than variedl from 12-24 in, the Ilid(ldefromti6-12 in, amidthe
in area B, 22TC to 170C, for the strong winds over B western from 6-18 in.
136 OWEN J. SEXTON' E(ologi(alMonlographls

About three-(lllarters of the ainimals caught iln the Iiles, 30.1% were subadult females, 24.4% were adult
easternil unit durilng the aestival season of 1953 re- females and 25.6% were adult males (Table 5). The
iiiainil ill the same a iea (lulirng the fall. One third sex ratio amongc all the sexually mature autumnal emi-
of tile turtles eal)tulled in the mid(lle section relmaille (,-rants was 0.85 Y Y / 1.0 S. This ratio is somllwhat
ther-e, but the other two thirds whi-ch lnove~d out were lower thaln that observed (luring the vernal emigra-
al( Ut eqJuallv divi(le(1 between those alloving eastward tions, 1.) Y Y /1.0 J iln 19554 aill(l ill 1915.5, lut it is
al4)d( tilose 11MVilig,westward. In the fall there was a higher than the ratio observed for .adult animmals in
very striking shift (721',) eastwar(l of those (Crq- the entire f)opulation. This latter ratio is 0.76 Y Y /
se k!/s which hadfo( toi ivly been (apture(l in the 1.0 S. One (liffterience between the emiiigrating turtles
weC Sten'il p)i rt of thle p)(id. of the vernlal and autuimnal seasons is that the juve-
Preeil)itatiOhi Wa15 iiiuch greater (luring 1954, amil lliles emaigrate relatively earci-ly ill the fall as contrasted
('Onis(neiUtyl the water level was higher. This to their time of emigration ill thel spri-ng (Table 6).
scemei(l to sfith)ilize tilhe (itilt)ution of those turtles ITe (liel pattern of Illovellient throutgh the inlet
which (li1 hlot eiiiigrate in the fall. There was no wxxas geiierally tile saiie (lurilig thle two sca:isols. Iln
lliovellileit out of tile eastern uiiit, laiid only reduced the fall (Table 8) tilhe grteatest rate of emitiigatioMnoc-
iovNte'Ient out of the western section iii the fall of urred(l about 110011 alndl shlortly thferefafter. lhe teiii-
1954. Ilowev-er, thle turtles from the idldlle unit left petmature of the water flowing1 through1 tilhe inlet wxIas
that -ai niost, going westwardl. Thle (lelptl of all units lOore' varailale thain the temiperattre of water at sta-
ill tle poiid was about 1X in greater than that of the tion's withill (Crame P11l(1, just as imi the vernal Season.
peievous faell. A iiiiiials winch elaiigrltel thlrough the lII a(ldltioll, therle was little muioveaent out of Crane
illet duruiglt that -;(vEsasoiin 19-54 are not iuieluded ill P.011(1 (11 those (la's whllen the illet templ)ertatture (li(l
TIal}e 14. iot exceed that of thle pond.
Ili 1954, 40.8 ii out preeipitatioil
Were( recor(dd at Ilie sarfac e vegetatiolon within CranmePond becat-me
the Gerorge Reserve weather station, ald, whilet the.'re i11u1(ldate(l with the infflux of wa (iter into tile pond~i
was no flow ol xvater 1)etweehI Crane Po1d(l and the duriing thle Ifll of 1_954. Taliking the (lepthl of xwter
Suloll Ol0llillo bodihis of walter1' (lurilig the atestival in (ira me Pond o(n August 2'3 ais aii a rh-itramy stalnd-
Sea-soi, tllheS(elattelr areas (lidl cotitatflin1sonic water 81(1( withI ai xvalue of 0 ciii, tile in(crea11iseill (lepthl can
thiroutgohout tile suiiuiiicr. Rainfall, totali.ig 8.5 in, he foll(owed thiough the course of the ('emligra.tioll.
flooded aIll of the balsilns (Uling Oc(.tober, aldl water On October 8 the depth of the watterl 1lld( incre'a1sed to
b)gt'o i1 to flow ilito (raiie PoIId ftroiii the west. This -3.5 cmi i. OnI October .1.0 it was 8.('Ill. Fllligration
influx of' wa tei was followed by, an1 emiiigationI of startell ill full onl (.)ctober 1.1. By October 15 the
( rlqsco'' s ( Table 15). (leultll
dem ()
of the it xv-ater'
\ltliff lE1'1t(
Ilad illrelased to
O'(.- 20 .13 ciiialo,
1'11,(1l it
relllaimle(l at tlitat level until Mla, .1.955. Tl'lis imlerease
T'1i.LIK, 15. The iiiiilumersof Clhry1siys emigrating ill tlle (leptl of thle wa't( 1 in (CIlll( P0o11( covered(l aa-ny
tli rough tlhe ilet to (fllne POIl(1 daring the aUtumn1lllof Ot thle b(lS of su-fla(ce vee(Y(tItitoll so) that thle Illat Il()
I 954. lon-er' floatted O11 tOpJ Of tlle, XX lt( 1. I ll ad(itioll, the
lhetlv rails 11151) l)eat t tll( ('t -1t1itll (tI IX'ol. Tlhe end

Number Number i'esult XX'11sthllxt the sui f(ac (If the

fpo)ll(l, Wili c' it still
Date Emigrating Date Emigrating conblltaille(l so)1111. siirfa tce v\({ttlilIl ap)l)t'cil1r(ld s(IIlX'Wllat
like (C'ane Pon(d (lurill, tilhe spiln before the 1a181ts
October November attaield the surface in late May.
3 ....... 1 7 ...... 4
9 ........ 4 10 ...... 1 The autuninal emlligratioIn indicattes that thle p)hle-
10 ....... 1 11 ...... 3 nomuenon of emoigration is miot limited temIllpoIrally to
11 ....... 9 13 ...... 1 the spring 11l1(1that, if the )1'roper (comistellation of
12 ....... 63 17 ...... 2
13 ....... 31 18 ...... 2 factors are present, it mllay occur at any tiue. TIme
14 ....... 16 19)....... 11 autumllnal emigration also su--ests rather strongly
19 ....... 1 thlat the exodus is not under the influence of some
22 ....... 6 sealsllnal factor, such as inc'reasing length of (laylight,
23 ........ i
24 7....... which is Illediate(l through the endocrine system.
25 ....... 2
DISCUSSIOMN
The distances llmoved by Cllrysenlys dumimig any
The auttuiiinal ciiiigatioii is vexry similar to the seasoIln seem to follow (Table 16).
certaimi )atterIls
v-enuial ones. The p)eak in tile number of aninials The average (listance traversed (luring the prevernal
emigIrating is rapi(ldv reached in both eases ( Tables seasomi is relatively constammt,41 in for 5 turtles in
4 amind15), an(ld miost of thle animals exit via the inlet 1953 anld 45 alm for 7 turtles in 1954. Admittedly,
(Table 3 ). The p)creenltae omiil)ositioni of thie (lifter- the samples are small, but they tend to bare out the
calt (lasses of turtles seems to be sinilar to the (0o11- general observations in the field. There is also little
position of the animiials emoigratinig during the spring vam'iation in distance traversed during the aestival
of 1954 and 1955 when substamitial numbers of enmii- season. The average distance between farthest points
grantits were capture(l. l)uring the fall app)roximnately of ctapture of imidividual turtles was 89.5 Ili for 68
19.91> of the emiiiramits thtrough the inlet were juve- turtles in 1953, 91-2i mn for 50 animals iii 1954, and
April. 1959 POPULATION OF THE. PAINTED TURTLE 137
TABLE 16. The extent of the spatial niovenients of the paiiited turtles, Chrgysenys picta mnarginala, caught two or
more tities
duriiig any one of the aniinual seasoiis. The (listallces refer to the ninimum dlirect distance over water
Iietweeii the two inost (listaInt points of capture for each iiidividual aiiinial.

Average Average time

Number of Range of minimum between first
animals minlimum distance Standard Length. of and last capture
Season and year captured distance in m in n error in r season in davs in days Comments
Prevernal 1954. 5 3.4-124 41 .... 21 days + 12.4 Data collected before
16 days with emigration
ice cover
Prevernal 19555.... 9. 2-92. 3 45 ... 20) 1 .4 Data collected before
eniigration
Vernal 193539....... ( ).0-301 144 ... . 46 13.1 Data collected after
emigration
Vernial 1954....... 8 1. 0-279 63 .... 46 2.5 Data collected after
emigration
Aestival 1953. 68 3.5-300 89. 5 15.7 92 28.5
Aestival 1954...... 50 6. 0-292 91.2 16.7 92 30.3
Aestival 1957..7 6 -276 85.8 . ... 92 32. 1
Autumnal 1953 .. . 54 () -336 130.0) 21.8 1)4 29.7 No emigration
Autumnal 1954 .... 1 11 -306 88.1 ... . 92 15.9 Emigration

1S .5 fo 7 (7/
C'rI/s/i/i Is ill 195i. TIie /1//ta foi10 tue Viioiiiielliit lreiaiiis ifavora)le, will remiiaini ii on)ie Jo-
vel'/a:ll 11-SIei/50/
I/ autlIll
t als Cire I/Olle V/Iiaxl)Ie. III allitv bmr \t'eais." Lake eIil(lota is probably al much
the velii1i l seas/lll of 193h, 9 (h r/se/i/s tlavelsed an more staible hiabitabt thimi Crane Pond.
ave
V/ae (/istal/e/ o if .144 whlile iIi 1 954, 8 traveledi Cagle (1944) eOnelud(1I on the basis of homillng
a ii avciCtia of 63 lin. Tef lil/mitedl samIples all/I the eXp)eriimtents aid(l by following the iiiovements of
litt'l'ellee ill theaverag"'".e lellgth's of tille betweenll first ike (I C /lsenos;
mIiia in 1I.lliiios a111(d i elilglan that
ali last ( 1n)tlllues, 1 3.1 (days and 2.1 da) s makes a it im.liiited turtle(s (a-s Well as( other species) inliabit a
(liffieullt to illtelp)let ti/c (liscrel ic11( v ill (listallice. A 1). Irtieula r area to whieh they wNill lreturn it released
sil jIilal disc lCptnlleypreva' ii ill tie falIl. F1iftv-fOUr elsexx-liei-e. Th1 data fior tlis eoiielusioii eoneernnigi
(Chl//sel 1/love/i 1111aVer/ge of 1:30(ill dui'iig the (Ch5rsemoqs aire rather ,sketehy siiiee thler e w-elre few
nuti/llumi/il sec/soil of 1953, hut 10 Illove/i only 88.1 ill ara(su Sitable for. Chrilseos8 inI the vicillitv. (agleI
hlullilng tie sal/e pemodl in 19'54. The fact ti/at /I/Il/V i ec ignizeel that the return of the painted(l tuiitles to
(Il/i/s 0I/S m/1//Irat fe/i ini thle fll of 1954 aln/d I/ot il thlese aiea Is 810ld1(1 n(ot lbe caiistrued as nil(tlial'ified
195:3 uillnohuhte(div aecounts fioI ti/c (disclrep'll(/y in evi(lenee for homlliang. Ie also found that somiie speeies
a V(1'al'x (list/Iiilce all/i tillie o})serwedi for tile tWl ye''ars. o)f turtles inlclu(linig C(5ysemyos maike seasoiial liiove-
(Wt c/ l's(, there is molet Valiati/ll ill various ell- ients aiXwayftromi the hoille rang1e,0s during each spring
Xl11oll/I///l.t/Ii taI'ct/s (luring these latter tw/o se/asosll ailot lbte fall.
than1/ ill the preverna I al/d aesti val1 seasolls. For il/- (ahln (19,37) mentioned observing Ch)r!'mscoy pictl
stIll le(, IllInilig ti/e pr)lexv/11al selIso/ls there I're inever ioor(]inoto in Illinois on1 laii(l (luring the summer and
/11V ili//lts //t the surface, aln/i the/lre //re / IIW/lw soe fall aind stated that their miiovemients had 1o(0 connee-
p)reselt /lul'ing the //estival. I-i contrast, X/lll/' ph)l/Uts tion with age', sex, breeding, Or hibernation.
ll/ lisc to the surf/ace (luring tue late velrnll s5e1son0 Williams (1952) observed the movenments of
(/s ill 19 )4) /1/ /llay (lisappear (lululig tile1 f/ll r11s )aiinted(l turtles at Fish Lake in southeastern Ali(higani,
(1/s ill 1954) InI addition, the w/ater level rl/(l/illS and his data show that so501 p)inIte(l turtles return
inlgIl (illling the plevesrlal seasonl/, is colnstantly (Irop- to their hoiiie range after reimioval from it. However,
pinlg dlurilng thie summ1//I/er, but fluetuates (llrin/g the WTilliamiiis nleglects to iiiention that imiost of the a reais
S l il/ / 111/ f /laI. in which he eollecte(l were the best turtle habitats in
Alti/ou-h the p)aintedl turtles inhabitil/g Crane the lake. The present author is aware of this faet
Ponlll l/Iove' f1o/1/ onIe/ area to al/oti/er hetween seas/sIll beeause lie eolleete(l tulrtles at Fish Lake ()11 several
/11 Witi/ill a sil/gle sCe/a/1 sO l le l)tpuiati(olns of Chr/f- occasions himself. It is not surprising that the Chrq-
,sCln//s observedl for various lelngthis of till/e by other senmJs returnle(l to those areas. Williams found that
WOllke/ls /Ippe//r to he imlore se/leldt/lry. Pe/Irse (1923) other Chrqlseaijs wail(lered about the lake and had no
stu/lie/i ti/e (C/ sc f/in/ys inhahitilng /i1/ arlea of 547 acres alpliarenlt hom)1e raiig(.
I/(/te// hehinId/ /i1 s/llld h/lM ill /1 bhay (f Lake Aleldota, The overall conclusion to be drawn fromi the work
Wiscollsil/. The turtles were
il/dili/(luallvn/ette/l, of these four investigators who studied Chrypsemys
i/laIke/i /11//d rele/asedl. turtles
()ine huln(lred sixtv-six )itoo 'argotiato is that indlividuals of this subspecies
wel e le//I l)tured/ /lurilng thie c(Iouse of the il1Vestigl - may or miay not restrict their activities to limited
tiO/l l)btw/ele 1917 alld 1921, al//i of these 30('ic/ had/ areas.
o/vedl /Iwav frol/I thie ilnitial poilnt of capllture /1n1d It is now essential to provide an integrate(l view
70' /I/ia l/ot. Pe/ll'ses c/licludedl ti/at ". . . painted of the factors (leternmining the spatial an(d temporal
r /Iti/er seledI/tarv anIil/I/is, and, if their en-
ttlletis /11' distrlibution of the painted turtle as represeiited by
138 OWEN J. SEXTONVo.2.N.
the Crane Pond population. The importance of cer-
tain factors, such as the need for water for a semi-
aquatic organismn,will not assume large proportion in
the following scheme. This is not because such fac-
tors are trivial, but because they are so essential to
the physiology of the ariiiials that it is understood
automatically that they figure large in determining
the distribution of painted turtles. The following dis-
cussion assumes that sufficient water is present to
support a population of Chrysemys.
The factors most constantly associated with shifts
in location by Chr!,semxs is the presence or absence of
vegetation or other structures which can support the
animals in the water. The spring emigration occurs
before the plants have attained the surface in Crane
Pond, and the fall eiiigiration occurs only when
autumn rains raise the water level to such a degree
as to inundate the miats of vegetation. During the
aestival season the turtles in Crane Pond are prac-
tically confined to those Ureas occupied by certain
species of plants. Typically these plants are those
which have a thick mat of vegetation floating at the
surface with stemnsrising from the substrate to the
mat. Shifts in activity areas of Chrysentys can be
correlated with the disappearance of some plant beds
or the appearance of others. All in all, it may be
concluded that the structure of the plant beds, their
organization in space, is of overwhelming importance
in determining the distribution of Chrysemtys. The
possible role of vegetation will be discussed shortly.
It is first necessary to stress that factors other
than vegetation can and do influence the behavior of
Chrysemys. Amiong these factors are currents, such
as that flowino into Crane Pond through the inlet,
and the temperature gradients of such currents.
Other investigators suggest that the behavior and
movements of turtles in general are influenced by
other factors. Parker (1922) found that young
loggerhead turtles ( Carettt (pretta) inoved towar(ds
an open horizon and away from interrupted ones.
Since Chrysernys can distinguish between lines which
differ as little as 1 mmiiin width (Casteel 1911), it
is certainly conceivable that the painted turtles can
orient themselves in relation to the hills north and
south of Crane Pond afl(1 to the lower points to the
east and west. Noble & Br'aslovskv (1935) showed
that hatchling Chrysemnys picta and Chelqldra serpen-
tina can detect light reflected from water surfaces
even if they cannot see the water. Terrapene carolina
can orient by using the sun (Gould 1957).
It is possible that past experience is important in
influencing the behavior of Chrysemyl. Casteel
(1911) and Tinklepaugh (1932) demonstrated learn-
ing in Chrysemijs, and the experience of emigrating
or of shiftino activity ranges could modify activities
in successive periods.
Although the factors affecting the shifts in loca-
tion of Chrljsemys have heen tentatively identifiedl, it
is essential to present a conceptualization of the in-
teraction of such extrinsic factors with intrinsic ones
as a basis for explaining the movements of the painted
turtles in the Crane Pond area. The overall behav-
Ecological Monoographs
ioral pattern is composed of appetitive and consuninia-
tory acts (Tinbergen 1951).
need
Turtles, as secondarily semni-aquaticaiiiima_-ls,
physical support to remain at the water surface. This
support is offered by the water and by aquatic plants
(or partially submerged rocks, stumps, etc.), or by a
combination of the two. It is clear from the evidence
provided in this paper that turtles will concentrate in
areas having surface vegetation. Here they canl
crawl about at the surface of the floating niats. They
can obtain air easily, yet are in water so that they
(an swallow their food. Turtles oIntop of thiesuurface
mat are exposed to the sun, and the insolation of the
water surface by the mat fromi the cold wateri pro-
vides an area thermally favorable for eating in cool
weather. Contrariwise, the cooler depths beneath the
mat can serve as refugia during torrid weather.
The behavior of a Chrysernys supporte(d by these
mats of surface vegetation differs considerably fromt
one in water containing no surface vegetation. The
supported turtles move over the mat surface half sub-
merged, half emerged, snapping at and into various
clumps of vegetation as they feed. Air is readily
available for breathing, and the animial is in the water
it needs to swallow its food. Some turtles remain
relatively motionless, evidently sunning in the shai-
low water. In contrast, Chrysemys in deep open
water rarely swim at the surface but dive down and
swinmjust above the substrate or above the submerged
vegetation. They generally move directly to areas
of surface vegetation.
This difference in behavior suggests that thie (0o11-
plete lack of support by plants or the wron- tvpe of
support plants induces a formi of appetitive behavior
in which the animal moves from place to plhcmeuntil
presefnte(l with a stimulus which rele.lses eisum-
(o
matory act or acts, generally those alluded to above.
The nature of consummatory a ivl app)etitive acts
have been discussed by Tinbergen (1951). This stimu-
lus is of a general nature in contrast to the specific re-
leasers identified in consuimmatorv acts of a social
nature. The T-structure mats of Plaints, or P)altly
submerged logs, would be examples of such generalizeJ
releasers. Such imats of mearly plant species, I? utnl-
ciulus longirostris, Potamogeton folilosus, filanllentolus
algae, all induced the (onsuLnhllatorv acts, and the
turtles in such plant beds would reniain withinyi're-
striete(d areas until the character of the area (heln-ed.
As is well known, field studies of miiostvertebrates,
in-
especially those dealing with birds and nmainmiiaRls,
dicate that the young animals have a girea-lter tenf(-
enevy to wander than do mature adults. That the
opposite is true in painted turtles is evident froni the
observations of Williams (1952) and from the p)resenlt
study. Williams found in his homing studies of the
painted turtle that the larger turtles did not return
to the site of their original capture as readily as did
the smaller ones. It was observed in the Crane Pond
studv that there was a greater tendency for the adults
to emigrate than for the young aninials to do so. This
odd exception can be explained oIn the basis of the
present interpretation of Chryseriiqs behavior by as-
April, 1959 POPULATION OF THE PAINTEI) TU RTLE 189
suiling that the heavier aniiiials need structural sup- T with thick imats of leaves or thalli representiag the
port iiiore than do the lighter ones and that the horizontal bar. The plants with this structure are:
threshhold at which the consuimiiatory behavioral pat- I'otamogeton foliosus, Ranunculus longirostris, and
teiii of traveliiig is elicited is lower in the large various species of filamentous allgae. The turtles
aaillamals than in the small ones. move about as such types appear or disappear. It
D)uiing the eiiiogrations of the vernal and autuiiinal was shown that the distribution of Chrysemiys was
.seasons this varliablel, al)p)etitive, sea1rching, pattern is clumped even within a bed of one particular plant
Se(on(larilv infilueined by other factors-thermal speeies. The activity area of individual lpainted
0i1r(heiints, water (cUrrenslts, an1n topograph)ly to name a turtles was not restricted to one locale. During any
few. The overall result is that a ninalds' niovemiients one season, Chrysemys may or may not return to
alre oriented iather thlaii random. areas inhabited during similar seasons of previous
The final conclusion is that the stability of the years.
activityv llrange of plainted turtles is (lepenldent upon a 7. The autumnal season extends from September
wide va rietv of faetors. Under the relatively un- 1 until Decemnber 1. It is not as well characterized
stable coniditionis of Crane P'ond, the turtles frequent- as the other seasons. The animals tend to move into
lv shift the locatiomi of their activities, and while areas of (leeper water. Emigration may also occur
their travels are all confined to the area from South- at this sealsoi un(der the proper conditions.
west Swallip to the easterii end of the 1)it*h, it is
S. Other authors have concluded that Cbiy'i>lscm Iys
iiil)nossible to saw that this tsentire ar11lea (constitutes
is rather sedentary. The fluid nature of the move-
the activityvtviaage, of a single aniiiial. U'n(ler more ments of the turtles in the Crane Pond area suggests
stable eoniditioIis, as is presuimiied to be the case in.
that this species is rea(lily adaptable to varying conl-
Peaie'.1st, studlv, thmeactivitv ranges of the animiials are
li tionis.
restrictedl to very limiteil areas.
9. The lack of suitable vegetation ind(luces an aj)pe-
titive behavioral pattera in. which the turtles move
aibout until they comane to an area containing the proper
1. A Iopulatioi of' apl)i)nximiatelvly1()0 indhivi(luals veget'ation ty)e.
of the painted turtle (Ch'risemils picta moaryinot Agas-
Siw watis studlie(l iln anmd aroumi(l Crane Pond on the IJI TERATURE CITED
E~dwin S. (George Rleserve near Pinckney, Michiganll. Beecher, W. J. 1-942. Nesting birds and the vegetationl
2. Turtles were cal)ture(l by various iaethods. The substralte. (Chi(ago: Chliea1go Ornotlbological Society.
following, (lata Were recor(d(l: i(lntification number, Breckenridge, W. J. 1955. Observations on the life
seX, (late, tinie, location. The turtles were released at Ihistory of thle soft-slhelled turtle Triotiyx 'crVox, w\itl
the poi at of cap turc as soon as possible. esp)ecial referenice to growvth. Copeia: 5-9.
:3. The a minimalactivitv eycle of Chr qseuol. was (Ie- Breder, R. B. 1.927. Turtle trailing: a lnewv teclhique,
Scbibd(l omi the basis of seasotial (litfeeies in be- for studying thle life Iial)its of certain testudniata.
ha viol. The followingI wIerletme Seasons studied: pie- Zoologica 9: 231 -243.
velrnal~ venia1'11,aestival, and autuiinllal. Cagle, F. R. 1944. Home range, homing belhavior and
4. The preverma inigratioll ill turtles. Mise. Publ. Mils. Zool. IUlliv.
season, Mabich 15 to Ma1c1clh,25,
Mlich. 61: 1-34.
w1as the period' durinig wxhich ChsIna out of
/s1,11/.s ca(1Inet< . 195). The life history of the slider turtle,
hliehriiatioii. There wais little activity ait this tilie.
P'e/Isctdcmys .scrip~ta troo~sti (Holbrlook). Ecol. Mlollog.
5. TlIm'vermiaI saisomi, extendiling fromI AL-rch 26 20: 831-54.
tMa :31, is tIme })peliod ill Which (h llr(/s( (/5 (liii- Cahn, A. R. 19837. The turtles of Illinois. I ll. Biol.
gra-<ltes from CranmelPOmi(to outlyinig bodies of water. MIlouog. 16: 1-21 8.
Tme miiigra:ti)mi OCCis oiilv(dlirimgl periods of highl
Carr, A. & D. K. Caldwell. 19,5(. The ecology amlll nmi-
ater, III)ost of thmea'n<limalls leatvilng, the pond through
gratioms of seai turtles. I. Results of field work iii
tie ildet. ()llte () the im-till factors orienlting em10iimCgr- Florida, 1955. Amier. Mus. Nov. No. 1
7983: 1-283.
tion through thIe inlet a1ppea'ms to be the teliipera' ture
Carr, A. & Leonard Giovannoli. 1957. The ecology and
reimni01t'elof the inlet. More a ldults e'niigr-ate aIlld mlIore
migrations of sea turtles, 11. Results of field \ork in
jilxeniles reimtin inI thmepond tlra a would be expected Costa Rica, 1955. Amer. Mus. Nov. No. 18835: 1-32.
Oni the ba-tsisof malmice. There is a1 teidenen for more
a (lilIt teaitles to (emiii Casteel, D. B. 1911. The discriminative abilitV of the
irate thliai imldes. Turtles Which
pailitel turtle. .Jour. Aniimal Behaivior 1: 1-28.
T'cIIIairid( (I in i ii1e P)ond (lurilig the vernil ' lseason
beca;iiie (olicenitra ted a1bout the four av-ailhble sunning Clark, P. J. & F. C. Evans. 1954. l)istance to iiearest
s)ots. neiglhbor ats ame1asure of spatial relationslips in
ol)opllltioiis. Ee ology 35: 445-453.
6. The aestix <1 sea soi eXtends(l fromii Jumme 1 to -. )1955. Oi some as)pects of spatial p)attern in
Au-igst :31. I)urinmithis seasoni the vernal emlimigriants biological l)olulatioils. Science 121: 397-398.
ra(~lua liv filter hack iiito (mImine Pond and become
Dansereau, P. 1951. i)escription and recordinig of vege-
(-olicentra1tc(l iII areas in which the vegetation reaches,
taltioii tio)01 ai structural b)alsis. Ecology 32: 172-2239.
hmt (oes iot p)ro Jct, beyond the surlafce. The pre- -. 1952. The varieties of evolutionary op)povtuiiity.
ferred v-egetation has at structure in the fori of a Rev. Cainaid. (le Biol. 11: 305-88.
 140 OWENJ.
OWEN J. SEXTON
SEXTON }Ecological Monographs
14() ~~~~~~~~~~~~~~~~
29, No. 2

Gould, E. 1957. Orientationl inl ')ox turtles, Terrapene Pearse, A. S. 1923. The abundance and migration of
c. carolina (Linnaeus) . IBiol. Bull. 112: 3 36-348. turtles. Ecology 4: 24-28.
Raney, E. C. & E. A. Lachner. 1942'. Summer food of
Heape, W. 1932. Emigration, migration, and nomlaldism.
ChryIserys picta marginata iii Chautauqua Lake, New
(Calmbidge: W. Ileffer n111(Sons Ltd. 369 pp.
York. Copeia: 83-85.
Lagler, K. F. 1.943. Food lhibits and economic rela- Sexton, 0. J. 1957. Notes (concerning turtle hatebllings.
tions of thie turtles of Alichigani withi special reference Copeia: 229-230.
to fish mniaiigement. Amer. Midland Nat. 29: 257-312.
Stickel, L. F. 1950. PoI)lnlltions and homne range rela-
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turtle, Cirysciys pincta bollii (Grray). Ilerpetologiea (Linnaeus). Ecol. Monog. 20: 351 -378.
lo: 137-144. Tinbergen, N. 1951. The Study of Instinet. Oxford:
Clareudon Press. 2'28 pp.
Nichols, J. T. 1939. Range and homing of individual
l)ox turtles. Copeia: 125-127. Tinklepaugh, 0. L. 1932. Mize learning of a turtle.
Jour. Conip. Psychol. 13: 2'01-206.
Noble, G. K. & A. Braslovsky. 1935. The sensory
Williams, J. E. 195-2. Ilomiuhg behav-ior of the painted
meclhaniism involved in the migration of newly hatched turtle and nmusk turtle in a lake. Copeia: 76-82.
freshi-waiter turtles. Aunt. iRec. (suppl.) 64: 88.
Woodbury, A. M. & R. Hardy. 1948. Studies of the
Parker, G. H. 1 922. Time crawling of young logger- desert tortoise, (Gophoers (aa/(Issizi. Ecol. MAtonog. 18:
lme<(1X towar(1 time sea. Jour. Exp. Zool. 36: 323-331. 145--200.