HIERARCHIAL BEHAVIOR IN THE
SOUTH AFRICAN CLAWED FROG,
Xenopus laevis Daudin

ROBERT RICE HAUBRICH

A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF

THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA
August, 1957

She vrl-ter vlshes especially to aekncnrled^ Ms ladebtedaess to the
late Dr. W. C. Allee and his vlfe, Mrs. W. C. Allee (Ann Silver), uhose ideas,
encourageiaent aad advice vere Instrumental in the productioEi of this vork.
fhe close personal relationship with both of than remains one of his most
cherished oeBories.

She writer also vlshes to Idiank Sr. B. Soffin Jbnes for taking on
the hurdsn of this program after the death of Dr. Allee, and vlshes to
adsxovled^ his indebtedness to him for eneouragem«nt and suggestions.

Qratitude also goes out to Dr. H. M. Uallbrunn -vho gave advice on
statistics. Dr. C. J. Gk>in vho suggested the e3q;>erltDental animal, Drs. L*

*

Bemer, J. H. Gregg, R. H. Waters and S. S. Wiaiberly vho criticized the
text and to many fellow studaats Including C. D. Wilder, W. C. Sloan, C. D.
Hynes, J. f . Howill and others for their discussions, critlclams and ideas
relating to the developnent of this vork.

11

■ . t'* * • •••

...«« t i • 1

, k

MKtezlals and IfethodA ^

Ssperlaattt X .«♦.•••••••••••♦♦••••*•• 3

Esperlaent H«,,##ji##fl#»«li(«»»» •• ^

Results « « 7

MSCUSSIOH OF laSSOiSBSB ..*••••••*••*««•#•• 30

gMMKY MD CQHCUJSIOii 38

l33CEtA3WKB CISBD • . • f. t # » i •♦•««#»♦**••

1»0

BIOCBRAPHKAL I3B8

INTROIUCTIOR

Hlerarchlal b«faavlor is a phenoaeam In indlvldu&ls display

certain aggressive actlcns toward one another such as pecking or nipping.
Differences in the frequency of these actions are often consistent enou^
to enable cue to rank individuals as to aggressive potential or position
in relation to each other. Bvidaace of hiexwrcMal behavior Is fomd la
such divergent parts of the animal kingdom as Arthropoda and Oiordata.

Among invertebrates such behavior has been reported by Allee and
Douglls (191*5) in the hermit crab, Pagurus longLcarpus. by Dou^s (l^)
In the lobster, Hoaarus anerlcanus, by Bardli {19^) in the Tiasp, Pollstes
galllGus, by Bwrjberg (1953) in the crayfish, Orooectes vinUs and by
Lowe (1956) in the cre^yflah, Caaibarell\is shufeldtii.

Aaong vertebrates evidence of hieararchial behavior is abundant in
the class Osteichthyes. Braddock (19^5) demanstrated it In Platypqeellna
aaculatus, as did Allee, Greenberg, Rosenthal and fraxik {19^) in the gireai
eunfish, Leponis cyanelltxs. Hewnan's (195^) work gives infonnation on intrar
and interspecific conrpetltion as well as hlerarchlal behavior in two trout
■pedes, aaloelinns fontinalls (Mitchell) and Salao gairdaeri Richardson.
Evidence of hlerarchlal behavior in the class Reptilla was reported by
CSreenberg and Noble (19^*) in the American CSxameleon, Anolls carolinensis.

Members of the class Aves probably have been studied with the
greatest frequency and have yielded most in the vay of facts and principles
of hlerarchlal behavior. Papers by Schjeldenq>-a!bbe (1913* 1935) aad Quhl

and AllM (X^) on th« dooestic tcnrl, Eeuoks (195^) m ,}tBigIe fowl,
Mtibs^hlag (19^1) on plgeoas, B«mett (1939) on ring dores, Hovard and
Anlen (19^2) on <|uall, HauBaerstram (l9'^2) on chickadees and Sabine (19^)
m JtDieoa and tree aparrova Indicate the vide cov«M0t in -^a elasa on
Tjoth dame sti Gated and vild forms. . .j, . .

Wbrk on rats "by Uhrich (1938)^ aqolrrels "by Gordon (l9kO), infra-
htanaa primates by Maslov (I9U0), chlarpanssees by Crawford (19^2) and Yerkea
(19^ ^ Rhesus laanlaay by Carpenter (19^2) all demonstrate the social
hlerardiy in the class MaBiaalla* A form of hlerarehial organlssatlaa In
htman Indlviaaal and grottp l)rtucvlar Is Indicated by HasMns (1951)* Allee
(1951), Keith (l9*^9), Sherlf (1956) and other*.

Hhile 'Uiis phenotaenon reaches Its grtfttwit eotoplexlty aoong verte-
brates (Allee, 1952) mtll recaitly no hlerarcMal behacslor had be«i re-
corded far the classes Agoatha, Chondrlchthyes or An^pliibla. Bie first breajt
111 this Bttuation occiirred vhen Allee and lUcldLnaon (l95*^) reported finding
a primitive form of this behavior in the dogfish ehark, Masteliis eaalSi a
member of the class Chmdrlehthyes. !Qais leaves A^gaa-ttxa, regarded as the
most primitive vertebrate daaa, «aA ih^phlbla, a claaa of medlm coBq?lexlty
as the oaaly two elaaaes of Vertebrates in which there has been no report of
hlerarehial b^unrior,

Prtfvious works have Indicated aggresslcn end territoriality in tSie
class Aatphibia, but not hlerarehial behavior. Martof (1953) reported on a
sort of territoriality observed among breeding males of the gre«i tros, Bana
clamltapis, based on Isolation, a type of intolerance, advertiseaoent and
fixation. He augcgeated tbat a relatively aggreaaive perioa is followed by

one of stability, lAloh ColllaB (19^) cmsldered to b« the case In oth«r
vertebrates. Pearson (19^^) found in the spadefoot toad, Scaphiopus
boIbroQki holbroolgl (Harlan), avoidance reactions smoag Individuals and a
type of tmdef ended home site territory vhlch seldom <9rerlapped others even
in crowded areas. ^Test (195^) reported the first definite case of aggresslca
In the class Auzphlhla as displayed iA dtffnat o£ ttzxitarjr "by the frog,
Phylobates trinltatis Boulaiger. Bvldence of territoriality In the bull*
frog, Rana catesbelaoa, was reported by 0. B. Ctoln (1955 )• ^ ^& vas
observed to croak and jm^ at a smaller frog In the same general area vhlch
resulted in the retreat of the smaller animal. Jn addition, other indi-
viduals vere consistently fo^{nd in certain areas located around a pool*
Qrant (l955) also reported on a case of territoriality in two species of
salamanders, Eurycea bislineatja and Hemidactyllum scutatua, based upon a
#»haniytep end a defended territory, tlie smm of defense included direct
biting attack in B. bisllneata and challenge only in H. scutatum.

Preliminary experlofints were attempted on certain species of frog«
but the problaa of classifying the actions of the animals yblle under obser-
vation was difficult. Use of Xenopus laevis Daiadin as the e^qperimentzQ.
animal was suggested by C. J. Goin (1953) of Ifalversity of Florida
because of its habits and mode of life. !Ilhe purpose of this study was four-
fold: to determine ^diether hierarchial behavior was present in Xenopus, to
analyze the phencnKUXi of kLeocarchlal behsvloor, to fill a gap in r«qi>oart8 of
hierarchial b^vlor In dlfflerent ph^ogeneHo groops and to add to 13m»
information on aggression with possible applications to human behavior.

MfcterlaXs and Methods

Frogs tised in ^bi» study weare obtained from the sttpoply house of Jay
E. Cook, CoclmeyBvllle, Maaryland and were received as adults by Cook from
South Af^ca (see Walker, 19^2)* Parlor to the observatioa period animla
vere kept in five gallon aqpiaria filled with water to a depth of ten inches.
Ihey vere kept out of direct sunlight and t«iiperatures vere maintained
between 6cPand 75° F. Bie vater was changed after each feeding or vh«i it
became doudy. At first city water which had been conditioned by four days
storage was xised, but after sane miwAl » died Xooal spring <xc stream water
was used. Bie above methods for maintaining adults are discussed in detail
by Parker, Bobbins and Loveridge (19^7)« - * " •

3Eenoms laevis was first described by Daudin (1803) and is primarily
an aqpmtic frog. Adaptations to this a^tic habit are evident in the large
webbed >'^r'H feet that peimlt rapid B»veineat in any direction, a well-developed
lateral line systeni (Slkan and Ifurray, 19^ araionia excretion in adult
(Muaro, 1953), and nasal cavities lined with sensory epithelixm that functions
In olfsotozy perception ^en submerged (Bat«r«oa^ 19$l)* ^ spite of the
aquatic habit at times the is seen ntcnrlng orerlmd or is found In clods

of BWd near dried up ponds (Leslie, I89O; Rose, 1950).

Two groups of twelve female frogs each were used in this study and
were kept in 8 - 10 inches of water at all times. ObservBtions on frogs
1 > 12, desigoated as Ebgoeriment I or Ckroup I took place in the Marine Bio-
logical Laboratory at Wbods Hole from June to August 195^* Observations on

frogB ion - 112, desismted as Sxperiment II or Oroizp II, were coorpleted duiing
the smmr aad fall of 1955 ^'t the University of Floricla. - .

. Ibqperlinent I

ftLx frogs iwre kei^t betwMSi observatioaa la MOh of two tea-galloa
aquaria and vere wlsShed once a veek duriag the eaperiaeatal period (Table l).
When animals were fir»t received certain digits on the froait limbs were clipped
to facilitate reeogaltiott. Further clipping vas unnecessary, even thou^
digits grev l>ack, ■beca\i8e characteristics like size, color pattern, behavior
and body shape varied nldaly anong iadlvidualB and awds it easy to recognise
then alter they had been observed for a time.

Pairs vere observed separately in five -gallon aquaria. Before each
series of pair observations vater -was changed and t«fo different animals vere
used. During all observations cxibes of liver, about l/h Inch cn a side, vere
plB«ed in aquaria oae at a tine as the animals ate them. Feeding occurred
cBily during time of dbserration.

^ . Activity differ«ices aiaong individuals were noted by pairing frogs In
all possible combinaticnB. Each pair vas oibaarved a total of 200 ainutes
divided Into ten twenty minute periods. Biese ofbserration periods nwre con-
ducted in two sets of five each. Periods vithin each set followed each other
consecutively, excepting short breaks between periods for lib* «aiperlmenter to
rest, but the two sets Involving one pair of animals vere not run consecutively.
3Dbie followlag infoimation vas recorded during the pair observations for each

1. Hips given. A nip vas recorded when one frog nipped or bit anotiier
and caused the other to retreat*

2* Pushes glvei. A puBh was recorded vhea oaae frog appcroacbed another
and displaced It throu^ contact. ^

3. J^^oaches* Ifunber of times each frog approached other aotloaLess
rwBiber of pair aad touched -without further activity.

k, I&toowa. Number of times the two aalaals made contact, vhlle in
maticn, vlth uo dbserTable dlffereatial betwem th«BU

5. Cuib«B of liver eaten "by eaeih ftfog and th* ordBr in iiblch each was
obtained.

6. !Ehe troQ obtaining food first at the start Of tioh 8«ri«8.

Activity differences were also noted vhaa each frog was isolated a
total of 200 minutes ^ttvlded as in liie paired sltuatlcn into two series of
ten twenty-Bdnute periods each. Olhe following infonaatiOBi was recorded for
each firog while isolatedi

1. KtBober of cttbes of liver satsn^ :

2. Hoe in seconds talaen to obtain each piece of liver.

. ■ Ejqperlment IZ ■

2h IMs eaqperlment, in order to provide better control and prevent
possible carryover of hierarchial behavior, the frogs were kept isolated in
OM ffallGBi jttrt bstuMR obMOvations. i^alred and isolated animals were ob«
sttred using nsthods similar to tluMM in Ixperlment Z. Temperatures varied
between 71° - 7^° ?. and the aniioals were welgjxed weekly during pair obser-
vatioas (OUile lA). In Experiosnt ZZ^ to dAterain* ^-Oisr poeitlcn in the
hierarchy was consist^t under different eondltlQaB, observatlaDis wer* asds
on sach "Tnima^ vhen in a groig> of twelve as well as \dien in a pair cooiblnatiai.

Since it vae li^posslble to record the activities of all the frogs in the
aguariuffi simultaneously the behavior of eaeth frog with reference to the
other eleven vas observed for a total of 200 minutes. Each flrog "was mdnr
ohservatlcEi for a total og 200 wln^iteii 9oA OfHif •ffbf.vities involving the
tingle frog "being observed at -ttie moBtent irere reecaraed. For instance, no
record was kept of idiieh animals were nipped by or received nips from the
frog under observation. Observations vere made like those on the paired
situation, namely, in ten i20HBinute periods at the rate of five periods a
(laar* 3he animals vere laolated a mtnlTiniin of twelve hours after each dally
observation •

•- •• • ■• ..■ '. • ■■ ' ' ■ ■• -

Besaltg

' ' febles 1 and lA indicate restdts of weekly weijjiings in number of
grams. Individuals in Group I varied in average veldts frca ^0 to 168
gnats, while individuals in Group II varied frcoi 37 to lUo grazas. All of
Qratx^ I (frogs 1 • 12) showed some gEdn during the experim^ital period as
did all but three frogs in Gtck^ II (frogs 101 - 112).

Vhen animals were placed together in pairs one of four different
tjGPes of ac-U.vity wu <^s«rved9 1. e. the nip, puah, approach or unknown.
These types of acttrlty ftccrlng emtacts wee recorded for each pairing and
the results are shown in t&ables 2 throii^ 5«

Besults in the nip category are recorded in Sable 2 for frogs 1 - 12,
and in liable 2A for frogs 101 • 112. Animals were ranked in order of total
nips given in all pairings. In Baperlment I this varied from UQ to 279;
while In Experim^t II it varied from 3 to ^77* lianklng of frogs for total
nips givei (totals along the bottom row) was correlated to ranking for total

nips recelred (totals along the left coUna}* Frogs X • XS v»ri«d in total
maaber of nips received from 76 to S^h. Hfhm. individual reoikl&gs in naaber
of nips given were cooipared to ranldngs in nuiaber of nips received the
results showed a ccarelation of -O.96. Frogs 101 - 112 varied in total
maiber of nips received dftom 57 to iSk, and -when individual rankings in
nuniber of nips glwi veare compered to twnliingB In nips received the result
was a correlation -<J.50. A statistically significant differeice at the
5 per c&xt level in nuniber of nips giv«i for any particular pairing is
Indicated in the tables "by an asterisk following the greater value. 3a
T&hle 2, 28 of the 66 pairings showed such a difference while In Kble 2A,
3^ of the 66 showed this differaoLce.

ItfbleB 3 and 3A axe similar in foim to the prewlous tahles "but daal
with the ^ish category. Individuals in Eaperioent I varied in total nuniber
Of poshes given fxm 85 to 5H. idxile those in Experla«t II varied in this
category from 20 to Wf» Also, IndlvlAials in Qroop 1 varied in total
pushes received trm. IO3 to 381, ^ftiile frogs in the other group varied frca
71 to 262, When individual rankings in nunbtar 9f yoabas given were contpared
to rankings in number of pushes received in RjqperliBent I Uie result was a
eoarrelation of -O.87. In 'Eaqperijma.t II when rankings were cooipared the
result was a correlation of -O.76. In Table 3, 38 of the 66 pairings showed
statistically sigaificant differences between individuals in number of pushes
glT«x (shown by an astarlBk fcOlowing the Xarsar total) while In Sable 3A,
30 Of the 66 palHngii showed mudh «ffta?«nee8, .

fobles k and kk deal with the approach category and are similar to
those described above. Individuals in Esi^rimaBt X varied in total woiber of

approaches made frm. klO to l^S vhile those in Bs^terlmeat II varied In this
category from l80 to 767 ♦ Ihdlvlchials in Group I varied in the nuaiber of
times each vas airproached £rcm a minimum of 1^23 to a wndniinn of 1207; -while
animals in Oroup n v«rle4 from 277 "to 773 in same category. When individ-
ual ranMngs In nuniber of approaches made vere conipea:>ed to rankings in appro-
aches received in Experiment X tdiae result vas a ccnrrelatioaa of -0*83, and in
Ibqperimesat II the same ccmpartsons produced a correlation of -0.88. Id.
Tfehle k, 1*2 of the 66 pairings showed statistically sigiificant differences,
indicated "by asterisks, "betWMn individuals in nianber of approaches made,
while in Table kA, 32 of the 66 pairings showed this* l . - ...

SHe untoaoua category, represented hy Tables ^ and ^ indicated the
otBlber of ttms trofft nade cmtacts that -were of equal or of no aggressive
potmtial as nearly as the observer could determine. The nttmbers of con-
taots vere the same for each meciber of each pair heoauae both Bniml.s vere
given credit vhaiever a ccaatact of this type occurred. Tbe total nvoiber of
ccmtacts in the unknown category among individuals varied from 667 to 2525
in Sscperlment I and from 511 to 2li-ll in Experiment II«

■..^> In Tables 9 and 9A total food consUKcption, as measured by cubes of
liver eatea, is recorded far each frog in each pair combination. Ihe animals
vere rsnked according to total ntaaber of evOftes eaten ov«r all pairings.
Sable 9 gives results for frogs 1-12 and shows an individual variation in
food consuaQytian from I5I to 327 unite. ISible 9Jl gtves results for frogs
101 - 112 and the food ccnstmiptlan in tMs group varied fjram 75 to 803 units.
When the rankings in total amount of food consuoed by the other eleven fjrogs
labile paired vlth each frog vere compared to rankings in total amount of

too^ eaten "by each £rog^ correlatloas q£ -0*62 atid •<Q*^ Wir* recorded,
respectively, in Sxperimaits I aad 11. A statistically «lgn±ficaiit diff-
erence "betveea £rogs ija azaouat of food ^ten in any particxilar pairing vas
indicated in TIables 9 and 9A "by an asterisk follovlng the greater valiM.
In !£able 9, 19 out of the 6C pairings showed such a difference. In Table
9A, 26 out of the 66 pairings shovtd this difference* ''^ '

Table 6 indbLoates nuadber of alps azid ptu^es that vbm given and re-
ceived by each anlTn&l vhile in a group of twelve. A total of 313 nips were
given by the 12 frogs during -tiie observation period and Individuals varied
in total number of nips given from 0 to I68. A total of 193 nips vere re-
ceived by the 12 frogs during the period of observation, and individuals
varied in the total nxaatoer of nips received flrom 7 to 36. 2he total number
of nips given by the group did not necessarily eqpial the total received
beoauae each f^ vas oS^aerved singly for 200 ni1nut<ea and only nips givan
oar rtteeived by that mSaal wv reeorAed during this pariod. Bo reeorA iMui
kept of ^dilch ftogs nipped or were nipped "bj the frog under observation. A
negative correlatioa of O.k^ was found between total number of nips given
amaag the 12 Individuals and total nips received by these same individuals.
' ' .. A total of 537 pushes were given by the group of 12 frogs and
individuals varied froci 5 to I36 in this categcury. A total of 735 pushes
vere received by all troga and individuals varied f^rcci 19 to 1^2. Sie total
number of pushes given did not equal the total ntanber of pushes reoaived for
the reasooB given above. A ne:3Eitive correlation of 0.62 was found tuttwen
total number of pushes glvaa among the 12 individuals and total pushes ra-
calved by these aame individuals. !Zhe total ntmiber of nips (given and

ree«iv«A) vas 506 uhlle the total niaober of pushes glreii emd received vas

1272. • ■ ■ " - ■ ■•

Saibls 7 soBMrlEes totsLU for tbe differeat lac^vlaraJL c«t«0orle8
ftfoa the paired sltuaticai. Freqaaicies of the categories Increased em-
sisteatly in both ea^perliaKits from a minimum at the nip to a maxtnnin at th«
unknown. Itable 8, nblch suBoarizes ftrequencies of nips and pushes for groups
of 12, also indicates an Increase from nip to push (506 nips, 1272 pushes).
SUal«s 7 and 3 indicate that an inverse relationship e^sts 'betveen £re^[uency
and aggressive iateaslty 0^ oontacts*

CoBiparisms in total amount of food consumed by frogs -when isolated,
paired and in groups of 12 are shoun in Seblea 10 and lOJL, She total amount
of food eaten by all frogs in 200 minutes Increased frm Mil ttoits \diile
isolated to UQk ^dien paired to 1199 vh'^ groups of 12. Ihe latter figure
is glgnlficantly lar^ than iiie other two valxws (P *<0«01 end<O.Of>,
respectively). ■

The number of times each f^g obtained food first at the start of a
series of observations wts rtcorded in OSable 11. la the paired situation
individuals varied from 4 to I6 in this category. When the group of twelve
animals vera compered a variation £raa 0 to U nas observed.

Individual variation in av«rage anount of tine taken to eOytalA food
•while isolated was recorded in teble 12. !I5ie frogs in Bxperiment I varied
txaa, 72 to 768 seconds vhile those animals in Esperlment II which ate varied
from to 13^ seconds, but two of the animals in Experiment H ate no food
idiiXe Isolated.

'QBable 13 shows rankings in each of the categories for frogs 1-12

These rankiass to be lised in correlatioai tests (Table l^i-) were obtained frcm

the following tables!

Hips vhile pair«d • . . . • Tables 2, 2k

Poshes ^^lils paired * fables 3j 3^

Appcroaches -while paired Tables k, kk

UaJfflown wMle paired ..*••••«»«* Sables 5> 3k

Etps while in groups of twelve ««•••* Table 6 .....

Pushes -while in fiewtps of -tarelve ..... Table 6 -

Awage -wei^t •»•••••••••••• Tables 1, lA

Food consump-tion -lAile in pairs Tables 9t 9k

Urn taken -to locate f ood vhile Isola-bed . Table 12

Table ik shovs the c<nTelR-tioa aaong all of the categories ranlaed in
T&blfi 13 for both se-ts of frogs. A measure of the sta-tistical siffiificence
of -these values also is indicated. - .

^^^^^^^^^^^

MBELOSBf IN GRAMS, MEASQRBD WKSSG EXFERIMERZAL EERIOD
FB0G3 1-12, EXFERIMSHT I (19^1^)

Wel^t. In Grams « Trog Ruaiber

Bate

1

2

3

5

6

7

8

9

10

u

12

30 Jua

131

170

105

• » •

90

66

• •

66

« •

30

k2

6 Jul

128

168

117

96

69

* «

70

• *

k6

kl

kk

13 Jul

127

172

109

* • •

98

70

• •

67

• •

k6

hi

kl

20 Jul

127

259

l0l^

128

95

68

72

72

53

k6

50

50

27 Jul

135

168

112

131*

93

73

77

71

5^

k&

3 Aug

136

172

107

137

97

69

77

67

50

55

51

10 Aug

ikz

168

109

iin

101

75

82

69

58

56

61

57

17 Aug

Ikl

167

113

lOl^

78

85

72

a

58

69

a

2k Aug

135

168

112

99

78

86

68

56

67

57

30 Aug

Vjk

112

137

103

76

Bk

70

67

63

70

60

kr, Vt,

13h

168

110

138

98

72

80

69

59

50

56

52

XABI£ U

HEZCET, IS G»AMS, MEASURED SUBIKG PAIR OBSERVAZZOSS
FROGS 101 - 112, EXPERIMENT U (1955)

101

102

103

10k

105

106

107

108

109

no

in

112

19 Jun

105

96

132

ek

7k

6k

65

k2

38

• •

SSS Jun

103

101

136

82

82

66

68

k3

kl

• *

i

3 Jul

101

106

133

83

77

60

68

k3

ko

• •

10 Jul

lOl*

no

1^3

82

75

66

78

k3

39

• •

17 Jul

101

108

139

93

77

68

78

k2

36

• •

2k Jul

98

106

138

93

78

62

76

kk

g

• •

k9

31 Jul

102

no

138

90

75

62

62

79

k9

36

55

51

7 Aug

98

116

138

lOl^

79

62

79

81

kl

33

• •

ka

Ik Aug

99

120

ll»6

96

79

71

85

85

k5

33

7 Sep

101

128

160

86

83

65

82

85

k2

31

58

JEv* NX*

101

no

ll»0

89

78

76

82

kl

37

56

kk

UN H o\H otflO «^ir\

Ql tr>CO i»t r4 b- 1- 0\ t» * 01 tn

* u\

coco t-vo rooivo-* •<*?o. ^

• H H

irwO trxirv t— OS

VO J* ONJt O • ONCO
H •

*. t

• H H H W H

57

H-sf 01 >o iiNO| t<-^ o\oo (n

5 .

I

15

1|

a

I

5

I
1

n

I

g "

I

. f ft

<3 X O ^ fl^

t

I « t

t- ^-^J> t>-^ »^ vo CU trv O O
u>v£) H <j\cb ON <5 t«-vq if\ on

H r4 H fi r4 H ri

OHOOOOO^HOO • UN

OOHH«nOOOOO • r* VO

H H H 01 01 to<n^ H ^01 O

OIMC&NOVO 'VO

* H

t-ON ijJgWOJgOjgJ^O

1^ f4 p*4 <^ ^"1 •'"4 1^

5 . •;

S| '

-I

is

8 1 :

ft
I si

I

■

I
If

I

I I I

JO 0 -

I

0\

?1

H i-l H i-l 1-4 • CO

^4 H H CM *

vo 01 «>• ON r|j ij^***;:}

#*4

iA<no\0\Oco

H

CVJ

H ^ Oi r-j irvVO 0| fO t-OO ON o.

o _

ta

i o

I

II 8

** ID

o

a

u

I

I I I
.o 041

17

01 coco \o t~ I— W

POCO CO O r-t O ^ CO ^

HHHCUHCUHHW

OOHOOmHO OJ^ ON •

: 8

3
3

I

2

H H H r-t CU H UN .^<»J ^

HOi'HHCVJH H ^

CU H VO •00 tl\ J* _ .
H H H • «OH H ^ CVi

8

I

hhhhhhhhShhh

1^

1^

^5

■i

4i

p o«

|g| «
IH I

Ml

SI

9^

o

5§

51

I I i

a

3

I

O C^ O On
^ if\Jsp r-

* H

5l :iftl^^l(8^ll;^S; 15

•« UNO

. 3- !>-^ ^ t- ITv CO v5

ra CO

I

« a >•
o o a

5

^1

« §

•d

IP

S ^

2l ^

o

1

I

I I I

1

I

2

HHHWH H * H

^ *v 5i '^^ 'lOt-itnH tn

UMfWO

o

TABES 6

mSSBBR OF NIPS AUS posses giver ASD KECEZVED BI each mXi VSES

Frog

Nips
QivmL Pteoeived

Pushes
CbLven Secttlv*cl

Rtps-Pushes
Giveu

nps-Pushtts

Becelved

101
102
103
10^
105
106
107
108
109

no
111

112
Sbtal

13

39

12

a6

7

19

35

8

53

19

13

38

Ik

89

33

1^

36

Ik

5

19

193»

' ' 537*

735*

58
19»t
30«*

g

ll

850

''.5

iff

928

# • Total nips or pushes given in this case do not necessarily ecfoal total
I nips or pushes received laecause in the grouped sitoEition fro^s were
observed one at a time. Ho record vas kept of vhicb frogs gave nips
or pushes to or received aipe or puahea ixoa. tJie particular aninal
being observed.

Ck»rx«latlax (r^y) b«tw»ca nuogibcr o^* aijpfl given and nips received vas -0,k3
Onnrtlatlon (r^y) between mafter of poaibes etvea txdt poshes received vas -0.62

SABXS 1

TREQMEHCX OF EACH CQRZACT TXPE WBEIT fROGS WERK IS PAIRS

Hugiber Observed Per Cent of Total
Type of Cmtact Experiment E:!iperlment

I n I EC

Mp
Push

Approach

1552
2691

9033
lllM

im
1890
5239
9527

6A

37.0

>5.6

28.9
52.5

mai

^ik20

laiuo

100.0

100.0

■ V.

naftlMJi OF EACH C(MEACT TZfE WHEN FROGS WERE IN GRCXJP OF THELVS

lype of Ckmtact JSmSoer Observe

.■ Hip

506

Ptuh

1272

Ibtal

1778

CNON

HH H H H H H CW • « <n 0\

^1

9

i|
11

|l

I'd II

I III

4 V*

«n cnco £^ irv o ift^ inj* • i*\

co *n cu o\eo »»i t- 0\ if\ H ♦ ro co
H HHH H*H r-j

• to

• 00

I

11

«1

|1

I

a5

I III

•EABIE 10

TOTAL rarras op tood EAaaar by frogs dukekj oBSERVATioaf time of ^

200 HINUTiSS VSSS IS0IAS3ED, PAIRED* AND ZH CSOUP OF TMELVE

Frog

Total l&dts Satea

Isolated

Paired*

Grouped

loa.

IS

3

IIS

Total

1X0

# « Average value of 200 minutes talcen from total observatioQ time of 2U00
minutes for each ftog.

•5» ' -

XABI£ IQJl

flSEASISnCAL !QSST OF DIFFERENCES AtOTG TEOS I£K>LA!IED^ PAIRED
AHD GROUPED Sllf lATICMIS COHSDERED IN TAH2 10

Ccnrparlsoci

t

P<

Isolated -Paired

0.k2

0.7

Isolated-Grouped

3.9T

0.01

Paired-Grouped

2.69

0.05

TABOE U

nUHBSR OF a?IMES EACH TBOG OBTAIMD FOOD FIRST WHE MBQB OP
A PAIR - EXPERIMBHTS I MP 11 - AND WHILE IN A GROUP OF THEL7S

SXFXRIMDiT II

Frog

I
I
I

9

10

u

12

Paired

Frog

101
102
103
10i»
105
106
lOT
108
109

HQ

112

10
13
9
8
U
10

17
Ik

15
6

3
Ik

6
i»
3
7

6

3
0
1
0
«
#

u

Total

132

130

k5

TABtB 12

ATEBAGB HlMBaSR OF SECONDS TAKES BY FROGS WHSTJ IS0LA3EEID
(TO OSCAIK EACH OF FOOD > EXFSRIXEIfIS I AHD H

Frog

Average Tlste

Frog

h.

. . -ii^: ■ . .

101

e

7»

102

3

768

103

k

68

l(A

5

7k -

105

6

157

106

I

1^

107

220

108

9

137

109

10

130

no

n

83

HI

12

118

112

Average Time

lEABES 13

RAHEDIGS OF FE«X}S IH YARIOOS CAEEGORISS
SmLRIMESTS X MB 11

Frog Hip Puah Ajfproaeh Haknown Hip Push

Grouped Food Food

Eat«i Tijae Weltftt

%

I
I
I

9

U

3.
t

I

X

I

i

I

9 9

n 20

10

9

i

U
T

1
i

s

5
9
10

u

8

6
7

EXnOPJMEHT I

8

2

12

12

1
5

1

f

1

19

u

11

10

8

9

1^

7

12

3

5

10

7

6

U

in

109
lU

i i

I
I

10 u
U 12

9 9

t

I

6
2

7
5
1
l^

9
12
11
10

EXPSRIMEaiT n

f

i

u

12
10

I

5
9

!

k

u

12
10
8
9

7
X

I
f
9

3

k

10

11
12

a

2

I

2

2

12

5
6

i

lABlX Ik

BARE CORBELmORS 07 TSS CMEOOIRZSS VTIB IBSfS IIIDICAIIlia
PBOBABILm OF CORBELATIOH Mim AS LARCS OR LABGSB
SUE TO CHANCE AUSM (Edwards, 195k)

CoQcparlsoaxs

Frogs 1-12

from 101

- 112

Correlatioa

P<

Correlatlm

P <

O.QL

0.8l

0.01

.70

.01

,ek

.01

.ko

•05

•32

.3

.77

.€0.

.IS

•a
*

•Ql

• • • •

• • • •

.71

.01

• • • •

.76

• • • •

• • • «

.97

.oe

.85

.1

.90

,6k V

.09

.87

•03.

.56

.1

.76

.8e

.03.

.27 ■

.73

.OL

.36

.83

.01

.52 :

.1

.06

.52

^

.91

.67

.02

.80

' m

,1

.80

.08

.d

.63

m

« .73

•01

.26

.10

.8

.70

.38

.3

.28

.1

KLp/Puali
Nip/Appapoada
Push/Approach
Nip/ltotaiowa
Push/lSikncnm
Approach/UbloioHii

Paired Grouped y
ItLp Hip
Puah Push

Hp '

Grouped Nlp/Ghra>i^d Push

Nip/FOod Cmsvmtptiaa
Posh/Food Consiairptiaa
Approach/Food Cosisisiipticxi
tfaJmown/Food Coasuntptloa

Hip/Food Zine
Push/Food Time '
Approach/Food SiiM
tiaiaxown/Tood !Eba«

Hip/WBiejxt

Push/Weig^it
Approacli/Wei
Uokaown/Veic^t

Food Consuniptioai/Wel^t
Food Slme/Wei^t

©ISCUSSIOH OF KESUiaS

'iOils study is coQcexned vlth three basic probLns all dealing vlth
hierarohial 'bsbxTiors the behavioral display of aggrsssion in fjrogs, indi-
vidual differences as to aggressive behavior and factors related to indi-
vidual differences in agsnMisiQti.

She statistically sigoificant eorrelatiocs that are se«a in flible
1^ vhen ranMngs of frogs in tlie nip^ push and approach categories are com-
pared support the assimptiaa that thett |j)^x«e categories represcsxt different
expressions of the same thing, namely, aggression. In addition, the
hierarchial standing of frogs 101 - 112 In -Qie nip and push categories as
indicated tiaa pali^Lngs are higjily cozrelated vlth hierarchial positians of
the MBM frogs \di«a observed in a group of twelve (Tlables 6 and lU). fhus,
frogs nipping the laost also posh most nbether they are observed in successive
pairs or as mflsxbers of a group of twelve.

Correlations between rankings in the uaknoun and aggressive cate-
gories are generally much lower than correlations among razUdngs in each of
the aggressive categories (Hkble l^i'). Hhis is evidence tlxat the unlsnown
category is different and It laay include aggressive activity of a low

level, it also indtides aotivltieB of a noai-ftggr«ssive nature.

suable 7 shovs the frequencies of the nip, push, approach and uixknoun
categories t^en frofs wur* saired. 3Sm unkncwn oatetfary oalEM up about ^0
per cent of all contaets, ■wifliat ovttr 30 per eent of all eoataets shov a
veak foim of aggression referred to as the approach, a little more than 10
per cent shov definite aggression in the push and sonetdiat less than 10 per

emt lndle«t« the aost IntaaiM farm of aggressloa, namely, the nip. Sables
6 and 8 give frequencies of the nip and push oategories vhen frogs vere in
a grcn;^ of twelve* Tbm relative frequoicles of nips end pushes are of the
MM order ^diether animals are paired or In a group of twelve. A negative
relationship exists between intensity of aggression and its freq^ieney of
eapresslon i^ther trogfi are observed in a series of pairings or in a groi^
of twelve. .

Each group of firogs dlcqplaysd a rather loose foim of hierarchy based
tQKBx differences ooDcag individuals in aggressive pot«itlal as measured by
the nip, push and approach. Sables 2-3 show the structure, arrangonent
•ad extsnd of these individual differences. Wide variaUons among frogs
are Indieated y/hm all aggresslTS aetirity is totaled for each amlnal.
XUfferenoes in aggressive activity between individuals in any one pair are
noted by cooqpering totalis for each grog within the chart* flftgr pcx* cent of
the differttxces between members of pairs are statistically significant as
indicated by asterisks in the squares of the greater totals. Ttae per cent
of sl0ilfleaatly dlfferwit totals remains consistent for both groups of
frogs tested and also for the thzve aggressive categories.

Bank oorrelation between total aggressive activity esipressed and
total aggressive activity permitted in this study is negative and statistically
significant. Frogs with the bluest aggressive potential, as measured by
sudaer of nips, pushes and approaches administered, eaperleooe the tmm9%
a^resslve advances from other frogs and vice versa. IMs sugsssts IStat
aggression Is a distinct foxm of behavior and is based upon certain behavioral
cues not detected by the observer.

c-;,.,;' ' Oi^er iafoxnatloti recorded during periods of Q^wrmfeiais Included
the follotfiug: (l) wei^ts, (2) amounts of food eaten aaid coBrparisons vh«i
fJrogs imre isolated, paired and grouped, (3) time taken to obtain food
virile iscdbted and (k) nximber of times food v&s obtained first vhen indi"
viduals were in pairs or in a grovq? of twelve. . ^.

TablA Ik aHaam correlatloas between all of the faxrbors. It Is
trca -tiwse eorrelatlaas, Ixued upoa rankings of the frogs (Table that
most interpretations regarding relatlonsJaipe are made. • <

" t Weigjit dlffereoces sd^t be thou^t of as one ejigpressloa of physio-
logical variation. When rankings in wel^t, obtained fraa Zeble 1, are
correlated with rankings in the nip, push, approach and imknown categories
the resvdts are variable (Skble 1^}. Differences noted previously betwMm
aggressive bdsavlor and the unknown category carry throu^ whezi the wei^t
factor is considered. Four of the six correlations between the aggressive
i«tgtgoirles and wel^t are statistically sl^ilfleant and the othar two are
almost 80. All are positive. Velght is considered to be a factor la
bierarchlal behavior of Umam frogs and heavier froigs tend to rank hlgber
in the three aggressive categories, niBMly, vlp, puih. and approach, yet do
not necessarily rank higgler in the uzxknown category.

ffehles 9 and 9A show how frogs rank In individual food caisuDptlaa.
differences in food consvpption are not as clear-cut as those within each
of the aggressive categories. Less than ^0 per cent of the pairings show
significant differences between frogs as Indicated by asterisks within the
tables. Also, tAien rankings of frog? in total food consvaoed (totals in
bottoa row) are caqpared to ranldnga in total food coasuned by others in

their preseace (totals In rl^t coltnm) Mbe results are Inconclusive. Group
I shows a slgEdfleantly hlgjti eoarrelatloaa, Grotrp II does not, althouf^, all
correlations are negative.

, : ^ ISmq renMngs in vel^t are ccccpared to rankings in aiaount of food
eaten the results are variable. ^Bm heaviest animals In Group II tend to
•ftt Urn most vherefts the heanrlest iu Qrouip I do not neeasflarlly •at tb* aost.
^hen ranMngs In tooA emstnptlQn are eorrelstttd tdlb thyote In three
aggressive categories and the unlmown the results are statlstiC6Llly signif-
icant at the 5 per cent level In six out of el^t cases and close In the
raaalnlng two. dere Is no T>reak In degree of correlation observed here, aa
In previous cocrparlsans, "between the three aggressive categories and the
vcbkamti, ■which indicates the more aggressive as well as the niore active
frogs eat more than the less aggressive and less active animals.

Sie srezase tljae ta]fiea l>y isolaiied aaiOAls to Iocftt« food varies
vldely anong inttvlduala (Wble 12). Qmm dlffWrwices May ^ to
variations In the chemoreceptor systems and in ability to respond to -ttiese
stimuli. When rankings in this category are cooipared to rankings in the
aggressive categories (Table ill-) the correlatians, while all positive and
significant at the 5 per cent level In Orovq} I, are positive "but of
questloaaa'ble sigalflcenee in Group H. When rankings in aioount of time
taken to locate food are ccccpared to rankings In amount of food consmed
th« coi7elation« are not consistently hi£^. Sh^refore, the xoore aggressive
firogs do not neeesswrlly find fbod more galekly, nor do the frogs eating
more while paired necessarily take the least time to locate food \iben. iso-
lated, t.; ■ , . . ■■■

IxifQaxmtim aa the nusber of tljaes each fjrog obtained food first
>)btle a noftier of a pair and vhlle In a grotq) of tvel'V« Is gtrsa In Table
U. Whidx animal obtains food first at the start of each series of ohser-
vatloos is apparently a chance phenorattion not noticeably related to any
factors previously discussed. . 'v • -

It Is possllale that the three aggressive categories represent
attempts by frogs to eat one another. Cannibalism Is veil knona In this
species especially -whan size differences are great and availability of food
Is limited (Rose, 1950). 2he behavior resulting in a alp vas observed to
be very sladlar or Identical to that leading to constnspticEi of Uver. Osily
the size and movement of the other frog semed to prevent the coanpletion
of the act of eating In the case of the nip. Ihe Increase in frequencies
of nips, pushes and approaches in the presaice of food is further evidence
sufg^orting this asstsiptloii* If the observations noted above are valid the
atteorpt to obtain food laay censtltute a camnoa basis for aggression in laaagr
vertebrates. ;•. ,• ■ ■ .

;'; A short tis» afber a tult of food Is plao«4 in an a<fiBriua vith a
group of twelve frogs the frequency of sweeping motlctts with front Iteert and
the amount of sulusnlng activity both increase as do frequencies of all types
of contacts. Hvrnx after food has been eaten the Increased activity caatinues
for 10-30 minutes. A searching type of behavior by one or two frogs in a
grovg? initiates the same sort of activity afflong the otlicrs even in the
absence of food. Sms, aetlvlty in the group can be Initiated by the behavloar
of one or two members, and this factor enhances the chances of locating food.
79^d ftLfiO la found atwi^ftr by ^harnw vlth siora tiIdmiI b ooGuj>|yis^ a oMrtaln

roluae of vater, mA In adflltion iSbm greater amount of Infiiviaaal activity
no doUbt requires tbe use of more energy. Tiahle 10 shows the compjurative
aoounts of food eaten by frogs viaea isolated, vbesa. paired and •whan In a
group of twelve. Hhe total amount of food consumed by frogs varies directly
Kith the nuniber of animals plafted togetlxir. Differences in total food con-
suinpUm betweei the isolated and pairiA iltuatians are not staUstically
Biffiificant, but tihen ftogs are observed in groups of twelve the total con-
sanption is slgilflcanUy greater than in either of the other two situaUons.
If increased food cansumptlcn is actually of poslttve value to the anJaal,
this may be an esaiQtle of cooperation or the Allee Effect (Odum, 1953) •

•Che present work has indicated the existence of certain b^wviaral
«itlties by the method of analysis. It has also attcaapted to relate these
entities one to aaother and relate thsm to certain broader aspects of
living things ouefe as iihyelology, agg»»iim, copperaUon and hlerarchlal
behavior. An atteimrt at controlled observation is used here to facilitate
an mderstandlng of behavior both In one species and as a ganeral phenomenon
of life. While one person has been studying the brtiavlor of these frogti
the data as tabulated In a asnse have reflected certain aspects of human
behavior. On the whole, correlations are hlijier in (kna^ II taaan they are
in Gxoiq* I, when rankings in the three aggressive categories and -ttie unknow
«re coogpared to each other and to rankings in other factors. It could be
that frogs in Qmapa 1 and II vere behavlorally dlffenmtj on the other hand,
as a result of continued work vlth the animals, the experimenter may have
learned to fit certain aspecte of behavior into categcxries with greater ease
in the second groi^). Assuaalng iatie latter, this probably represents greater

accuracy tn dstexminlng wt^efseSjta In Qroup II. Hofverar^ there still is a
source of danger in the metbod of analysis used in this study. Blvlslcsi of
the bdiKvioral yboHA Into gui^ increasingly aGn>Bx«at categories or entities
■i^t eneeniMgi m eibeerver to igaore relatiaBdilp? between peu^s and the
bduYioral vfaole. Cie parts versus the vhole dichotomy is one that science
luMi AM9ed £rm the ,pt«rt, because an inqportant feature of the scientific
ttetliod is the analysis of vholes into paarts. Often diehotaoies are eliffl*
Inated as more is known about a subject. For Instance, the study of heredi-ty
moA suTiroament has becosie not so imich a pcroblen of two separate things as
different aspects of the same thing. ItLkeviBe, future vork may shov that
hairarehial behavior is a special case of territoriality or vice versa.

, . Mooh of this vort: bai esntemd around liw study of tadlvldual dif «
ferences in aggression, i. e. the determination of the presence of and form
Of biararchial }»tBmriM iA this £rog. An analysis of hierarchlal behavior is
also attenipted. An increasing number of studies In animal behavior are not
concerned vlth an analysis of both the phenomenon of aggression and of the
hierairohy (Allee, Foreoan, ftuiks and Holablrd, 19?^; Braddodc, 19^(91
Bovjberg, 1956 j Crawford, 19l»0i Colllas, 19^3i Douglis, 19^) Gre«iberg, 19^7)
and Delany, 1955)* Such studies njjmamt att«spts to analyze the broader

aspects Of behavior. . -

Many European workers, especially theebhologlsts, emphasize the
miilywLa of more basic betuvrioral eatitios m is evident in the studies of
releaser mechanisms (Tinbergen, 19^^^ 19^^ 19?3), social faeilitation
(Barling, 1938j Davles, 1953)^ displacement activities (Lorenz, 19^1j
Wnbergen, 1953) and imprinting (lorenz, 1952K . ; ,

Sbie cLLff erences vhlch exist betveea these tvo schools iu their
mpfproaches to the geaeral study of behavior are also evident in their
^fffffcmahies to the study of bdiavloral evolutioa. Sils is clearly shovn
tilum the Su3^^pean viewpoint discussed by tOEtRiz (19^0) Is coopered with the
American airproach as Indicated by Allee (1952). ^Ebe Europeans attempt to
coorpare oore basic bn^Msviorttl entities aiacxig dif£to«&t «gp«ol«8 KiiyBreas the
Americans attenq^t a ccepearison of more general phenoBMna. Both astproachas
are of course useful in any study of evolutiacL be it organic or behayloral.

A study «M mdi of two groups of twelve frogs In an ftttesapt to
amalyie «sgrossicin and to drtemLne ^Aetlier inOividaaU wlIMn the gjroi^
y [lY^an iggressive differeaces resultlxig in hlerarchial behavioor.

1» Aggressioo in Xgoopus laevis Tln^^^n aiJ^Ojaditid ■behavioraUy ia one
of three ways, namely, "by the ajjproach, tflie pttflh sod the nip. These
Obtteories also represeat iaoreasing intensities of aggressive behavior
in tSie oraer given.

2* Contacts repires«iting the aore intense categories of aggresslTe
1»^hearlor occur le«9 ftsfMatiy ttm do tHoee r«(pres«atin6 Hio less inteoae
ones in 'bath the paired and grcuped situatione.

3. Large differences aoang frogs are indicated vhen aggressive
ecfcivity is totalled for each anlaal. Progs displaying 1^ aost aggressive
activity in one category tend to display the aost In the others*

k» Differences in «ount of aggressive "behavior displayed betwMtt
airftan of pairs are statistically slgplfloant in ^ per cent of the cases.
!Ihls is ccQsistent for each of Ha* aggressive categories.

ft A category '^rispn*^ as unkoown repres«a.ts contacts in "which, no
a^resslve SUfxm&m MmMI mOmXa could be detendned. She total
frequency of these kinds of contacts is as great as all other types cooibined.

6. Frogs vlth greater aggressive yoteatials, M Msured by nuaber of
nips, pushes and approaches, experieaace fewer aggressive advances flrom otiier
fro^ and vice versa.

7« Seavler frogs display aggressive behavior aoore frequoatly than

lighter caaes, yet are xxot necessarily 'tiM aore active anlasls (aotivity
referrin^ to raoMixg in the untooua catee^ry).

8. Sbe more aggressive as imU as oore active £rog& eat more thaa the
lees aggressive and less active anlneifl, ytt lAiiii renMngs in total food
coasuoed are compered to raaldngs In total food consumed by others In their
presence tiie results are sot necessarily Tal^oly correlated*

9. K> cousistoit relatlonehlps exist amoig rankinfls in fdod can-
suB^tioti, tim l&ken. to locate food viiile Isolated and vel^t*

IQ, As the nualber of anljoaals placed togttixer is increased to twelve,
the amount of food ccxisuaed "by each troQ increases. !Ihls may "be an eTBMiila
of cooperation or the AUm fiffeot*

XI* It is possl^ thttt aggresiiiit« 'behavior in these fSrogs reflevto
an attatrpt to eat (me anotberj as "behavior resulting in a nip is nearly
identical to that vhich leads to ooiisuBQitim of liver*

Allee, V. C, 1951. Cooperatlm amaag AnlTnols, vi-fch Hvinan SgpHcatloos.
IfeifwwOT, Tcnrk. 233 PP*

— , 1952. Dnmi nance and Merar<day In societies of verteTsrates; cited
in Stsnicture et Hiyslologj' des Soc?.etes Aolnales, ed. Pierre Qrasse,
(Coll. Ihtemat. Centre* Bat. Beah* Sd.i ^} 157-183- ).

and J* C. UcMnson^ 195^> Dominance and eiibordlnatlQQ. In the
amfoHi. aogflj3h, Mustelus cauls (mtchill). Pfar^'Biol. Zool., ^{k)t3^'-3l6h.

— — , and M. Doviglla, 19^5* A dotalnaace order in the hermit crab,
fliygaa IfUfi^ipHDW aey. Bcology, ^tkllAlZ*

Focrenen, E. M. Banks and C. H. Holabird, 1935* Sffeets of an

androgen an dominance and siibordlnanco In six ccoman "breeds of Galltts
ffOlus. Phyalol. Zool., 28(2)1 89-U5.

B. Gbreenlwrg, a. N. BoMithal and P. f!rank, Sons effeets of

social orgBnlzatioci on growth In the green simfiah, I«ponls cyanellTia.
Jour» Eacp» Zool., 108:1-19.

Banks, E. H., 19^6. Social orggoilzatlca ift Md twl hma [(hUxm

gRilus sub^. ), Scolosy. ^i239-2.kQ,

Bennett, M. A., 19^* Tbe social hlerardiy In ring donres. Ecology, 20(3)<
337-357.

Bovjberg, R. V. , 1953. ilcnlnanoe co-dar In Hm en^fiah, Qronectes vlrllla
Hagsn. fhyslol. Zool., SjSiXl^l^

1956. ScDMi factors affecting a^resslre "behaivlar In cra^ah.
Phyalol. Zool., ^:127-136.

Biraddock, J. C, 19i^5. Soma aspects of the dnmlnanee-st&ordtnatlon
relationship in the fish, Platypoecllus aaculatns. Riyslol. Zool.,
2^:176-195.

, 19'^9> Bie effect of prior residence vipcai danlnanee In HtM flah,
Platypoecilus aaculatus. Physiol. Zool., 22:l6l-l69.

Carpenter, C. B., 19l|-2. Sexual behavior of free ranging Rhesus iBODkeys

(Maeaca aulatta). I. Speelmms, procedures and Isehavloral eharaeteriatloa
of estrus. J. Comp. Psychol., 33;113-1^>3*

ko " i

eolilasj Ifi 1., 19^3. Statltstleal aaalywls of f&ctors -Hfalch make for sixccew
in Intitlal «icount«r8 Txitween hens. Aa« Hat»« 21: 519-5 38«

— 19Mt^. AggresBlve behavior amoog vertebrate animals. Phyalol* Zool«>
1X:83-123.

Crawford, M. P., 19^^0. Tbe relation between social doalnance and the

BMnstrual eyele tn fesoule chl^panzeea. J. Ckanp* Peyebol*, 30:1*83-513.

1911^2* DcBdnanee and social behavior, for chimpanzees, in a nm
competative situation. J. Cflpp. Psychol., 321:267-277.

Darling, F, P., I938. Bird Flocks and the Breeains Cycle. 2he Iftilverslty
Press, Cafflbridge. 12^4- pp.

Daudln, F. M., I803. Historie Ifaturelle des Rainettes. des Grenouilles et
d6S Crapauda. Paris, p. Q5»

Davies, J. L., 1953. Colony size and rao?roduction in the grey seal. Piroo.
Zool. 80c. Londfln., 123(ll): 327-332.

Delaney, M. J., 1955. ^ erolution of aggressive behavior in vertebrate

animals (tMpiibllahed). Dept. of Zool., Glasgov, Scotland. (How at
Ifetiv.of Southharaptcn, England).

Dlebschlag, E., 19^1. Psychological observations on the rank order in the
domestic pigeon. From the Zool. Inst, of the Ifelv. of Marburg.
(PsychologLsche beobachtungsn uber die xtuigrardnimg bei der haiistaube.
Zeitachr. f. Tlerpsychol.^ 4:(H.2) 173-188).

DougLis, M., 191*6. Seme evidence of a flomlnnnce -subordinanee relationship
onang lobsters. Anat. Rec., 2i'553» (Abstract).

— — , 191*8. Social factors inflvienclng the hierarchies of aoall flx>ck8 of
dOB«stic hen: lha interaction between residtat and part time loembers of
or^tfiized flocks. Physiol. Zool., a:ll*7-l8a«

Edwards, A. L., 1951*. Statistical Methods for the Behavioral Sciences.
Rhinehart and Coorpany, Inc., Binr York. 5^*2 pp.

Bllan, E., and R. W. Murray, 1951. New laterlal line sensory organs in
Xenopus laevis. Nature, 168(1*272:1*77*

Ck>in, Coleman J., 1953. Parsonal Conanunication. -^''^

Oola, Olive B., 1955 . World Outside >ty Poor. MacMlllaa Co., New York. p. 1*3.

Gordon, K., 19l*0. Territorial behavior and social dooinance asiong Sculridae.
J. Mwwaal.. 17:171-172» . .

Grant, W. 1955 • •Mtrrltorlallam in two species of salaajaad^s. Science,
1S:(33.35)J137»

Oreenberg, B., 191*7. Some relations between territory, social hierarchy

and leadership in the greeri suailsli, Lepotnls eyan^OJLus* Riysiol. Zool«
^: 267-299.

— — , and G. K. Koible, IS^th, Social behavior of the Aa»rloan chameleon,

Anolis carol jnensls Volgb. Physiol. Zool». 17:-^2-4^,

Oi&l, A. M., and ¥. C. Allee, 19hh, Seme measureable effects of social
organization In nocks of hens, fhyglol. Zool.. 18:320-31*7.

BsnMcrstroni, F., 191^2. Sondnanee in vlnter flocks of chickadees. Wilscn
Bull., 54:32-1*2.

Hasklns, CP., 1951 • Of Societies and Mai. Horton, ITev York. 2^ pp.

Edward, W. E., and J. T. Salem, Jr., 19l»2. Intercovey social relationships
in the valley quail. VUson BuU., §4:162-170.

Kslth, A., 191*9. A .Tfev !Iheory of Hianan Eyolutlon. Philosophical Ubrary,
New York, 1*51 pp. • ■ ; : ;.. - . -

lAslle, J. M., 1890. Notes on the habits and oviposition of Xaiopus laevis«
Froc. Zool. Soc.. London. 69-7I.

lorenz, K., 191*1. V«*glelohttide bewegungsstudlen an anatinon'. Jour. f«'

Oroithol.. 82(Pestschrlft Belnroth):19l*-29l*,

——-1 1950. The conparative method in studying innate behavior patterns.
Physiological Mechanisms in Animal Behavior. Acadsmic Press, New York.
221-268.

— — , 1952. King SolcEm's Ring. Ihoinas Y. Growell (Jo., New York. 202 pp.

lorn, M. S., 1956. Dominance-subordination relationships in the crawfish,
CambareU.us shufeldtli. Tulane Studies in Zoology. 1*(5):1^-170.

Martof, B. S., 1953' Tisrrltorlallty in the greeox fSrog, Rana clnwltj^nw.

Ecology^ 2it'l56-174.

Maslow, A. H., 19l*0. Doodnance qfuality and social behavior in infra -human
primates. J. Soc. Psychol., 11:313-321*.

Munro, A. F., 1953* ^e anmonia and urea exeretion of dlffer«it species of
amphibians during develosment and metaBorphosls. Biochem. Jour..
5l*:29-36. . i '- '

Wmmei, M. a., 1956. social "behavior and Inter^ciflc canrpttltlon In two
trout species. HiyaiQl. Zool.j 22,^611-81.

Odbaa, I. P., 1953. FuoaacientalB of Icology* V* »• Semders Co., Philad«lpiil*.

Pardl, t., 191*8. DoBiiaance order in PolisteB VAsps. Physiol. Zool., gLtl-lS*

Pirker, F. Jr., S. L. Kobins sad A, LaTreriage, 19**?. Breedijas, rearlag and
care of the South African clawed frog (Xeoogjs laevls). Baa Aawr. Mat*,

ja: 38-^9.

PBiteraoa, Bellie P., 1951. aasal cavities of "ttbe toed Haaiplpe caryalhoi
Mlr-ril) and otiber Plpidae. Proc. Zool. 80c., Loadoaa, l^t2JJ3ai-iH5.

Peargon, P. G., 1955 . Population ecology of the Bpedefoot toad, Seaphiopus
\ Hblbroold. (Harlan). Ecological Monoscanhs, 2^:233-207.

Bose, W., 3^0. The Reptiles and Actplii'biana of Southera Africa. HMfcMr
Miller Ltd., Cape Town. pp. 1-3^.

Sabine, W. S., 191*9. Doainance in vinter floclcs of Juncos and tree ^perrowa.

Riysiol. Zool., ^:61v-85.

Sdijeiaerup-EbTae, T., 1913. Honsenes stsDBie fidrag til Honsanes peyiwlogl
aatiaea, 2L»2^-276.

— — * 19^» Social behavior of birdsj cited In C. A. Murchlnaon, A aandbook
of Social Psyciioiogy. Worchester. Pp. 9^7-972.

Sharif, M., 1956. Experimeats in ffccnxp cooflict. Scientific American,
121(5) »5^-58.

l^st, F. H«, 195l». Social aegresslveneas in an aacEMbian. Sci«ice,
120(3100) »1^>

Tittbergan, H., 19lt2. An ob.iectivi3tic study of the innate beba-/iour of aniaals.
Series D. Bibllothica Biothcoretica. V.l pars 2. 1. J. Brill, london.
pp. 39-98.

19li8. Social releasers and the experimental xaethod required for their
study. Wilson Bull., 60:6-52.

1953. Social Behaviour in Animals, vith Special Beferences to
Vertebrates. Methuen and Co., Ltd., Lcndoi. J<^ Wiley and Soas, Inc.,
liev York. 150 pp.

Uhrl6h, J., 1938. The social hlerawShy In aXMno mice. Jbur. Cocg). Faychol.,
g5.»273-^3.

MetrogravTge. BBltiaiore^ Ud^p ApriLL ^i*

Ytix^e, R. NU, 1939* Social <3anlnanoe and seaenaT itatns in the nMnrpan:
ghe Quarterly Beviev of Biology. l^t»115<436«

BIOGSRAPHICAIi X9SM8

Bobert Bice Baubrlcli vas Ixxax In CilscraBiaat, Sew VSaxagOilxtif Hay k,
19S% aad gracbuaiwd txm St«r«u EL^ fichool^ GLaxmaeat, ija. June, 19^. H»
catered the t]Dlv«r8l% ot Wn Vtupi&sILn in iSm ftiSX of 19^ and after thrott
MMsters entered the Axny Air Force lAiere he sp«it three years.

In ibrch^ 191^, he entered ItLchlgaa 8tate IMTer^i^ and receiwA
a Bachelor of Science degree in Forestry (l9*^9) and a Master of Science
degree In Zcxtlogy (19^2) £roa that inatituticn. Ihe sisimer of 19^2, iias
spent at Point Barrov, ilLaska, vith the Office of Iforal BesMxtfli. He \m§m
his dcxrtoral program at the IMversity of KLorlda in September, I952 and spent
the ■ini II of 19^3 and 19^ at the ifarine Biological Laboratory, Mooda Hole,
ilHHnchusetts. VhUe at the Oiiversity of florlda he tan£^ biology
Inlxn-atory and ganeral biology (C-61, C-^} and also vas Assistant Baaiteit
Adrlsor In one of the housing areas on canqpus*

Haubrloh Is a neaber of XL Si0Mi fl, Rd BL^m, Blgna XI, fhe
Association, foor the Study of Aoifflal Behaviour and the Ecological Society
of Aaerloa*

ahls dlasortatioQ "was prepared under the direction of the chairman
of -aift candidate ' s supervisory committee and has "be«i approved by all
msmberB of the conunittee. It was suhmitted to the Dean of the College of
Arts and Sciences and to the Graduate Cornell and vaa apjaroved as partial
fulfillaent of the reqvUreoi^its for the degree of Doctor of Philosophy.

Au^ist 10, 1957

i-'<'

"Smmf CkxUefte of Arts azid Sciences

Been, Graduate School

SUHSR7IS0KY