Supplemental Table 2. Evidence that males use contest competition to monopolize access to females. Bold font with gray background indicates species that demonstrate
prominent precopulatory features: Z=sexually dimorphic size, P=sexually dimorphic shape and/or appendages, T=teeth/tusks, S=singing, (see Methods and Fig. 1). Group size and
social structure follows May-Collado et al. (2007): solitary (strong bonds limited to time calf is dependent, may aggregate for breeding), fluid (group living with weak associations),
fission-fusion (fluid group living with some long-term associations) and family-based (long-term associations among kin). Evidence of male contests that likely reflect the ability of
winners to monopolize access to females is indicated using ** for the "conservative" criteria and * for "relaxed" criteria (see Methods). Column 3 (conservative criteria) is based on
behavioral observations of male contest and column 4 (relaxed criteria) is based on these data in addition to genetic studies, life history data and what is known about the
distribution of females during the breeding season, group size and social structure. See column 5 for the data used for classifying species. Following Lüpold et al. (2013),
classification of the opportunity to monopolize females differed among species based on the best available information as follows:
 if male fights or contests observed, or sex-specific scarring indicative of past combat observed, infer that there is opportunity for the winners to monopolize access to females:
 if females observed copulating with more than one male, infer that the opportunity for males to monopolize access to females is low;
 if groups are of mixed sex and large or very large in number, infer that the opportunity for males to monopolize access to females is low (note: eastern spinners seem to be an
exception, Perrin and Mesnick 2003);
 if females occur singly or in small groups that are dispersed, or highly dispersed, during the breeding season, infer that the opportunity for males to monopolize access is low.
Exceptions include forms of sequential female defense polygyny (e.g., Dall's porpoise, Willis and Dill 2007; Indian Ocean bottlenose dolphin, Connor et al. 1992, 2000; common
bottlenose dolphin, Wells 2003);
 if groups are mixed sex and stable, mating may occur when groups meet (e.g., resident killer whales, Baird 2000) or if adult males move between groups (e.g., long-finned pilot
whales (Amos et al. 1993), infer that the opportunity for males to monopolize access to females is low. Mixed sex groups can also be mix-sex breeding groups and if there is
evidence of multiple mating within groups and no evidence of male contests, infer that the opportunity for males to monopolize access to females is low (e.g., short-finned
pilot whales, Kasuya et al. 1993; see Möller 2012).
Species
Common name
Balaena mysticetus
Bowhead whale
Balaenoptera acutorostrata
Minke whale
Opportunity for
males to
monopolize
access to
females
inferred from
observations of
male contests,
"conservative
criteria"
no
Opportunity
for males to
monopolize
access to
females
inferred from
observations
of male
contests,
genetic
studies, life
history data
and social
structure,
"relaxed
criteria"
no
no
Evidence on opportunity for males to
monopolize access to females used to
classify species
gregarious during mating season; sexual
activity observed in active mating groups
with multiple males and maybe multiple
females; little known opportunity to
monopolize access to females
dispersed in open waters, little known
opportunity to monopolize access to
Group
size
(mean or
range)
Social structure
Citations
1
Solitary
1, 2
1 to 3
Unknown /
structured by age
3, 4, 5
females
Balaenoptera borealis
Sei whale
no
Balaenoptera edeni
Bryde’s whale
no
Balaenoptera musculus (S)
Blue whale
no
Balaenoptera physalus (S)
Fin whale
no
Berardius bairdii (T)
Baird's beaked whale
no
Caperea marginata
Pygmy right whale
no
Cephalorhynchus commersonii
Commerson’s dolphin
no
Cephalorhynchus heavisidii
Heaviside’s dolphin
no
Cephalorhynchus hectori
Hector’s dolphin
Delphinapterus leucas
Beluga whale
no
no
no
dispersed in open waters, little known
opportunity to monopolize access to
females
dispersed in open waters with extended
breeding season, little known opportunity
to monopolize access to females
dispersed in open waters, if and how song
functions as a male mating display is
unknown; singers are solitary; while it is
reasonable to assume that songs function
to advertise the species and location of the
singing whale, it is not known to what
extent song is also under selection as a
form of inter- and/or intra-sexual display;
little known opportunity to monopolize
access to females but function of song
warrants further investigation
dispersed in open waters, male song may
serve to attract females from great
distances to aggregations of patchily
distributed prey; little known opportunity
monopolize access to females but song
warrants further investigation
mixed sex groups and some sexual
segregation; some evidence to suggest that
females mate with multiple males; both
sexes are heavily scarred (males may be
slightly more so) but difficult to ascertain if
the scars are from males, females or both
sexes; little known opportunity to
monopolize access to females
little known, dispersed with extended
breeding season, little known opportunity
to monopolize access to females
generally dispersed and fluid, little known
opportunity to monopolize access to
females
generally dispersed and fluid, little known
opportunity to monopolize access to
females
generally dispersed and fluid, little known
opportunity to monopolize access to
females
large mixed schools; little known
and sex?
2 to 5
Solitary
1, 5
1
Solitary
1, 5
1
Solitary
1, 6, 7
1.55
Solitary
1,8
7.2
Fission-fusion
1, 9, 10
1
Solitary
1, 11
6.9
Fluid
1, 5, 12
3.2
Fluid
1, 5, 13
8
Fluid
1, 14
32.9
Fluid
5, 15
opportunity to monopolize access to
females
Delphinus capensis
Long-beaked common
dolphin
Delphinus delphis
Short-beaked
common dolphin
Eschrichtius robustus
Gray whale
no
no
no
no
very large mixed schools; little known
opportunity to monopolize access to
females
411.69
Unknown
5
no
very large mixed schools; little known
opportunity to monopolize access to
females
230.38
Fluid, Fission-fusion
5
1
Solitary
5, 16
1 to 2
Unknown
5, 17
30.12
Long-term
associations in
mixed sex groups
1, 18
41.1
Family based
1, 19, 20
84.3
Family based
1, 21
63
Fluid, Fission-fusion
1, 22, 23
no
Eubalaena japonica
North Pacific right
whale
no
Feresa attenuata
Pygmy killer whale
no
Globicephala macrorhynchus (Z) (P)
Short-finned pilot
whale
no
no
Globicephala melas (Z) (P)
Long-finned pilot
whale
no
no
Grampus griseus
Risso’s dolphin
no
no
gregarious groups during mating season
and females observed mating with multiple
males; little known opportunity to
monopolize access to females
little known; gregarious groups may form
during mating season (behavior inferred
from the better known sister species in
which females known to copulate with
multiple males in mating groups)
stong long-term associations in mixed sex
groups; no known observations of
aggression among males for access to
females
multiple mating for females thought to
contribute to group cohesion; strong, longterm associations with male dispersal from
natal groups; groups contain a few large
males unrelated to females in group; no
known observations of aggression among
males
strong, long-term associations in which
both males and females remain with natal
group; sexually mature males may move
among groups; no known observations of
aggression among males
strong, long-term associations of males and
females in medium to large groups; both
sexes are heavily scarred (males may be
slightly more so) but difficult to ascertain if
these scars are from males, females or
both sexes; no known observations of
aggression among males
Hyperoodon ampullatus (P), (T) **
Northern bottlenose
whale
yes
yes
Inia geoffrensis **
Boto
yes
yes
Kogia breviceps
Pygmy sperm whale
no
Kogia sima
Dwarf sperm whale
no
Lagenorhynchus acutus
Lagenorhynchus obliquidens
Atlantic white-sided
dolphin
Pacific white-sided
dolphin
no
no
no
no
no
no
Lagenorhynchus obscurus
Dusky dolphin
Lissodelphis borealis (Z)
Northern right whale
dolphin
no
Lipotes vexillifer
Baiji
no
Megaptera novaeangliae (S) **
Humpback whale
yes
yes
Mesoplodon carlhubbsi (T) **
Hubbs' beaked whale
yes
yes
Mesoplodon europaeus (T) *
Gervais' beaked
whale
yes
head-butting observed between males
apparently contesting dominance
aggregations during breeding season;
intense aggression between males may
lead to lethal injury; males display with
objects to females
dispersed, little known opportunity to
monopolize access to females
dispersed, little known opportunity to
monopolize access to females
large groups of mixed sex; little known
opportunity to monopolize females
very large groups of mixed sex; little known
opportunity to monopolize females
Females observed to mate with multiple
males; mating groups typically consist of a
small group of adult males chasing one
sexually mature female who mates multiple
times, males do not appear to interfere
with each other during chases, although
some evidence that males receive more
overt aggression than females
very large groups of mixed sex; little known
opportunity to monopolize females
dispersed, little known opportunity to
monopolize access to females
aggregations on breeding grounds; male
competitive groups aggressively pursue
females, sometimes with spectacular fights
and bloody injuries among males during
chases; complex song displays by males
contests between males inferred based on
male-specific scarring pattern; adult males
are heavily covered with long parallel tooth
rake marks likely acquired during fights
with other males; small dispersed groups;
composition possibly includes adults of
both sexes
contests between males inferred based on
male-specific scarring pattern likely
acquired during fights with other males;
scars are singular tooth rake marks and not
heavy, indicating that contests do not seem
to be of the same intensity as in some
other ziphiids; small dispersed groups;
composition possibly includes adults of
7
Fission-fusion
1, 24
6.22
Solitary, fluid
1, 25, 26
2
Unknown
1, 27
1.87
Unknown
1, 27
53.2
Fluid
1, 28
127.38
Fluid
1, 29
86
Fluid, Fission-fusion
1, 30, 31
110.2
Unknown
1, 32, 33
3.4
Solitary, fluid
1, 34
1
Solitary, fluid, longterm associations
between females &
escorts
1, 35 -38
2 to 10
Unknown
39 - 42
Unknown
39 - 43
(group size
is mean for
genus)
2.9 (1 to
15)
both sexes
Mesoplodon ginkgodens (T) *
Mesoplodon mirus (T) *
Ginkgo-toothed
beaked whale
yes
True's beaked whale
yes
Mesoplodon perrini (T)*
Perrin's beaked whale
yes
Monodon monoceros (T) (Z) **
Narwhal
Neophocaena phocaenoides
Finless porpoise
Orcinus orca (Z) (P)
Killer whale
Peponocephala electra
Melon-headed whale
Phocoena phocoena
Harbor porpoise
Phocoena sinus
Vaquita
no
Phocoena spinipinnis
Burmeister's porpoise
no
Phocoenoides dalli (P) **
Dall’s porpoise
yes
yes
no
no
no
no
no
yes
no
yes
contests between males inferred based on
male-specific scarring pattern likely
acquired during fights with other males;
scarring not heavy indicating that contests
do not seem to be of the same intensity as
in some other ziphiids
contests between males inferred based on
male-specific scarring pattern likely
acquired during fights with other males;
few, light, closely-spaced parallel scars on
males indicate that contests do not seem
to be of the same intensity as in other
ziphiids; small dispersed groups
contests between males possible based on
male specific scarring pattern likely
acquired during fights with other males;
scarring not heavy indicating that contests
do not seem to be of the same intensity as
in other ziphiids
males engage in tusk displays during
summer months; male contests during the
late winter/early spring (unobserved)
mating season are unknown; male-specific
scarring
dispersed, little known opportunity to
monopolize access to females
strong, long-term associations; for many of
the known ecotypes, both males and
females remain in stable, long-term mixed
sex groups; mating thought to occur when
groups meet; no known observations of
aggression among males
very large groups of mixed sex; little known
opportunity to monopolize access to
females
dispersed, little known opportunity to
monopolize access to females; highly
seasonal and synchronous reproduction;
mating likely promiscuous
dispersed, little known opportunity to
monopolize access to females
dispersed, little known opportunity to
monopolize access to females
intense aggressive interactions observed
Unknown
39 - 43
(group size
is mean for
genus)
Unknown
39 - 43
(group size
is mean for
genus)
2.9 (1 to
15)
Unknown
39 - 43
(group size
is mean for
genus), 44
3
Fluid, family based
1, 45, 46
3
Solitary
1, 47
12
Family based
1, 48
257.7
Family based
1, 49
5.7
Fluid
1, 50
2
Fluid
1, 51
4.5
Unknown
1, 52
7.4
Fluid
1, 53
2.9 (1 to
15)
2.9 (1 to
15)
Physeter macrocephalus (Z), (P)
(sounds) **
Sperm whale
Platanista gangetica
South Asian river
dolphin
Pontoporia blainvillei
Franciscana
no
no
Pseudorca crassidens
False killer whale
no
no
Sotalia fluviatilis
Tucuxi
no
no
Sotalia guianensis
Guiana dolphin
no
no
Stenella attenuata
Pantropical spotted
dolphin
no
Stenella coeruleoalba
Striped dolphin
no
Stenella frontalis
Atlantic spotted
dolphin
no
Stenella longirostris orientalis (P) *
Eastern spinner
dolphin
yes
yes
yes
no
between males while engaged in mate
guarding; sequential female defense
polygyny
fights observed between adult males and
adult males often heavily scared from the
teeth rake marks of other males and have
broken teeth and jaws attributed to
intraspecific fighting; roving strategy for
adult males, presumably searching for
receptive females
dispersed, little known opportunity to
monopolize access to females
monogamous; dispersed pairs; no known
observations of aggression among males
strong long-term associations in mixed sex
groups; no known observations of
aggression among males
female multiple mating observed;
dispersed; little known opportunity to
monopolize access to females
female multiple mating observed;
dispersed; little known opportunity to
monopolize access to females
very large groups of mixed sex; little known
opportunity to monopolize access to
females; observations of small groups of
strongly bonded males warrants further
investigation
very large groups of mixed sex; schools
stratified by age and sex and mature males
may rejoin female groups during the
breeding season; no known observations of
aggression among males
dispersed, little known opportunity to
monopolize access to females;
observations of small groups of bonded
males warrants further investigation
very few males with testes full of sperm
indicates that a few sexually and socially
dominant males may monopolize mating
within groups; very large groups of mixed
sex; observations of small groups of
strongly bonded males warrants further
investigation regarding possibility of male
competition for access to females; both
fluid and stable associations observed
22.1
Adult males solitary,
family based
1, 54
2.45
Solitary
1, 55
7.1
Fluid
1, 56
36.16
Family based
1, 57, 58
5.8
Weak, Fissionfusion
1, 59, 60,
61
17.6
Weak, Fissionfusion
1, 61, 62,
63
360
Fluid, Fission-fusion
1, 64
302
Fluid, Fission-fusion
1, 65
10
Fluid, Fission-fusion
1, 66
147.74
Fluid, Fission-fusion
1, 67
depending on location
Steno bredanensis
Rough-toothed
dolphin
no
Tasmacetus shepherdi (T) *
Shepherd's beaked
whale
yes
Tursiops aduncus **
Indo-Pacific
bottlenose dolphin
Tursiops truncatus *
Common bottlenose
dolphin
Ziphius cavirostris (T) **
Cuvier's beaked whale
1
2
3
4
yes
yes
yes
yes
yes
medium-size groups of mixed sex with no
known evidence of contests among males
for access; observations of one adult male
in groups warrants further investigation
regarding possibility of male competition
for access to females
contests between males inferred based on
male specific scarring pattern apparently
caused by fighting with other males;
scarring is light so contests do not seem to
be of the same intensity as in some other
ziphiids; small dispersed groups;
composition possibly includes adults of
both sexes
alliances and higher level coalitions of adult
males coercively herd females and
compete aggressively for access to them
with other male coalitions
scramble competition to find mates and
some evidence of contest competition to
gain access to them; females consort with
multiple males during a breeding season;
males, singly or in alliances, accompany a
female (presumably in estrus); light, sexspecific scarring patterns; sex ratio in older
age classes is skewed (males have greater
mortality than females)
contests between males inferred based on
intense male-specific scarring pattern
apparently caused by fighting with other
males; small dispersed groups; group
composition possibly includes adults of
both sex with one adult male
40
Unknown
1, 68, 69
3 to 6
Unknown
70
4 to 7
Fluid, Fission-fusion
1, 71 - 74
2 to 15
Fluid, Fission-fusion
74 - 79
2.9
Family based
1, 80, 81
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