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Photo-identification Catalogue and Status of the
Northern Resident Killer Whale Population in 2019
Jared R. Towers, James F. Pilkington, Brian Gisborne, Brianna M. Wright,
Graeme M. Ellis, John K. B. Ford, and Thomas Doniol-Valcroze
Fisheries and Oceans Canada
Science Branch, Pacific Region
Pacific Biological Station
3190 Hammond Bay Road
Nanaimo, BC
V9T 6N7
2020
Canadian Technical Report of
Fisheries and Aquatic Sciences 3371
Canadian Technical Report of Fisheries and Aquatic Sciences
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Canadian Technical Report of
Fisheries and Aquatic Sciences 3371
2020
PHOTO-IDENTIFICATION CATALOGUE AND STATUS OF THE NORTHERN
RESIDENT KILLER WHALE POPULATION IN 2019
by
Jared R. Towers, James F. Pilkington, Brian Gisborne, Brianna M. Wright,
Graeme M. Ellis, John K. B. Ford, and Thomas Doniol-Valcroze
Fisheries and Oceans Canada
Cetacean Research Program
Pacific Biological Station
Nanaimo, BC
V9T 6N7
E-mail: Jared.Towers@dfo-mpo.gc.ca
ii
© Her Majesty the Queen in Right of Canada, 2020
Cat. No. Fs97-6/3371E-PDF ISBN 978-0-660-34745-5 ISSN 1488-5379
Correct citation for this publication:
Towers, J.R., Pilkington, J.F., Gisborne, B., Wright, B.M., Ellis, G.M., Ford, J.K.B., and
Doniol-Valcroze, T. 2020. Photo-identification catalogue and status of the
northern resident killer whale population in 2019. Can. Tech. Rep. Fish. Aquat.
Sci. 3371: iv + 69 p.
iii
TABLE OF CONTENTS
ABSTRACT ..................................................................................................................... iv!
RÉSUMÉ ......................................................................................................................... iv!
1.0 INTRODUCTION ...................................................................................................... 1!
2.0 MATERIALS AND METHODS ................................................................................. 2!
2.1!DATA COLLECTION .............................................................................................................. 2!
2.2!DEFINING AGE, SEX, AND DEATH ..................................................................................... 2
2.3!NAMING ................................................................................................................................. 2
2.4!DATA ANALYSIS ................................................................................................................... 3
2.5!DATA PRESENTATION ......................................................................................................... 3
3.0 RESULTS AND DISCUSSION ................................................................................. 3!
4.0 ACKNOWLEDGEMENTS ........................................................................................ 6!
5.0 REFERENCES ......................................................................................................... 6!
APPENDIX ..................................................................................................................... 11!
PHOTO-IDENTIFICATION CATALOGUE OF NORTHERN RESIDENT KILLER WHALES ........ 11!
iv
ABSTRACT
Towers, J.R., Pilkington, J.F., Gisborne, B., Wright, B.M., Ellis, G.M., Ford, J.K.B., and
Doniol-Valcroze, T. 2020. Photo-identification catalogue and status of the
northern resident killer whale population in 2019. Can. Tech. Rep. Fish. Aquat.
Sci. 3371: iv + 69 p.
Censuses of the northern resident killer whale population using photo-
identification have been conducted annually since 1973. These studies are based on
photographic recapture of permanent natural markings on every individual within the
population. In this report, we summarize northern resident killer whale population trends
over the time series of this study and provide a photo-identification catalogue of all
individuals considered to be alive in 2019. This population has grown at a mean annual
rate of 2.2% since 1973 and in 2019 contained a minimum of 310 individuals. Continued
annual photo-identification censusing is a key strategy to accurately document the
abundance, genealogy, sociality, demographics, and health of this threatened
population.
RÉSUMÉ
Towers, J.R., Pilkington, J.F., Gisborne, B., Wright, B.M., Ellis, G.M., Ford, J.K.B., and
Doniol-Valcroze, T. 2020. Photo-identification catalogue and status of the
northern resident killer whale population in 2019. Can. Tech. Rep. Fish. Aquat.
Sci. 3371: iv + 69 p.
Des recensements de la population d’épaulards résidents du nord ont été menés
tous les ans depuis 1973. Ces études reposent sur l'identification de chacun des
individus à partir de photographies de marques naturelles permanentes. Dans le
présent rapport, nous résumons les tendances démographiques de la population
d'épaulards résidents du nord pour toute la durée de l’étude, et présentons un catalogue
de photographies d'identification des individus considérés comme étant en vie en 2019.
Le taux d’accroissement moyen de cette population est de 2,2% par an depuis 1973,
avec un minimum de 310 individus en 2019. Le maintien d’un recensement
photographique annuel constitue une stratégie essentielle pour documenter
l’abondance, la généalogie, la vie sociale, la démographie et la santé de cette
population menacée.
1.0 INTRODUCTION
Killer whales in the coastal waters of British Columbia were first found to be
individually recognizable in 1970 (Spong et al. 1970). Field research using photo-
identification began on these killer whales in 1973 (Bigg et al. 1976) and continues to
the present. Over this time period, photo-identification data has been fundamental in
defining killer whale populations in British Columbia (Bigg 1982; Ford and Ellis 1999;
Ford et al. 1994, 2000), determining their social structure (Bigg et al. 1990; Olesiuk et
al. 1990), distribution and abundance (Ellis et al. 2011; Ford et al. 2014; Towers et al.
2019), and facilitating studies on their behaviours (Morton 1990; Baird and Dill 1995;
Barrett-Lennard et al. 1996; Ford and Ellis 2006; Ford et al. 1998, 2011; Deecke et al.
2010; Wright et al. 2017; Towers et al. 2018), cultures and evolution (Riesch et al. 2012;
Foster et al. 2012; Wright et al. 2016; Whitehead and Ford 2018).
Three ecotypes of killer whale occupy coastal waters off western Canada –
residents, Bigg’s, and offshores. Each of these ecotypes are unique in their morphology
and ecology (Ford 2014). Although their ranges overlap, they do not mix socially and as
a result, are reproductively isolated and genetically distinct (Barrett-Lennard 2000). In
BC, the resident ecotype is composed of two populations that do not intermingle.
Named northern and southern for their usual distribution in relation to each other along
the BC coast, both populations range from southeastern Alaska to outer coast waters
off the continental US (Ford et al. 2017).
The northern and southern resident killer whale communities are composed of
several groups of maternally related individuals that maintain social cohesion over long
time periods (Ford et al. 2000). These social groups, initially referred to as pods and
subpods (Bigg et al. 1987), are now more commonly referred to as matrilines (Towers et
al. 2015). They can contain up to five generations of living individuals (see R5 matriline
in Appendix) and are typically composed of an older female, all her offspring, all the
offspring of her daughters and so forth. Every individual within each matriline shares a
socially learned dialect. The northern resident killer whale community contains three
acoustic clans, each containing matrilines with acoustic similarities and thus, common
maternal ancestry (Ford 1991).
Northern resident killer whales have a strong preference for large salmon,
primarily Chinook and secondarily, chum (Ford et al. 2010; Ford and Ellis 2006). Their
health, survival, and fecundity are correlated with the availability and size of these prey
(Ford et al. 2010; Groskreutz et al. 2019; Murray et al. 2019). Other historic, current, or
emerging threats include live-captures (Bigg and Wolman 1975), intentional shootings
(Hoyt 2019; DFO unpubl. data), acute and chronic acoustic disturbance (Morton and
Symonds 2002; Williams et al. 2013), vessel strikes (Williams and O’Hara 2010; Murray
et al. 2019), entanglement and ingestion of fishing gear (Fisheries and Oceans Canada
2018), and bioaccumulation of contaminants (Ross et al. 2000). These threats,
2
combined with relatively small population size and low reproductive rates (Olesiuk et al.
2005), led to this population being listed as Threatened by the Committee on the Status
of Endangered Wildlife in Canada (COSEWIC) in 2001 and subsequently under
Canada’s Species at Risk Act (SARA). Objectives for the improvement of our
understanding and conservation of this population are provided in the Resident Killer
Whale Recovery Strategy (Fisheries and Oceans Canada 2018). Among them, the
annual photo-identification census is listed as a key technique because most efforts
taken to recover the population are underpinned by knowledge of population health and
size.
Photo-identification data from annual censuses of northern residents have been
published periodically in books and technical reports to provide up-to-date working
documents that present details on the evolving appearances, demographics, size, and
growth trends of the population (Bigg et al. 1987; Ford et al. 1994, 2000; Ellis et al.
2007, 2011; Towers et al. 2015). In this report we provide an update on the size of the
northern resident killer whale population in 2019, its annual growth rate since 1973, and
an updated photo-identification catalogue of all individuals considered alive in 2019.
2.0 MATERIALS AND METHODS
2.1 DATA COLLECTION
Digital identification photographs of northern resident killer whale dorsal fins and
saddle patches were collected perpendicular to the whales’ left sides from small boats
using methods first described by Bigg et al. (1976, 1986) and later updated by Ellis et al.
(2011).
2.2 DEFINING AGE, SEX, AND DEATH
Birth years for northern resident killer whales first documented as adults were
estimated based on known or inferred ages of their known or inferred kin (Bigg et al.
1990). Years of birth for individuals first documented as juveniles or calves were
assigned based on the size of the whale when it was first documented. Individuals were
only considered dead after being absent from several encounters with their closest kin
or preferred travel partners (Olesiuk et al. 2005).
Sexes were confirmed visually in the field from observations of skin pigmentation
on the underside of the body, dorsal fin growth (sprouting males), close association with
newborn offspring (reproductive females), or DNA from genetic samples (DFO 2019). If
an animal of unknown sex reaches 15 years of age without sprouting or producing a
calf, it is assumed to be female until otherwise confirmed.
2.3 NAMING
3
Individuals are named with an alphanumeric designation based on the letter of
the pod they belong to and the order in which they were first identified. Matrilines are
named after the eldest living reproductive female within a group of maternally related
individuals, although if she has a surviving brother or uncle, the matriline is named after
the deceased mother of the eldest male (Towers et al. 2015). Pod names refer to the
most distinctive whale documented in each social group when they were first identified
in the early 1970s (Bigg et al. 1987). Each clan bears the same letter as the most
abundant pod within it.
2.4 DATA ANALYSIS
Identification metadata were individually applied to all images of northern resident
killer whales on an annual basis as per data analysis and management techniques
provided in Towers et al. (2012). Population size was calculated as the minimum
number of individuals alive each year, assuming that all whales missing from censused
matrilines were dead and that all uncensused individuals were alive (DFO 2019).
2.5 DATA PRESENTATION
The appendix includes identification photographs of whales known or presumed
to be alive at the end of the 2019 field season with the inclusion of two individuals (I26
and I42) considered missing in 2019. Individuals are laid out by generation, in order of
birth, and in relation to maternal kin. Clusters often represent socially cohesive groups,
however, some individuals may or may not exhibit social cohesion with others in their
matriline (Ford and Ellis 2002; Towers et al. 2015). Tabs are provided on the outside
edge of each page indicating the acoustic clan to which each group belongs. Schematic
diagrams portray matrilineal genealogical relationships that have been inferred from
long-term observations of social associations (Bigg et al. 1987; Ford et al. 1994, 2000;
Ellis et al. 2007, 2011; Towers et al. 2015). They include every individual documented
over the course of the study with the exception of matrilines that no longer contain any
living members. Clear boxes represent individuals known or presumed to be alive in
2019 and shaded boxes represent deceased individuals. Known or estimated birth
years are listed above each identification photograph and below each shaded box. If
known, sexes are indicated with symbols above each identification photograph and
within each schematic box. Lines linking mothers and offspring are solid if the
relationship is positive (i.e., known with certainty). This includes individuals that have
been documented since birth. Relationships between whales born before the study
began in the early 1970s are not known with certainty and are either probable, indicated
by a dashed line, or possible, indicated by a dotted line.
3.0 RESULTS AND DISCUSSION
Since the last northern resident killer whale photo-identification catalogue and
population update was published in 2015 (Towers et al. 2015) we have traveled several
4
thousand kilometres each year in search of northern resident killer whales. We located
them an average of 60 times per year (range: 39-82) and analyzed a total of 51,696
identification photos between 2015 and 2019. These are similar levels of effort,
encounters, and data as in previous census years (Bigg 1982; Ellis et al. 2011; Towers
et al. 2015).
Complete accounting of all northern resident killer whales within a census year is
rare. As a result, there is occasionally some uncertainty in the total population size in a
given year. Between 2015 and 2019 we accounted for all northern resident killer whales
at an average of 93% (range: 83%- 99%) of the population each year. Any uncertainty
in statuses of individuals in a matriline were typically clarified whenever that matriline
was next encountered. In 2019, one matriline (R17) was not encountered, but assuming
all 12 individuals within it were alive, the minimum population size for northern resident
killer whales was 310 individuals (range: 310-314).
Figure 1. Abundance trend of the northern resident killer whale population from 1973-
2019. Uncertainties in minimum and maximum population sizes are represented
with shading.
Between 1973 and 2019, the northern resident killer whale population
experienced periods of both growth and decline (Figure 1). The 7% decline in
population size between 1998 and 2001 coincided with a reduction in the abundance of
5
primary prey for this population, Chinook salmon (Ford et al. 2010). Immature
individuals that survived this period experienced constrained growth and have
significantly shorter maximum body lengths than individuals that matured before this
decline (Groskreutz et al. 2019). Since 2002, no annual decline in total numbers was
documented until 2018 when the northern resident population showed a growth rate of -
0.3% (DFO 2019).
Figure 2. Abundance trends of the northern resident killer whale clans from 1973-2019.
Uncertainties in minimum and maximum population sizes are represented with
shading.
The mean rate of annual growth for the northern resident population over the time
series of this study is 2.2%. G clan had the greatest mean annual growth rate of 2.8%,
followed by R clan at 2.4% and A clan at 1.9% (Figure 2). However, over the last five
years G clan has shown the lowest annual mean growth rate (-0.2%), followed by A clan
(2.2%) and R clan (4.0%). These trends are likely influenced by a suite of factors from
changing demographics within each clan to the availability of preferred prey and the
ability of individuals to withstand both chronic and acute threats within their
environment. Continued population censusing using photo-identification will continue to
improve our understanding of northern resident killer whale population health and the
impacts that anthropogenically and environmentally influenced changes may have on it.
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4.0 ACKNOWLEDGEMENTS
Over the years, people too numerous to list have contributed to this long-term
study. We thank them all. In particular, we express our gratitude to the following
individuals who have provided images used in this report: Robin Abernethy, Lance
Barrett-Lennard, Mark Malleson, Christie McMillan, Miguel Neves dos Reis, Stephen
Page, Nicole Robinson, Lisa Spaven, Eva Stredulinsky, and Sara Tavares. Other
individuals who have assisted with or contributed to this study since the publication of
the last photo-identification catalogue include: Jim Borrowman, Ali Bowker, Troy Bright,
Miray Campbell, Ivan Dubinsky, Elysanne Durand, John Durban, Archie Dundas, Dave
Ellifrit, Holly Fearnbach, Sarah Fortune, Katy Gavrilchuk, Mike Greenfelder, Muriel
Hallé, Karen Hansen, Jackie Hildering, Stan Hutchings, Lisa Larsson, Jeff Litton, Bill
and Donna Mackay, Dena Matkin, Kai Meyer, Albert Michaud, Hermann Mueter, Linda
Nichol, Bruce Paterson, Kathy Peavey, Tasli Shaw, Nick Sinclair, Dylan Smyth, Paul
Spong, Gary Sutton, Helena Symonds, Sheila Thornton, Andrew Trites, Jane Watson,
and Janie Wray. We are also grateful for support and cooperation from the following
organizations: BC Cetacean Sightings Network, Canadian Coast Guard, Center for
Whale Research, CetaceaLab, Coastal Ocean Research Institute, Gitga’at First Nation,
Langara Fishing Lodge, MERS Marine Education and Research Society, Marine
Mammal Research Unit (UBC), North Coast Cetacean Society, North Island Marine
Mammal Stewardship Association, Ocean Wise, OrcaLab, and the Strawberry Isle
Marine Research Society. Finally, we thank Elysanne Durand for conducting a review of
the appendix in this report and apologize to anyone we missed. Since 2001, major
funding for this research has been provided by the Species at Risk program of Fisheries
and Oceans Canada and most field work was conducted under DFO Marine Mammal
Research License 001.
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status of the northern resident killer whale population in 2014. Can. Tech. Rep.
Fish. Aquat. Sci. 3139: iv + 75 p.
Towers, J.R., Ford, J.K.B., and Ellis, G.M. 2012. Digital photo-identification dataset
management and analysis: Testing protocols using a commercially available
application. Can. Tech. Rep. Fish. Aquat. Sci. 2978: iv + 16 p.
Towers, J.R., Hallé, M.J., Symonds, H.K., Sutton, G.J., Morton, A.B., Spong, P.,
Borrowman, J.P. and Ford, J.K.B. 2018. Infanticide in a mammal-eating killer
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Towers, J.R., Sutton, G.J., Shaw, T.J.H., Malleson, M., Matkin, D., Gisborne, B., Forde,
J., Ellifrit, D., Ellis, G.M., Ford, J.K.B., and Doniol-Valcroze, T. 2019. Photo-
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Whitehead, H. and Ford, J.K.B. 2018. Consequences of culturally-driven ecological
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Williams, R., Erbe, C., Ashe, E., Beerman, A. and Smith, J. 2013. Severity of killer
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Williams, R. and O’Hara, P. 2010. Modelling ship strike risk to fin, humpback and killer
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11
APPENDIX
Photo-identication catalogue of northern resident killer whales
12
A55 ♂ 1990 A62 ♀ 1994
A91 ♂ 2009 A107 ♂ 2013
A83 ♂ 2005
A-Clan
A1 Pod,
A34 Matriline
13
A67 ♀ 1997
A102 2012
A80 ♂ 2004 A96 ♀ 2010
A34 ♀ 1975
A-Clan
A112 2016
A12 ♀
A33 ♂
1971
A34 ♀
A62 ♀A55 ♂A74
A31 ♂
1960
A67 ♀
2000
A96 ♀A80 ♂
A107 ♂
A91 ♂
A83 ♂A112A92
2009
1943
A102
14
A50 ♀ 1984
A84 ♂ 2005
A72 ♀ 1999 A99 ♂ 2011
A108 ♂ 2014
A-Clan
A1 Pod,
A50 Matriline
A2 ♀
A3 ♂
1954
A30 ♀
A39 ♂ A38 ♂A50 ♀A40 A54 ♀
1930
A6 ♂
1965 1981
A84 ♂A86 ♀A75 ♀A93
1949
A72 ♀A115 ♀
A113 ♀
A118 ♀
A108 ♂
2009
19751971
2017
A99 ♂
A101 ♀
A106 ♂
A120
15
A54 ♀ 1989
A86 ♀ 2006
A75 ♀ 2002 A118 ♀ 2018
A113 ♀ 2016
A-Clan
A101 ♀ 2012 A120 2019
A106 ♂ 2013
A1 Pod,
A54 Matriline
16
A70 ♀ 1999
A52 ♀ 1987
A105 2013
A81 ♂ 2004 A111 2016
A-Clan
A4 Pod,
A35 Matriline
A100 2011
17
A35 ♀ 1974
A90 2008A77 ♂ 2003
A56 ♀ 1990
A4 Pod,
A56 Matriline
A117 2017
A-Clan
A11 ♀
A70 ♀A59 ♀
A35 ♀
A77 ♂
A13 ♂A56 ♀
A48 ♀
1992
1983
A65
1996
A87
1978
A117
A52 ♀
A82
2004
A105
2007
A97
A111A76
2002
A81 ♂
1959
2010
A100
A90
18
A73 ♀ 2000
A116 2017
A4 Pod,
A24 Matriline
A-Clan
A104 2013
19
A-Clan
A64 ♀ 1995
A110 ♂ 2015
A94 ♂ 2009
A78 ♂ 2003
A89 ♂ 2008
A45 ♀
A41 A53A49
1988
A58
1992198519831981
A68
1997
A73 ♀
A116
A10 ♀
1942
A4 ♂
1954
A19
1973
A24 ♀A47
1983
A64 ♀A71 ♂A94 ♂A78 ♂
A110 ♂
1967
A104
A89 ♂
1999
20
A60 ♂ 1992
A43 ♀ 1981
A69 ♀ 1997
A95 ♂ 2009 A109 ♀ 2014
A16 ♀A27 ♂A21
1972
A43 ♀A29 A60 ♂
197719671964
A63
1995
A69 ♀
A95 ♂
A7 ♀
1927
A23 ♀
1948
A109 ♀
A5 Pod,
A23 Matriline
A-Clan
21
A-Clan
A61 ♂ 1994
A85 ♀ 2005
A5 Pod,
A25 Matriline
A121 2019
A17 A25 ♀A18 ♀
1972
A15 ♂
197919691964
A14 ♀
1948
A51 ♀ A61 ♂
A85 ♀A98
1986
2010
A121
22
A-Clan
A42 ♀ 1980
A79 ♀ 2004
A66 ♂ 1996 A88 2008 A114 ♀ 2017
A5 Pod,
A42 Matriline
A103 2013
A119 ♂ 2018
A9 ♀
A5 ♂
1960
A26 ♂
A42 ♀A28 ♀
A8 ♀
1959
A66 ♂A79 ♀
A57 ♀
1942
1972
A88
1991
1974
A114 ♀
A119 ♂
A103
23
A-Clan
B1 Pod,
B7 Matriline
B14 ♀ 1991
B17 2008
B16 ♂ 2004
B7 ♀ 1949
B19 2014
B10 ♂ 1979 B13 ♂ 1987
B3 ♂
B1 ♂B6 ♂B5 ♂
1973196419601953
B7 ♀
B10 ♂B12 ♂B14 ♀B13 ♂
B2 ♂
1954
B11 ♀
1929
B8 ♂
1965 1984
B15 ♂
B17 B18
1995
B16 ♂B19
2008
24
C16 ♂ 1989
C31 2009 C27 ♂ 2006
A-Clan
C1 Pod,
C6 Matriline
25
C8 ♀ 1975
C22 ♂ 1997 C34 2013
C14 ♂ 1985
A-Clan
C4 ♀C7 ♂C3 ♂
1954
1934
C1 ♂ C6 ♀
1953
C14 ♂C18 ♂C12
1979
C8 ♀C9 ♂
1972
C22 ♂C25C21 ♀
1994
C19 ♀
2004
C31 C27 ♂
C16 ♂
1953
1956
1991
C34
1991
26
C13 ♀ 1985
C32 2013 C36 2019
A-Clan
C1 Pod,
C10 Matriline
C28 2007
27
C10 ♀ 1972
C23 ♀ 1998
C35 2015
C26 ♀ 2004 C30 2009
C33 2014
A-Clan
C29 2009
C2 ♂
C11 ♂C10 ♀C15
196419581946
C5 ♀
1924
C20 ♂C17 ♂C23 ♀C26 ♀
C13 ♀
C24 ♂C28
C30
1993
C35
1989
C36 C33
2000
C32 C29
28
D12 ♀ 1982
D20 ♀ 1999
D29 ♂ 2014
D9 ♀ 1972
A-Clan
D1 Pod,
D12 Matriline
D1 Pod,
D9 Matriline
Since the death of C19,
C31 has been travelling
with D9.
29
D12 ♀
D20 ♀
D29 ♂
D9 ♀D8 ♀D4 ♂
1960
D16 D18
1995
1968
1987
D15
1987
D13 ♀ 1984
D23 ♀ 2005D19 ♀ 1998 D32 2018
D7 ♀
1943
D10 ♀D13 ♀
D24
2006
D19 ♀D27
1978
D32
A-Clan
D1 Pod,
D13 Matriline
D30
2015
D23 ♀
D27 2012
30
D17 ♀ 1990
D26 2010D21 ♂ 2005
D11 ♀ 1975
D25 ♂ 2007 D28 ♀ 2013
A-Clan
D1 Pod,
D11 Matriline
D31 2015
D3 ♀D1 ♂
1938
D2 ♂D11 ♀D5 ♂
19641956
D22D17 ♀D25 ♂D14
20051987
D26 D31
1954
D28 ♀
D21 ♂
31
H9 ♀ 1988
H16 2012H12 ♀ 2002
H5 ♀ 1973
H13 ♂ 2004
H17 2013
H15 2008
H6 ♀H1 ♂
1943
H3 ♀H5 ♀
H2 ♂
19661960
H7 ♂H10H4 ♂
19861975
H11
1981
H8 ♀
1994
H14
2005
H9 ♀
H15
H12 ♀
2000
H13 ♂
1954
H16
H17
A-Clan
H1 Pod,
H5 Matriline
32
I97 ♂ 2002
I131 2010I112 ♂ 2006
I54 ♀ 1983
I71 ♀ 1993 I142 2013
A-Clan
I1 Pod,
I19 Matriline
e I19 matriline does not
always travel as a cohesive
group.
33
I92 ♀ 1999
I155 2017
I117 ♂ 2005 I132 2010
A-Clan
I125 2008
I19 ♀
I70I56 I92 ♀I117 ♂
I54 ♀
I97 ♂I114
I132
1993
I155
I23 ♂
1974
1986
I71 ♀
I112 ♂ I131
I81
1997
I142
2006
1969
I125
34
I116 ♀ 2006
I69 ♀ 1991
I140 2013
I113 ♂ 2006
I40 ♀
I156I90 ♀I82 ♀
I116 ♀
I69 ♀
I140
I1 ♀I3 ♂
1953 1956
I139I113 ♂
I115
20051994
A-Clan
I1 Pod,
I40 Matriline
I133
I162
I162 2019
35
I40 ♀ 1980
I133 2010I90 ♀ 1999
I139 2012
I156 2017
I22 ♀ 1966
I121 ♂ 2007
I2 Pod,
I22 Matriline
I14 ♂I5 ♂I22 ♀I8 ♂I28 ♂
I39 ♂I55
1965 1974
I2 ♀
1929
1987
I121 ♂
1956 1956
1980
A-Clan
Individuals in the I40
matriline are not always
found together.
36
I91 ♂ 1996
I26 ♀ 1975
I118 ♂ 2006
I93 ♂ 1992 I148 2014
I18 Pod,
I17 Matriline
A-Clan
37
I26 ♀
I91 ♂I118 ♂
I93 ♂
I60
1987
I148
I50 ♀ 1983
I100 ♀ 2003 I130 2010
I89 ♂ 1996 I147 2014
I50 ♀
I100 ♀I130I89 ♂I147
I38
1980
I50 and her ospring are
not always found with the
rest of the I17 matriline.
I151
2015
A-Clan
38
I17 ♀
I57 ♀I99 ♀
I158
I87
1996
I88
2000
I127
2008
I149
I99 ♀ 2003
I57 ♀ 1989
I158 2016
I17 ♀ 1960
I149 2014
I124 ♀ 2007
I163 2019
I163
I124 ♀
I18 Pod,
I17 Matriline
continued
A-Clan
39
I134 2010
I21 ♀ 1979
I119 ♂ 2006 I157 2018
I18 Pod,
I18 Matriline
I150 2014
A-Clan
40
I20 ♀ 1965
I101 ♂ 2003I83 ♀ 1997
I141 2013
I126 ♀ 2008
A-Clan
I18 Pod,
I18 Matriline
continued
I52 ♂ 1986
I160 2019
41
A-Clan
I48 ♂ 1983
I24 ♀ 1980
I96 ♀ 2002
I53 ♂ 1986
I24, her progeny, and
I53 do not oen travel
with others in the I18
matriline.
I152 2015
I20 ♀
I66 I83 ♀I21 ♀
I119 ♂I126 ♀I95
I84
1996
I52 ♂
1991
I101 ♂
2001
I134 I141I157
I59I48 ♂
I7 ♀
1969
1988
I18 ♀
I24 ♀I53 ♂
I120
I49 ♂
1976
I58
1989
I73
1992
I96 ♀
1948
2007
I152
I160
I150
42
G37 ♀ 1984
G63 ♀ 1999 G80 ♀ 2006
G109 2015
G105 2012
G-Clan
G1 Pod,
G3 Matriline
G93 2010 G118 2019
G37 ♀
G106
G105
G80 ♀
G48 ♀
G109
G63 ♀G116 ♂G83
G93 G118
43
G48 ♀ 1990
G106 2013 G116 ♂ 2018
G-Clan
G83 2007
G61
1998
G56
1994
G20 ♀
G69 ♀G73 ♂G92
G3 ♀
G22 ♀G32 ♂
G51 ♀
G19 ♂
1976
G44
1989
G68
G1 ♂
G81 ♂
G86
G74
2005
2001
1954
G107 G108
G114 ♀
G104
G37, G48, and their
progeny are sometimes
observed separate from
others in the G20 lineage.
44
G51 ♀ 1992
G81 ♂ 2006 G108 2014
G-Clan
G1 Pod,
G3 Matriline
continued
G20 ♀ 1972
G69 ♀ 2001 G73 ♂ 2004 G92 2009
G107 2014
45
G22 ♀ 1979
G114 ♀ 2017 G32 ♂ 1982
G3 ♀ 1957
G104 2012G86 2008
G-Clan
G22 and her progeny do
not always travel with
others in the G3 matriline.
46
G52 ♀ 1993
G39 ♂ 1986
G1 Pod,
G16 Matriline
G46 ♀ 1991
G72 ♂ 2005
G1 Pod,
G46 Matriline
G5 ♂
1959
G29 ♀
G90
G45 ♂
1988
G46 ♀
G72 ♂
G30 ♀
1917
G24 ♀
1942
2008
1971
G-Clan
47
G31 ♀ 1981
G54 ♀ 1994 G84 ♂ 2007 G96 2011
G103 2012
G11 ♂
1963
G16 ♀
G39 ♂G52 ♀
G94
1972
G91
2008 2010
G18 ♀
1947
G31 ♀
G54 ♀G66 G84 ♂
G85
1999
2008
G103
G96
G-Clan
G1 Pod,
G31 Matriline
48
G60 ♂ 1998 G78 2004
G-Clan
G1 Pod,
G17 Matriline
G65 and G38 are oen
found separate from
others in the G17
matriline. Similarly, G60
and G78 normally travel
together independent of
their other maternal kin.
G25 ♀G9 ♂
1966
G78 G87G40 ♂G57 G60 ♂
1975
200819921987
G7 ♂G17 ♀
G38 ♂ G23 ♀
G59 ♂G50 ♀G65 ♂G79
1950 1954
1992
G102
2013
G97
2011
G110
49
G59 ♂ 1995 G79 2005G65 ♂ 2001
G50 ♀ 1991
G38 ♂ 1986
G-Clan
G110 2015
50
G70 ♀ 2003 G89 2008
G71 ♂ 2005 G98 2011
G-Clan
G12 Pod,
G2 Matriline
G34 ♀
G70 ♀G89
G49 ♀
G71 ♂
G55
1994
1977
G98
G2 ♀
G36G28 ♂G43 G53 ♀G77 ♀
1985
G117
1981 1989
G101
1990
G82 G113
51
G2 ♀ 1963
G77 ♀ 2005
G53 ♀ 1994
G101 2012 G117 2019
G-Clan
G82 2007 G113 2017
Individuals in the G2
matriline are not always
found together.
52
G62 ♀ 1999
G-Clan
G12 Pod,
G62 Matriline
G88 2008
G12 Pod,
G27 Matriline
G112 2017
G8 ♀
1971
G35 ♂G62 ♀
19901985
G47
G33 ♂ G27 ♀
G58 ♀
G41 ♀G64 ♀G76 ♀
G88 G111
2016
1979
G42
1986
G12 ♀
1956
G115 ♀G99
G100 ♂
G112 G75
2005
1987
G95
1974
53
G76 ♀ 2005G64 ♀ 2000G58 ♀ 1996 G100 ♂ 2012
G115 ♀ 2017 G99 2012
G-Clan
G95 2011
e G27 matriline does
not always travel as a
cohesive group.
54
I12 ♀ 1971
I105 ♂ 2004 I138 2012
G-Clan
I11 Pod,
I11 Matriline
I42 ♂ 1983
I13 ♀ 1974
I108 ♂ 2005
I12 ♀
I78 ♂I105 ♂
I47 ♂I138
I11 ♀
I13 ♀
I108 ♂I75
1995
I37 ♂
1979
I64 ♂
I42 ♂
I135
2011
1955
1990
1985 1997
55
G-Clan
I51 ♀ 1986
I129 2009
I106 ♂ 2004
I128 2009
I98 ♂ 2002
I144 2014
I11 Pod,
I16 Matriline
I16 ♀
I72 I85 I43 ♂
I129
I106 ♂
I128I109I51 ♀
1993
I98 ♂
200619981983
I144
I154
2016
1969
56
G-Clan
I27 ♀ 1974
I77 ♂ 1997 I107 ♂ 2004
I27 ♀
I63 ♀I77 ♂I107 ♂
I143
2014
1990
I11 Pod,
I27 Matriline
I4 ♀
I76 ♂ I102 ♀
I15 ♀
1953
I44 ♂I41 ♂I65 ♀
1985
I103
1980
I122 ♂I145
I153
2016
I161
2019
2003
57
G-Clan
I4 ♀ 1980
I76 ♂ 1997 I102 ♀ 2003
I11 Pod,
I4 Matriline
58
G-Clan
I65 ♀ 1990
I122 ♂ 2007 I145 2014
I11 Pod,
I65 Matriline
59
I62 ♂ 1988
I33 ♀ 1971
I110 ♀ 2006
I45 ♂ 1985
I33 ♀
I79 I94
I45 ♂I62 ♂
1997
I110 ♀
2001
I32 ♂
1964
I31 Pod,
I33 Matriline
I123 ♂I104 ♀
I35 ♀
I61 I67 ♂I74
1988 1994
I86
1998
I146 ♀
I137
I46 ♂I68 ♀
I80
1997
I36 ♀
1980
I111
2006
I31 ♀
1948
1985
I136
I159
2019 1991
G-Clan
60
I35 ♀ 1974
I123 ♂ 2007
I104 ♀ 2002 I137 2012
I146 ♀ 2014
I31 Pod,
I35 Matriline
I68 ♀ 1991
I136 2012
I31 Pod,
I68 Matriline
G-Clan
61
R73 2019
R29 ♀ 1994
R63 2015
R1 Pod,
R5 Matriline
R54 2010R48 ♀ 2006
R72 2019
R-Clan
62
R39 ♀ 2001
R35 ♀ 1998
R55 2010 R69 2018
R74 2019R65 2015 R59 2013
R1 Pod,
R5 Matriline
continued
R-Clan
63
R22 ♀ 1984
R50 ♀ 2007R44 ♂ 2004 R58 2011
R22 and her progeny are
seldom observed in the
presence of others in the
R4 lineage.
R68 2018
R66 2015
R73
R63 R55
R44 ♂R58
R29 ♀R39 ♀R35 ♀
R65
R50 ♀
R54
R48 ♀
R72
R74 R59 R69
R22 ♀
R66
R68
R-Clan
64
R36 ♂ 1998
R4 ♀ 1965
R41 ♀ 2002
R33 ♂ 1995
R26 ♂ 1988
R61 2014
R4 ♀
R33 ♂R36 ♂R41 ♀R28 ♂R26 ♂
R61
R-Clan
1992
R1 Pod,
R5 Matriline
continued
65
R18 ♀ 1968
R46 ♂ 2004
R37 ♂ 1999
R25 ♂ 1987
R19 R20 ♀
19791975
R18 ♀
R25 ♂R37 ♂
R21 ♂R46 ♂
1982
R31 ♂R32 R40 ♀R45
1997 2004
R51 R56
R62
R-Clan
R18 and her ospring are
usually found travelling
independent of others in
the R5 matriline.
R31 ♂ 1997
R30 ♂
R5 ♀
R24 ♀
R64
66
R24 ♀ 1987
R56 2011
R40 ♀ 2001
R30 ♂ 1994
R5 ♀ 1949
R51 2008
R62 2014
R-Clan
R1 Pod,
R5 Matriline
continued
R64 2015
67
R13 ♀ 1979
R47 ♀ 2005
R7 ♀
R11
R8 ♂
1962
R13 ♀
R47 ♀
1973
1946
R57 2011
R57
R-Clan
R1 Pod,
R13 Matriline
R71 2019
R71
68
R42 ♀ 2002 R49 2006R38 ♀ 2000
R60 2013
R17 ♀
R34 ♂R43 ♂
R27 ♂
1990
R23 ♀
R49 R67R38 ♀
R60
R42 ♀
R52 2009
R52
R-Clan
R1 Pod,
R17 Matriline
R70 2018
R70
R53
69
R23 ♀ 1985
R67 2016
R34 ♂ 1996
R17 ♀ 1971
R43 ♂ 2002
R-Clan
R53 2009
R17 and her male
ospring do not always
travel with R23 and her
progeny.
