MARINE MAMMAL SCIENCE, 33(1): 389–406 (January 2017)
©2016 Society for Marine Mammalogy
Louis M. Herman
“Do not go where the path may lead; go instead where there is no path and leave a
Ralph Waldo Emerson
Emerson’s words epitomize much of the journey, career, and remarkable life of Dr.
Louis M. Herman, a researcher and emeritus professor at University of Hawaii at
Manoa. Lou followed his own path and was a true trailblazer in the ﬁeld of marine
mammal science. To many, his name is synonymous with pioneering studies of both
dolphin cognition and humpback whale behavior. He was a charter member of the
Society for Marine Mammalogy and developed the world-renowned Kewalo Basin
Marine Mammal Laboratory (KBMML). At KBMML, Lou worked with dolphins
Keakiko, Nana, Puka, Akeakamai, Phoenix, Elele, and Hiapo, and scores of students
and colleagues, to conduct groundbreaking studies of dolphin sensory perception,
cognition, and language abilities. He initiated one of the longest continuous and
most productive scientiﬁc investigations of humpback whales in the world. Lou’s suc-
cess can, at least in part, be attributed to his tenacity and resilience perhaps best
illustrated by his ability to overcome the criminal theft of the dolphins Puka and Kea
in 1977. Like the phoenix bird, for which he named one of his new dolphins, Lou
emerged from this tragedy with even greater energy and determination, leaving a
legacy of nearly 60 yr of innovation and discovery, with a publication record extend-
ing from the late 1950s to the present day.
Over his distinguished career, Lou authored or coauthored 181 scientiﬁc publica-
tions, including 161 on marine mammals, and was an invited and keynote speaker at
countless conferences. His many discoveries on dolphins and whales were featured in
more than 230 media articles, television and radio programs, and documentary ﬁlms.
Along with a voracious appetite for research, Lou was a dedicated mentor. As a profes-
sor at University of Hawaii, he supervised 40 graduate students (see Appendix S1),
many of whom are now prominent ﬁgures in the ﬁeld of marine mammal science.
Lou created innovative internship programs that provided hands-on opportunities for
undergraduate students from around the world to get their start with marine mam-
mals. KBMML’s doors were also opened to hundreds of Earthwatch Institute volun-
teers and Dolphin Institute participants to assist in research efforts, as well as to
thousands of Hawaii’s school children to learn ﬁrst-hand about some of the latest dis-
coveries on dolphins and whales.
Lou also cared deeply about the conservation of marine mammals, serving for
many years as chair of the conservation committee on the Hawaiian Islands Hump-
back Whale National Marine Sanctuary Advisory Council. In many of his televi-
sion, ﬁlm, and radio interviews he discussed how his laboratory’s discoveries could
serve as a launch pad for inspiring individuals to care about and protect dolphins,
whales, and their marine habitats. In 1993, Lou, together with former student and
long-time colleague Adam Pack, founded The Dolphin Institute (TDI), a nonproﬁt
organization dedicated to dolphins and whales through education, research, and
In March 2015, Lou’s family, former students, and colleagues gathered in Hono-
lulu to celebrate his academic career and achievements at a surprise Festschrift. At
that event, Lou’s inspiring journey was retraced. Below, we provide a synopsis of
Lou’s many contributions to marine mammal science, as well as a short chronicle of
the adventures that led him there. A more complete biography of Lou’s early career
may be found in Herman (2012a).
Louis Marvin Herman was born in Queens, New York, to Jewish immigrants, the
youngest of four children. At an early age, Lou developed a life-long passion for
swimming and the ocean. Family outings in the 1930s were spent at the beach. Lou
was a competitive college swimmer at City College of New York, where he earned
his Bachelor’s and Master’s degrees in Psychology. Lou also spent his college summers
lifeguarding at New York City’s largest and busiest beach, Far Rockaway. Years later,
after moving to Hawaii, Lou swam in the Masters Nationals and Aloha State Games,
and frequently competed in Hawaii’s famous 2.4 mile open-ocean Waikiki Roughwa-
ter Swim beginning in 1972 and as recently as 2014.
In 1953, Lou enlisted in the Air Force where he served as an intelligence ofﬁcer
during and after the Korean War, interviewing repatriated Air Force pilots who had
been subject to Chinese interrogation techniques. In 1957, after spending 9 mo at
Emory University in Atlanta studying concept learning in rhesus monkeys, Lou
entered the graduate program at Penn State University. In 1961, Lou earned his
390 MARINE MAMMAL SCIENCE, VOL. 33, NO. 1, 2017
Ph.D. in Experimental Psychology, focusing on human information processing. Penn
State was also where Lou met and married fellow graduate student Hannah Schattner.
After graduation, Lou and Hannah headed to Columbus, Ohio, where Lou took up
his ﬁrst job at the North American Aviation Company. One of his projects involved
working on the problem of how to improve a sonar operator’s ability to correctly clas-
sify echo returns as “submarine” or “whale,” foreshadowing his future research study-
ing whales and dolphin echolocation.
The type of creativity and ingenuity that many of Lou’s students and colleagues
witnessed over the years in his studies of dolphins and whales was evident early in his
career. In 1962, Lou’s dissertation, which investigated how people process informa-
tion when confronted with competing demands from two simultaneous auditory
tasks, won the ﬁrst “Creative Talent Award” from the American Institutes for
Research (Harlow et al. 1962). Given the signiﬁcance of the award, Lou was inspired
to pursue a path in academia. In 1963, he became an Assistant Professor at Queens
College in New York City where he studied information processing and human per-
formance (e.g., Herman 1965, 1969; Herman and Bahrick 1966; Herman and Kan-
towitz 1969, 1970; Herman and McCauley 1969). In 1966, Lou and Hannah
traveled across the country to Hawaii where Lou took up a position as an Associate
Professor in the Psychology Department at the University of Hawaii at Manoa.
In addition to teaching classes in experimental psychology, Lou set up a new
undergraduate lab course to teach students the principles of conditioning using white
rats. However, he quickly turned his attention towards the study of learning and cog-
nition in dolphins after some encouragement from a student and realizing that little
had been reported about the subject. After spending the summer of 1967 conducting
a dolphin research project with his students at Hawaii’s Sea Life Park (Herman et al.
1969), Lou located an abandoned shark display facility adjacent to Ala Moana Beach
Park. In early 1969 he welcomed his ﬁrst dolphins to the property and in 1971 he
formally named the facility the Kewalo Basin Marine Mammal Laboratory. His vision
was to create a laboratory fully devoted to the scientiﬁc study of dolphin sensory per-
ception, cognition, and communication abilities. During KBMML’s tenure from
1971 to 2004, and under Lou’s direction and careful guidance, that vision was real-
At KBMML, Lou set out on a mission to characterize what he called the “cognitive
characteristics” of the bottlenose dolphin, including its abilities, specializations and
limitations (Herman 1980a). Together with his students and colleagues, Lou began
exploring visual and auditory sensory perceptual characteristics, auditory working
memory capacity, and concept formation abilities in the dolphin. With an innovative
apparatus of his own design he and his ﬁrst marine mammal graduate student Frank
Beach Jr. demonstrated that a dolphin could learn and apply a “win-stay, lose-shift”
rule across numerous novel problems after a single trial with arbitrary sounds, and at
levels of performance comparable to those obtained with nonhuman primates (Beach
and Herman 1972, Herman and Arbeit 1973). Lou also forged new ground in dol-
phin sensory perception, showing the dolphin’s ability to detect small degrees of fre-
quency modulation, as well as its sensitivity to other types of sounds (Herman and
Arbeit 1971a,b, 1972; Thompson and Herman 1975), and its sensitivity to temporal
differences in sounds (Yunker and Herman 1974). He made new discoveries about
dolphin short-term memory abilities for single novel sounds as well as lists of sounds,
and the mental processes involved in retaining these memories (Herman and Gordon
1974; Herman 1975; Thompson and Herman 1977, 1981; Herman 1980a; Herman
and Thompson 1982). In the 1970s, Lou and his students also mapped the dolphins’
visual acuity underwater and in-air, revealing its sensitivity in different areas of the
visible spectrum, and characterizing its limitations in discriminating different colors
(Herman et al. 1975, Madsen and Herman 1980).
Lou pioneered the scientiﬁc study of Hawaii’s humpback whales in the 1970s. In
1976, when most people were not even aware of the presence of humpback whales in
Hawaiian waters, Lou conducted the ﬁrst all-island aerial surveys of the population.
At that time, there were an estimated 500–800 individuals of this endangered whale
species migrating to Hawaii each year compared to the more than 10,000 individuals
that now make this journey. From small Cessna airplanes he and his students docu-
mented the whales’ presence, abundance, and distribution as well as the areas of high
calf density. These ﬂights were complemented by a boat-based effort to photograph
the whales at close range and to begin to identify individuals from the unique mark-
ings on their tail ﬂukes. In 1977, Lou published his ﬁrst seminal article on humpback
whales describing their numbers, distribution, and behavior in Hawaiian waters
(Herman and Antinoja 1977; also see Herman et al. 1980). It was here that he coined
the term “escort” to identify the adult whale often seen accompanying a mother-calf
pair. These escorts are now known to be males competing for access to the females
but at the time it was considered possible that they were female “aunties” helping to
guard the calf. Always moving in new directions, Lou also brieﬂy entered the world
of historical ecology to investigate whether humpback whales had a historical pres-
ence in Hawaiian waters (Herman 1979).
During the winter breeding season of 1977, Lou, Ron Antinoja and graduate stu-
dent Paul Forestell set up KBMML’s ﬁrst whale research ﬁeld camp on the island of
Lanai. Inﬂatable boats were launched daily to document humpback whale identities,
social organization and behavior. In 1978, young marine mammal scientists Randy
Wells and Giuseppe Notarbartolo di Sciara joined the humpback whale project, now
launched from the islands of Molokai and Maui. In addition to helping with more
aerial surveys and underwater behavioral studies, Randy, Giuseppe, and Paul assisted
Lou in the development of what has now become one of the largest long-term archival
catalogs of individually identiﬁed humpback whales.
In 1980, Lou ventured north to Sitka, Alaska, with Hannah, their 1-yr-old daugh-
ter Elia, and students Paul Forestell, Scott Baker, and Bill Stifel, to inaugurate
KBMML’s studies of North Paciﬁc humpback whales in their feeding grounds.
KBMML’s early research in Alaska continued through 1986, providing fundamental
information on the migratory movements of individually identiﬁed humpbacks
between winter and summer grounds across the North Paciﬁc, mapping the long-
term associations within cooperative feeding groups, and investigating the impact of
vessel trafﬁc in Glacier Bay National Park and Preserve on humpback whale behavior
(Baker et al. 1982, 1983, 1985, 1986; Baker and Herman 1984a, 1985, 1989; Perry
et al. 1985). In 2007, Adam Pack resumed KBMML’s studies of humpbacks in
Alaska and continues this work today photographing many of the same individuals
ﬁrst identiﬁed by Lou and his team in 1980.
Whether he was investigating whales or dolphins, Lou’s scientiﬁc approach was
always rigorous and creative. In the late 1970s and early 1980s, he again broke new
ground with a study investigating the dolphin’s ability to comprehend sentences
within two artiﬁcial language systems. One system was acoustically based, with dif-
ferent sounds representing different objects, agents, actions, modiﬁers, and relation-
ships. The other system was visually based, with these same items represented by
different gestures provided by a human. The languages also had sets of grammatical
rules that governed the order in which different symbol types could appear to create
392 MARINE MAMMAL SCIENCE, VOL. 33, NO. 1, 2017
different meanings. Phoenix, who was taught the acoustic language, and Akeakamai,
who was taught the visual language, excelled in their abilities to comprehend novel
sentences, importantly showing that they could understand that the same words
placed in different orders created entirely different meanings. As Lou often liked to
explain it: “A Venetian blind is not the same as a blind Venetian.” Thus, Lou showed
for the ﬁrst time that a dolphin could take into account both the semantic and syntac-
tic components within sentences, the two fundamental features of human language
(Herman et al. 1984, 1993a,b; see also Herman 1986, 1987; Shyan and Herman
1987; Herman 1988, 1989; Herman and Morrel-Samuels 1990; Morrel-Samuels and
Herman 1993; Herman 2002a, 2009a; Herman and Uyeyama 1999). Further work
showing that Akeakamai could report on the presence or absence of named objects
from her habitat, reinforced the idea that the dolphins understood the referential
function of their language symbols (Herman and Forestell 1985, Herman et al.
1993b). Lou’s whole approach to working with a dolphin was to tutor and educate it
the way you would a child to reveal the full ﬂower of its intellect, and always to be
alert and responsive to what the dolphin’s behavior indicated about what it under-
stood. He passed this wisdom on to his human students, who he taught to be ﬂexible
and willing to modify their approach to any research project based on the dolphins’
responses, which so often were unanticipated and remarkable.
The early 1980s also marked the publication of Lou’s ﬁrst edited book “Cetacean
Behavior: Mechanisms and Functions” (Herman 1980b). In addition to his own chap-
ters on dolphin cognitive characteristics, communication systems in cetaceans (with
William Tavolga) and the social and ecological correlates of cetacean vision and visual
appearance (with Carolyn Madsen), Cetacean Behavior contains chapters by notable
researchers Bill Dawson, R. H. Defran, Ken Norris, Art Popper, Karen Pryor, and
Randy Wells. It is considered by many to be a classic text in marine mammal science.
As noted in a recent letter to Lou by Andy Read: “[Cetacean Behavior] turned out to
be formative for me...it also introduced a new and exciting world of social behavior
and cognition, underpinned by the concept that whales and dolphins were individuals
with rich social lives.”
Throughout the 1980s, 1990s, and 2000s, Lou, his students, and colleagues con-
tinued to pursue the answers to key questions about dolphin sensory perception, cog-
nition, and communication. New light was shed on dolphin memory, abstraction
abilities, concept formation, sound perception, echolocation, social cognition, self-
awareness, and creativity. In many cases, the dolphin’s cognitive skills rivaled those
of apes leading Lou to suggest, “The major link that cognitively connects the other-
wise evolutionary divergent delphinids and primates may be social pressure...” (Her-
man 1980a, p. 421).
Some notable discoveries with the dolphins were:
•An ability to imitate both arbitrary novel sounds and motor behaviors, a rarity in
the non-human animal kingdom (Richards et al. 1984, Xitco 1988, Herman
2002b), and one of the ﬁrst studies of vocal mimicry in nonhuman mammals.
•An ability to apply a concept of “sameness” that was shown earlier with auditory
stimuli (Herman and Gordon 1974) to a wide variety of visual stimuli, thus
demonstrating an invariance of cognitive performance across visual and auditory
modalities (Herman et al. 1989 1994; Herman 1990; Mercado et al. 2000).
•An ability to spontaneously understand television displays as representing the
real world; a skill that some ape species ﬁnd difﬁcult without human assistance
(Herman et al. 1990).
•An ability to categorize different melodic sequences and spontaneously recognize
the same sequence shifted across octaves; a skill difﬁcult for some songbirds (Ral-
ston and Herman 1995; also see Richards et al. 1984 for spontaneous octave
shifts in vocal mimicry).
•An ability to spontaneously recognize complex shapes across the senses of echolo-
cation and vision (Pack and Herman 1995; Herman et al. 1998; Pack et al.
2002b, 2004), thus demonstrating that through echolocation, dolphins can
appreciate the spatial structure of objects.
•An ability to make ﬁne angular discriminations by echolocation (Branstetter
et al. 2003, 2007).
•An ability to demonstrate an awareness of its own actions (Mercado et al. 1998,
1999), thus revealing new dimensions of dolphin self-awareness beyond mirror-
self recognition (Reiss and Marino 2001).
•An ability to understand human directed referential pointing at distally placed
objects; a skill which comes easily to the dolphin (Herman et al. 1999; Pack and
Herman 2004, 2006, 2007a; also see Xitco et al. 2001 for spontaneous dolphin
productive pointing), but which appears difﬁcult for some ape species (e.g., Povi-
nelli et al. 1997, Call and Tomasello 1994).
•An ability to understand symbolic references to its own body parts via human
gestures, expanding further our understanding of dolphin self-awareness (Her-
man et al. 2001).
•An ability to be vigilant either visually or acoustically, by reporting amidst dis-
tractor stimuli each occurrence of either key images appearing on a television
monitor or key sounds projected underwater (Hoffmann-Kuhnt 2003).
•An ability to perform self-created behaviors in tandem with another dolphin
(Braslau-Schneck 1994, Herman 2006), a skill that demonstrates both creativity
and intraspecies coordination.
In parallel with the dolphin work, KBMML continued ﬁeld research efforts to
address key issues in humpback whale behavior, ecology, and communication, focus-
ing primarily on the waters off Hawaii Island and Maui. In addition to KBMML’s in-
house work, several of these studies were collaborations spearheaded by other research
groups. Both efforts yielded new insights into fundamental features of humpback
whale ecology such as migratory movements within breeding grounds and between
breeding and feeding grounds (Baker and Herman 1981; Gabriele et al. 1996; Craig
and Herman 1997; Cerchio et al. 1998; Salden et al. 1999; Calambokidis et al. 2000,
2001; Craig et al. 2003); reproductive histories, crude birth rates, and calving rates of
females (Baker et al. 1987, Perry et al. 1990, Craig and Herman 2000, Herman et al.
2011); population size increases and distributional changes in the breeding grounds
(Baker and Herman 1987; Calambokidis et al. 1997; Mobley et al. 1999); and calf
mortality and adult survival rates (Gabriele et al. 2001, Mizroch et al. 2004).
Other studies focused on describing and understanding the behaviors, social inter-
actions, and communication systems involved in the humpback whale mating sys-
tem. Notable among these were studies:
•Revealing the nuances of male-male aggressive behavior during competition over
females (Baker and Herman 1984b, Helweg and Herman 1993, E. Herman et al.
•Documenting that afﬁliations between humpback whales in the breeding
grounds (other than mother-calf pairs) tend to be transient (Mobley and Herman
394 MARINE MAMMAL SCIENCE, VOL. 33, NO. 1, 2017
•Documenting the effects of vessel trafﬁc on humpback whale behavior (Bauer
and Herman 1986).
•Showing that playback of humpback song to pods in Hawaiian waters resulted in
few whales approaching the sound source compared to playback of social sounds
from highly active competitive groups or a humpback feeding call recorded in
Alaskan waters by Baker (1985). Importantly, neither mother-calf pods nor
known females approached during song playback, indicating that unlike male
song in some songbird species, male song in humpbacks does not appear to
attract females to an individual singer (Mobley et al. 1986, 1988).
•Showing both shared and distinct themes in comparisons of male song across
Japan, Hawaii and Mexico (Helweg et al. 1990).
•Characterizing various aspects of humpback whale song and proposing sound
production mechanisms (Helweg et al. 1992; Mercado et al. 2003, 2005, 2010;
Green et al. 2007, 2011).
•Demonstrating diurnal and seasonal variations in behavior and pod characteris-
tics (Helweg and Herman 1994, Craig et al. 2002, Pack and Herman 2007b).
•Characterizing the distribution, movements and spacing of male singers (Frankel
et al. 1995).
•Reporting the unusual nonagonistic behaviors by a male towards another male
that died within a competitive group (Pack et al. 1998).
•Showing that individual females when with calf tend to prefer waters off Maui
and when without calf tend to prefer waters off Hawaii Island (Craig and Herman
•Describing residency characteristics of individual whales in the breeding grounds
(Craig et al. 2001).
•Showing that males preferentially associate with females without calf (i.e., those
with a higher reproductive potential), rather than females with calf (i.e., those
with a lower reproductive potential), and that individual females tend to attract
more escorts when they are without calf than when they are with calf (Craig et al.
•Describing male sexual behavior in a variety of contexts (Pack et al. 2002a).
•Showing that male body size confers an advantage in competition (Spitz et al.
•Showing that individual females vary their migratory timing across years,
depending on their reproductive status (Craig et al. 2003).
•Showing that maximum source levels of humpback song vary between units from
151 to 173 dB re 1lPa, and high frequency harmonics extend beyond 24 kHz
(Au et al. 2006).
•Showing that larger females tend to attract greater numbers of escorts and pro-
duce larger calves than do smaller females (Pack et al. 2009).
•Showing that in male-female dyads, mature females preferentially associate with
large mature males, mature males are less discriminatory among females, and
immature males and females tend to pair together (Pack et al. 2012).
•Showing that pursuit by male escorts is energetically costly to maternal females,
and thus maternal females tend to segregate themselves and their calves in shal-
low waters to avoid male harassment (Craig et al. 2014).
Along the way, Lou coauthored a catalog of North Paciﬁc humpback whale tail
ﬂuke images and life-history data (Perry et al. 1988); coedited a book, “Language and
Communication: Comparative Perspectives” (Roitblat et al. 1993); traveled to Ecua-
dor to carry out ﬁeld research on the Amazon river dolphin in the Rio Largarto Cocha
(Herman et al. 1996); published various syntheses of his work (Herman 1991, 2000,
2002c, 2006, 2009b, 2010; Herman and Pack 1994, 2001; Herman et al. 2008);
worked with his daughter Elia to deploy National Geographic’s Crittercam (suction
cup mounted video, audio, and data logging tags) on humpback whales for the ﬁrst
time in a breeding ground (E. Herman et al. 2007); was awarded with Adam Pack
and Matthias Hoffmann-Kuhnt the American Psychological Association’s Division 6’s
F. A. Beach Comparative Psychology Award for the Best Paper published in 1998 in the
Journal Comparative Psychology (Herman et al. 1998); and saw KBMML’s work fea-
tured in the United States and internationally in scores of newspaper and magazine
articles such as National Geographic (1979, 2008, 2015), People Magazine (1979),
the New York Times (1980), Daily Telegraph (1988), Femina (France) (1991), Bart
(Japan) (1994), TIME (1996), The Economist (1996), National Wildlife (2002,
2003), BBC Wildlife (U.K.) (2003), Smithsonian (2008); as well as in numerous tele-
vision and ﬁlm documentaries, including NOVA’s “Signs of the Apes: Songs of the
Whales” (1983), the MacGillivray Freeman IMAX ﬁlm “The Discoverers” (1993),
the PBS documentary “Dolphins with Robin Williams” (1997), BBC’s Animal
Minds (U.K., 1999), BBC’s Wildlife on One with Sir David Attenborough: Dol-
phins–Deep Thinkers? (U.K., 2003), and National Geographic’s “Humpbacks: Inside
the Pod” (2008).
In December of 2005, Lou retired from the University of Hawaii, but not from
writing and research. In tribute to the dolphins, Lou returned to the question he
started with all those years ago: What is the large dolphin brain capable of? This led
him to publish insightful works on dolphin rational behavior (Herman 2006), dol-
phin cognition (Herman 2010), and dolphin awareness of body and self (Herman
2012b). He also tackled the 32 yr of accumulated data on individually photographed
humpback whales in Hawaii and in 2011 produced a seminal paper on long-term
resightings of humpbacks (Herman et al. 2011). Two years later, Lou published a
paper entitled “Humpback whale song: Who sings?” in which he showed that both
immature and mature males participate in the chorus of humpback whale song on
the breeding grounds (Herman et al. 2013). Lou’s last two solo publications, com-
pleted during the ﬁnal months of his life, represented the culmination of all that he
had learned and taught others about dolphins and humpback whales. The ﬁrst paper
(Herman 2017) is an updated chapter in the third edition of the “Encyclopedia of
Marine Mammals”that brings together the current state of knowledge on dolphin
language abilities and cognition. The second paper, entitled “The multiple functions
of male song within the humpback whale (Megaptera novaeangliae) mating system:
Review, evaluation, and synthesis” (Herman, in press), brings together the historical
and most recent information on humpback whale song and discusses its place within
the humpback whale mating system. This paper was also an opportunity for Lou to
emphasize with new insights, his original lek theory of the humpback whale mating
system, which he ﬁrst described in his 1980 book (Herman and Tavolga 1980). In
recognition of his achievements, in 2008, Lou’s work was listed among the top 100
pioneering accomplishments at the University of Hawaii during its 100 yr history.
And in 2012, he was awarded the University of Hawaii College of Social Sciences
Award for Distinguished Retired Faculty.
Over the course of his research career, Lou continually demonstrated his gifts for
innovation, keen insight, and creative problem solving, whether it was designing a
language system to communicate with dolphins (Herman 1980a, Herman et al.
396 MARINE MAMMAL SCIENCE, VOL. 33, NO. 1, 2017
1984), or working with graduate students to develop a new technique, called “under-
water videogrammetry,” to measure the body sizes of humpback whales at sea (Spitz
et al. 2000). Perhaps one of the best examples of his never-ending ability to think cre-
atively occurred in 1985, when “Humphrey the Wrong Way Whale” lost its way up
the Sacramento River. In a conference call of whale experts, it was Lou who suggested
that instead of the aversive approach that had been tried unsuccessfully to redirect
Humphrey, a feeding call that Lou’s graduate student Scott Baker had recorded in
Alaska (Baker 1985), and which had proven attractive to individual whales when
played back in Hawaii (Mobley et al. 1988), could lure Humphrey back to sea. Lou
and his staff put together the tapes and instructions and shipped them off to Califor-
nia, and on cue Humphrey turned and traveled to the source of the sound and out to
the ocean over a 2 d period.
Lou leaves behind a professional and personal legacy. KBMML was where many
students ﬁrst “cut their research teeth.” honing their skills in dolphin training and
whale ﬁeld observations, and learning the art of research design, hypothesis testing,
data analysis and scientiﬁc writing. Lou challenged his students to follow his model
and think outside the box when it came to designing studies and problem solving.
But perhaps most importantly, he had a profound impact on the lives of so many
people—students, interns, volunteers—because of their exposure to all that he cre-
ated. “Lou changed my life” is the most consistent theme echoed in the countless
messages that have been received over the last couple of years from those who worked
with him in various capacities and who themselves have followed a wide range of per-
sonal and professional paths. Even the experiences professional colleagues were
afforded by Lou had profound personal impacts as noted by the late Stan Kuzcaj who
wrote: “The invitation to collaborate with him (Lou) on dolphin cognition and com-
munication literally transformed my research career.”
In one recent letter, Lou’s former Ph.D. student, Paul Forestell, currently Provost
and Vice President for Academic Affairs at Keuka College in Keuka Park, New York,
wrote the following to Lou:
...(one) way we can gain immortality is through our ideas, and our intellectual
“genes” that we pass on through our ability to teach. You should take great
pride and solace in knowing there are so many of us wandering around promot-
ing ideas and actions that are rooted in your cognitive “family tree.” I am
honored to be part of your intellectual heritage.
And as Phil Clapham wrote:
Lou was one of the early pioneers of research on living whales and dolphins,
and those of us who followed in his footsteps owe him a great debt. Since the
advent of non-lethal methods to study cetaceans, numerous scientists have con-
tributed to our knowledge of these remarkable animals; yet it is unlikely that
many of us could claim that the ﬁeld today would be signiﬁcantly different,
and considerably less advanced, had we not worked within it. That is not true
of Lou Herman.
Indeed, it is difﬁcult to imagine what our understanding of dolphin intelligence
and whale behavior would be like without the wealth of knowledge revealed from
Lou’s work. His discoveries will live on through the marvelous trail he has left and
through those whom he inspired to carry on with his quest to understand the com-
plexities of cetacean cognition and behavior.
To continue to support burgeoning minds in the ﬁeld, the Herman family has set up the Louis
M. Herman Scholarship Fund to support students engaged in cognitive and behavioral research
of whales or dolphins. A. A. Pack may be contacted by e-mail for more information about the
scholarship fund and how to make a contribution.
Au, W. W. L., A. A. Pack, M. O. Lammers, L. M. Herman, M. H. Deakos and K. Andrews.
2006. Acoustic properties of humpback whale songs. Journal of the Acoustical Society of
Baker, C. S. 1985. The population structure and social organization of humpback whales
(Megaptera novaeangliae) in the central and eastern North Paciﬁc. Doctoral dissertation,
University of Hawaii, Honolulu, HI. 306 pp.
Baker, C. S., and L. M. Herman. 1981. Migration and local movement of humpback whales
(Megaptera novaeangliae) through Hawaiian waters. Canadian Journal of Zoology 59:460–
Baker, C. S., and L. M. Herman. 1984a. Seasonal contrasts in the social behavior of the
humpback whale. Cetus 5:14–16.
Baker, C. S., and L. M. Herman. 1984b. Aggressive behavior between humpback whales
(Megaptera novaeangliae) wintering in Hawaiian waters. Canadian Journal of Zoology
Baker, C. S., and L. M. Herman. 1985. Whales that go to extremes. Natural History 94:52–
Baker, C. S., and L. M. Herman. 1987. Alternative population estimates of humpback whales
(Megaptera novaeangliae) in Hawaiian waters. Canadian Journal of Zoology 65:2818–
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whales to vessel trafﬁc: Experimental and opportunistic observations. Final Report to the
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Baker, C. S., L. M. Herman, B. G. Bays and W. F. Stifel. 1982. The impact of vessel trafﬁc on
the behavior of humpback whales in Southeast Alaska: 1981 season. Final Report to the
National Marine Mammal Laboratory, National Marine Fisheries Service. Contract No.
Baker, C. S., L. M. Herman, B. G. Bays and G. B. Bauer. 1983. The impact of vessel trafﬁc on
the behavior of humpback whales in Southeast Alaska: 1982 season. Final Report to the
National Marine Mammal Laboratory, National Marine Fisheries Service, Contract No.
Baker, C. S., L. M. Herman, A. Perry, W. S. Lawton, J. M. Straley and J. H. Straley. 1985.
Population characteristics and migration of summer and late season humpback whales
(Megaptera novaeangliae) in Southeastern Alaska. Marine Mammal Science 1:304–323.
Baker, C. S., L. M. Herman, A. Perry, et al. 1986. Migratory movement and population
structure of humpback whales (Megaptera novaeangliae) in the central and eastern North
Paciﬁc. Marine Ecology Progress Series 31:105–119.
Baker, C. S., A. Perry and L. M. Herman. 1987. Reproductive histories of female humpback
whales Megaptera novaeangliae in the North Paciﬁc. Marine Ecology Progress Series
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Beach, F. A., III, and L. M. Herman. 1972. Preliminary studies of auditory problem solving
and intertask transfer by the bottlenose dolphin. The Psychological Record 22:49–62.
Branstetter, B. K., S. J. Mevissen, L. M. Herman, A. A. Pack and S. Roberts. 2003.
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structure of humpback whales in the North Paciﬁc. Marine Mammal Science 17:769–
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orangutans (Pongo pygmaeus). Journal of Comparative Psychology 108:307–317.
Cerchio, S., C. M. Gabriele, T. F. Norris and L. M. Herman. 1998. Movements of humpback
whales between Kauai and Hawaii: Implications for population structure and abundance
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humpback whales (Megaptera novaeangliae) to the Hawaiian Islands. Canadian Journal of
Craig, A. S., and L. M. Herman. 2000. Habitat preferences of female humpback whales
Megaptera novaeangliae in the Hawaiian Islands are associated with reproductive status.
Marine Ecology Progress Series 193:209–216.
Craig, A. S., L. M. Herman and A. A. Pack. 2001. Estimating residence times of humpback
whales in Hawaii. Report to the Hawaiian Islands Humpback Whale National Marine
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migration of a North Paciﬁc humpback whale. Marine Mammal Science 12:457–465.
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membership of humpback whales (Megaptera novaeangliae) wintering in Hawaiian waters.
Helweg, D. A., L. M. Herman, S. Yamamoto and P. H. Forestell. 1990. Comparison of songs
of humpback whales (Megaptera novaeangliae) recorded in Japan, Hawaii, and Mexico
during the winter of 1989. Scientiﬁc Reports of Cetacean Research 1:1–20.
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in Tursiops truncatus. Pages 79–87 in Proceedings of the Eighth Annual Conference on
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Herman, L. M., and P. H. Forestell. 1985. Reporting presence or absence of named objects by
a language-trained dolphin. Neuroscience and Biobehavioral Reviews 9:667–691.
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Herman, L. M., and B. H. Kantowitz. 1970. The psychological refractory period effect: Only
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explanation in the study of behavior. Volume 1. Interpretation, intentionality, and
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recognition of veridical and degraded video displays of an artiﬁcial gestural language.
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and develop abstract concepts. Marine Mammal Science 10:70–80.
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perceive the spatial structure of objects through echolocation. Journal of Comparative
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Pack. 1999. Dolphins (Tursiops truncatus) comprehend the referential character of the
human pointing gesture. Journal of Comparative Psychology 113:1–18.
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representations of its body parts. Animal Learning and Behavior 29:250–264.
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2011. Resightings of humpback whales in Hawaiian waters over spans of 10-32 years:
Site ﬁdelity, sex ratios, calving rates, female demographics, and the dynamics of social
and behavioural roles of individuals. Marine Mammal Science 27:736–768.
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Hoffmann-Kuhnt, M. 2003. Visual and auditory vigilance in the bottlenosed dolphin.
Doctoral dissertation. Freie Universit€at Berlin, Berlin, Germany. 175 pp.
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appearance. Pages 101–147 in L. M. Herman, ed. Cetacean behavior: Mechanisms and
functions. Wiley Interscience, New York, NY.
Mercado, E., III, S. O. Murray, R. K. Uyeyama, A. A. Pack and L. M. Herman. 1998. Memory
for recent actions in the bottlenosed dolphin (Tursiops truncatus): Repetition of arbitrary
behaviors using an abstract rule. Animal Learning and Behavior 26:210–218.
Mercado, E., III, R. K. Uyeyama, A. A. Pack, and L. M. Herman. 1999. Memory for action
events in the bottlenosed dolphin. Animal Cognition 2:17–25.
Mercado, E. M., III, D. A. Killebrew, A. A. Pack, I. V. B. Macha and L. M. Herman. 2000.
Generalization of same-different classiﬁcation abilities in bottlenosed dolphins.
Behavioural Processes 50:79–94.
Mercado, E., III, L. M. Herman and A. A. Pack. 2003. Stereotypical sound patterns in
humpback whale songs: Usage and function. Aquatic Mammals 29:37–52.
Mercado, E., III, L. M. Herman and A. A. Pack. 2005. Song copying by humpback whales:
Themes and variations. Animal Cognition 8:93–102.
Mercado, E., III, J. Schneider, A. A. Pack and L. M. Herman. 2010. A source-ﬁlter model of
sound production by humpback whales. Journal of the Acoustical Society of America
Mizroch, S. A., L. M. Herman, J. M. Straley, et al. 2004. Estimating the adult survival rate of
central North Paciﬁc humpback whales (Megaptera novaeangliae). Journal of Mammalogy
Mobley, J. R., Jr., and L. M. Herman. 1985. Transience of social afﬁliations among humpback
whales (Megaptera novaeangliae) on the Hawaiian wintering grounds. Canadian Journal of
Mobley, J. R., Jr., L. M. Herman and A. S. Frankel. 1986. Sound playback experiments with
humpback whales in the Hawaiian wintering grounds. Sea Grant Quarterly 8:1–6.
Mobley, J. R., Jr., L. M. Herman and A. S. Frankel. 1988. Responses of wintering
humpback whales (Megaptera novaeangliae) to playback of recordings of winter and
summer vocalizations and synthetic sound. Behavioral Ecology and Sociobiology
Mobley, J. R., Jr., G. B. Bauer and L. M. Herman. 1999. Changes over a ten-year interval in
the distribution and relative abundance of humpback whales (Megaptera novaeangliae)
wintering in Hawaiian waters. Aquatic Mammals 25:63–72.
Morrel-Samuels, P., and L. M. Herman. 1993. Cognitive factors affecting comprehension of
gesture language signs: A brief comparison of dolphins and humans. Pages 211–222 in
H. R. Roitblat, L. M. Herman and P. Nachtigall, eds. Language and communication:
Comparative perspectives. Lawrence Erlbaum Associates, Hillsdale, NJ.
Pack, A. A., and L. M. Herman. 1995. Sensory integration in the bottlenosed dolphin:
Immediate recognition of complex shapes across the senses of echolocation and vision.
Journal of the Acoustical Society of America 98:722–733.
Pack, A. A., and L. M. Herman. 2004. Dolphins (Tursiops truncatus) comprehend the referent
of both static and dynamic human gazing and pointing in an object choice task. Journal
of Comparative Psychology 118:160–171.
Pack, A. A., and L. M. Herman. 2006. Dolphin social cognition and joint attention: Our
current understanding. Aquatic Mammals 32:443–460.
Pack, A. A., and L. M. Herman. 2007a. The dolphin’s (Tursiops truncatus) understanding of
human gaze and pointing: Knowing what and where. Journal of Comparative Psychology
Pack, A. A., and L. M. Herman. 2007b. Using electronic maps of real-time whale locations to
mitigate vessel collisions with whales. Final Report to the National Fish and Wildlife
Foundation, Washington, DC. 78 pp.
Pack, A. A., D. R. Salden, M. Ferrari, D. Glockner-Ferrari, L. M. Herman, H. A. Stubbs and
J. M. Straley. 1998. Male humpback whale dies in competitive group in Hawaii. Marine
Mammal Science 14:861–873.
Pack, A. A., L. M. Herman, A. S. Craig, S. S. Spitz and M. H. Deakos. 2002a. Penis extrusions
by humpback whales (Megaptera novaeangliae). Aquatic Mammals 28:131–146.
Pack, A. A., L. M. Herman, M. Hoffmann-Kuhnt and B. K. Branstetter. 2002b. The object
behind the echo: Dolphins (Tursiops truncatus) perceive object shape globally through
echolocation. Behavioural Processes 58:1–26.
Pack, A. A., L. M. Herman and M. Hoffmann-Kuhnt. 2004. Dolphin echolocation shape
perception: From sound to object. Pages 288–298 in J. Thomas, C. Moss and M. Vater,
eds. Advances in the study of echolocation in bats and dolphins. University of Chicago
Press, Chicago, IL.
Pack, A. A., L. M. Herman, S. S. Spitz, S. Hakala, M. H. Deakos and E. Y. K. Herman. 2009.
Male humpback whales in the Hawaiian winter grounds preferentially associate with
larger females. Animal Behaviour 77:653–662.
Pack, A. A., L. M. Herman, S. S. Spitz, et al. 2012. Size assortative pairing and discrimination
of potential mates by humpback whales in the Hawaiian breeding grounds. Animal
Perry, A., C. S. Baker and L. M. Herman. 1985. The natural history of humpback whales in
Glacier Bay, Alaska. Report to the National Park Service.
Perry, A., J. R. Mobley Jr., C. S. Baker and L. M. Herman. 1988. Humpback whales of the
central and eastern North Paciﬁc: A catalog of individual identiﬁcation photographs. Sea
Grant Miscellaneous Reports, UNIHI-SEA GRANT-NR-88-02.
Perry, A., C. S. Baker and L. M. Herman. 1990. Population characteristics of individually
identiﬁed humpback whales in the central and eastern North Paciﬁc: A summary and
critique. Report of the International Whaling Commission (Special Issue 12):307–317.
Povinelli, D. J., J. E. Reaux, D. T. Bierschwale, A. D. Allain and B. B. Simon. 1997.
Exploitation of pointing as a referential gesture in young children, but not adolescent
chimpanzees. Cognitive Development 12:423–461.
Ralston, J. V., and L. M. Herman. 1995. Perception and generalization of frequency contours
by a bottlenose dolphin (Tursiops truncatus). Journal of Comparative Psychology
Reiss, D., and L. Marino. 2001. Mirror self-recognition in the bottlenose dolphin: A case of
cognitive convergence. Proceedings of the National Academy of Sciences 98:5937–5942.
Richards, D. G., J. P. Wolz and L. M. Herman. 1984. Vocal mimicry of computer generated
sounds and vocal labeling of objects by a bottlenosed dolphin, Tursiops truncatus. Journal
of Comparative Psychology 98:10–28.
Roitblat, H. R., L. M. Herman and P. Nachtigall, eds. 1993. Language and communication:
Comparative perspectives. Lawrence Erlbaum Associates, Hillsdale, NJ.
Salden, D. R., L. M. Herman, M. Yamaguchi and F. Sato. 1999. Multiple visits of individual
humpback whales (Megaptera novaeangliae) between the Hawaiian and Japanese winter
grounds. Canadian Journal of Zoology 77:504–508.
Shyan, M. R., and L. M. Herman. 1987. Determinants of recognition of gestural signs in an
artiﬁcial language by Atlantic bottle-nosed dolphins (Tursiops truncatus) and humans
(Homo sapiens). Journal of Comparative Psychology 101:105–119.
Spitz, S. S., L. M. Herman and A. A. Pack. 2000. Measuring sizes of humpback whales
(Megaptera novaeangliae) through underwater videogrammetry. Marine Mammal Science
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Spitz, S. S., L. M. Herman, A. A. Pack and M. H. Deakos. 2002. The relation of body size of
male humpback whales to their social roles on the Hawaiian winter grounds. Canadian
Journal of Zoology 80:1938–1947.
Thompson, R. K. R., and L. M. Herman. 1975. Underwater frequency discrimination in the
bottlenosed dolphin (1–140 kHz). Journal of the Acoustical Society of America 57:943–
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ADAM A. PACK,Departments of Psychology and Biology, University of Hawaii at Hilo, 200
West Kawili Street, Hilo, Hawaii 96720, U.S.A. and The Dolphin Institute, PO Box 6279,
Hilo, Hawaii 96720, U.S.A.; ELIA Y. K. HERMAN,State of Hawaii Department of Land and
Natural Resources, 1151 Punchbowl Street #330, Honolulu, Hawaii 96813, U.S.A. and The
Dolphin Institute, PO Box 6279, Hilo, Hawaii 96720, U.S.A.; C. SCOTT BAKER,Marine
Mammal Institute and Department of Fisheries and Wildlife, Hatﬁeld Marine Science Center,
Oregon State University, 2030 SE Marine Science Drive, Newport, Oregon 97365, U.S.A.;
GORDON B. BAUER,Division of Social Sciences, New College of Florida, 5800 Bay Shore
Road, Sarasota, Florida 34243, U.S.A.; PHILLIP J. CLAPHAM,Cetacean Assessment and Ecol-
ogy Program, NOAA Marine Mammal Laboratory, Alaska Fisheries Science Center, 7600 Sand
Point Way NE, Seattle, Washington 98115, U.S.A.; RICHARD C. CONNOR,Biology Depart-
ment, University of Massachusetts Dartmouth, 285 Old Westport Road, Dartmouth, Mas-
sachusetts 02747, U.S.A.; ALISON S. CRAIG,School of Applied Sciences, Edinburgh Napier
University, Sighthill Court, Edinburgh, EH11 4BN, Scotland, United Kingdom; PAUL H.
FORESTELL,Keuka College, 141 Central Avenue, Keuka Park, New York 14478, U.S.A.;
ADAM S. FRANKEL,Hawaii Marine Mammal Consortium, PO Box 6107, Kamuela, Hawaii
96743, U.S.A. and Marine Acoustics, Inc. 2 Corporate Place, Suite 105, Middletown, Rhode
Island 02842, U.S.A.; GIUSEPPE NOTARBARTOLO DI SCIARA,Tethys Research Institute, Via
Benedetto Marcello 43, 20124, Milan, Italy; MATTHIAS HOFFMANN-KUHNT,Acoustic
Research Laboratory, Tropical Marine Science Institute, National University of Singapore, 18
Kent Ridge Road, Singapore 119227; EDUARDO MERCADO III, Department of Psychology,
University at Buffalo, The State University of New York, Buffalo, New York 14260, U.S.A.;
JOSEPH MOBLEY,School of Nursing and Dental Hygiene, University of Hawaii at Manoa,
2528 McCarthy Mall, Honolulu, Hawaii 96822, U.S.A. and Marine Mammal Research Con-
sultants, 520 Lunalilo Home Road #6106, Honolulu, Hawaii 96825, U.S.A.; MELISSA R.
SHYAN-NORWALT,Department of Psychology, McMicken College of Arts & Sciences,
University of Cincinnati, 7148 Edwards One, Cincinnati, Ohio 45221, U.S.A.; SCOTT S.
SPITZ,Marine Mammal Research Consultants, 520 Lunalilo Home Road #6106, Honolulu,
Hawaii 96825, U.S.A. and The Dolphin Institute, PO Box 6279, Hilo, Hawaii 96720,
U.S.A.; MOBY SOLANGI,Institute for Marine Mammal Studies, 10801 Dolphin Lane, Gulf-
port, Mississippi 39503, U.S.A.; ROGER K. R. THOMPSON,Department of Psychology &
Biological Foundations of Behavior Program, Franklin & Marshall College, PO Box 3003,
Lancaster, Pennsylvania 17604, U.S.A.; LORENZO VON FERSEN,Tiergarten Nuernberg, Am
Tiergarten 30, D-90480 Nuremberg, Germany; ROBERT UYEYAMA,The Dolphin Institute,
PO Box 6279, Hilo, Hawaii 96720, U.S.A.; RANDALL WELLS,Chicago Zoological Society’S
SARASOTA DOLPHIN RESEARCH PROGRAM,℅MOTE MARINE LABORATORY, 1600 KEN
THOMPSON PARKWAY,SARASOTA,FLORIDA 34236, U.S.A.; JAMES P. WOLZ,The Dolphin
Institute, PO Box 6279, Hilo, Hawaii 96720, U.S.A.
The following supporting information is available for this article online at http://
Appendix S1. Names of graduate students Lou Herman mentored, including degree
(s) achieved and year(s) of completion.
406 MARINE MAMMAL SCIENCE, VOL. 33, NO. 1, 2017
Appendix S1. Names of graduate students Lou Herman mentored, including degree(s) achieved and year(s) of
Masters Degree Students
M. E. McCauley (1968) Stacy Braslau-Schneck, (1994)
D. Bailey (1969) Richard J. Coleman (1994)
A. Llacuna (1969) Matthias Hoffmann-Kuhnt (1994)
Esme Hoban (1983) Susan H. Reeve (1994)
Adam S. Frankel (1987) Alison S. Craig (1995)
Gregory A. Hunter (1988) Eduardo Mercado III (1995)
Hilary L. Maybaum (1988) Kristen B. Taylor (1995)
Adam A. Pack (1988) Amy Cutting (1997)
Brian J. Tarbox (1988) Robert Uyeyama (1999)
Mark J. Xitco (1988) Brian Branstetter (2001)
David A. Helweg (1989) Mark H. Deakos (2002)
Kathy A. Sdao (1990) Rebecca Cowan (2003)
Melissa Shaw (1990) Siri Hakala (2004)
Christine M. Gabriele (1991) Amy Miller (2004)
Christopher G. Prince (1993) Kira Goetschius (2006)
Frank A. Beach III (1969) Adam S. Frankel (1994)
Ross L. Pepper (1969) Adam A. Pack (1994)
Carolyn Madsen (1976) Eduardo Mercado III (1998)
Roger K. R. Thompson (1976) Scott S. Spitz (1999)
Joseph R. Mobley (1984) Alison S. Craig (2001)
C. Scott Baker (1985) Matthias Hoffmann-Kuhnt (2003)
Melissa R. Shyan (1985) Robert K. U. Uyeyama (2007)
Gordon B. Bauer (1986) Mark H. Deakos (2010)
Paul H. Forestell (1988)