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MARINE MAMMAL SCIENCE, 33(1): 389–406 (January 2017)
©2016 Society for Marine Mammalogy
DOI: 10.1111/mms.12387
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
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 500800 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
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 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
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-
phinsDeep 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 Mammalsthat 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.
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
peoplestudents, interns, volunteersbecause 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.
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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 5:1416.
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
National Park Service, Alaska Regional Office, Anchorage, AK.
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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 1:304323.
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 31:105119.
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
Bauer, G. B., and L. M. Herman 1986. Effects of vessel traffic on the behavior of humpback
whales in Hawaii. Final Report to the National Marine Fisheries Services.
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 22:4962.
Branstetter, B. K., S. J. Mevissen, L. M. Herman, A. A. Pack and S. Roberts. 2003.
Horizontal angular discrimination by an echolocating bottlenose dolphin (Tursiops
truncatus). Bioacoustics 14:1534.
Branstetter, B. K., S. J. Mevissen, A. A. Pack, L. M. Herman, S. P. Roberts and L. K. Carsrud.
2007. Dolphin (Tursiops truncatus) echoic angular discrimination: Effects of object
separation and complexity. Journal of the Acoustical Society of America 121:626635.
Braslau-Schneck, S. 1994. Innovative behaviors and synchronization in bottlenosed dolphins.
M.S. thesis, University of Hawaii, Honolulu, HI. 134 pp.
Calambokidis, J., G. H. Steiger, J. M. Straley, et al. 1997. Population, abundance, and
structure of humpback whales in the North Pacific Basin. Draft final report to Southwest
Fisheries Science Center, La Jolla, CA by Cascadia Research Collective, 218 1/2 West
Fourth Avenue, Olympia, WA. 37 pp.
Calambokidis, J., G. H. Steiger, K. Rasmussen, et al. 2000. Migratory destinations of
humpback whales that feed off California, Oregon, and Washington. Marine Ecology
Progress Series 192:295304.
Calambokidis, J., G. H. Steiger, J. M. Straley, et al. 2001. Movements and population
structure of humpback whales in the North Pacific. Marine Mammal Science 17:769
Call, J., and M. Tomasello. 1994. Production and comprehension of referential pointing by
orangutans (Pongo pygmaeus). Journal of Comparative Psychology 108:307317.
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
estimation in the Hawaiian Islands. Marine Ecology Progress Series 175:1322.
Craig, A. S., and L. M. Herman. 1997. Sex differences in site fidelity and migration of
humpback whales (Megaptera novaeangliae) to the Hawaiian Islands. Canadian Journal of
Zoology 75:19231933.
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 193:209216.
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
Sanctuary, Office of National Marine Sanctuaries, National Oceanic and Atmospheric
Administration, U.S. Department of Commerce and Department of Land and Natural
Resources, State of Hawaii. 22 pp.
Craig, A. S., L. M. Herman and A. A. Pack. 2002. Male mate choice and male-male
competition coexist in the humpback whale (Megaptera novaeangliae). Canadian Journal of
Zoology 80:745155.
Craig, A. S., L. M. Herman, C. M. Gabriele and A. A. Pack. 2003. Migratory timing of
humpback whales (Megaptera novaeangliae) in the central North Pacific varies with age,
sex and reproductive status. Behaviour 140:9811001.
Craig, A. S., L. M. Herman, A. A. Pack and J. O. Waterman. 2014. Habitat segregation by
female humpback whales in Hawaiian waters: Avoidance of males? Behaviour 151:613
Frankel, A. S., C. W. Clark, L. M. Herman and C. M. Gabriele. 1995. Spatial distribution,
habitat utilization, and social interactions of humpback whales, Megaptera novaeangliae,
off Hawai’i, determined using acoustic and visual techniques. Canadian Journal of
Zoology 73:11341146.
Gabriele, C. M., J. M. Straley, L. M. Herman and R. J. Coleman. 1996. Fastest documented
migration of a North Pacific humpback whale. Marine Mammal Science 12:457465.
Gabriele, C. M., J. M. Straley, S. A. Mizroch, et al. 2001. Estimating the mortality rate of
humpback whale calves in the central North Pacific Ocean. Canadian Journal of Zoology
Green, S. R., E. Mercado III, A. A. Pack and L. M. Herman. 2007. Temporal structure in
humpback whale (Megaptera novaeangliae) songs. Aquatic Mammals 33:202213.
Green, S. R., E. Mercado III, A. A. Pack and L. M. Herman. 2011. Recurring patterns in
the songs of humpback whales (Megaptera novaeangliae). Behavioural Processes 86:284
Harlow, H. F., J. G. Miller and T. M. Newcomb. 1962. Identifying creative talent in
psychology. American Psychologist 17:679.
Helweg, D. A., and L. M. Herman. 1993. Observations of an S-shaped posture in humpback
whales. Aquatic Mammals 18:7478.
Helweg, D. A., and L. M. Herman. 1994. Diurnal patterns of behaviour and group
membership of humpback whales (Megaptera novaeangliae) wintering in Hawaiian waters.
Ethology 98:298311.
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 1:120.
Helweg, D. A., A. S. Frankel, J. R. Mobley Jr. and L. M. Herman. 1992. Humpback whale
song: Our current understanding. Pages 459483 in J. R. Thomas, R. A. Kastelein and
A. Ya. Supin, eds. Sensory processes of marine mammals. Plenum Press, New York, NY.
Herman, E. Y. K., L. M. Herman, A. A. Pack, G. Marshall, C. M. Shepard and M. Bakhtiari.
2007. When whales collide: Crittercam offers insights into the competitive behavior of
humpback whales on their Hawaiian wintering grounds. Marine Technology Society
Journal 41:3543.
Herman, L. M. 1965. Study of the single channel hypothesis and input regulation within a
continuous, simultaneous task situation. The Quarterly Journal of Experimental
Psychology Volume XVII:3746.
Herman, L. M. 1969. Effects of second signals on response time to first signals under certainty
and uncertainty. Journal of Experimental Psychology 80:106112.
Herman, L. M. 1975. Interference and auditory short-term memory in the bottlenose dolphin.
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Herman, L. M. 1979. Humpback whales in Hawaiian waters: A study in historical ecology.
Pacific Science 33:115.
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New York, NY.
Herman, L. M. 1986. Cognition and language competencies of bottlenosed dolphins. Pages
221251 in R. J. Schusterman, J. Thomas and F. G. Wood, eds. Dolphin cognition and
behavior: A comparative approach. Lawrence Erlbaum Associates, Hillsdale, NJ.
Herman, L. M. 1987. Receptive competencies of language trained animals. Pages 160 in J. S.
Rosenblatt, C. Beer, M. C. Busnel and P. J. B. Slater, eds. Advances in the study of
behavior. Volume 17. Academic Press, Petaluma, CA.
Herman, L. M. 1988. The language of animal language research. Psychological Record
Herman, L. M. 1989. In which Procrustean bed does the sea lion sleep tonight? Psychological
Record 39:1949.
Herman, L. M. 1990. Cognitive performance of dolphins in visually guided tasks. Pages 455
462 in J. A. Thomas and R. A. Kastelein, eds. Sensory abilities of cetaceans: Laboratory
and field evidence. Plenum Press, New York, NY.
Herman, L. M. 1991. What the dolphin knows, or might know, in its natural world. Pages
349364 in K. Pryor and K. S. Norris, eds. Dolphin societies: Discoveries and puzzles.
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108 in C. Nehaniv and K. Dautenhahn, eds. Imitation in animals and artifacts. MIT
Press, Cambridge, MA.
Herman, L. M. 2002c. Exploring the cognitive world of the bottlenosed dolphin. Pages 275
283 in M. Bekoff, C. Allen, and G. Burghardt, eds. The cognitive animal: Empirical and
theoretical perspectives on animal cognition. MIT Press, Cambridge, MA.
Herman, L. M. 2006. Intelligence and rational behaviour in the bottlenosed dolphin. Pages
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edition. Academic Press, New York, NY.
Herman, L. M. 2010. What laboratory research has told us about dolphin cognition.
International Journal of Comparative Psychology 23:310330.
Herman, L. M. 2012a. Historical perspectives. Aquatic Mammals 38:102125.
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Herman, L. M. In press. The multiple functions of male song within the humpback whale
(Megaptera novaeangliae) mating system: Review, evaluation, and synthesis. Biological
Herman, L. M., and R. C. Antinoja. 1977. Humpback whales in the Hawaiian breeding
waters: Population and pod characteristics. Scientific Reports of the Whales Research
Institute, Tokyo 29:5985.
Herman, L. M., and W. R. Arbeit. 1971a. Auditory frequency discrimination from 136 kHz
in Tursiops truncatus. Pages 7987 in Proceedings of the Eighth Annual Conference on
Biological Sonar and Diving Mammals. Stanford Research Institute, Menlo Park, CA.
Herman, L. M., and W. R. Arbeit. 1971b. Complex learning in the dolphin: Discrimination
of successively presented auditory patterns and formation of discrimination learning sets.
Final Report to the Naval Undersea Research and Development Center, Hawaii Division
Contact N66001-70-C-0939. 39 pp.
Herman, L. M., and W. R. Arbeit. 1972. Frequency difference limens in the bottlenose
dolphin: 170 KC/S. Journal of Auditory Research 2:109120.
Herman, L. M., and W. R. Arbeit. 1973. Stimulus control and auditory discrimination
learning sets in the bottlenosed dolphin. Journal of the Experimental Analysis of
Behavior 19:379394.
Herman, L. M., and P. H. Forestell. 1985. Reporting presence or absence of named objects by
a language-trained dolphin. Neuroscience and Biobehavioral Reviews 9:667691.
Herman, L. M., and J. A. Gordon. 1974. Auditory delayed matching in the bottlenosed
dolphin. Journal of the Experimental Analysis of Behavior 21:1926.
Herman, L. M., and B. H. Kantowitz. 1969. Effects of second signals occurring after response
selection on responses to first signals. Journal of Experimental Psychology 81:344350.
Herman, L. M., and B. H. Kantowitz. 1970. The psychological refractory period effect: Only
half the double-stimulation story? Psychological Bulletin 73:7488.
Herman, L. M., and M. E. McCauley. 1969. Delay in responding to the first stimulus in the
“psychological refractory period” experiment: Comparisons with delay produced by a
second stimulus not requiring a response. Journal of Experimental Psychology 81:344
Herman, L. M., and P. Morrel-Samuels. 1990. Knowledge acquisition and asymmetries
between language comprehension and production: Dolphins and apes as a general model
for animals. Pages 283312 in M. Bekoff and D. Jamieson, eds. Interpretation and
explanation in the study of behavior. Volume 1. Interpretation, intentionality, and
communication. Westview Press, Boulder, CO.
Herman, L. M., and A. A. Pack. 1994. Animal intelligence: Historical perspectives and
contemporary approaches. Pages 8696 in R. Sternberg, ed. Encyclopedia of human
intelligence. Macmillan, New York, NY.
Herman, L. M., and A. A. Pack. 2001. Laboratory evidence for cultural transmission
mechanisms. Commentary on Rendell and Whitehead. Behavioral and Brain Sciences
Herman, L. M., and W. N. Tavolga. 1980. The communication systems of cetaceans. Pages
149209 in L. M. Herman, ed. Cetacean behavior: Mechanisms and functions. Wiley
Interscience, New York, NY.
Herman, L. M., and R. K. R. Thompson. 1982. Symbolic, identity, and probe delayed
matching of sounds by the bottlenosed dolphin. Animal Learning and Behavior 10:22
Herman, L. M., and R. K. Uyeyama. 1999. The dolphin’s grammatical competency:
Comments on Kako (1998). Animal Learning and Behavior 27:1823.
Herman, L. M., F. A. Beach III, R. L. Pepper and R. B. Stalling. 1969. Learning-set formation
in the bottlenose dolphin. Psychonomic Science 14:9899.
Herman, L. M., M. F. Peacock, M. P. Yunker and C. Madsen. 1975. Bottlenosed dolphin:
Double-slit pupil yields equivalent aerial and underwater diurnal acuity. Science
Herman, L. M., P. H. Forestell and R. C. Antinoja. 1980. Study of the 1976/77 migration of
humpback whales into Hawaiian waters: Composite description. Final Report to the
U.S. Marine Mammal Commission, Report No. MMC-77/19. United States National
Technical Information Services, Arlington, VA.
Herman, L. M., D. G. Richards and J. P. Wolz. 1984. Comprehension of sentences by
bottlenosed dolphins. Cognition 16:129219.
Herman, L. M., J. R. Hovancik, J. D. Gory and G. L. Bradshaw. 1989. Generalization of
visual matching by a bottlenosed dolphin (Tursiops truncatus): Evidence for invariance of
cognitive performance with visual or auditory materials. Journal of Experimental
Psychology: Animal Behavior Processes 15:124136.
Herman, L. M., P. Morrel-Samuels and A. A. Pack. 1990. Bottlenosed dolphin and human
recognition of veridical and degraded video displays of an artificial gestural language.
Journal of Experimental Psychology: General 119:215230.
Herman, L. M., S. Kuczaj III and M. D. Holder. 1993a. Responses to anomalous gestural
sequences by a language-trained dolphin: Evidence for processing of semantic relations
and syntactic information. Journal of Experimental Psychology: General 122:184194.
Herman, L. M., A. A. Pack and P. Morrel-Samuels. 1993b. Representational and conceptual
skills of dolphins. Pages 273298 in H. R. Roitblat, L. M. Herman and P. Nachtigall,
eds. Language and communication: Comparative perspectives. Lawrence Erlbaum
Associates, Hillside, NJ.
Herman, L. M., A. A. Pack and A. M. Wood. 1994. Bottlenose dolphins can generalize rules
and develop abstract concepts. Marine Mammal Science 10:7080.
Herman, L. M., L. von Fersen and M. Solangi. 1996. The bufeo (Inia geoffrensis) in the Rio
Largarto Cocha of the Ecuadorian Amazon. Marine Mammal Science 12:118125.
Herman, L. M., A. A. Pack and M. Hoffmann-Kuhnt. 1998. Seeing through sound: Dolphins
perceive the spatial structure of objects through echolocation. Journal of Comparative
Psychology 112:292305.
Herman, L. M., S. L. Abichandani, A. N. Elhajj, E. Y. K. Herman, J. L. Sanchez and A. A.
Pack. 1999. Dolphins (Tursiops truncatus) comprehend the referential character of the
human pointing gesture. Journal of Comparative Psychology 113:118.
Herman, L. M., D. S. Matus, E. Y. K. Herman, M. Ivancic and A. A. Pack. 2001. The
bottlenosed dolphin’s (Tursiops truncatus) understanding of gestures as symbolic
representations of its body parts. Animal Learning and Behavior 29:250264.
Herman, L. M., R. K. Uyeyama and A. A. Pack. 2008. Bottlenose dolphins understand
relationships between concepts. Behavioral and Brain Sciences 31:139140.
Herman, L. M., A. A. Pack, K. Rose, A. Craig, E. Y. K. Herman, S. Hakala and A. Milette.
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 27:736768.
Herman, L. M., A. A. Pack, S. S. Spitz, E. Y. K. Herman, K. Rose, S. Hakala and M. H.
Deakos. 2013. Humpback whale song: Who sings? Behavioural Ecology and
Sociobiology 67:16531663.
Hoffmann-Kuhnt, M. 2003. Visual and auditory vigilance in the bottlenosed dolphin.
Doctoral dissertation. Freie Universitat Berlin, Berlin, Germany. 175 pp.
Madsen, C. J., and L. M. Herman. 1980. Social and ecological correlates of vision and visual
appearance. Pages 101147 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 26:210218.
Mercado, E., III, R. K. Uyeyama, A. A. Pack, and L. M. Herman. 1999. Memory for action
events in the bottlenosed dolphin. Animal Cognition 2:1725.
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 50:7994.
Mercado, E., III, L. M. Herman and A. A. Pack. 2003. Stereotypical sound patterns in
humpback whale songs: Usage and function. Aquatic Mammals 29:3752.
Mercado, E., III, L. M. Herman and A. A. Pack. 2005. Song copying by humpback whales:
Themes and variations. Animal Cognition 8:93102.
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
Zoology 63:762772.
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 8:16.
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 25:6372.
Morrel-Samuels, P., and L. M. Herman. 1993. Cognitive factors affecting comprehension of
gesture language signs: A brief comparison of dolphins and humans. Pages 211222 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 98:722733.
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 118:160171.
Pack, A. A., and L. M. Herman. 2006. Dolphin social cognition and joint attention: Our
current understanding. Aquatic Mammals 32:443460.
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 14:861873.
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 28:131146.
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 58:126.
Pack, A. A., L. M. Herman and M. Hoffmann-Kuhnt. 2004. Dolphin echolocation shape
perception: From sound to object. Pages 288298 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 77:653662.
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
Behaviour 84:983993.
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):307317.
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 12:423461.
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 98:59375942.
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 98:1028.
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 77:504508.
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 101:105119.
Spitz, S. S., L. M. Herman and A. A. Pack. 2000. Measuring sizes of humpback whales
(Megaptera novaeangliae) through underwater videogrammetry. Marine Mammal Science
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 80:19381947.
Thompson, R. K. R., and L. M. Herman. 1975. Underwater frequency discrimination in the
bottlenosed dolphin (1140 kHz). Journal of the Acoustical Society of America 57:943
Thompson, R. K. R., and L. M. Herman. 1977. Memory for lists of sounds by the bottlenosed
dolphin: Convergence of memory processes with humans? Science 195:501503.
Thompson, R. K. R., and L. M. Herman. 1981. Auditory delayed discriminations by the
dolphin: Nonequivalence with delayed matching performance. Animal Learning and
Behavior 9:915.
Yunker, M. P., and L. M. Herman. 1974. Discrimination of auditory temporal differences by
the bottlenosed dolphin (1140 kHz). Journal of the Acoustical Society of America
Xitco, M. J., Jr. 1988. Mimicry of modeled behaviors by bottlenose dolphins. M.S. thesis,
University of Hawaii, Honolulu, HI. 124 pp.
Xitco, M. J., Jr., J. D. Gory and S. A. Kuczaj III. 2001. Spontaneous pointing by bottlenose
dolphins (Tursiops truncatus). Animal Cognition 4:115123.
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
Institute, PO Box 6279, Hilo, Hawaii 96720, U.S.A.
Supporting Information
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.
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)
Doctoral Students
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)
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Following a 1st signal to respond (S1), a 2nd signal (S2) to which no response was required was inserted during response execution, i.e., after an overt response to S1 had begun but before the entire response had been completed. No significant decrements in response latency to S1 or in errors in responding to S1 were found by comparison with an independent control group never experiencing S2. This was in contrast to prior findings in which significant response decrements had been found when S2 occurred during response selection, i.e., prior to initiation of any overt movement. Ss were 18 male undergraduates. The set of findings considered compatible with an interpretation of independence of response-selection and response-execution stages and with response-conflict theory. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
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Obtained frequency difference limens (DL) for an Atlantic bottlenose dolphin, Tursiops truncatus (Montagu) for 1, 6, 12, 19, 27, 36, 50, and 70 kHz. Ss were required to select the pure tone signal from a sequential pair consisting of a pure tone signal and a frequency-modulated (FM) signal of the same center frequency. Using a threshold-tracking procedure, the percent-frequency (EP) the FM signal was reduced following correct responses and increased following errors, providing estimates of the minimum deviation required for correct discrimination. The effects of modulation rates of 1 and 2 Hz were also studied. Thresholds and threshold variability were lowest for the 2-Hz rate. Relative DLs (DF/F) were roughly constant (.002-.003) for 6-50 kHz but were elevated for 1 kHz and 70 kHz. Results indicate exceptionally fine frequency discrimination by T. truncatus between 6 and 50 kHz. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
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2 Atlantic bottlenose dolphins were tested with 2 simultaneous sound stimuli as discriminanda in successive reversal training (SRT) and discrimination learning set (DLS) paradigms. The first S was tested in the order SRT-DLS, and the second in the order DLS-SRT. Both demonstrated progressive improvement over successive reversals under SRT, with the DLS-SRT S showing fewer errors per reversal and achieving almost twice as many reversals within the fixed number of trials given. Although neither S clearly formed a discrimination learning set within the 180 different auditory problems administered, there was evidence that the prior SRT experience of one S facilitated some aspects of its subsequent DLS performance. The failure in DLS was discussed in terms of the marginal levels of within-problem learning evidenced.
Technical Report
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The behavior of humpback whales summering in southeastern Alaska was observed in the presence and absence of vessel traffic. During the first study year (1981), small and medium-sized vessels were directed to operate within 400 m of whales according to an experimental plan. The second study year (1982) concentrated on observations of generally greater than 400 m. Whales showed predictable behavior responses to vessels operating at distances of less than 4,000 m. Changes in whale behavior were correlated with the speed, size, distance, and numbers of vessels within the proximity.
Responses of Atlantic bottle-nosed dolphins (Tursiops truncatus) and of humans were collected and analyzed in order to determine the features required for recognition and discrimination of signs (hand signals) in an artificial gestural communication system. Subjects responded to systematically modified signs in which sign components were contrasted for competitive feature salience. One dolphin, with 6 yrs of training in the language, was shown these modified signs intermixed with normal signs in a linguistic, sentence-comprehension context. A second dolphin, familiar with action signs only and with no sentence-comprehension training, served as a nonlingual control. Human subjects were tested in two parallel tasks. The dolphin with sign-language experience attended to (in order of importance) location, completed temporal pattern, gross motor motion, and direction of motion, as salient features. Fine motor motion, hand shape, and hand orientation were less salient. The non-sign-language dolphin attended to all sign features equally and was unaffected by temporal pattern changes. Humans tested in a linguistic context attended to (in order) gross motor motion, location, and an interaction of fine motor motion, hand shape, and hand orientation. Direction of motion and temporal pattern were not salient. Nonlinguistic-context humans attended to all sign features equally and were unaffected by temporal pattern changes. Results indicate that language experience and/or testing context affect feature salience for sign recognition.
Humpback whales (Megaptera novaeangliae) are seasonal breeders, annually migrating from high-latitude summer feeding grounds to low-latitude winter breeding grounds. The social matrix on the winter grounds is a loose network of interacting individuals and groups and notably includes lone males that produce long bouts of complex song that collectively yield an asynchronous chorus. Occasionally, a male will sing while accompanying other whales. Despite a wealth of knowledge about the social matrix, the full characterization of the mating system remains unresolved, without any firm consensus, as does the function of song within that system. Here, I consider and critically analyse three proposed functions of song that have received the most attention in the literature: female attraction to individual singers, determining or facilitating male–male interactions, and attracting females to a male aggregation within the context of a lekking system. Female attraction suggests that humpback song is an advertisement and invitation to females, but field observations and song playback studies reveal that female visits to individual singers are virtually absent. Other observations suggest instead that females might convey their presence to singers (or to other males) through the percussive sounds of flipper or tail slapping or possibly through vocalizations. There is some evidence for male–male interactions, both dominance and affiliative: visits to singers are almost always other lone males not singing at that time. The joiner may be seeking a coalition with the singer to engage cooperatively in attempts to obtain females, or may be seeking to disrupt the song or to affirm his dominance. Some observations support one or the other intent. However, other observations, in part based on the brevity of most pairings, suggest that the joiner is prospecting, seeking to determine whether the singer is accompanying a female, and if not soon departs. In the lekking hypothesis, the aggregation of vocalizing males on a winter ground and the visits there by non-maternal females apparently for mating meet the fundamental definition of a lekking system and its role though communal display in attracting females to the aggregation, although not to an individual singer. Communal singing is viewed as a form of by-product mutualism in which individuals benefit one another as incidental consequences of their own selfish actions. Possibly, communal singing may also act to stimulate female receptivity. Thus, there are both limitations and merit in all three proposals. Full consideration of song as serving multiple functions is therefore necessary to understand its role in the mating system and the forces acting on the evolution of song. I suggest that song may be the prime vector recruiting colonists to new winter grounds pioneered by vagrant males as population pressures increase or as former winter grounds become unavailable or undesirable, with such instances documented relatively recently. Speculatively, song may have evolved historically as an aggregating call during the dynamic ocean conditions and resulting habitat uncertainties in the late Miocene–early Pliocene epochs when Megaptera began to proliferate. Early song may have been comprised of simpler precursor sounds that through natural selection and ritualization evolved into complex song.