Case
Study of a Novel Ritualized Motor Pattern, the "Neck-bite," Accompanying Human
Ejaculation*
Clara B. Jones
5/27/2011 (posted to blog, 5/22/2020; slightly edited & revised, 5/23/2020)
Abstract
Communication forms
the “glue” in all societies, and species-typical traits (e.g., morphology, behavior) have
evolved by natural or sexual selection to
enhance communication between a signaler and receiver(s).
Over evolutionary time, the process of "ritualization" may simplify and
exaggerate conserved morphological characters, modifying their
original function(s), yielding stereotyped phenotypic features with
signaling function(s) differing from those of the conserved traits.
Employing analyses of photographs and videos as well as a modified
version of phenetic methodology, the present case study dissects an
apparently rare motor pattern of the human behavioral repertoire
occurring at ejaculation during dorsal-ventral intercourse between
consenting heterosexual partners. Dissection of the case material
led to the diagnostic inference that this response, the “neck-bite”,
represents a newly-described ritualized morphological feature for
Homo
sapiens with morphological and functional similarities to the "neck-bite" behaviors reported for other mammals.
Key
Words: Communication; Ethology; Ritualization; Neck-Bite; Copulation;
Carnivores; Primates; Qualitative Research, Ejaculation
Introduction
Communication
is the “glue” of any society (Wilson, 1971). When “true
communication” occurs, both Sender (Actor) and Receiver (Recipient) benefit from the
exchange of information (Bradbury & Vehrencamp, 1998). For
humans and many other taxa (e.g., birds and mammals), communication
may be both auditory (e.g., Marler, 1976) and non-auditory (e.g.,
gestures: Morris, 1995). As discussed by Maynard Smith & Parker
(2003; also see Enquist et al., 2010), elements of communication
function as signals (traits modified by selection for communication
function), and, in some species (e.g., many primates, including
humans), these elements may be combined and recombined, producing
complex, multimodal patterns (e.g., Jones & Van Cantfort, 2007),
functionally referential communication (signals about external
events: e.g., Seyfarth et al., 1980), and context-dependent meaning
(e.g., Quattara et al., 2009).
Ritualized and
Stereotyped Signals and Displays
Tinbergen
(1952; also see Morris, 1957; Lorenz, 1981; Eibl-Eibesfeldt, 2007)
describes the process of “ritualization” whereby features of the
conserved (“primitive”) species-typical repertoire become
“emancipated” or disassociated over evolutionary time from their
previous function(s) leading to involuntary, exaggerated, simplified
motor patterns known as “stereotyped” signals often termed
postures, displays, or “fixed action patterns.” It is through
the process of "emancipation" that stereotyped signals are said to be
“derived” from earlier, conserved morphological elements.
Stereotyped motor patterns are characterized by “typical
intensity,” relatively invariant and relatively immobile
morphological features characteristically similar across individuals
of the same sex and species across space and time. Finally,
ritualized behaviors may present as “intention movements” whereby
the stereotyped action pattern is identifiable but appears truncated
and incomplete in form.
Employing
or not employing an exaggerated, stereotyped signal is not, by
definition, an intentional or conscious and aware “decision”
since the evolutionary process leading to ritualization of phenotypic
characters yields “hard-wired,” involuntary responses that are "canalized," resistant to environmental modification or "plasticity" across time and space (Morris, 1957;
Lorenz, 1981; Eibl-Eibesfeldt, 2007). As a caveat,
although there are certain mechanistic similarities, stereotyped
motor patterns should not be confused with "reflexes" which are not
ritualized, derived signals but action patterns induced at the level
of the spinal cord controlled by the autonomic nervous system. In
ethological terms, "displays" have evolved in response to particular
stimuli termed “releasers,” while reflexes have no particular
communicative function and are not stimulus-specific (e.g., rapid
jerking of a body part from an aversive stimulus may be exhibited in
response to a wide range of different stimuli such as heat, electric
shock, or physical disgust). Ritualization leading to stereotyped
phenotypic elements yields discrete (digital), discontinuous signals
rather than graded (analog) signals transmitted as a continuous
sequence of information-transmission (e.g., Eibl-Eibesfeldt, 2007;
Maynard Smith & Parker, 2003; Enquist et al., 2010).
Most
motor patterns in the behavioral repertoire of humans are graded
(Eibl-Eibesfeldt, 2007). Although discrete vocal signals may not be
rare in Homo
sapiens
(Marler, 1976), this article addresses only stereotyped motor
patterns or displays, not discrete auditory or other sensory signals.
Stereotyped and ritualized behaviors in humans have been documented
and discussed by Eibl-Eibesfeldt (2007), in particular, the
“eye-flash” motor pattern (stereotyped lifting of the eyebrows).
The same author conducted cross-cultural research including cryptic
filming of the eye-flash, demonstrating that, in cultures throughout
the world, the "eye-flash" occurs between males and females while
flirting and in apparent courtship and other bonding contexts. Based
on many years of research, Eibl-Eibesfeldt (2007) concluded that
stereotyped and ritualized action patterns are rare in the human
behavioral repertoire, and it seems reasonable to postulate that this
observation obtains because, compared to other organisms, phenotypes
of Homo
sapiens are
shaped to a large degree by learning. Learning mechanisms (e.g., "association," "social learning") are
thought to have evolved in response to rapidly changing
(“stochastic” or predictable relative to "generation time") environments permitting flexible, resilient, and
“plastic” behavior, effects with the potential for rapid
adjustment of the phenotype to environmental heterogeneity (e.g.,
Jones, 2005, 2012; Weber & Depew, 2003), possibly decreasing the
selective advantage of some hard-wired responses such as those
dedicated by the process of ritualization to stereotypy over time and
space (e.g., as appears to have occurred in human evolution).
The
present case study discusses a human motor pattern that is, based
upon detailed analysis, inferred to be stereotyped. This response,
the “neck bite,” is observed across numerous taxa, both
invertebrate (e.g., Buskirk et al., 1984) and vertebrate
(Saint-Girons, 2006). The behavior functions intraspecifically in a
variety of modes as evolutionarily modified signals conditionally
displayed in a range of contexts and states. To my knowledge, the
stereotyped "neck-bite" as a component of the human sexual repertoire,
has not previously been described in the technical literature.
Methods
Qualitative Research
Design and Procedures
Information
about the sexual encounter dissected in the present case study was
availed to the author (hereafter, “the investigator”) by one of
the participant-observers (hereafter, “the subject,” "receiver," or, "recipient"). On the
basis of her training in Ethology emphasizing the study of sexual and
aggressive behaviors, the investigator suspected that certain details
of the described events deviated from patterns of behavior generally
attributed to human dorsal-ventral intercourse, appearing, on the
basis of the subject’s verbal report, to include action patterns
often described as elements of non-human animal copulation and/or ejaculation. Motor
patterns comprising the highly variable sequences of behavior
observed for human sex (e.g., Roughgarden, 2009), generally appear
intentional, fluid, and continuous unlike certain of the fixed and
discrete responses described by the subject (Morris, 1957; Lorenz,
1981; Eibl-Eibesfeldt, 2007). Based on the subject’s report of one
heterosexual experience with a long-term partner, the investigator
decided, with permission of the subject but without knowledge of the
subject’s partner (the "signaler" or "actor"), to further explore the described events on the
condition that identities of both participants remain anonymous.
Rather
than limit her behavioral analysis to narrative and interpretive
methods, the investigator chose to employ "phenetic analysis," a
standard procedure employed in systematic biology commonly applying
diagnostic criteria to museum specimens to determine phylogenetic
proximity among species (Schuh & Brower, 2009). This qualitative, descriptive method relies upon subjective assessments by an expert inspecting
similarities of morphological traits, in the present case, behavioral
characters, to construct "phylogenetic trees." The construction of
trees was not a goal of the present study; though comparative observations to other mammalian taxa, especially, felids and canids, were conducted and are discussed. Rather, phenetic methods
were used to increase reliability of diagnosis by matching motor
patterns described by the subject to a template of morphological
features generally accepted by ethologists as diagnostic criteria for
identification of ritualized motor patterns. Deviations from
phenetic practice entailed, in particular, diagnosis based upon a
sample size of one (N= 1), constituting a Case Study, prohibiting application of descriptive
statistics base upon, say, analysis of film or photographs yielding sequential responses, possibly, coordinated ones, by actor and recipient. Thus, case material employed for diagnosis
deviated significantly from phenetic diagnoses because it was based
upon the subject’s verbal, participant report rather than direct
inspection of case material. Ethological methods often entail
direct observation in laboratory or in nature using frame-by-frame
analysis of images (Eibl-Eibesfeldt, 2007; Lehner, 1998; Jones, 1986)
permitting calculation of “transition probabilities“ or
assessment of results with descriptive and/or inferential statistical tests (Lehner, 1998; Jones,
1983).
Phenetic
analysis, like all qualitative research, does not permit
generalization beyond given case material, limited, instead, to
subjective inferences by a trained investigator’s dissection of
comparative morphological traits (in the biological literature, a "character" or "trait" is generally assumed to be genetically correlated, thus, a potential target of selection; this may apply to both "plastic" or "canalized" traits). Phenetic analysis, also, does not
permit distinctions between traits derived from a common ancestor
(plesiomorphies) and convergent traits evolved independently in one
or more lineages (apomorphies). However, phenetic methods, like all
scientific procedures, requires a description precise enough to
permit other trained researchers to independently describe and
replicate the investigator’s dissection and diagnosis. The
specific research question posed in the present study was: Subsequent
to dissection and prior to diagnosis, do the motor patterns described
by the subject justify their classification as ritualized,
stereotyped morphological traits similar in form but not, necessarily, in function
to conserved (primitive) action patterns in the target species (in the present case, humans) or in
other taxa?
To
supplement the modified phenetic analysis, the investigator studied
>250 photographs and videos of mammals [available on the internet, including, YouTube], including humans, in order
to familiarize herself with the variable forms, contexts, and
apparent motivations and functions of motor patterns targeting the
neck. The diagnostic criteria used to infer stereotypy of the motor
patterns displayed in the survey of images and the case material
described herein are the same ones that would be employed by any
other ethologist evaluating possible ritualization of morphological
traits (Lehner, 1998) . A major difference was that this study’s
case material consisted of subject report based upon memory of events
rather than direct or indirect inspection by the investigator.
The
dissection and diagnosis reported in this paper probably yielded
inferences with a higher degree of confidence and a lower level of
error than procedures frequently used by fieldworkers collecting
narrative data from indigenous or other local individuals in habitat
countries (e.g., to obtain information about rare or difficult to
locate species or recalled events "in the world") since this study’s subject was a participant
observer providing first-hand case material and because a unique sexual encounter is likely to be consolidated relative reliably in memory due to its intense emotional, associational, and cognitive effects. This investigator
decided not to employ interview methods with the subject because this
procedure has the potential to introduce additional error due to
prompting, increasing the likelihood that a subject will embellish,
exaggerate, or otherwise bias information (e.g., Berkowitz et al.,
2008). Certain personal details are omitted from this case study to
shield the identities of the subject and the subject’s partner.
Results
Mammalian Motor
Patterns Targeting the Neck
The "neck-bite" is common to mammalian hunt and kill, and predators and
their prey are presumed to have co-evolved (Franks & Nobel, 2004),
with predator hunting strategies a function of prey defense responses
(Sih, 1985; Alcock, 1998). Several sources note that selection will
be particularly sensitive to the relative sizes of predator and prey,
favoring flexible responses since hunting and defense are species-
and condition-dependent and, frequently, opportunistic (Mc Coy et
al., 2011). Documenting the history, likely origin(s), forms,
distributions, and functions of the mammalian neck-bite motor pattern
formed a critical component and first stage of this report’s
results, permitting historical and comparative dissection and
diagnosis. The following background assembles information on motor
patterns targeting the neck in extant mammals, forming a body of
evidence from the scientific literature (see, especially, Estes,
1992; Eisenberg, 1981). In the section on primates, Paleotropical
non-human species are emphasized because of their systematic
proximity to Homo
sapiens and
because, in general, these families are more completely described
than Neotropical monkeys.
Information
in the Results section was extracted from: Estes, 1992; Ewer., 1968; Morris, 1957; Moynihan,
1970; Napier, 1970; Poole, 1972; Klieman & Eisenberg, 1973;
Eisenberg, 1981; Lorenz, 1981; Buskirk et al., 1984; Boesch &
Boesch, 1989; Martin, 1989; Estes, 1992; McKenna & Bell, 1997;
Smuts, 1998; Millon et al., 1999; American Psychiatric Association,
2000; Skinner & Chimimba, 2005; van der Dennen, 2005; Bauchot,
2006; Surbeck & Hohmann, 2008; Roughgarden, 2009; Swedell &
Schrier, 2009; and, Enquist et al., 2010.
Carnivores
My
literature search of mammalian motor patterns targeting the neck
primarily discussed these responses as they function during the kill
in carnivores. These morphological traits have been studied,
particularly, in canids (e.g., wolf, fox, coyote) and felids (e.g.,
lion, tiger, leopard). Fossil Carnivores are dated to the Eocene,
~55 mbp, and their common ancestor can be traced to the early
Paleocene, ~60 mbp. According to my sources, the felid-canid split
was completed in the Oligocene-Miocene, ~20 mbp, with felids
splitting from the common ancestor earlier than canids. These
carnivore families, on whole, evolved different patterns of hunting
and killing though both groups primarily target the neck. Canids
generally hunt in groups, usually killing with a “death-shake”
rather than the suffocating “neck-bite” employed by felids.
Extant felids are solitary hunters generally larger than canids,
character states likely to have favored a selfish and efficient,
though individually riskier, kill tactic. Compared to group hunting,
solitary hunting is probably more sensitive to relative size
differences between predator and prey. It seems reasonable to
suggest, additionally, solitary hunting and the neck-bite are almost
certainly the derived state compared to group-hunting and the
death-shake since the neck-bite is more energetically efficient than
the death-shake. These propositions are consistent with reports that
the felid line is older than that of canids since felids would have
had a temporally wider opportunity for evolutionary changes to occur.
While
characteristic hunting and killing strategies have been documented
for carnivores, there are exceptions to observed patterns requiring
analysis. For example, among felids, jaguars and cougars, both
Neotropical taxa but not particularly closely related to each other,
preferentially kill prey by using their teeth and jaws to crush the
temporal lobes of their prey’s skulls, a method dispensing of prey
virtually immediately compared to the methods of killing used by
canids and felids. Again, based upon relatively increased energetic
efficiency, killing prey with the method employed by jaguars and
cougars suggests that it is a derived response. It has been
suggested that this motor pattern is an adaptation favored during the
Pleistocene’s great mammal extinctions when many available prey
items were armored reptiles, a hypothesis seemingly supported by the
observation that, while not a particularly large cat, cougars feed
upon large, dangerous prey such as moose, elk, and bighorn sheep.
With
the exception of humans and bonobos, the dorsal-ventral sexual
posture is ubiquitous among mammals. Male canids and felids
generally employ a stereotyped "neck-bite" during copulation
interpreted as a character derived from a conserved action pattern.
Though stereotyped, the neck-bite appears as an intention movement
generally causing no harm to the female in
copula
because the force of biting to damage or to kill is inhibited. During
sex, some mammalian males appear to use the "neck- bite" to maintain
balance during intromission, and it is also likely that the motor
pattern is employed multi-modally to communicate dominance and
control over the female. This topic is discussed in greater detail
below.
Other
ritualized "neck-bite" responses, apparently derived from conserved
motor patterns, occur in association with maternal “mouth transport”
and play. Females in many mammalian families move their young from
one location to another using a method of transport employing mouth
and teeth with some force but inhibited sufficiently to prevent
harming young (Fig. 1). The stereotyped "play-bite" most likely
derived, also, from a conserved motor pattern and most often
exhibited by immatures, is a common behavior, particularly among
social mammals. A number of functions have been attributed to play,
but most reports emphasize its role in social bonding, as an
interaction permitting assessment of relative strength, communication
of non-aggressive motivation, solicitation of non-aggressive
response, and/or practice of behaviors characteristic of adult
behavioral repertoires.
Primates
With
the exception of three apes (bonobos, chimpanzees, and humans),
coordinated hunting and killing of conspecifics and contraspecifics
is rare or absent among primates. The three hominins are primarily
terrestrial and widely acknowledged for their expert use of tools [Note added 5/23/2020: could this be related to frugivory?].
In the non-human apes and a few monkey species (e.g., capuchins),
tools may be employed to break shells of nuts or crustaceans, to
extract termites or ants from their colonies, and/or to extract
marrow from bones and brains from skulls. Where killing has been
observed in monkeys (e.g., Rhesus macaques, capuchins), an
opportunistic capture and kill strategy was employed, and food items
included both invertebrate and vertebrate taxa, including ants,
spiders, birds, small mammals, as well as conspecific and/or
contraspecific species of monkey. Bonobos, chimpanzees, and
carnivorous monkeys generally kill prey by dismemberment and/or by
disembowelment, using jaws and teeth, facilitated by hands, in both
contexts. However, it has been communicated to the investigator
anecdotally that chimpanzees sometimes kill in the manner of cougars
and jaguars, by crushing the skulls of prey with the force of their
mouths and teeth.
When
primates kill, action patterns appear, ceteris
paribus,
intentional and graded rather than discrete and stereotyped as
described for canids and felids. Humans are noteworthy for employing
all manner of methods to kill conspecifics and contraspecifics, not
only for food but, also, legitimately (e.g., policing, self-defense),
by accident (e.g., vehicles, sports events), as well as for other
reasons frequently classified as illegal acts (e.g., murder) or
paraphilias (e.g., consensual or non-consensual practices leading to
death exhibited for sexual gratification of one or more
participants). These practices may be displayed independent of
whether they are (e.g., attacks by formal street gangs, prison
revolts, death resulting from impulsive aggression) or are not (e.g.,
offensive or defensive attacks by police, state warfare) considered
illegal or pathological. Cultural rituals incorporating and/or
leading to death (e.g., cannibalism, human sacrifice) are not
uncommon practices in human societies and were not classified as
paraphilias in the references inspected by this investigator.
In
bi-sexual contexts among non-human primates, the stereotyped
neck-bite or, similarly, a stereotyped shoulder-bite (Fig. 2), may or
may not be displayed, even in species characterized by a detectable
male dominance hierarchy. With the exception of bonobos and humans [Note added 5/23/2020: also, unless I am mistaken, gorillas in zoos] that frequently intromit in ventral-ventral orientation, primates
copulate in the dorsal-ventral position with males typically
restraining a female’s ankles with his feet in order to maintain
balance. Both of these sexual postures, particularly dorsal-ventral,
exhibit steoreotyped elements (e.g., "typical intensity" of bodies and
arms during orgasm, often accompanied by vocalizations [signals?]). If a neck-bite occurs in combination with
sexual egress, it is generally non-damaging. Despite the variety of
patterns reported for class Mammalia, the non-human primate
literature is dominated by studies of the "neck-bite" employed by males
of a single species, Hamadryas baboons, to aggressively coordinate
and control females in their harems. Although the teeth of these
males may pierce the skin of females’ necks, the motor pattern is
considered to be ritualized because the act of biting to kill is
incomplete [Note added 5/23/2020: though, Hamadryas are among the few mammal species, the others being, humans, agouti, Northern elephant seals, walrus, chimpanzees, lions, domestic cats (Estes, 1992), in which adult males have been documented to kill mates or prospective mates].
Where
it has been reported to occur, the neck-bite is “gentle” in
prosimians (e.g., greater bushbabies), most of whom are solitary and
nocturnal. In general, males of "conserved" or "primitive" primate species are
reported to be “patient” with females in estrus (e.g., lesser
bushbabies), not attempting to mount and intromit until the female
communicates receptivity behaviorally (e.g., by approach and/or by
presenting to the male with a stereotyped rear-present posture)
and/or with other signals or displays (e.g., vocalizations). In
humans, the stereotyped neck-bite has not, apparently, been reported,
though “love bites” are common, particularly as components of
flirting, play, foreplay, or other intimate contexts. "Love-bites," even those causing erythema (“hickies”) (Fig. 3) and, further,
the infliction of mild pain including digging fingernails into a
sexual partner’s back during foreplay or intercourse, are not
considered pathological unless they are employed for sexual
gratification in sado-masochistic or other contexts including, for
example, humiliation, beating, restraint, cutting, hypoxia, and/or
death, again, whether or not one or more participants act
consensually.
Play
focused on the neck is virtually ubiquitous among primates and other mammals, and,
similar to carnivores, may occur as stereotyped play-bites,
particularly in non-human species. Primate play bouts are often
reciprocal, with interactants soliciting play by vocalization,
exposure of neck or other areas of the body, or touch (e.g.,
tickling). My inspection of photographs and videos of play in
simians suggested that play in anthropoids, especially among
cercopithecines, frequently includes apparently stereotyped
play-bites (Fig.4). The stereotyped play-bite was not, in general,
apparent in the hominid images investigated. Stereotyped and
non-stereotyped components of anthropoid and hominid play appeared
primarily as solicitations or, possibly, appeasement, sometimes involving a "bared-neck" display,
or vulnerable display of ventral region. These responses may
function, also, to communicate subordinance or non-aggressive
motivation or to solicit a non-aggressive response from the receiver.
Bouts of play in hominids did not appear biased in favor of the neck
region. My visual inspections of grooming, the most common social
behavior among non-human primates, indicated that anthropoids groom
most of the surface of the recipient’s body, though hand postures
appeared to include stereotyped elements. If play bouts were skewed
in favor of the neck region, it was not apparent during investigation
of images of anthropoids. Non-human hominids (orangutans, gorillas,
bonobos, and chimpanzees) frequently targeted the neck during
grooming. However, though solicitations leading to grooming appeared
to be stereotyped components of these interactions, grooming itself
did not appear to be ritualized.
Human
play directed at the neck and other bodily regions appeared primarily
intentional, although certain of these action patterns exhibited
during play, flirting, or in association with lovemaking may
represent “intention movements” (Fig. 5), exhibiting elements of
a stereotyped response (e.g., kissing, in the image depicted, or
stroking, gentle scratching). All of these behaviors share
similarities with both play and grooming in other primates. It seems
possible that the phenotypes of many primate taxa, including humans,
may incorporate both ritualized and non-ritualized action patterns
for flexible, condition-dependent use. Intentional acts of play in
the neck region are numerous in humans, including snuggling, mild
nuzzling, and kissing, and all of these motor patterns appear to have
some stereotyped elements (Fig. 6). Similarly, stereotyped elements
in adult humans appear to occur as gentle biting in intimate contexts
including mothers or other caretakers directing mild biting to
infants or children, or nursing mothers stimulating an infant’s
neck with fingers or mouth to induce sucking at the breast.
Most
of the photographs and videos of human play that I reviewed were
images of children. Many of these images depicted children playing
in (apparent) solitary mode, standing side-by-side with one or more
children at play [Note added 5/23/2020: see Piaget on this stage of so-called "parallel play"]. Quite a few images showed children playing
face-to-face without threat display or other signs of negative
arousal, and there seemed to be no bias in smiling which sometimes
was exhibited and sometimes not. Considering the many images
inspected depicting children at play, only one photograph employed
features appearing to unambiguously fit diagnostic criteria for a
ritualized action pattern. In this image (Fig. 7), a female child ~3
years old exposed and directed her bared-neck to her male playmate of
about the same age whose head, oriented away from the girl, was,
likewise, positioned with a bared-neck. Finally, the hypothesized
functions of play in primates are similar to those advanced for
social carnivores, although there is a strong predeliction in the
primate literature to interpret human play and other behaviors as
consequences of conscious and aware brain processing. Primates, like
most mammalian species, employ similar behavior patterns during play
or grooming.
Behavioral
Observations
The
subject’s description of the action pattern being dissected and
diagnosed occurred in conditions not atypical for sexual encounters
between human partners. In other words, copulation occurred in a
known location on a comfortable surface (in this case, a bed), in
this case, in dorsal-ventral posture. Description of the servicing
partner’s (the Actor or Donor) sexual behavior was unremarkable up to the point of
ejaculation. In other words, intromission was followed by thrusting
increasing in intensity to the point of ejaculation. The servicing
partner’s head was turned to the right above the receiving
partner’s right clavicle. As commonly displayed, the servicing
partner’s body became rigid during ejaculation, with hands and,
partially, forelimbs pressed on surface of mattress on either side of
the receiving partner’s (Subject, Recipient) trunk. The remarkable feature, and the
motor pattern of note, concerned a response by the servicing partner
accompanying the end-stage of ejaculation. During this phase of the
sexual sequence, the servicing partner involuntarily, seemingly
reflexively by subject report, head-turned further to the right and,
with open mouth and bared teeth, bit the receiving partner’s
epidermis between nape of neck and clavicle in “typical intensity,” exhibiting a “frozen” (immobile) open-mouth motor pattern. No
element of the completed sexual sequence was reported to have been
painful or discomforting to the receiving partner [subject], including the
neck-bite that did not pierce the epidermal layer of skin. According
to the subject, characteristics of the servicing partner’s
end-stage ejaculation were observed in detail because the receiving
participant’s head was already turned to the right, and, also,
because, the response, unique in the receiving partner’s sexual
history triggered (released?) a startle reflex directing receiving
partner’s [subject] eyes to face of the servicing partner [actor], an action by
receiving partner [subject] that did not deflect the attention of the servicing
partner [actor].
Table
1 displays action patterns by the servicing partner as reported by
the subject, matched to the diagnostic criteria conventionally
accepted by ethologists to describe and define ritualized behaviors.
The behavioral sequences detailed by the subject are consistent with
most of the diagnostic criteria defining ritualized behavior by
ethologists. The case material subsequently dissected was diagnosed.
Another focus of the dissection was to demonstrate clear and precise
differentiation between the case material and the intentional and
graded “love-bite” commonly elemental to situations involving
intimate human contact between two or more adults. Inspection of the
results exhibited in Table 1 support the inference that the neck-bite
detailed by the subject was stereotyped, classifiable as a discrete
phenotypic character trait in the human behavioral repertoire.
It
is submitted that the case material described by the subject and
dissected by the investigator ranks with the “eye-flash” as
another, previously undescribed, involuntary and, most likely, genetically-correlated ["hard-wired"] stereotyped action pattern [trait or character]. As reported, the receiving
partner’s body was relatively stationary and rigid with head lying
on the mattress and turned to the right. It seems of interest to
note that, in this posture, the receptive partner’s neck would not
be exposed to the servicing partner, the neck being at least partly
sheltered by chin. Unfortunately insufficient material was provided
by the subject to permit a diagnostic judgment regarding the
likelihood that the receiving partner’s posture exhibited
stereotypy.
Demographic
Information
The
subject and partner were both individuals of color in their early- to mid-thirties, one partner
divorced and the other married and living with spouse. Both
participants were parents of children born to a single mate. The
partners were long-term, distant acquaintances, beginning a sexual
relationship approximately six months prior to the events discussed
in the present case study. The partners’ sexual relationship
terminated approximately four months afterwards. The sexual
relationship occurred in a major urban location in the United States,
and encounters took place in either the home of one partner or a
location associated with one partner’s profession. Partners met in
public rarely and, in addition, had only occasional communication by
phone. Based upon subject report, the sexual relationship was
mutually satisfying, both physically and emotionally. Encounters
were primarily sexual, rarely involving other activities (e.g.,
sharing meals, having extended conversations). At one point, the
partners considered conceiving a child together; however, for
practical, logistic, and other reasons this plan was rejected, a
decision that was not mutual. Neither partner assumed primary
financial responsibility in the relationship, although, one partner was significantly more secure than the other and provided transportation exclusively. The spouse of the married
partner was reported to have had no knowledge of the extra-marital
affair, and, according to the subject, both partners denied having
conflict or guilt about the relationship. Per
subject report, the relationship terminated by mutual agreement
associated with but not limited to obligations and other restrictions
and limitations imposed by one partner’s marital status and
increasing anxiety and conflict about the relationship by the other
partner. Apparently, no deception occurred concerning the married
partner’s intent not to seek a divorce. Subsequent to ending their
relationship, the partners maintained no contact with each other. Throughout the relationship, both partners lived in separate rented
domiciles.
Developmental
History
One
partner’s developmental trajectory through high school took place
in a stable, lower-middle-class home, with married, ostensibly
monogamous, parents, siblings, and minimal conflict among family
members. The maternal parent of this participant was a full-time
homemaker. The family income did not permit home ownership, and the
family inhabited an adequately-sized apartment in a depressed though
relatively safe neighborhood in a major urban area. The second
partner was raised in a middle-class family residing in their own
home in a small city near a major metropolis. Although parents of
this participant remained married and were ostensibly monogamous, the
household was a zone of intra- and inter-individual conflict, including maternal
neglect. The mother of this family unit worked outside the home,
with a live-in housekeeper tending to children and household duties. Age of first sexual
encounter for one partner was unknown to the subject whose first
intercourse occurred after the age of sixteen. One partner married
before the age of twenty, divorcing soon after the age of thirty.
The other partner married after the age of twenty-five. When the
relationship described in the present case study terminated, the
divorced participant had more than one child while the other had one
with one expected in ~7 months.
Cognitive and Mental
Status Summary
Based
upon subject report, both partners’ I.Q.s measured in the normal-high
range though, as noted, one partner’s psychological trajectory was
unstable relative to the other. No learning disabilities were
reported for either partner or children.
Work and Academic
Summary
Both
partners were professionals with post-graduate training with one
partner having work of significantly higher status, socially and
financially, compared to the other. The professions of both partners
were highly specialized.
Symptoms and
Challenges
As
reported by the subject, from childhood, the life trajectory of the
unstable partner was single-mindedly focused on studies interrupted
by episodes of serious mood disorders accompanied by anxiety
(including bouts of agoraphobia). This partner’s status, compared
to the other, was described as “unbalanced,” both
psychologically, financially, socially, and energetically. The more
stable partner apparently exhibited a noteworthy capacity for
compassion and friendship; however, this partner’s sexual habits
might be termed serially "polygamous." According to the subject, the cognitive,
mental, and behavioral status of each partner’s children appeared
to be within normal range.
Diagnostic
Impressions
The
investigator is not a psychological/psychiatric clinician; thus, the
following interpretation is not supported by training or expertise
using diagnostic criteria for psychopathologies. However, as noted
above, the technical literature makes clear, gentle biting, focusing,
in particular, on the "love-bite," is not classified as pathological
behavior. The present case material was tentatively evaluated,
further, with reference to the DSM-IV-PR Axes delineating relative
degrees and levels of severity of presenting behaviors. According to
these diagnostic criteria, the investigator concluded that the
partner described as “unstable” would be classified in Axis I
(severe presenting characteristics requiring immediate attention from
a professional) and in Axis IV (significant work and interpersonal
stress). The partner described as “stable”, was classified in
Axis IV based on moderate to severe work stress and, despite this
participant’s denial, likely multiple stressors associated with
infidelity. Following the diagnostic criteria for Axis IV’s Global
Assessment of Functioning Scale (GAS), the less stable partner
received two ratings based on separate criteria detailed for the
scale: 41-50 (difficulty with social and occupational symptoms) and
31-40 (some impairment in reality-testing, interpersonal
relationships, judgment, and mood). The more stable partner also
received two ratings: 41-50 (same criteria as above) and 61-70
(generally functioning, some meaningful interpersonal relationships).
Discussion
Using
qualitative phenetic methods, this paper compiles evidence to test
the hypothesis that a stereotyped "neck-bite" motor pattern is a
component of the human behavioral repertoire. Employing ethological
descriptions of the motor pattern by a participant observer matched
to diagnostic criteria for ritualization and by inspecting scores of
photographs of behavior targeting the neck in a variety of mammal
species in diverse contexts, the investigator concluded that the
described behavior exemplifies a stereotyped trait in the behavioral
repertoire of humans. Comparing features of the present case
material with structures and functions of the "neck-bite" postural response [action pattern, motor pattern] across mammals provided critical data for the present inferences.
After Estes (1992), these comparisons revealed significant
variability in structure and function of the neck-bite, not only
between but also within Orders. For example, in carnivores, the
neck-bite posture used during copulation is very similar when canids
and felids are compared. However, modes of hunting and killing
between these units have evolved differently, with canids, in
general, hunting in groups and killing with the death-shake while
felids are, on whole, solitary hunters killing with a ritualized
neck-bite motor pattern.
Estes
(1992) provides other examples of variability in the appearance of
the neck-bite posture between the closely-related lesser and greater
bushbabies. The "neck-bite" is absent from the behavioral repertoire
of the first species but evident in the latter, more recently
evolved, species. This pattern indicates that the "neck-bite" is a
derived character state, evolved from solitary taxa lacking the
morphological character. When studying this and other stereotyped
traits, however, it is important to recall as a caveat
that evolutionary biologists and behavioral ecologists conclude that "ritualized" motor patterns have the potential to evolve rapidly
(Enquist et al., 2010), presumably in response to changing
environmental conditions. Thus, systematic patterns of these derived
signals over time are not necessarily expected to be linear or
predictable within or between taxa. The latter issue should apply to
investigations of both structure and function.
The
ethological literature advances the idea that ritualized behaviors
signal motivation in a variety of conditions, frequently functioning
to decrease chances of direct confrontation between conspecifics
and/or the likelihood of damaging aggression (Eibl-Eibesfeldt, 2007;
Lorenz, 1981; Eisenberg, 1981; Morris, 1957). Researchers
investigating signals and displays in non-human animals have
generally followed the diagnostic criteria and conclusions by
ethologists as well as treatments relying upon contemporary methods
and theoretical treatments (Maynard Smith & Harper, 2003; Enquist
et al., 2010). An extensive literature exists discussing and
analyzing differential forms and functions of these morphological
traits (Maynard Smith & Harper, 2003 and references; Enquist et
al., 2010 and references). While thorough review of the ethology of
mammals is beyond the scope of the present paper, Eibl-Eibesfeldt
(2007), Eisenberg (1981), and Estes (1992) provide early, important
summaries, with additional information located primarily in technical
papers. Since, to my knowledge, no comprehensive evaluation exists of "ritualized" action patterns in mammals (but see, Eibl-Eibesfeldt, 2007; Ewer, 1968), this topic remains open for
investigation.
Unfortunately,
discussions of stereotyped responses by primatologists have been
biased by early research on Hamadryas baboons (see Swedell &
Schrier, 2009 and references therein). Because Hamadryas males employ the ritualized
neck-bite aggressively for coordination and control of harem females
in sexual and other contexts, the motor pattern has generally been
interpreted in the framework of male coercion and force. Despite the
aggressive behavior exhibited towards females by males of this
species, it would be erroneous to generalize from this single case to
all simian species displaying the neck-bite since most primate
studies do not precisely discuss stereotyped and/or ritualized features of sexual
behavior. Furthermore, field experiments have not been conducted,
and sex and aggression have not been studied systematically in many
primate species.
According
to Estes (1992), aggression (including rape) by adult mammalian males toward
reproductive females is likely to be favored by selection in two
conditions. First, females of harem species generally don’t
copulate outside of their receptive periods. This state leads to a
female-biased "operational sex ratio" if more than one female is in
estrus at the same time and, also, if females breed seasonally (Emlen
& Oring, 1977). Since a male will have difficulty monopolizing
more than one female at a time (Emlen & Oring, 1977), this
condition restricts a harem male’s temporal window for copulation
and successful fertilization. Thus, in these conditions, coercion and
force by males may be favored by selection, possibly representing a
time-saving strategy benefiting a male’s “fitness budget” and
minimizing mating investment. Estes (1992) points out, further, that "multimale-multifemale" ("polygynandrous") societies are primed, as well, for selection to
favor male coercion and force toward females since females may mate
with multiple males (sometimes referred to as “female promiscuity”
or, less commonly in the mammalian literature, “polyandry," see elephant seals, "incitation of 'male-male competition'": Cox & LeBoeuf, 1977) during estrus and because females
in these socio-sexual groups are pregnant or lactating for most of
their lives. Both of these conditions lead to a male-biased "operational sex ratio" inducing intense male-male competition for
cycling mates (e.g., chimpanzees).
On
the other hand, Estes (1992) continues, when tactics of
multiply-mating females successfully prohibit a single male from
obtaining exclusive access to one or more females, aggressive
behavior by males toward the opposite sex may be rare or absent. For
example, female mantled howlers may rebuff male advances with a
stereotyped, bared-teeth, open-mouth display (Jones, 1985), and
female bonobos may form coalitions to overpower males (Vervaecke et
al. 2000 [Note added 5/24/2020: bonobo females are dominant to males, uncommon among mammals]). Estes (1992) states further, that, as exemplified by
bushbabies, solitary male primates rarely exhibit the neck-bite or
aggression toward females in sexual contexts. Where the ritualized
neck-bite is displayed in these species, however, it is generally
expressed mildly and non-aggressively. Importantly, Estes (1992)
shows that male to female aggression is, also, rare or absent where
females are matrilocal (e.g., vervets), where females are dominant to
males (e.g., bonobos), and where females exhibit strong "female-choice" of mates (e.g., mantled howlers, many cercopithecines). Following
the criteria advanced by Emlen & Oring, 1977 who discuss “female
emancipation,” it might also be suggested that male aggression
toward females is not likely to be favored in heterogeneous regimes
in which resources, especially food, are highly variable, making it
energetically expensive for males to monopolize females (muruquis,
mantled howlers, bonobos). It seems likely, as well, that male
aggression towards females would be rare or absent in species
characterized by female same-sex mate choice (e.g., Japanese
macaques); however, to my knowledge, research has not been conducted
or published on the use of stereotyped responses in species
exhibiting same-sex mate choice or on coercion and force directed at
females by males in these same societies.
Theoretical
work advances the idea that males and females are in an evolutionary "arms race" to impose greater costs on the opposite sex than that sex
imposes upon the other (Rice, 2000). The treatment by Rice (2000)
assumes that males initiate the co-evolutionary race, a position that
has been criticized in the animal behavior literature. Though Rice’s
(2000) ideas are widely accepted by scientists studying the evolution
and behavioral ecology of socio-sexual behavior, they have not often
been applied by primatologists, or other social scientists, possibly because of the significant
influence of feminists and humanists in this field. As the brief
review in this paper indicates, sometimes males appear to have “won”
this race (mantled howlers, Hamadryas baboons, chimpanzees), sometimes females appear
to have won (lemurs, bonobos), and sometimes power relations between
the sexes seem to vary by context (some capuchins, mantled howlers,
gorillas, humans). Differential social influence between primate
males and females is probably sensitive to limiting resource dispersion (distribution and abundance) over
time and space; thus, it may be difficult to easily categorize these
groups within and between taxa. Estes’ review (1992; also see
Eisenberg, 1981; Martin, 1989; McKenna & Bell, 1997; Skinner &
Chimimba, 2005) demonstrates noteworthy variability in male-female
relationships within and between mammalian orders, dispensing with
the prejudice that male to female behavior in sexual contexts can be
generally characterized as coercion or force. Estes’ (1992) and
Eisenberg’s (1981), among others’, important contributions to the
literature on these topics in mammals require updating and
comprehensive treatments, additional research programs in need of
attention. As a final observation, it is likely that, when males
direct a stereotyped "neck-bite" to females, the motor pattern
expresses multiple motivations such as dominance, sexual intent,
and/or coordination and control, motivations that may or may not be
exhibited aggressively, other topics awaiting updates.
Limitations and
Recommendations
Perhaps
the primary criticism to be made of the present dissection and
diagnosis of case material is the high probability that
intraindividual and interindividual error within and between subject
and investigator significantly impacted the study’s reliability and
validity. Interview methodology was rejected in an attempt to
minimize potentially biasing effects repeating case material solely
to create a more vivid mental representation of events. Subject
report was not sufficiently detailed to permit identification of
possible "releasers" (a stimulus or stimulus pattern associated with
the reported behaviors) inducing the ritualized neck-bite.
During
a sexual encounter, it is likely that a mosaic of releasing stimuli
arise from the physical (e.g., colors, shapes of objects,
similarities and/or differences between familiar and less familiar
contexts) and biotic, including social (e.g., mood, familiarity of
sexual partners, general or specific anxiety or conflict about
activities, number of concurrent sexual partners, olfactory and other
perceptual stimuli) contexts. Precise delineation and identification
of releasing stimuli would possibly pose an insurmountable challenge
even under controlled conditions (e.g., by field experimentation or
direct observation and measurement in the laboratory). However, it
is clear that direct, sequential observation and measurement of the
morphological and other characteristics of the sequence of behaviors
comprising the sexual encounter and its context, possibly by video or
related imaging techniques, would, at minimum, be required to permit
informed speculation about releasing stimuli as well as precise
delineation of morphological patterns. If these are idiosyncratic,
specific to individuals, time, and space, the enterprise, even with a
robust sample size, becomes much more challenging.
In
this report, case material relied upon a single encounter (N= 1),
obviating within-species comparative analysis and generalization.
Thus, inferences made in the present study only achieve the status of
propositions made by an expert investigator-observer. It is unclear whether
the presumed absence of published reports describing the stereotyped "neck-bite" motor pattern delivered at ejaculation by the servicing
partner to a receptive sexual partner represents the rarity of this
response in Homo
sapiens
or reluctance to reveal such information (e.g., for reasons of
reputation, embarrassment, shame, guilt). Another possibility would
be failure to recognize details of a behavioral variant by one or
more participants. A first step towards demonstrating the "neck-bite" as a component of the stereotyped behavioral repertoire of (most
likely male) humans would require collection of anonymous
questionnaire or other survey data using a large sample of sexually
active adults. Stereotyped motor patterns appear to be rare elements
of human phenotypes, and their proximate and evolutionary causation
as well as their frequency, intensity, variability, forms, and
functions are poorly understood. As suggested throughout this
report, additional research is indicated.
Conclusions
The
present case study used a modified phenetic methodology to dissect a
participating subject’s verbal report detailing events occurring in
one sexual context. Because certain features of the situation
appeared to deviate from published reports of intercourse between
heterosexual partners, elements of the events were scrutinized by an
investigator with expertise in ethological theory, methods, and
research. The morphological feature of note entailed co-occurring
and post-ejaculation behavior by the servicing partner who was
diagnosed to have exhibited a stereotyped "neck-bite," a posture
observed to function in a broad range of mammalian taxa in a variety
of conditions. In the ethological literature, the ritualized "neck-bite" motor pattern is generally advanced to function as a signal
communicating motivation and/or intent, and numerous experts suggest,
as well, that stereotyped patterns of behavior often function to
decrease the likelihood of conflict or damaging aggression between
conspecifics. Limitations of the program reported herein are
discussed and recommendations are made to decrease error and refine
procedures in future studies.
Stereotyped
motor patterns targeting the neck reported for extant mammalian taxa,
including the case study treated in the present report, display
features most similar to the felid "neck-bite" rather than the "death-shake" of canids. Thus, the felid "neck-bite" may have been a
generalized developmental template in mammals giving rise to a broad
range of ritualized motor patterns targeting the neck. Another
evolutionary scenario would advance the idea of convergence of motor
patterns, possibly induced by a fundamental mammalian behavioral
“toolbox”. Jaguars and cougars, for instance, may exemplify the
relatively complete disinhibition, loss, and substitution of an
ancestral feature, a topic studied by ethologists, particularly,
Moynihan (1970), but awaiting a comprehensive comparative
investigation. Importantly for topics of current interest to
evolutionary biologists, these processes may pertain to the
generation of behavioral novelty (West-Eberhard, 2003) and to
evolvability (Kirschner & Gerhart, 1998). Also, as several
sources have indicated, it would be significant to study relative
predator-prey sizes and other characteristics within and across
orders and families, including but not limited to, cranial anatomy,
odontology, socio-sexual systems, ecology, climate, and other factors.
Finally, biophysical and behavioral research, including, field experiments, mathematical
modeling, simulations, and other theoretical treatments, could compare the
dorsal-ventral with other sexual postures (e.g., the “missionary”
position) to assess whether these positions differ in their
likelihoods of inducing the stereotyped "neck-bite" and/or whether the
receiving partner is differentially vulnerable to harm by the
servicing partner. Data such as these might provide insights into
the evolution of mammalian and other vertebrate sexual postures,
including reasons that ventral-ventral sex is so rare in nature [Note added 5/23/2020: the dorso-ventral sexual position would seem to benefit male interests if it benefits male coordination and control compared to the ventral-ventral posture]. It
is hoped that the dissection and diagnosis exemplified by the present
case study will generate intense scrutiny, probing questions, and
future research on ritualized signals and displays in humans and
other species.
Acknowledgments
I
thank Jason Epperson for technical assistance and Dr. Kenneth J.
Zucker for encouragement. Rick Murphy generously shared his
photograph of a “shoulder-bite” displayed during copulation by a
lowland gorilla silverback during ejaculation. I am very grateful
for Murphy’s permission to publish this image. The accessibility
and broad coverage of topics discussed in this report were
significantly aided by the photographs available at Wikimedia
Commons, by You Tube’s video collection, and on Google.
References
Alcock,
J. (1998). Animal behavior: An evolutionary approach (6th
Ed.). Sunderland, MA: Sinauer.
American
Psychiatric Association. (2000). Quick reference to the diagnostic
criteria from DSM-IV-PR (1st
Ed.). Arlington, VA: American Psychiatric Association.
Berkowitz,
S.R., Laney, C., Morris, E.K., Garry, M., & Loftus, E.F. (2008).
Pluto behaving badly: False beliefs and their consequences. American
Journal of Psychology, 121, 643-660.
Boesch,
C., & Boesch, H. (1989). Hunting behavior of wild chimpanzees in
the Tai National Park. American Journal of Physical Anthropology, 78,
547-573.
Bradbury,
J.W., & Vehrencamp, S.L. (1998). Principles of animal
communication. Sunderland, MA: Sinauer Assoc., Inc.
Buskirk,
R.E., Frolich, C., & Ross, K.G. (1984). The natural selection of
sexual cannibalism. American Naturalist, 123, 612-625.
Cox, C. and Le Boeuf, B..J. (1977). Female incitation of male competition: A mechanism in sexual selection. The American Naturalist, 3(978), 317-335.
Eibl-Eibesfeldt, I. (2007). Human ethology. Piscataway, NJ: Aldine Transactions (Aldine De Gruyter).
Cox, C. and Le Boeuf, B..J. (1977). Female incitation of male competition: A mechanism in sexual selection. The American Naturalist, 3(978), 317-335.
Eibl-Eibesfeldt, I. (2007). Human ethology. Piscataway, NJ: Aldine Transactions (Aldine De Gruyter).
Eisenberg,
J.F. (1981). The mammalian radiations. Chicago: University of Chicago
Press.
Emlen,
S.T., & Oring, L.W. (1977). Ecology, sexual selection, and the
evolution of mating systems. Science, 197, 215-223.
Enquist,
M., Hurd, P.L., & Ghirlanda, S. (2010). Signaling. In D.E.
Westneat & C.W. Fox (Eds.), Evolutionary behavioral ecology
(266-284). Oxford: Oxford University Press.
Estes,
R. (1992). The behavior guide to African mammals including hoofed
mammals, carnivores, and primates. Chicago: University of Chicago.
Ewer, R.F. (1968). Ethology of mammals. Logos, London.
Ewer, R.F. (1968). Ethology of mammals. Logos, London.
Franks,
D.W., & Nobel, J. (2004). Warning signals and predator-prey
coevolution. Proceedings of Biological Sciences, 271, 1859-1865.
Jones,
C.B. (1983). Social organization of captive black howler monkeys
(Alouatta
caraya):
“Social competition” and the use of non-damaging behavior.
Primates, 24, 25-39.
Jones,
C.B. (1985). Reproductive patterns in mantled howler monkeys: Estrus,
mate choice, and copulation. Primates, 26, 130-142.
Jones,
C.B. (1986). Infant transfer behavior in humans: A note on the
exploitation of young. Aggressive Behavior, 12, 167-173.
Jones,
C.B. (2005). Behavioral flexibility in primates. New York: Springer.
Jones,
C.B., & Van Cantfort, T.E. (2007). Multimodal communication by
male mantled howler monkeys (Alouatta
palliata)
in sexual contexts: A descriptive analysis. Folia primatologica, 78,
166-185.
Kirschner,
M., & Gerhart, J. (1998). Evolvability. Proceedings of the
National Academy of Sciences USA, 95, 8420-8427.
Kleiman,
D.G., & Eisenberg, J.F. (1973). Comparisons of canid and felid
social systems from an evolutionary perspective. Animal Behaviour, 4,
637-659.
Lehner,
P.N. (1998). Handbook of ethological methods (2nd
Ed.). Cambridge: Cambridge University Press.
Lorenz,
K. (1981). The foundations of ethology. Berlin/New York:
Springer-Verlag.
Mc
Coy, M.W., Bolker, B.M., Warkentin, K.M., & Vonesh, J.R. (2011).
Predicting predation through prey ontogeny using size-dependent
functional response models. American Naturalist, 177, 752-766.
Mc
Kenna, M.C., & Bell, S.K. (1997). Classification of mammals. New
York: Columbia University Press.
Marler,
P. (1976). An ethological theory of the origin of vocal learning.
Annals of the New York Academy of Sciences, 280, 386-395.
Martin,
L.D. (1989). Fossil history of the terrestrial carnivores. In J.L.
Gittleman (Ed.). Carnivore behavior, ecology, and evolution. Ithaca,
NY: Comstock Pub.
Maynard
Smith, J., Parker, D. (2003). Animal signals. Oxford: Oxford
University Press.
Millon,
T., Blaney, P.H., & Davis, R.D. (1999). Oxford textbook of
psychopathology. Oxford: Oxford University Press.
Morris,
D. (1957). “Typical intensity” and its relation to the problem of
ritualization. Behaviour, 11, 1-12.
Morris,
D. (1995). Bodytalk: The meaning of human gestures. New York: Crown
Publishers.
Morris,
D. (1997). Intimate behaviour. Tokyo: Kedansha Globe.
Moynihan,
M. (1970). The control, suppression, decay, disappearance, and
replacement of displays. Journal of Theoretical Biology, 29, 85-112.
Napier,
J.R. (1970). The roots of mankind. Washington, DC: Smithsonian
Institution Press.
Poole,
T.B. (1972). Some behavioral differences between the European
polecat, Mustela
putorius,
the ferret, M.
furo,
and their hybrids. Journal of Zoology, 56, 25-35.
Quattara,
K., Lemasson, A., & Zuberbuhler, K. (2009). Campbell’s monkeys
concatenate vocalizations into context-specific call sequences.
Proceedings of the National Academy of Sciences USA, 106,
22026-22031.
Rice,
W. (2000). Dangerous liaisons. Proceedings of the National Academy of
Sciences USA, 97, 12953-12955.
Roughgarden,
J. (2009). Evolution’s rainbow. Berkeley: University of California
Press.
Saint-Girons,
M.-C. (2006). Snakes’ enemies. In R. Bauchot (Ed.). Snakes: A
natural history (172-183). New York: Sterling Pub. Co., Inc.
Schuh,
R.T., & Brower, A.V.Z. (2009). Biological systematics: Principles
and applications (2nd
Ed.). Ithaca, NY: Comstock Pub. Assoc.
Seyfarth,
R.M., Cheney, D.L., & Marler, P. (1980). Monkey responses to
three different alarm calls: Evidence of predator classification and
semantic communication. Science, 210, 801-803.
Sih,
A. (1985). Predation, competition, and prey communities: A review of
field experiments. Annual Reviews of Ecology and Systematics, 16,
292-311.
Skinner,
J.D., & Chimimba, C.T. (2005). The mammals of the southern
African sub-region. Capetown, SA: Cambridge University Press.
Smuts,
B.B. (1996). Male aggression against women: An evolutionary
perspective. In D.M. Buss & N.M. Malamuth (Eds.). Sex, power,
conflict: Evolutionary and feminist perspectives. New York: Oxford
University Press.
Surbeck,
M., & Hohmann, G. (2008). Primate hunting by bonobos at
Luikotale, Salonga National Park. Current Biology, 18, R906-R907.
Swedell,
L., & Schrier, A. (2009). Male aggression toward females in
Hamadryas baboons: Conditioning, coercion, and control. In M.N.
Mubler & R.W. Wrangham (Eds.). Sexual coercion in primates and
humans: An evolutionary perspective on male aggression against
females (pp. 244-269). Cambridge, MA: Harvard University Press.
Tinbergen,
N. (1952). Derived activities: Their causation, biological
significance, origin, and emancipation during evolution. Quarterly
Review of Biology, 27, 1-32.
Van
der Dennen, J.M.G. (2005). Ritualized primitive warfare and rituals
in war: Phenocopy, homology, or….? Retrieved 16 April 2011 from
http://rechten.eldoc.ub.rug.nl/files/root/algemeen/overigepublicates/2005enouder/ritual/ritual.pdf
Vervaecke,
H., de Vries, H., & van Elsacker, L. (2000). Distribution and
functions of coalitions in captive bonobos (Pan
paniscus).
Primates, 41, 247-263.
Weber,
B.H., & Depew, D.J. (Eds.). (2003). Evolution and learning: The
Baldwin Effect reconsidered. Cambridge, MA: MIT Press.
West-Eberhard,
M.J. (2003). Developmental plasticity and evolution. Oxford: Oxford
University Press.
Wilson,
E.O. (1971). The insect societies. Cambridge, MA: Belknap.
Table
1. Table exhibits Ethological Features of ritualized motor patterns,
Diagnostic Criteria as defined by ethologists for features of
ritualized motor patterns, and investigator’s subjective diagnosis
of this report’s case material matched to factors in the first two
columns. This table provides a summary of the results employed for
inferences, and, in the final analysis, the motor pattern described
by the subject was diagnosed as a stereotyped neck-bite by the
investigator. See text for greater detail, discussion, examples, and
references.
ETHOLOGICAL
FEATURES
|
DIAGNOSTIC CRITERIA | CASE MATERIAL MATCH |
| Ritualization |
Process
leading to discrete rather than graded morphological feature of a
conserved character trait “derived” and “emancipated” from
primitive form
|
Case report described discrete motor pattern comparatively similar to behaviors targeting the neck in other mammals, including primates |
| Stereotyped morphological feature |
Character
trait with signal function(s) exaggerated, simplified, and
involuntary compared to conserved feature; stereotyped signals and
displays appear “fixed” or “frozen” in time and space;
“fixed action patterns” are condition-dependent, induced by
“releasers” (subjective and/or environmental “releasing
stimuli”)
|
Report of case material described motor pattern characteristic of stereotyped responses, including displays, and “fixed action patterns”; morphological features very similar to felid and other primate neck-bite displays; precise function(s) of case material and releasing stimuli could not be determined |
| Typical intensity |
For
given age and sex, stereotyped motor patterns, ceteris
paribus,
exhibit the same or similar morphological features within and
across conditions
|
Unknown since N= 1; however, as described by subject, typical intensity exhibited for duration of event |
| Intention movement | A stereotyped motor pattern similar to other ones but failing to fully complete, carry out, or terminate a given ritualized morphological feature | As described by subject, motor pattern did not progress to uncomfortable biting nor did it puncture epidermal layer of skin of neck |
Figure Legends
Figure
1. Felid “neck-transport” employed by females to move young from
one location to another. Most canids, also, exhibit this motor
pattern. Though not as incomplete or truncated as an intention
movement, this response is inhibited or restrained compared to the
neck-bite employed to kill. For this and following images,
additional detail is provided in text of paper. Photo accessed from
Wikimedia Commons.
Figure
2. A modified neck-bite, the “shoulder-bite”, displayed by a
silverback lowland gorilla. According to the researcher providing
this image, this display always occurs during ejaculation. The
female in
copula may
respond with screams or may chase the servicing male; at other times,
however, the female remains silent and passive. The shoulder-bite,
in addition to the neck-bite, also occurs in sexual contexts in
Hamadryas baboons, with varying responses by receiving females.
©Rick Murphy
Figure
3. Example of erythemas (“hickies”) resulting from human “love
bites”. Love-bites appear to represent intentionally motivated
behavior and not stereotyped motor patterns. Love-bites are not
classified as paraphilias unless they escalate to sado-masochistic or
other behavior harmful, physically or otherwise, to one or more
sexual partner. Photo accessed from Wikimedia Commons.
Figure
4. An example of anthropoid (cercopithecene) “play-bite”,
diagnosed as generally stereotyped based upon investigator inspection
of photographs and videos. Similar analyses of hominid play did not
reveal any apparent biases to target the neck. Photo accessed from
Wikimedia Commons.
Figure
5. Movement by human male to female appearing to target female’s
neck. Displays such as this one have been termed “impulsive” and
may represent intentionally incompleted kiss or play. It is not
possible from this image to determine whether the motor pattern
includes stereotyped elements and whether it is classifiable as an
intention movement by ethological criteria. Photo accessed from
Wikimedia Commons.
Figure
6. The human kiss may have characteristics consistent with
stereotypy, note, especially, tongue movement (the “French kiss”)
that may at times be involuntary, thus, arguably, ritualized. Mouth
motor patterns similar to the human kiss are displayed by bonobos,
chimpanzees, and, possibly, other primates. Some other taxa (e.g.,
horse) exhibit similar behavior. Photo accessed from Wikimedia
Commons.
Figure
7. The sole image of (human) children at play clearly depicting motor
patterns oriented in the manner of stereotyped play postures employed
by many mammalian species to solicit grooming or other behavior
targeting the neck. These morphological features may also
communicate subordinance and/or non-aggressive motivation in a broad
range of vertebrate taxa. Despite appearance, it is not possible to
confidently determine whether these motor patterns in this image
conform to ethologists’ diagnostic criteria for stereotyped
responses. Photo accessed from Google.
*This MS was submitted to journal, Archives of Sexual Behavior ~2011 but rejected on the basis of "ethical" concerns.
