SCHEMATIC
NOTE ON MAMMALIAN SIGNALING
Table
1: Sensory
modalities of mammals (“Type of Signal or Display”), with
examples, including, “Feature” or “Capacity” of Signal or
Display in each of six domains. Signals or Displays may be emitted
or advertized to one or more Receiver, and one or more animal
(non-human or human) may be actively Eavesdropping (Figure 1).
Depending upon physiological and behavioral traits of species, signal
and display types represent a sensory “toolbox” with potential
for multimodal (“complex”) transmission of information (Fig. 1).
Both unimodal and multimodal communication are subject to temporal
regulation, yielding varying sensory elements subject to statistical
encoding, decoding, interpretation, and “prediction”. Despite
potential for regulation of signal and display features, accurate
signals are characterized by “redundancy”, a property that is
highly correlated with reliability of communication (Wilson 1975).
On the other hand, messages are not necessarily selected
for maximal accuracy*# in a single or a few contexts or domains since
it may benefit individuals to utilize a signal or display in multiple
contexts and for multiple functions, leading to one or more signals
optimally effective across conditions. Furthermore, highly accurate
messages may be easier to escape or avoid compared to “fuzzier”
ones because they may be more detectable than (optimally) inaccurate
signals. Thus, it may not benefit a Signaler to optimize production
of a Receiver’s “pattern detection” mechanisms.
Signals
and displays of “solitary” compared to social mammals are not
necessarily less “complex” since different selection pressures
may have favored different population structures and traits.
However, excepting traits related to courtship and mating, signals
and displays associated with within-group coordination and control
(“integration”) are expected to be more differentiated,
elaborate, and “complex” among social mammals because of higher
local population density and, especially, higher rates of
interaction. (©Clara
B. Jones, based on Goodenough et al. 2009)
TYPE OF SIGNAL OR DISPLAY | |||||
FEATURE OR CAPACITY | Visual (horns, proboscis, sexual swellings, mimicry, “natal coats”, stotting) | Auditory (howls, echolocation, ululation, “contact calls”, “alarm calls”, language) | Chemical (pheromone, excretions, venom) | Tactile (grooming, “neck bite”, copulation, spines, defensive integuments) | Electrical (monotremes) |
Effective distance | Medium |
High
|
High | Low | Low |
Localization | High | Medium | Changeable | High | High |
Ability to go around obstacles | Low | Medium | Medium | Medium | High |
Detection of change | High | High | Low | High | High |
Complexity | High | High | Low | Changeable | Low |
Durability | Changeable | Low | High | Low | Low |
*Recent research shows that inaccurate signals promote phenotypic flexibility. If this is generally the case, inaccurate signalling may be one alternative response that promotes coexistence, mutualism, and/or social behavior because the response may mitigate chances of competitive exclusion or aggression [aversive responses, punishment].
#See Sultan SE, Spencer HG 2002 Am Nat 160: 271-283, p 280, column 1, paragraph 2...
#See Sultan SE, Spencer HG 2002 Am Nat 160: 271-283, p 280, column 1, paragraph 2...
Figure 1: This figure displays a Signaler-Receiver schema in optimality (cost-benefit) context as follows. The classical, Ethological, view, adopted by behavioral ecologists, holds that “the actor [Signaler] is selected to manipulate the behaviour of the reactor [Receiver]”, and communication “makes sense only in the context of an exchange of information” (Dawkins and Krebs 1978) whereby a signal or display (Table 1) is employed by a Signaler to induce a self-interested, beneficial response. A corollary of the latter perspective would be that, as a recipient of information (the “signal” or “display”), a Receiver “decides” (Hebbian decision) to respond or not, becoming a potential “Sender”, responding in a manner, presumably, biased by the original “message” but, like the original transmission, biased by self-interest. Proximate and ultimate (reproductive) benefits or losses to Signalers will be a function of statistical averages, and Signalers whose messages, on average, produce a threshold-level of benefits “propagate their genes more efficiently” than Signalers who do not. Following Dawkins and Krebs (1978), “communication results in a net average benefit to the actor”.
Fig. 1 displays differential, initial benefits and costs to Signaler
and Receiver; however, it remains unclear whether or to what extent,
average benefits may accrue to Receivers, particularly, where signals
or displays are “dishonest” (e.g., female mimicry of male
genitalia, fake orgasm by females). The present discussion assumes
that the condition-dependent, “fitness optima” of Signaler and
the intended Receiver (or, unintended Receiver[s]) conflict.
However, theory holds that some threshold of conflict will favor
costly (“exaggerated”, “complex”), honest signals and
displays, maximizing the transfer of “accurate” information and
minimizing likelihoods of aggression (Tinbergen 1952). In these
regimes, interactions between Sender and Receiver should be, “at
least statistically, predictable from…past behaviour” (Dawkins
and Krebs 1978). As Dawkins and Krebs (1978), Maynard Smith and
Harper (2003), and Eisenberg (1981) point out, signals and displays
may derive from
changes in biological information emitted by a Signaler
(“cues”), including, proximate or ultimate events associated with
thermoregulation (e.g., “huddling”) and excretion (urination,
defecation: e.g., orgasm). Calculations of differential benefits and
costs are complicated by the recognition that such calculations
(“decisions”) by “ego” are estimates of future gains
and losses. ©Clara B. Jones (after Bradbury and Vehrencamp 1998)