What Is "Behavioral Ecology"?: A White Paper (by Clara B. Jones, 12/28/2017)

What Is "Behavioral Ecology"? A White Paper (by Clara B. Jones, 12/28/2017) 

Definition of Behavioral Ecology: Variations in behavior relative to ecological [economic] factors, in particular, spatial & temporal dispersion [distribution & abundance] of limiting resources; Ways in which Dispersion [Distribution & Abundance in Time & Space] of organisms "maps" onto Dispersion of limiting resources [in T & S in a given population]--the [John Hurrel] Crook-ian Model of Behavioral Ecology [Behaviour Supplement X, 1964]...limited by energetics x sex [on average & ceteris paribus]--males expected to be Time-Minimizers, females expected to be Energy-Maximizers

FIRST PRINCIPLES OF BEHAVIORAL ECOLOGY:: E[nergy]: Acquisition->Consumption->Allocation====> Worker &/or Reproductive &/or Dependent...(Males, T[ime] Minimizers; Females, E[nergy] Maximizers)

The organizing principle of this White Paper is that "Behavioral Ecology" is a sub-field of Ecology, not a sub-field of Animal Behavior, Comparative Psychology, Ethology, or Anthropology.

As such, Behavioral Ecologists will study behavioral, including, social*, traits as they operate/function at population, community, and ecosystem levels, incorporating concerns for scale, mechanisms, development, tradeoffs, mediating factors, and filtering, among other related issues.

Students of Behavioral Ecology will demonstrate an awareness of the roots of their field, including, but, not limited to, the early work of John Eisenberg, John Hurrell Crook, Stephen Emlen, Jack Bradbury, and Sandy Vehrencamp.

Many of the traits of interest to Behavioral Ecologists will be genetically correlated; thus, genetic and genomic studies will be employed to identify genes, gene complexes, and/or circuits underlying behavioral, including, social*, traits--relative to abiotic and biotic environmental factors and interactions.

The journal, Behavioral Ecology, will be viewed as an Ecology journal on par with the journals, Functional Ecology, Journal of Animal Ecology, Ecology and Evolution, and Journal of Applied Animal Ecology.

Behavioral Ecology will reflect the intimate links between Ecology and Evolutionary Biology (EcoEvo)**.

Behavioral Ecology will become a predictive discipline, not only a project of descriptive work. As such, a truly predictive Behavioral Ecology will be a hypothetico-deductive enterprise based on First Principles.

Like its parent discipline, Ecology, Behavioral Ecology methodology will incorporate modeling and simulation, as well as, field and laboratory experiments and will investigate tradeoffs and alternative hypotheses. Practitioners can conduct experiments with agent-based [individual-based] methods.

Behavioral Ecologists will be trained by Ecologists and Evolutionary Biologists (EcoEvo) from Departments of Ecology and Evolution and, in addition, will study Ethology, Animal Behavior, & Population Genetics.

Behavioral Ecology will be characterized by strong theory, and students will be trained in quantitative methods, at minimum, statistics, biostatistics, coding, calculus, agent-based [individual-based] modeling. Higher-order quantitative skills might incorporate Fisher's Fundamental Equation, the Price Equation, inclusive fitness ("kin selection") & Hamilton's Rule, as well as, the Nash Equilibrium. As in other sub-fields of Ecology, theory will take the form of Mathematics, though verbal formulations will often be a preliminary step. Marshall's book, Social Evolution and Inclusive-Fitness Theory, might be incorporated into any graduate student's program:

https://www.amazon.com/Social-Evolution-Inclusive-Fitness-Theory/dp/0691161569/ref=sr_1_1?keywords=james+marshall+social+biology&qid=1558917758&s=books&sr=1-1-catcorr

The practitioner of Behavioral Ecology will study virtually any topic investigated by other Ecologists. A good exercise is to peruse the contents of the journals mentioned above, interpolating and/or reframing most any paper into a study of Behavioral Ecology, including, Social* Biology. Once the practitioner gets the knack of doing this, s/he/they can advance to other topics generated by books such as The Princeton Guide To Ecology or any good Ecology textbook. In 2013, the British Ecological Society identified "100 fundamental questions in Ecology" that can be re-framed as questions for research in Behavioral Ecology and Social Biology: https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1111/1365-2745.12025

Behavioral Ecology will include a new sub-field, Applied Behavioral Ecology, that may be of particular interest to students of Human Behavior and Conservation Biology.***

Behavioral Ecology will embrace a new sub-field, Behavioral MacroEcology, that will, in part, investigate ecosystem, regional, and global patterns of diversity in Behavioral Ecological factors and traits (including Sociobiological* factors and traits) and that may require assembly of large databases (as per a new sub-field, Computational Behavioral Ecology).

Behavioral Ecology will be an active special interest group of ESA****.

*Group-formation, Group-maintenance, Group-living, Intraspecific/Interspecific interactions, Cooperative and/or Altruistic traits, Facilitation, and Co-existence. Intraindividual traits ["behavioral syndromes"] will be studied as they may influence group and/or population effects.

**"...tending, in the course of generations, to modify organic structures in accordance with external circumstances, as food, the nature of the habitat, and the meteoric agencies...." Charles Darwin, Origin of Species, 1861 (3rd Edition)

***See, for example, Palkovacs EP, Moritsch MM, Contolini GM, Pelletier F (2018) Ecology of harvest-driven trait changes and implications for ecosystem management. Frontiers in Ecology and the Environment, 16(1): 20-28, doi: 10.1002/fee.1743

****An organism's use of energy (E) is the essence of Behavioral ECOLOGY [1st Principles of Ecology= Acquisition, Consumption, Allocation (e.g., to Behavior]. Similarly, a group-living organism's use of energy (E) is the essence of Social Biology [a sub-field of Behavioral ECOLOGY]. All Behaviors [action patterns, motor patterns] are a function of the laws of thermodynamics.

Primary Citation: John Hurrel Crook, Behaviour. Supplement No. 10, The Evolution of Social Organisation and Visual Communication in the Weaver Birds (Ploceine) (1964)

Are Humans Cooperative Breeders? [2]

Are humans cooperative breeders? [2]

"Given the high diversity of ecological and social environments within which humans are currently found, specific human groups, populations, or cultures, rather than the human species as a whole, should be considered as units for comparison, analysis, and discussion." (Crespi 2014; also see Jones 2011)

Based upon Sarah Blaffer Hrdy's influence, it has become fashionable in the literature of Anthropology, to term [and, by implication, to classify] humans as "cooperative breeders". The purpose of this blogpost is to highlight Crespi's (2014) recent treatment of this topic and to provide, hopefully, constructive criticism of the topic's most recent discussion (Kramer & Russell 2015). In particular, I will make explicit certain untested assumptions underlying the classification as well as an apparent misunderstanding and/or mis-application of Hamilton's rule. This diagnosis is not intended to be exhaustive; however, it is my opinion that the concerns raised here provide a framework for necessary research to be conducted before the variety and range of conformations of human reproductive units can be described, including, traits associated with different conformations within and between populations.

In particular, I wish to stress that allomothering/alloparenting may be imposed by exploitation in the form of, for instance, coercion or social parasitism by reproductives [mothers] upon Donors [Actors, "helpers", "allomothers"] and that, rather than benefiting reproductive interests of Recipients [mothers] AND Donors ["helpers", "alloparents", "allomothers"], as per Cooperation, apparent "cooperation" may be deleterious to Donors/"allomothers" who may exhibit Altruism and/or apparent "cooperation" may be induced/imposed by Selfish behaviors on the part of Recipients [mothers], e.g., via coercion, including, in some cases, manipulation via financial reward.

Kramer & Russell (2015) recently addressed monogamy as a possible precursor to "cooperative breeding" in humans, explicating, clearly, and without qualification, the current, accepted argument[s] in Anthropology for classifying, without qualification, the human species as "cooperative breeders" [see epigram to this blogpost]. The following issues of concern pertain to the aforementioned paper and to Hrdy's earlier claims (Hrdy 1999, 2005, 2009 cited in Crespi 2014 and Kramer & Russell 2015 [subsequently, K&R]).

1. K&R define "social [sic] breeding systems" as "cooperative breeders" exhibiting "two key characteristics" [pg 74]: (1) "those individuals who provide care to the offspring of others [and are] currently non-breeders" and (2) "helping behaviors may be directed to another's offspring or toward mothers or other caretakers, which facilitates increased investment in offspring now or in the future." There is little consensus among researchers about what characterizes "cooperative breeding"; thus, the criteria advanced by K&R require comparative analysis and may be considered tentative hypotheses. An obvious problem confronting the aforementioned two criteria is that many species not classified as "cooperative breeders" meet the two conditions [e.g., grooming or "allomothering" in many primate and social carnivore taxa such as macaques, baboons, wild dogs, and hyenas or numerous "communal" rodents].

2. Throughout K&R, and much [all?] of the literature on [apparent, ostensible] "cooperation" in the Anthropology literature, apparent, ostensible "cooperation" is assumed to be a positive interaction [facilitation]. This is a non-trivial, untested and implicit assumption. "Helping" cum "cooperation" requires measurement or estimation of differential [reproductive] costs and benefits associated with "allomothering" behaviors [traits] before we can label these interactions "cooperative". Further, Cooperation may be imposed upon Donors ["helpers", "allomothers"] by Recipients [mothers] by a wide range of exploitation and manipulation [negative interactions], such as coercion, force, persuasion, social parasitism, shaping, and/or financial reward.

3. Related to #2 above, K&R, and other Anthropologists, appear to assume that benefiting kin is generally, if not always, advantageous to Donor [mother]. This assumption suggests a fundamental misunderstanding of Hamilton's Rule which does not state that benefits to [potential] Donor will always be + since costs of doing so may be prohibitively high. This fundamental error highlights the importance of measuring or estimating differential costs and benefits to Donor ["allomother"] and Recipient [mother] alike.

On the other hand, induced or imposed [potential] costs to [potential] Donor may make it beneficial for Donor to "help" where it is not possible for [potential] Donor to escape costs, such as, where response[s] to [potential] Recipient [mother or other self-interested group member"/imposer" of costs] are not under [potential] Donor's ["allomother's"] control. [Reproductive] costs from NOT "helping"/"allomothering" may be prohibitively high for [potential] Donor if s/he can not escape control by [potential] Recipient [mother].

4. After K&R, "Humans live in multi-male, multi-female groups that include multiple breeding females who often reside within socially recognized, long-term monandrous unions." [p 75]. These conditions may characterize some other mammalian groups, populations, or taxa, a topic that remains unstudied systematically.

5. Two additional criteria that have not, to my knowledge, been addressed by Anthropologists relative to their classification of humans as "cooperative breeders" is the extent to which the moniker requires high "skew" in groups and totipotent "helpers" [non-reproductive "allomothers" who may revert, in a condition-dependent, situation-dependent manner, to reproductives]. Should the latter trait be agreed upon as a diagnostic trait for classification of a group or population to be considered "cooperatively-breeding", menopausal females [often assumed to be grandmothers] and pre-reproductive offspring would not be considered diagnostic for said classification. See, for example, literature on "cooperatively-breeding" callitrichids for diagnostic use of high "skew" and totipotency, in addition, to employment of these terms in literature on Social Insects (see, also, Crespi 2014).

6. I am not aware of any quantitative [especially, modelling or theoretical] treatments of the above topics nor of experiments [lab, naturalistic, agent-based modeling] on the aforementioned matters in the Anthrropology literature.

7. I intend for the previous comments to convince the reader of the need for additional research within and between human groups in order to systematically evaluate the range and varieties of reproductive units in humans, a species with noteworthy facultative behavioral responses (see Jones 2011 and "Are humans cooperative breeders? [1], this Blog, as well as epigram above from Crespi 2014; also see Crespi 2009).

8. Following the epigram to this blogpost and related literature cited, it is clear that it is ill-advised to label humans as a species characterized by a single "breeding" system. Furthermore, without further evidence, it is not clear that "cooperative breeding" is the norm in most human populations over time and space. Related to this, there is no consensus among mainstream researchers concerning diagnostic criteria for a cooperatively-breeding group or population.

9. In accord with the call for additional research, there is a need to evaluate the literature on deleterious effects of "allo"-carers upon dependent human young. For example, male caretakers may injure or kill young at alarming rates in human groups and populations. Presumed benefits of grandmothers may be exaggerated and their deleterious effects may be minimized.

10. In 1999 (Jones & McJetters 1999), I wrote the following as a Note [p 123, #s 2 & 3] in a paper about women on death row. The statement suggests ways to quantitatively address the Socioecology of Co[-]operative Breeding in Humans. The significance of the Notes is unrelated to the topic of the paper [women on death row]. One significance of the Notes is to highlight Crespi's (2014; Jones 2011) statement  above [epigram to this blogpost]...human mating systems are expected to vary, and the causes and consequences of such variation may differ as a function of, say, SES [class], race, etc. Another point of interest [Note 3] is expected to be the dynamics of interactions between mothers and potential or actual mates.

Note 2: "We believe that Emlen's (1995) ecological model of the family will facilitate the analysis of behavioral differences between Black and White women, including differences in violent behavior. Emlen's ecological model predicts cooperative family organization (e.g., extended families) in regimes where independent breeding is uncertain, as is the case for many Black females. Where independent breeding is more reliable, as it may be for most White women, individualistic strategies are predicted. Ray & Smith (1991, p 150) also note the importance of ecological models for analyzing the consequences of differential access to resources in time and space for homicidal behavior. Ecological models are potentially of value to the student of violence because they are sensitive to interindividual competition for limiting resources."

Note 3: "A complicating factor, however, is that the profile of Black female homicide offenders is expected to be tied to that of Black males, since ecological theory predicts that female behavior will be a function of their resource base, including potential investment by males in herself, her childre, and her kin. Since the social and economic status of Black males is expected to be lower on average than that of White males, the causes and consequences of criminal behavior by Black and White women may continue to differ significantly."

Crespi (2009)

http://www.sfu.ca/biology/faculty/crespi/pdfs/122-Crespi2009SkewTheory.pdf

Crespi (2014)

http://www.sfu.ca/biology/faculty/crespi/pdfs/168-Crespi2013HumanNature.pdf

Emlen (1978)

Emlen ST (1978) The evolution of cooperative breeding in birds. In: Krebs JR, Davies NB, Behavioural ecology: an evolutionary approach. Blackwell, UK.

Hrdy (1976)

https://books.google.com/books?hl=en&lr=&id=x_hfmoOaHpoC&oi=fnd&pg=PA101&dq=sb+hrdy++1976&ots=IrAlXVg1ae&sig=OZrcPETIUNcFezBBbxBVssldVR4#v=onepage&q&f=false

Jones (1986)

http://onlinelibrary.wiley.com/doi/10.1002/1098-2337(1986)12:3%3C167::AID-AB2480120303%3E3.0.CO;2-X/abstract

Jones (2011)

http://link.springer.com/article/10.1007/s10508-011-9741-5#page-1

Jones CB, McJetters Y. (1999) Gender, race, and homicide: a preliminary analysis. The Western J of Black Studies 23:2, 119-124.

Kramer & Russell (2015)

http://onlinelibrary.wiley.com/doi/10.1002/evan.21445/abstract

Solomon & French (1997)

Solomon NG, French JA (1997) Cooperative breeding in mammals. CUP, UK.

Are Humans Cooperative Breeders? [1...my 6 min video]

https://www.youtube.com/watch?v=Cvz3D3sKlJ8&feature=youtu.be

Terminology In Social Biology: Cooperation= Intraspecific...Mutualism= Interspecific?

My query to Trivers:

Currently, there is a discussion on Twitter regarding proper usage of the term, "mutualism". 

In my training, "mutualism" is reserved for Interspecific interactions, "cooperation" for Intraspecific interactions.

It seems that in the Ecology literature, "mutualism" is reserved for Interspecific interactions but that in the Behavior literature, "mutualism" and "cooperation" appear to be used interchangeably.

Please "weigh in" on this terminological issue, permitting me to quote your reply.

Thank you for your attention to this post.  

-----------------------------------------------------

Response from Robert Trivers:

yes i am used to your distinction but have paid no attention to the word
or its use for many years

mutualiism classically was between species, neither harm nor benefit
given, or when benefit not to the level of a symbiosis

Clara B. Jones Comments On Terminology In Social Biology (YouTube)

https://www.youtube.com/watch?v=o0cMyWzB33o&feature=youtu.be

Summary of Comments:
1. Terminology in Social Biology not standardized across taxa from social microbes to humans.
2. Want standardized terminology for comprehensive Science of Social Biology and for formulation of general models ("laws", principles, treatments, statements) of Social Biology.
3. Comments pertain primarily to empirical literature since theoretical treatments are usually clear about notation and assumptions.
4. Comments pertain primarily to vertebrate literature since social insect literature generally utilizes Hamilton's 4-way schema defining conspecific interactions based upon differential reproductive costs and benefits*.
5. Two primary concerns at this time. First, usage of word "social".
6. "Social" used in 4 ways in literature:
a. grouped or clustered Population Structure (Spatial Ecology or Population Genetics);
b. any interaction between/among conspecifics;
c. any "positive" interaction between/among conspecifics (e.g., helping, many non-damaging responses);
d. an interaction whereby an Actor facilitates the reproduction* of a conspecific Recipient; this is W.D. Hamilton's definition, the definition that I advocate.
[1] usage of "positive" interaction is problematic;
[2] most researchers assume "positive" interactions are induced by cooperation or altruism;
[3] Hamilton defined "cooperation" as an interaction in which both Actor & Recipient benefit reproductively, defining "altruism" as an interaction in which the Recipient benefits at the expense of the Actor's reproduction;
[4] in Hamilton's system, Cooperation & Altruism are the only forms of Social interaction among conspecifics;
[5] problematically, "positive" interactions may be induced not only by Cooperation or Altruism but, also, by Selfish responses which Hamilton defined as an interaction among conspecifics whereby an Actor benefits reproductively at the expense of a Recipient;
[6] thus, we cannot assume that "positive" responses are necessarily induced by Cooperation or Altruism as is generally assumed in the literature;
[7] researchers, then, must differentiate between "positive" interactions induced by Cooperation or Altruism and "positive" interactions induced by Selfish responses (e.g., by coercion, force, persuasion, or exploitation [e.g., manipulation, "social parasitism").

*Reproductive costs and benefits may be measured as, for example, differential offspring mortality, number, quality, inter-birth-interval. See Lehmann & Keller (2006, JEB).

Notes + lulu.com link to book, A mechanistic approach to studying ..., covering social parasitism in mammals. Clara B. Jones

Notes:: Terminology: Social parasitism in social insects [after Holldobler & Wilson, 1998]...

I. Types of Social Parasitism: Coexistence in the same nest of two species of social insects, one of which is parasitically dependent upon the other. The term can, also, be applied loosely to the relation between symphiles and their social insect hosts. Symbioses, Commensalism; no documented cases of Mutualism. [Symphilic: an amicably-accepted symbiont; Symbiont: An organism living in symbiosis with another--n.b. Not all researchers consider Social Parasitism to be, Symbiosis].

1. Inquilinism: The relation in which a socially parasitic species spends the entire life cycle in the nests of its host species. Workers are either lacking or if present, scarce and degenerate in behavior. This condition sometimes referred to as "permanent parasitism"
2. Dulosis: The relation in which workers of a parasitic [dulotic or slave-making] ant species raid the nests of another species, capture brood [usually pupae], and rear them as enslaved nestmates.
3.Xenobiosis: The relation in which colonies of one species live in the nests of another species and move freely among the hosts, obtaining food from them by regurgitation or other means but still keeping their brood separate.
4.Parabiosis: The utilization of the same nest and sometimes the same odor trails by colonies of different species which nevertheless keep their broods separate.
5. Cleptobiosis: The relation in which one species robs the food stores or scavenges in the refuse piles of another species but does not nest in close association with it.
6. Lestobiosis: The relation in which colonies of a small species nest in the walls of the nests of a larger species and enter the chambers of the larger species to prey on brood or to rob the food stores.
7. Plesiobiosis: The close proximity of two or more nests, accompanied by little or no direct communiccation between the colonies inhabiting them.

n.b. "intermorph;" compound nests; mixed colonies

II. Social Parasitism most likely to be observed during population expansion or colony foundation. [in mammals, during dispersal?, settlement?]

III. Evolutionary factors: Disruptive Selection; Emery's Rule [parasite & host closely related or similar in other ways--in social insects, many exceptions to this rule observed]; Allopatric or Sympatric speciation [rapid rate of speciation]; Competition: [-, - interactions between species (or individuals?)]

IV. Do the principles observed in social insects apply across scale [i.e., to the individual level] and/or to other taxa, including, humans?


-----------------------------------------------------------------------------------------------------------


https://www.lulu.com/shop/clara-b-jones/a-mechanistic-approach-to-studying-mammalian-populations/paperback/product-8dnw7q.html?q=clara+b.+jones&page=1&pageSize=4

Schematic Note On Mammalian Signaling

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...

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)