tag:blogger.com,1999:blog-46045567908620465882024-02-22T08:11:42.986-08:00Science (Clara B. Jones, Ph.D.)All aspects of thermal [tolerance] evolvability in group-living mammals and other group-living taxa (see Profile). For social taxa, I am primarily interested in differential reaction norms as they relate to relative reproductive success of individuals, and as RRS across individuals relates to shifting mean fitness of populations. I am interested in thermosensory responses, including, gene expression, protein regulation, & feedback [from molecular to phenotype levels].Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comBlogger51125tag:blogger.com,1999:blog-4604556790862046588.post-47478040149219810122022-11-15T07:42:00.004-08:002022-12-05T11:35:11.042-08:00MAMMAL SOCIAL EVOLUTION: MAJOR TRANSITIONS APPROACH--PROPOSED SCHEMA AND NOTES (order hard copy at lulu.com)<p><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">MAMMAL SOCIAL
EVOLUTION: MAJOR TRANSITIONS APPROACH—PROPOSED SCHEMA AND NOTES</span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></b></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Cite </span></b><span style="color: #0f1419; font-family: "Times New Roman", serif; font-size: 12pt;">[APA style]: Jones, C.B. (October, 2022).
Mammal Social Evolution: Major Transitions Approach—Proposed Schema and Notes. <i>Science
</i>Blog [vertebratesocialbehavior.blogspot.com]; lulu.com.</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Clara B. Jones<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Silver Spring, MD
20910, USA<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Cell: 828-279-4429<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Email: </span><a href="mailto:foucault03@gmail.com"><span style="font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">foucault03@gmail.com</span></a><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">; </span><a href="mailto:mapcbj@gmail.com"><span style="font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">mapcbj@gmail.com</span></a></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">“The major transitions approach provides a
conceptual framework that facilitates comparison across pivotal moments in the
history of life. It suggests that the same problem arises at each transition:
How are the potentially selfish interests of individuals overcome to form
mutually dependent cooperative groups? We can then ask whether there are any
similarities across transitions in the answers to this problem. Consequently,
rather than looking for different explanations for the succession of different
taxonomic groups, we could potentially identify a few key factors that have
been important again and again at driving increases in organismal complexity.
This approach would both unify and simplify our understanding of the evolution
of life on Earth.” West et al. (2015)<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">“[Each] activity performed by an individual
can be thought of as incurring a certain probability of death and a certain
probability of successful reproduction.” McCleery (1978)<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">“The key to the sociobiology of mammals is
milk.” Wilson (1975)</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">TABLE OF CONTENTS<o:p></o:p></span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Abstract<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Key Words<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Foreword<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Preface<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Introduction<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Terminology<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Interindividual Interactions and Hamilton’s
Rule<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">SCHEMA: Proposed Transitions to Complex
Sociality in Mammals [Pre-social; Sub-social; Social (Cooperation); Complex
Social]<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Notes [Major Transitions; Cooperation;
Division-of-Labor; Humans]<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Conclusion<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Acknowledgments<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">References<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><br /></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></b></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></b></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b></b></p><div class="separator" style="clear: both; text-align: center;"><b><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhnPsxlMP70y1v0G62lrI9MnXaBNbZ1yA7b8szvJiNdtwSJKmr2UU0WCtL0semdhJKGHjO-INJkxUva0CGxO7GR9JgZy-WPMRX1FZGLfQ2BB-L2xtuhcN-PJakRsSIIqGB2BAA_FPH86KxtjGmdBy35eklIqbJmFyKQD4c_CgZ1EfCYjj-CftPA4_kqhw/s1598/Naked%20mole-rats%20queen%20and%20pups%20in%20nest.jpg" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1053" data-original-width="1598" height="211" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhnPsxlMP70y1v0G62lrI9MnXaBNbZ1yA7b8szvJiNdtwSJKmr2UU0WCtL0semdhJKGHjO-INJkxUva0CGxO7GR9JgZy-WPMRX1FZGLfQ2BB-L2xtuhcN-PJakRsSIIqGB2BAA_FPH86KxtjGmdBy35eklIqbJmFyKQD4c_CgZ1EfCYjj-CftPA4_kqhw/s320/Naked%20mole-rats%20queen%20and%20pups%20in%20nest.jpg" width="320" /></a></b></div><b><br /><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></b><p></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b><b style="background-color: transparent;"><span style="font-family: "Times New Roman",serif; font-size: 12pt; line-height: 17.12px;">Figure 1</span></b><span style="background-color: transparent; font-family: "Times New Roman", serif; font-size: 12pt; line-height: 17.12px;">. Female naked mole-rat (<i>Heterocephalus glaber</i>) Breeder with Helpers and offspring [cf. Bennett & Faulkes (2000) for introduction to African mole-rats, Bathyergidae]. Naked mole-rats are classified, “eusocial,” and are the only species of mammal known to exhibit morphological specialization [dispersal morph, breeder morph?], constituting a “caste” system, comparable to the social insects. Overlap of generations, cooperative breeding, and reproductive division-of-labor [differentiation into specialized Breeder – Helper tasks, roles, and/or morphology] characterize eusocial taxa, and the three species of social mole-rats meet these criteria [<i>H. glaber</i>; <i>Fukomys damarensis</i>, the Damaraland mole-rat; <i>Cryptomys hottentatus hottentatus</i>, the common mole-rat]. Questions regarding classification of social mole-rats and cooperatively-breeding mammals are in some dispute. Some researchers have attempted to demote social mole-rats to the status of Cooperative Breeders. Furthermore, it is not uncommon for researchers to designate species with communally-breeding females who may demonstrate “helping,” as Cooperative Breeders [e.g., jackals, warthogs]. However, though social mole-rats may not prove to be the only Eusocial mammals [e.g., see, Cape porcupines], necessary conditions for Complex Social classification are “reproductive division-of-labor” and a group structure including, in addition to “Helpers,” only one or a few female Breeders [“Queen(s);” more or less “pure” Breeder[s)]. See text herein for further treatments. ©Chris Faulkes</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><br /></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">MAMMAL SOCIAL EVOLUTION: MAJOR TRANSITIONS APPROACH—PROPOSED
SCHEMA AND NOTES </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">ABSTRACT<o:p></o:p></span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">With this document, I attempt to persuade
mammalian Social Biologists to adopt the Major Transitions Approach to Social
Evolution as the standard model. Herein, I define the latter approach, clarify
terminology and definitions, discuss a standardized way to address
Interindividual Interactions, and incorporate Hamilton’s Rule as the general
law or principle for the discipline. I propose a Schema for classifying
mammalian social evolution, highlighting mammalian traits expected to benefit or
disadvantage female mammals’ “fitness” [lifetime reproductive success] via
mechanisms to reduce maternal costs. Throughout, I highlight the role[s] that
abiotic and biotic factors play as regimes with the potential for transitioning
from one “grade” to another, and it is paramount to keep in mind that sociality
is not an ideal end in itself. Female mammals may reduce maternal costs via a
number of pathways, including, remaining solitary breeders. Gregariousness or
sociality are not necessary or sufficient to benefit female “fitness.” There
are many “routes” to “fitness” that may create barriers to, disadvantage, or
benefit living in groups. In some environmental regimes, for example, and for
some maternal conditions, selection may favor energy-saving reduction of
nursing bouts or shorter periods of offspring development creating
opportunities that might be used for the acquisition of food convertible to
present and future reproductive demands. Throughout the text, arrows designate
comments and notes addressing pertinent questions, concerns, as well as, topics
for future investigation.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">KEY WORDS</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">:
mammal social evolution; major transitions approach; interindividual
interactions; Hamilton’s Rule; Social Paleontology; Therapsids; solitary
breeding; <i>bauplan</i>; group-living; aggregations; Pre-social; Sub-social; Interdependence;
Cooperation; Complex Sociality; reproductive division-of-labor; totipotency;
specialization; Primitive Eusociality; social mole-rats; humans</span></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">FOREWORD<o:p></o:p></span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Herein, I propose Transitions in Mammals from
Solitary [independent] breeding to Pre-social to Sub-Social to Social
(Cooperation) and to Complex Social [Division-of Labor (Cooperation between
Specialists); Primitive Eusocial (in Mammals, no sterile “castes,” but,
“totipotency;” Eusociality requires at least two overlapping generations,
Cooperative Breeding, Reproductive Division-of-Labor (Breeder-Helper): Wilson
(1971); Task, Role, and/or Morphological Specialization [morphological
specialization in mammals, apparently, only found in naked mole-rats, <i>Heterocephalus
glaber</i>, Fig. 1<i> </i>(disperser morph; breeding female morph)]. Currently,
among Mammals, only the social mole-rats [African mole-rats: Bathyergidae] are
classified, Primitively Eusocial [see Bennett & Faulkes (2000) for
scientific names, classification, and other details; also, see Faulkes &
Bennett (2009)]. Primitively Eusocial mole-rats are “totipotent” [capable of
both Breeding, in addition to, “Helping”]. Primitively Eusocial mole-rats lack
“sterile castes” characterizing some social insects. Social insects with more
or less “sterile castes” are classified, Advanced Eusocial. Transitions are not
inevitable.</span></b><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
Energy</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> and <b>Matter</b> came from the Big Bang
billions and billions of years ago, and we just keep recycling [moving,
transitioning] them around. It is useful to view sociality through this lens. A
transition favored by selection in given abiotic [e.g., climate, nutrients] and
biotic [e.g., nutrient sources, interindividual contexts] regimes are expected
to create an energy-savings that can be used for acquisition, consumption, and
allocation of limiting resources to reproduction, enhancing, maternal “fitness.”
For reference to “work,” see Schmid-Hempel (1990), and think in terms of the 1<sup>st</sup>
Principles of Physics and 1<sup>st</sup> Principles of Ecology [acquisition;
consumption; allocation].</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #222222; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
Complex</span></b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">: consisting of many
different connecting parts, such as, individuals [organisms (see, “major
transitions approach:” Maynard Smith & Szathmáry 1995] interacting in a
group or population.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
Complexity: </span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">see announcement and
descriptions of 2021 Nobel Prize in Physics—complexity emerges from simple
rules [algorithms]; also, see Lex Fridman Podcast #234 with Stephen Wolfram via
YouTube.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
Essential Background Reading and Videos</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">:</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">West et al. [(2015)
provides a conceptual overview of the major transitions approach applied to social
evolution; also, see Bourke 2011; Cooper & West 2018; JF Eisenberg 1981. Refer
to EO Wilson's chapters on non-human mammals and humans in 1975 <i>Sociobiology</i>;
Cooper & West 2018; Toth et al. 2007. DeJong 1976 [allocation of energy by
females], as well as, Schmid-Hempel 1990 [“work”= Force x displacement (<i>W= Fs</i>)] are useful. Dugatkin
(1998) is an outdated discussion of Cooperation; however, his chapters on
mammals provide interesting case studies of behaviors that may classify as
“cooperation.” Downing et al. (2020), using a major transitions approach, is a
good place to begin for comparisons with cooperatively-breeding birds. For an
introductory but comprehensive theoretical treatment of “inclusive fitness,”
see, Marshall (2015). <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Additionally, I
strongly recommend, as introductions to social insect eusociality and for
comparative purposes, as well as, terminology, Wilson (1971) and Hölldobler
& Wilson (1990). The latter book, <i>Ants</i>, all of which are eusocial,
demonstrates the wide variability of group architectures classified as, “eusocial.”
Hölldobler & Wilson (2009) treats the distinction, “Breeder,” “Helper,” as
the first “grade” of “caste.” Based on studies with social insects, Robinson
(1992) suggests that inter-individual behavioral variation [phenotypic
variation] may be considered a type of “specialization” and “caste.” Note that,
in mammals, & other vertebrates, Breeders exhibit some helping, and Helpers
are “totipotent” [capable of both breeding and helping] due to which
observation Keller & Perrin (1995) recommended employing a “continuum”
approach (cf. Sherman et al. 1995)]. In some mammals, Helpers may breed
“opportunistically” [e.g., meerkats], though, often [usually?], mammal Helpers
do not breed when “helping.” You will find a wealth of references in my 2014 Springer
Brief [e.g., see therein, Table 3.1 on prehistoric mammals] and in my 2020 and
2021 lulu.com monographs [Jones (2020) is attached to the Profile of my Twitter
feed, @cbjones1943]. All of these sources highlight the variability of social complexity,
as well as, flexibility and/or plasticity of many traits [cf. Gene Robinson
1992; cf. Libbrecht FINE video-lecture (see below)]. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Four FINE Seminar
Series video-lectures on social insects are highly recommended: C. Penick; R.
Libbrecht; R. Gadaghar; K. Kapheim that are available on YouTube or on the FINE
website—also, see FINE Twitter feed. Penick makes some comparisons to Mammals. Though,
one does not expect perfect correspondence[s] when one takes a comparative
approach [whether within and between Genera, Families, or Orders of a Class or
between Classes], broad patterns and, even, some analogies [e.g., dominance
hierarchies: see Jones 2020; Penick, as well as, Gadagkar FINE videos] may be
discovered, especially, for different species in the same environmental
regime[s]. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Since 1980, I have
found the Social Insect literature particularly useful and edifying for
assessing comparative social evolution employing a major transitions approach.
For example, Christine Nalepa (2010) found that altricial cockroaches gave rise
to eusocial termites. The altricial state is primitive in mammals (as per
Eisenberg 1981). Students of mammal social evolution must keep in mind that the
precocial state may give rise to group-living taxa, also—a topic requiring
systematic investigation. For researchers seeking bird or insect models [lab or
field] with mammalian traits, polygynandrous acorn woodpeckers, [some] termites
and/or [some] wasps are recommended. For many potential models for mammalian
social evolution [e.g., Cape porcupines; jackals; hystricognaths], see
Macdonald (2009, 2001</span></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">PREFACE <o:p></o:p></span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Mammalian Paleontology: </span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">As of this writing, Class Mammalia comprises
>6,000 species and ~29 Orders. Mammals evolved in the Late Triassic ~210 mbp
from Therapsids [Cynodont Clade, sometimes called, "mammal-like reptiles”
(they are not dinosaurs)]. Lactation, endothermy
[thermoregulation], hair [e.g., thermoregulation], some other mammalian
diagnostic traits [e.g., typical jaw, teeth, skin], evolved in Therapsids, and
their potential association with the evolution of sociality in the Class
requires systematic investigation [see “What is a Mammal?” in Macdonald (2001)].
Eisenberg (1981) advanced the view that marsupials remain mammals’ sole
“control group,” and, because both Classes are endothermic, social evolution in
birds and mammals should be compared and contrasted. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Therapsids were extant
throughout the Permian and the Triassic, and humans are both Cynodonts and
Therapsids. The huge initial investments mammalian females make in producing
offspring, particularly, lactation, might predispose them to give and/or receive
“help” or, on the other hand, not to give or receive “help,” depending upon
whether a transition from solitary to gregarious living yields a greater share
of lifetime reproductive benefits. These alternate paths to “fitness,” relative
to phylogeny, maternal condition, and environmental regime, need directed study.
The “litany” is that most mammals are “sexually segregated” [“solitary”];
however, [sexual] selection is expected to act on males and females
differentially. For example, in a number of mammalian taxa, such as bats and
rodents, males are “solitary,” females, communal [or, “social?”—see below],
with varying degrees of Interdependence [below].<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In addition to the
litanies that most mammals are “sexually segregated” [“solitary,” “dispersed”]
and polygynous, physical traits—in particular, exterior phenotypes—are said to
be generalized, presumably, in response to evolution in extremely heterogeneous
regimes (see Eisenberg 1981). The prior conclusions are consistent with the
idea that in highly variable environments where patterns of limiting resource
availability cannot be detected and tracked, as well as, where any patterns
that may exist [e.g., seasonal availability of food] occur in periods greater
than a population’s “generation time,” generalized traits will be the optimal and
most energetically efficient strategy (see Levins 1968). These topics require
systematic investigation to assess their accuracy and their relevance to mammalian
social biology.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
I propose a new sub-discipline of Paleontology to be called, Sociopaleontology
or Social Paleontology and/or Comparative Sociopaleontology or Comparative
Social Paleontology</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">. Much work needs to
be systematically undertaken on mammalian Social Paleontology; however, a fair
amount of evidence is available on climate and potential food availability
during the Triassic. Unless I am mistaken, the earliest mammals were likely to
have been omnivores, herbivores, or insectivores, and the current hypothesis
about why they survived the major extinction event is that these small
creatures had very “fast” life-histories (cf. Funston et al. 2022; see Stearns
& Koella 1986). Mammalian Social Paleontology has been conducted at least since
1902; for recent publications, see Ladeveze, et al. (2011); Jaswoski &
Abdala (2017); Weaver et al. (2021). </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">INTRODUCTION</span></b><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">“Major Transitions”
refers to biological transitions from simple to complex, as from single cells
to multicellular organization to organism to superorganism (Maynard Smith &
Szathmáry 1995). Transitions are not inevitable. Transitions to mammalian Interdependence or
Social “grades” (“grade” after Wilson
1971), if they occur, embody females breeding alone to females breeding in a
group—with or without a breeding male—and polygyny is generally accepted to be
the most common population structure in mammals (see, Hex, Tombak, &
Rubenstein 2021; Wittenberger 1980) Often adult males’ territories embrace the
locales of more than one group of female breeder[s]. In other words, “polygyny”
does not necessarily imply that an adult male cohabits with a breeding female
or a group of female breeders [i.e., a “harem”]. Population structure will be a
function of abiotic and biotic environmental regimes, including, the dispersion
of limiting resources (Crook 1964), including, mates, breeding sites, and food
availability.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Transitions embody
increasing degrees of interindividual interdependence as we transition from Sub-social
to Complex Social “grades, should these condition-, situation-, context- [and,
density-?] dependent transitions occur [e.g., groups without Cooperation among
group-members may be favored by selection in given selective regimes (see
below)]. Transitions in mammalian sociality from Sub-social to Complex Social,
also, embody Breeder[s] becoming increasingly dependent upon other group
members. If complex sociality evolves, breeding females depend upon “Helpers”—a
process associated with reduction of maternal costs and increasing de-coupling
between Survival and Fecundity (“trade-off:” Stearns 1989) for the breeding
female[s]. Recall that no transition is inevitable. If, when, and under what abiotic
[e.g., climate, nutrients] and biotic [including, interindividual context] conditions
transitions evolve will, necessarily, depend upon relative gains in
"fitness" [“lifetime reproductive success”] of individuals transitioning
from one "grade" to another [e.g., from a female breeding in a
solitary state to a female breeding in a group, a <i>bauplan</i> in vertebrates].</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 11.5pt; mso-fareast-font-family: "Times New Roman";">--</span><span style="color: #0f1419; font-family: Wingdings; font-size: 11.5pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 11.5pt; mso-fareast-font-family: "Times New Roman";"> In order for the
study of Major Transitions to Social Evolution to be a scientific endeavor, it
is necessary for the enterprise to be grounded in 1<sup>st</sup> Principles of
Physics and in the 1<sup>st</sup> Principles of Ecology: acquisition, consumption,
& allocation [i.e., allocation of Energy into Survival and, essentially,
Reproduction (life-history strategies: Stearns 1992)]. Transitions from one
“grade” to another are not inevitable but depend upon the abiotic and biotic
[including, interindividual] patterning of environmental regimes. A</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">n environmental, Evolutionary Behavioral
Ecology approach entails use of John Hurrell Crook’s (1964) schema whereby
patterns of individual and interindividual responses, as well as, the
architecture of group structures, are functions of dispersion [distribution and
abundance] of limiting resources [e.g., food, mates, breeding sites] across
Time and Space. With the assistance of quantitative methods, including,
models, simulations, and experiments, a major transitions approach simplifies
the Behavioral Ecologist's and Evolutionary Biologist's search for patterns and
principles [“laws”] within and between Genera, Families, Orders, and Classes.
Recall that transitions are not inevitable and depend upon propitious abiotic
[e.g., climate, nutrients] and biotic [e.g., limiting food resources;
interindividual interactions] regimes favoring the most efficient condition-,
context-, situation- [and density-?] dependent mutations arising in a given population.</span><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Terminology:</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Herein, “Group” is
defined as a Reproductive Unit. Thus, I dispense with the Social Scientists'
dichotomy between “social system” and “mating system” because, in gregarious
animals, breeding takes place within a group. West-Eberhard, in her classic 1979
paper, suggested that “social” and “reproductive” can be thought of
interchangeably.</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Further, I propose
that we employ the term, “group-living” rather than “social organization” or
“social group” unless, or until, Cooperation [reproductive benefits accrue to
both Actor and Recipient (+, +), as per Hamilton 1964] arises in a given mammalian
population—thus, for example, “group-living” breeders, not “social” breeders.</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Additionally, if or
until Cooperation (“Social,” as per Hamilton 1964; see below) arises among
individuals in a group-living mammal population, I propose that we employ the
term, “gregarious” rather than refer to all interindividual interactions as “social.”
“Gregarious” is the term employed in ornithology for group-living birds.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Thinking, in
particular, of the Complex Social “grade” in the Schema presented below, only
females are discussed as “Breeders” because: females produce large and few
gametes [eggs] relative to males producing small and copious gametes [sperm],
an observation denoted “Bateman’s Rule” (Trivers 1972); thus, females, make a
larger initial investment in reproduction compared to males (Trivers 1972 [for
optimal female metabolic strategy, see Schoener (1971)] and, in the context of
the present work, may be more predisposed to receive “help” where it is
beneficial to “fitness;” population-level life table parameters are a function
of female traits because population growth depends upon female productivity;
and, females are the limiting resource determining male reproductive success
[“fitness”]. As noted above, the very high metabolic and other costs of
lactation may, in some regimes, predispose female mammals to reduce maternal
costs via gains from group-living [e.g., predator defense, more efficient
foraging: cf. Alexander 1974].<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">INTERINDIVIDUAL INTERACTIONS
AND HAMILTON’S RULE:<b> </b>Behaviors are outputs, also, traits, of an
individual—not only action or motor patterns, but, also, cortical, olfactory,
auditory, etc., outputs of an individual in response to stimuli. Neurobiological
responses are sometimes classified as “behaviors;” however, these are not
directly exposed to the abiotic or biotic environments, thus, cannot be acted
on directly by selection. Brain structures and mechanisms may be favored by
selection via selection on phenotypic traits that are genetically-correlated
and associated with a given brain structure, module, or mechanism—relative to
environmental regime. I hypothesize that the brain is rule-governed via some
inherent Hamiltonian algorithm, probably, a straightforward property or mechanism
influencing all brain and behavioral operations.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Hamilton's Rule [<i>rb</i> – <i>c</i> >0
---> <i>rb</i> > <i>c</i>] is a general principle [law,
formulation] where <i>r</i>= coefficient of relatedness; <i>b</i>=
benefits to Recipient of an act facilitating reproduction; <i>c</i>=
reproductive costs to Actor. An Actor’s “inclusive fitness” is comprised of
reproductive benefits accrued from assisting the reproduction of
offspring—“direct fitness,” and reproductive benefits accrued from assisting
the reproduction of non-offspring relatives—“indirect fitness.” Hamilton’s rule
expresses when, and when not, to favor kin. It is a common error in the Animal
Behavior literature to assume that Hamilton's Rule directs an individual to
always favor kin. It does not. Under conditions of intense local competition,
for example, “kin may be ego's worst enemy,” whereby it would be deleterious to
Actor’s “fitness” to favor the reproductive interests of a relative. <a name="_Hlk113742354">Critical references: for drivers of Social Evolution, see
Schmid-Hempel (1990)<i>;</i> for general Social Evolution, see Bourke
(2011); for recent theoretical treatment of “inclusive fitness theory,”
see Marshall (2015); for Major Transitions Approach applied to social evolution
(cf. West et al. 2015; Cooper & West 2018; Toth et al 2007). Eisenberg
(1981) is a necessary introduction to general mammalian patterns, including,
group structures and gregarious behaviors. Some of the older work (e.g., Wilson
1975, Eisenberg 1981) tends to be characterized by group-level thinking. <o:p></o:p></a></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Herein, Hamilton's Rule (Hamilton 1964) is
followed whereby use of the term, “social,” is limited to Cooperation [Actor gains
reproductively; Recipient gains reproductively: +, +] or Altruism [Actor bears
reproductive Costs; Recipient gains reproductively: –, +]. Thus, interactions
whereby the Recipient gains reproductively are considered, “social.” Altruism
is, generally, reserved with reference to social insects exhibiting Advanced
Eusociality (more or less sterile "castes;" see below); although, most
social scientists would claim that humans exhibit Altruism. Specialists still
debate how, under what conditions, and whether Altruism [as well as, Spite: -,
-] can evolve. Currently, many researchers hypothesize that Altruism [and
Spite] can evolve if the long-term reproductive benefits outweigh the
short-term reproductive costs. In practice, especially, in the field, it may be
difficult to differentiate between Selfish and Cooperative interactions,
emphasizing the need for researchers to investigate interindividual asymmetries
and to specify operational definitions and/or assays for each behavioral
category (cf. Lehmann & Keller 2006). Reciprocity (Trivers 1971) and other
mechanisms [e.g., “group” selection, “byproduct” effects] that have been put
forward to explain and account for the evolution of Cooperation and Altruism
are assumed, herein, to be a function, instead, of Hamilton’s Rule. As stated
above, I take Hamilton’s Rule as a general principle of social evolution;
though, consistent with Information Theory, any series of interactions will
exhibit some degree of “noise” and “error.”<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Importantly, Hamilton’s Rule emphasizes the
fundamental property of individual-level selection because each partner, Actor and
Recipient, are affected differentially [reproductive Benefits in the case of a
Recipient, reproductive Costs in the Actor’s case]. Selection acts differentially
upon each individual—Actor and Recipient—not on the interaction itself
[sometimes termed a “bond,” especially, in the Primatology literature].</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In 2011, Bourke
stated, “No level of ecological benefit can bring about altruism if relatedness
is not above zero.” Although this author is speaking of Hamiltonian “altruism”
[-, +] in this quote, it raises the question of whether or not there is some,
situation-dependent, threshold that must be met before Actor assists the reproduction
of a Recipient. Clearly, actors interact with other group-members, and some of
these interindividual interactions appear to be collaborative [e.g.,
coalitions, grooming, “allomothering” (see “Interdependence” below)]. In the
Animal Behavior literature, such interindividual interactions are usually
termed, “Cooperation.” But, are they? <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">This question
confronts mammalian social biologists at every turn; and, as alluded to above,
it may be very difficult to distinguish between selfish [+, -] and cooperative
[+, +] interactions in practice. Unless actors are subject to error [to a
degree deleterious to their own “fitness”]; unless Actor is pathological;
unless, Actor is the victim of self-deception, or unless Recipient is
perpetrating deception to which Actor is susceptible (see Trivers 2014, Otte
1975); or, unless, in point of fact, Actors simply do not assist the
reproduction of non-relatives, we need to address when and under what
conditions Actor will Cooperate with an unrelated Recipient. A satisfying
answer to this apparent conundrum is provided by West et al. (2002) who suggest
that Actor [or, Actor’s algorithm] takes into consideration [not necessarily in
a conscious or aware manner] the reproductive effects of her/his actions, not
only on Recipient, but, also, on all of those affected by Actor’s action. Thus,
for example, it may benefit Actor to assist the reproduction of an unrelated
Recipient if the act’s effects reduce competition for his/her kin in the group
or population. Related to the aforementioned, it is useful to recall Maynard
Smith & Szathmary’s (1995) caveat: “The evolution of sociality does not
depend only on relatedness: there must be something useful that individuals can
do for one another.”<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Conceptualizing Social
Evolution requires that interindividual interactions may be positive [+],
negative [-], or neutral [0] relative to future reproductive success
[“fitness”] and that, except for clones [in mammals, monozygotes (“identical
twins”)], each individual's interests differ; in other words, each individual
of an interacting dyad has different, condition-/situation-/context-
[density-?] dependent “fitness optima.” Ubiquitous “genetic conflict[s]” and
other asymmetries between individuals [e.g., body size; age; sex; rank;
matriline] determine an entity's “fitness optima”—relative to other
individuals, as well as, environmental regime.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Recall that,
consistent with Hamilton’s (1964) general treatments, “Social”= Cooperation or
Altruism [the latter, seemingly rare or absent in mammals, including, humans:
see West et al., 2015]. Thus, interindividual interactions are not “social”
unless they have been demonstrated to be Cooperative [+, +]. Interactions
deemed to be “Social,” necessarily presume an interindividual interaction in
which the Recipient gains reproductively [+]. Interpreting and quantifying all non-agonistic
interindividual interactions as “social” interactions, is a ubiquitous error
employed in the fields of Animal Behavior and in the Social Sciences. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">SCHEMA: PROPOSED TRANSITIONS
TO COMPLEX SOCIALITY IN MAMMALS [Pre-social ---> Sub-social → Social (Cooperation)
→ Complex Social] <a name="_Hlk114011558"><o:p></o:p></a></span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Logic: Female Mammals bear a costly, initial
investment in reproduction, primarily due to lactation. Mechanisms and
processes favoring reduction of maternal costs may benefit mammal females in
some environmental regimes. Sociality is not the only route to “fitness” whereby
female mammals can reduce maternal costs. For example, selection may favor
females who “park” their young or who reduce the number of nursing bouts or who
rest more, presumably, freeing mothers to forage for nutrients convertible to present
and/or future maternal effort. “Mixed” cost-reducing strategies may, of course,
be selected. In all cases, transitions from one mode of breeding to another
will be a function of “fitness” benefits to females, as well as, intensity of
selection pressures under changing abiotic and biotic conditions. Transitions
are not inevitable. Whenever one notes a female trait, the observer should ask
whether the trait has the potential to or might be a pre-adaptation for
reduction of maternal costs. The present schema pertains to mammalian
populations and species in which transitions to social evolution may occur—may
have the potential to benefit female “fitness.” It is important to note that
most mammals are “sexually-segregated,” though breeding females may form groups
of interacting individuals, with or without Interdependence, a necessary
precursor for the evolution of Cooperation, the first “grade” of sociality (cf.
West et al. 2015, 2021; see below). Even where sociality is beneficial to
female mammals, the researcher must recall that there are, always, “tradeoffs”
in Survival and Fecundity. In a classic paper, Alexander (1974) discussed the
costs, as well as the benefits of sociality. Because specialized traits are
uncommon in Mammals compared to other Vertebrate Classes, and because
specialization is one feature of Complex Sociality, the Primitively Eusocial
naked mole-rats [<i>H. glaber</i>], exhibiting morphological specialization
like the Social Insects, stand out as the pinnacle of social evolution in
mammals.</span></b><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Pre-social “grade” 1</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">: Therapsid breeding independently [“solitary”
breeding] with maternal care (see Toth et al. 2007). Mothers provide all “work” associated with offspring
care before emancipation. Mother and offspring contact [e.g., “huddling” for
thermoregulation] may be a precursor to retention of some young in unit; the
very high energetic costs of lactation and nursing may set the stage for
selection to favor mechanisms to reduce maternal costs [e.g., precocial young] and
may have, but not necessarily, preadapted mammal breeders to conditions
favoring group-living and the benefits that may accrue that can be allocated to
Survival and Fecundity [e.g., increased offspring survival or quality; increased
predator protection; secure breeding sites; increased access to food or reduced
foraging costs]. Because of the high energetic costs of lactation, female
mammals might seem to be preadapted for social life; however, solitary breeding
may be the most adaptive mode (see Eisenberg 1981).<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Mammalian females
would seem to be pre-adapted for gregariousness and sociality because of the
high initial costs associated with gestation and, particularly, lactation and
nursing. But, keep in mind that relative costs and benefits to maternal
lifetime reproductive success [“fitness”] depend upon numerous factors
pertaining to abiotic and biotic environmental regimes, dispersion
[distribution and abundance in time and space] of limiting resources
[including, mates, breeding sites, food, etc.], as well as, local conditions
[e.g., competition and density-dependence]. Alexander (1974), in particular,
reminded us of the tradeoffs associated with social evolution.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Interindividual
interactions, including, mother-offspring interactions, are not deemed,
“social” or “cooperative” unless Hamilton’s criterion is met [+, +; also see,
Trivers 1972]. In this sense, mothers suffer reproductive costs from maternal
care and mother-offspring genetic conflict, explaining Trivers’ (1972)
proposition that mothers terminate maternal investment at some threshold level
of investment at which offspring have some minimum likelihood of survival until
reproduction at which “point” mothers redirect their allocation to preparing
for future reproductive events. In mammals, and other vertebrates, females may
resume breeding before a prior offspring or litter has reached emancipation.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Actors, should be
unwilling to cooperate with prospective unrelated Recipients unless
reproductive Benefits would accrue to Actor by way of kin [in the group or
population] affected by the Recipient (West et al. 2002). <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">As per Eisenberg
(1981), the altricial state is “primitive” [ancestral] in mammals. The roles of
lactation (cf. Pond 1977) and of altricial young [e.g., in mammals: marsupials;
most rodents; cats; dogs; primates, including, humans] and other ancestral
mammalian traits, for the evolution of Interdependence and Cooperation [sociality]
requires systematic investigation [e.g., fur (e.g., thermoregulation),
endothermy (e.g., thermoregulation)]. That gregariousness has been described in
species with precocial young [e.g., most ungulates], demonstrates that the
precocial state may be a route to sociality. </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Pre-social “grade” 2</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">:</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Aggregations</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">: Incipient group-formation that may transition
to group-maintenance (see Bourke 2011). Aggregations are unstable and impermanent
in Time and Space. Aggregations [clusters of conspecifics—as distinguished
from, “guilds”] may form around clumped, limiting resources [e.g., as typically
“solitary” deer in Spring may cluster around clumped food]. It seems an open
question whether or not Aggregations [or, opportunistic encounters] are a.
necessary precursor to group-maintenance and group-living (see Jones 2014,
Table 3.1). Based on the prehistoric evidence that I am aware of, aggregations
appear early in mammalian evolution which some researchers have interpreted as
evidence of "gregarious" or "social" groups. Weaver et al.
(2021) suggest that sociality is a "labile" trait in mammals. While “aggregations”
are not “groups” in the sense that they are not reproductive units [see above],
opportunities for mating, including, multiple mating, would, likely, present
themselves.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Sub-social “grade” 1</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">: As in Pre-social “grade” 1 except, females transition
from breeding independently [“solitary breeding”] to breeding in a group—without
Interdependence [below] (for critical references pertaining to this transition,
see Crook 1964 and Emlen 1982). In vertebrates, a transition from independent [“solitary”]
breeding to breeding in a group is a universal transition [a “<i>bauplan</i>”] if
group-living and breeding in groups evolves at all. Whether or not females can
skip this grade, going directly—evolutionarily—from, say, Pre-social “grade” 1 to
Sub-social “grade” 2 [Interdependence] is not clear to me. Systematic
comparative research is needed. Hystricognaths would be good candidate study models.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In Sub-social “grade” 1,
interindividual interactions within groups are, I suggest, undifferentiated but
interactions are expected to be non-random because of asymmetries between
individuals (e.g., by genotype; age; sex; body size). Asymmetries result in
Actor and Recipient having different “fitness optima,” with “genetic conflict”
being a necessary driver. As pointed out above, interindividual interactions
may be positive [+], negative [-], or neutral [0] with respect to Benefits or
Costs to either or both interacting individuals' future, inclusive,
reproductive success [“fitness”], including, mother → young interactions, adult
→ adult interactions, mother → offspring or non-offspring interactions, as well
as, offspring → offspring interactions with kin or non-kin.</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Thus, I suggest that, +,
+ interactions will be some sub-set of all interactions, but, interactions, whether
+, +; +, -; -, -; or, 0, 0 [+, -, or 0 with respect to reproductive benefits or
costs], will not be stable or structured in Time or Space [i.e., interindividual
interactions have not yet differentiated or become structured or iterated via
selection]. As noted above, transitions, if they occur, depend upon their being
favored in given, abiotic and biotic, environmental regimes and depend upon
energy-savings afforded to females representing gains in “fitness.” I propose
that, relative to abiotic and biotic conditions, selection may, for example,
favor cooperative [+, +] interindividual interactions, disfavor, selfish [+, -],
altruistic [-, +], or spiteful [-, -], or some combination of the possible
interactions. In some regimes, one of the other possibilities, or some
combination of their genes may be favored so that genes besides cooperative
ones may be retained in the population, functioning, possibly, according to
density- and/or frequency-dependent processes.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Sub-social “grade” 2</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> [<b>Interdependence]: </b>As in
Sub-social “grade” 1, except interindividual interactions are differentiated
[structured: e.g., evolution of grooming or “allomaternal” care (see Hrdy 2011,
Hawkes 2004)]. If a transition to this grade is favored, it is, like other
transitions, expected to reduce maternal costs. Interdependent interactions are
common in group-living mammal populations. Interdependence is recognized to be a
necessary precursor to the evolution of Cooperation [sociality], which is,
itself, the "gateway” to the </span><a href="https://twitter.com/hashtag/evolution?src=hashtag_click"><span style="color: black; font-family: "Times New Roman", serif; font-size: 12pt; text-decoration-line: none;">evolution</span></a><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> of Complex Sociality in Mammals and
other taxa (cf. West et al. 2021; also, see Cooper & West 2018). The FINE lecture by L. Ebensberger reporting
his results for South American Degus [related, as hystricognaths, to African
mole rats], seems to suggest that the transition from solitary breeding to breeding
in a group may be driven by assistance given to mothers by other females in
female-female but not female-male units. The paper by West et al. (2021)
highlights that the potential for and/or transition to Cooperation in mammals
evolves from monogamy or groups of female relatives (see, importantly,
Wittenberger 1980).</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Rather than view
differentiated interdependent interactions between an Actor and a Recipient as
necessarily positive for the “fitness” of each member of the dyad [i.e.,
Hamiltonian +, + interactions], it may be more accurate to hold that
differentiated, interdependent interactions may have some combination of
positive, negative, or neutral consequences for an Actor’s and a Recipient’s
“fitness.” In some environmental regimes, it is posited, +, + interactions will
be favored by selection, yielding a stable state in a group or population—relative
to environmental regime. I would speculate that genes with negative or neutral
outcomes relative to the “fitness” of individuals in differentiated,
interdependent interactions would be maintained with some frequency in the
given population since environments, including, interindividual contexts, are
expected to vary and change, implying that frequency-dependent mechanisms, in
addition to density-dependent ones, would be operating. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p><div class="separator" style="clear: both; text-align: center;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjQjVsXFi49icj_3XPrePTFLMGBDdb3zTVFSSEeg8YsOwivRRBFQ0GntX7LJyA-YkvBcG3p9XzMtej4GnL7sbScu9YKAUn60WzjPDuYm7Wy_OOKIkdxuhwkwBtrses2cZnESSM3HUWt8RXZB7gEktUQGnD-No9Br7s1HhSKOMxL6A1_vNXmnWnJcruGWw/s3072/a%20few%20families%20moving%20between%20camps.JPG" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="2048" data-original-width="3072" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjQjVsXFi49icj_3XPrePTFLMGBDdb3zTVFSSEeg8YsOwivRRBFQ0GntX7LJyA-YkvBcG3p9XzMtej4GnL7sbScu9YKAUn60WzjPDuYm7Wy_OOKIkdxuhwkwBtrses2cZnESSM3HUWt8RXZB7gEktUQGnD-No9Br7s1HhSKOMxL6A1_vNXmnWnJcruGWw/s320/a%20few%20families%20moving%20between%20camps.JPG" width="320" /></a></span></b></div><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></b><p></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><br /></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"></p><p class="MsoNormal" style="-webkit-text-stroke-width: 0px; color: black; font-family: "Times New Roman"; font-size: medium; font-style: normal; font-variant-caps: normal; font-variant-ligatures: normal; font-weight: 400; letter-spacing: normal; orphans: 2; text-align: left; text-decoration-color: initial; text-decoration-style: initial; text-decoration-thickness: initial; text-indent: 0px; text-transform: none; white-space: normal; widows: 2; word-spacing: 0px;"><span style="font-family: "Times New Roman", serif; font-size: 12pt; line-height: 17.12px;"><o:p></o:p></span></p><p></p><p class="MsoNormal" style="-webkit-text-stroke-width: 0px; background: white; color: black; font-family: "Times New Roman"; font-size: medium; font-style: normal; font-variant-caps: normal; font-variant-ligatures: normal; font-weight: 400; letter-spacing: normal; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto; orphans: 2; text-align: left; text-decoration-color: initial; text-decoration-style: initial; text-decoration-thickness: initial; text-indent: 0px; text-transform: none; white-space: normal; widows: 2; word-spacing: 0px;"><b style="background-color: transparent;"><span style="font-family: "Times New Roman", serif; font-size: 12pt; line-height: 17.12px;">Figure 2</span></b><span style="background-color: transparent; font-family: "Times New Roman", serif; font-size: 12pt; line-height: 17.12px;">. A moving unit of Hadza hunter-gatherers in Tanzania, part of a larger “band.” Mothers and their offspring are accompanied by other related and unrelated group-members who might serve as “allomothers” [caregivers, including, siblings, other than the mother]. “Allomothering” or “alloparenting” may take many forms, such as, “babysitting,” “grandmothering,” play, or transport. A complicating factor in <i>Homo sapiens</i> is that caregivers may be paid for their services, an economic transaction that may have different causes and consequences than “allomothering” in non-human animals. ©Brian Wood</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Social “grade”</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> <b>[Cooperation: +, + interactions]</b></span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #222222; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In some environmental regimes, Hamiltonian sociality [Cooperation:
+, +] may be favored if beneficial to breeders’ reproduction [if energy-savings
can be converted to offspring]. Recall that transitions are not inevitable. </span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Interdependence is a necessary precursor to
the evolution of Cooperation (West et al. 2015). Cooperation is the “gateway”
to Complex Sociality [below]. My tentative speculation for the evolutionary
transition from Interdependence to Social [Cooperation] is that, during Sub-social
“grade” 2, Interdependence, selection acts on +, + interindividual interactions
with greater selection intensity in some environmental regimes [e.g., following
models in Behavioral Ecology, a change in climate might effect greater
variability in rainfall leading to clumping or greater ephemerality of limiting
resources favoring group-formation and consequent interindividual interactions,
possibly, followed by group maintenance, possibly followed by interindividual Interdependence,
possibly, transitioning to Cooperation where conditions and selection favor +, + interactions]. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">These
mutually-beneficial [+, +] interactions [“relationships”] may become more
frequent [more structured; more predictable] in certain environmental
conditions [“environmental potential” for Cooperation] compared to selfish or
neutral interactions which may be favored in other [abiotic and/or biotic] regimes.
This "rugged" fitness incline must have been difficult to climb or
not advantageous to female mammals since there seems to be a consensus that
most members of the Class are “sexually segregated” (“solitary”). However, in
some regimes inhabited by mammals, communal or cooperative interactions,
particularly among females, have proven to be a stable and more energetically
efficient strategy than solitary breeding or non-cooperative interactions The
question of how common Cooperation [+, +] may be among mammal species,
including, humans, is unclear [see below]. Cooperative interactions and
genotypes may coexist with non-cooperative interactions and genotypes in the
same group or population.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">If transitions to Cooperation
[+, +] occur, it must be because selection has favored a mutation correlated
with a trait or traits [e.g., via pleiotropy] leading to this “grade,”
increasing efficiency, driven by energy-savings [“Selection always increases
efficiency.” DeJong 1976; for optimal female metabolic strategy, see Schoener
1971]. The present treatment assumes that a population must include C(ooperation)C,
CS(elfish), and SS genotypes. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In this grade, resulting
from interindividual asymmetries, some females find it advantageous to
concentrate upon breeding, some, on “helping.”
In mammals, Breeders may “help,” and “Helpers” retain the capacity to
breed [“totipotent” condition]. Where this differentiation arises, females may
be pre-adapted for Complex Sociality—Reproductive Division-of-Labor and Specialization.
Such interindividual asymmetries, then, may set the stage for Cooperation
between Specialists [Division-of-Labor between more or less exclusive Breeders
and more or less exclusive “Helpers” in the present case]. While this phenomenon
of incipient differentiation needs systematic investigation, one can detect
these features in “allomothering;” “grand-mothering;” and dominance hierarchies
whereby different individuals appear to be taking on different roles.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Complex Social
“grade:”</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> In some environmental
regimes [“clumped,” limiting food sources; intense local competition?; “poor”
habitats?; moderate to high environmental heterogeneity, such as, ephemeral
food?; seasonal environments?], sociality as per Cooperation [among members of
a group or population] may transition to Complex Sociality: Reproductive
Division-of-Labor [Cooperation among specialists—more or less exclusive Breeders,
more or less exclusive “Helpers”], which may lead, in Mammals, to <b>Primitive
Eusociality</b> [task, role, and/or morphological (naked mole-rats)
Specialization]. <b>Advanced Eusociality</b>, more or less sterile “castes,” is
absent in Mammals. According to Macdonald (2001), some naked mole-rat “helpers”
never have an opportunity to breed throughout their lifetimes.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Received wisdom is
that mammals are characterized by generalized traits (cf. Eisenberg 1981), a
condition that should have placed significant limits on mammals’ potential for
“specialization,” and, Complex Sociality, to evolve. In light of the foregoing,
naked mole-rats must be viewed as a remarkable, possibly, unique, example of Eusociality
in the Class, having evolved morphological specialization. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Again, in mammals, the
transitions, should they occur, entail transitions from Solitary Breeding by
Females --</span><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> Females breeding in a group without
Interdependence --</span><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> Females breeding in a group with
Interdependence --</span><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> Females breeding in a group with Cooperation
--</span><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> Complex Sociality [Reproductive Division-of-Labor; Specialization].
In mammals, reproductive division-of-labor is found among Cooperative Breeders,
as well as, the social mole-rats [see below]. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Naked mole-rats [<i>H.
glaber</i>] exhibit size “polyethism,” “temporal” division of labor [age
“polyethism”], reproductive division-of-labor, as well as, morphological
specialization [disperser morph; female breeder morph]. Damaraland mole-rats [<i>F.
damarensis</i>] exhibit “temporal” division-of-labor, as well as, reproductive
division-of-labor [for temporal division-of-labor in primates, see Jones 2020
and Hrdy & Hrdy 1976]. Social mole-rats [Bathyergidae] are, currently, the
only mammals classified, “eusocial” [for “cooperatively-breeding” mammals, see
below], and naked mole-rats are the only mammals recognized to exhibit
morphological “castes.” As pointed out previously, Breeding and Helping roles and
tasks in mammals are not obligatory in that Breeders usually do some work and
Helpers may breed in various proportions, lending support to the view that
Breeding and Helping may be conceptualized along a continuum (cf. Keller &
Perrin 1995; for an earlier treatment on the continuum concept, see ,Sherman et
al. 1995). Robinson (1992) suggested that interindividual variability [resulting
from differential intraindividual "personality" or "behavioral
syndromes"] may be viewed as primitive “castes,” and Hölldobler &
Wilson (2009) pointed out that the initial differentiation into Breede -Helper
roles represents an early “caste” stage.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In humans and some
other mammals exhibiting post-reproductively sterile females (see Cant et al.
2009), more or less, sterile “castes” [for a discussion of post-reproductive
sterility in humans, see Foster & Ratnieks 2005] have not been classified, “eusocial,”
because unambiguous, specialized, and stable reproductive division-of-labor [Breeder
and Helper roles and tasks] have not been identified—even if the species meet
all other criteria for eusocial status [overlap of generations and cooperative
breeding (for humans, see Crespi 2014)]. </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";">O</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">n the other hand, if
Breeding and Helping are considered as occurring along a continuum from,
proportionately, less to more as per Keller & Perrin (1995), this
diagnostic trait classification may need revision. Nonetheless, currently, cooperative
breeders and social mole-rats are the only mammalian taxa known to exhibit reproductive
division-of-labor, and, as a consequence, the only mammalian taxa exhibiting “complex"
social traits, as per a major transitions approach. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">On related matters, Jones
(2014) suggested that cooperatively-breeding mammals might be considered, eusocial,
with the social mole-rats, based on the criteria that both exhibit reproductive
division-of-labor, combined with, overlap of generations, as well as,
cooperative breeding. On the other hand, Clutton-Brock (2021, 2016) and his
colleagues, apparently, classify social mole-rats, cooperative breeders,
though, Markus Zöttl has informed me that the two social systems are considered
synonymous by some social biologists. Following the aforementioned treatments, Crespi
(2014) concluded that, while humans exhibit a number of “insectan” traits, the
species does not meet the criterion for exhibiting reproductive division-of-labor
(see sections on humans in Jones 2020, 2021; also, see Foster & Ratnieks
2005). </span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Superorganism “grade:” </span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">This “grade,” characterized by social insect
nests and mounds, may be absent in mammals. Unless I am mistaken, some
Anthropologists, students of “gene-culture co-evolution,” for example, claim
that “complex” human societies [e.g., organizations; nation-states] are “superorganisms,”
presuming that group-level selection is operating (for treatments of the topic
of “superorganism,” see Holldobler & Wilson 2009; for “superorganismality,”
see Boomsma & Gawne 2018).</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">NOTES</span></b><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Major Transitions:</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Evolutionary social theorists
studying mammals must model “Solitary,” Pre-social, Sub-social, Social
[Cooperation], and Complex Social “grades,” relying upon Hamilton's (1964)
formulations (see Marshall 2015 for an introductory mathematical treatment). Also,
it will be important for behavioral ecologists to quantify the relative
proportion of Selfish and Cooperative genotypes and phenotypes in given
populations of group-living mammalian [and, other group-living vertebrate] taxa
as well as, the abiotic [e.g., climate, nutrients] and biotic [including,
interindividual] regimes associated with group-living populations and species.
The possible “rule-governed” [lawful] nature of [demographic] ratios should be
evaluated (cf. Oster & Wilson 1978), as well.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Furthermore, genomic
studies can assess the degree of monomorphism or polymorphism for Selfish vs.
Cooperative phenotypes, as well as, what traits correlate with those genotypes
[see work of Gene E. Robinson's lab ("socio-genomics:" Robinson,
Grozinger, & Whitfield 2005)]. These studies will, also, identify genomic
“toolkits” and convergences (cf. Woodard et al. 2011).</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">A major component of a
scientist's job is to simplify; to build models and algorithms; to unpack
variation; as well as, to conduct laboratory and/or field experiments. The
Major Transitions Approach to Social Evolution provides a straightforward way
to proceed from independent [“solitary”] breeding with maternal care to Complex
Sociality; from Primitive [ancestral] to Derived; and, for group-living taxa,
from less to greater dependence of Breeders relying upon Helpers, including,
the process of uncoupling Survival and Fecundity during a breeder’s
life-history trajectory across transitions, should they be favored. In the Social
[Cooperation] and Complex Social [reproductive division-of-labor;
specialization] “grades,” females are not expected to breed successfully or not
able to maximize “fitness optima” without assistance from conspecifics,
usually, offspring and other female relatives. Transitions driven by the Costs
[to reproduction] of maternal care, may lead, in some regimes, to the evolution
of a range of mechanisms to reduce those Costs [see Fig. 2]. The FINE videos recommended
above help us to conceptualize what variables might play a part in some females
becoming more or less obligate [“pure”] Breeders, some females becoming, more
or less obligate “Helpers.” In mammals and birds [CB Jones, unpublished],
species exhibiting reproductive division-of-labor [a type of morphological
specialization or “caste,” as per Hölldobler & Wilson 2009], all Breeders
and Helpers are more or less non-obligate—in various proportions. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">We have the necessary
data to test and to revise the proposed schema of Mammalian Social Evolution
using a Major Transitions Approach. Indeed, treatments of mammalian sociality
advanced by Wilson (1975), as well as, Eisenberg (1981), hint at "major
transitions” thinking, a method not unrelated to comparative phylogenetic
studies. There are many existing descriptive and natural history data sets, in
addition to, museum work, as well as, lab and field research, whose data sets
permit testing of assumptions, conduct of experiments, and development and
application of models and simulations to address my proposed schema and other
topics herein. Experiments can be conducted with simulations. As a first
approximation, the present treatment yields numerous questions that lend
themselves to scientific methodologies.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In his 2019
book, <i>Genesis</i>, EO Wilson proposed a set of six transitions that
differ from the series employed by West et al. (2015) and, herein. Thus, "</span><span style="background-attachment: initial; background-clip: initial; background-image: initial; background-origin: initial; background-position: initial; background-repeat: initial; background-size: initial; color: #0f1111; font-family: "Times New Roman", serif; font-size: 12pt;">The centerpiece of <i>Genesis</i> is
a listing and discussion of the six so-called ‘great transitions of evolution’….
These stages are: (1) the origin of life; (2) the invention of complex
(“eukaryotic”) cells; (3) the invention of sexual reproduction, leading to a
controlled system of DNA exchange and the multiplication of species; (4) the
origin of organisms composed of multiple cells; (5) the origin of societies;
and, finally, (6) the origin of language." (L.D. Wilson, amazon.com review
of <i>Genesis</i>). The inclusion of language as the pinnacle of social
complexity is consistent with a Social Sciences, <i>Scala Naturae</i> Approach,
to evolutionary transitions. It is important to point out that, by 2019, Wilson
had become a dedicated group selectionist, though, according to Robert L.
Trivers, [personal communication] this tendency is present in Wilson’s whole
oeuvre.</span><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Cooperation:</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><i><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></i><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">How common is
Cooperation [Hamilton’s Social “grade”] in Mammals? We need systematic research
on this topic, including, the development of assays, in order to differentiate
Interdependence from Cooperation. There are many candidate traits to consider
in the literature [e.g., see Wilson’s 1975 and Dugatkin’s 1997 chapters on mammals
and humans. Also, Eisenberg (1981) should be consulted. The anthropologist, LH
Morgan’s discussion of human social traits may be useful, as well as,
economist, Adam Smith’s writings]. The “litany” and “received wisdom” are that
most mammals are “sexually-segregated” or “solitary.” <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">I suspect that
systematic work will show that Hamiltonian Cooperation [+, +] is less common
among group-living mammals than Hamiltonian Selfish interactions, though,
“alliances” would seem to be a possible candidate for Cooperation [see below].
My chance observations of the seemingly infrequent dyadic displacements of a
third group member by adult male or adult female dyads of the polygynandrous
mantled howler monkey (<i>A. p. palliata</i>), suggested that these events were
opportunistic in nature (see, Jones 1980). Rather than Cooperation, coalitions
and alliances may be types of Interdependence, such as, coordination, collaboration,
or facilitation, deserving systematic investigation. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Whatever the case may
be, it might be expected that there are various types and degrees of
Interdependence that might function as precursors to the evolution of
Hamiltonian sociality [Cooperation]. Of course, like all action and motor
patterns, as well as, behaviors, responses are expected to be
situation-dependent, and their classification may be expected to vary depending
upon varying abiotic and biotic environmental regimes.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">It will be necessary
for students of Social Biology to determine whether coordination and/or collaboration
that is <b>imposed</b> by a third-party or his/her representatives or that is <b>imposed</b>
by one Actor in the case of division-of labor, [<b>imposed</b> via persuasion,
coercion, force, or manipulation (e.g., exploitation)], so common in humans, should
be termed "Cooperation” and considered, “Social.” Probably not. To date,
it appears that little, or no, systematic investigation has been made of this
issue which, at once pertains fundamentally to terminological, as well as, social
biological questions. Related to this question for the human case, should
non-human mammals in which collaboration is imposed by the Breeder’s olfactory
emissions be considered, “Cooperation?”<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Transitions from
Selfish [+, -] to Cooperative [+, +] relationships, then, may be only apparent.
Interacting individuals may collaborate for one purpose or in one way that
appear to be cooperative without being so. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Regarding the
aforementioned, and related, conundrums, an economy based on money exchange may
be a mechanism for enforcing Cooperation, increasing the likelihood that
recipients will cooperate, and/or decreasing the likelihood that recipients
will “cheat” [be “free-riders”]. Adam Smith (2003) stated, “It’s not from the
benevolence of the butcher, the brewer, or the baker that we expect our dinner,
but their regard to their own self-interest.” Smith’s idea may indicate that,
before an economic medium of exchange beyond barter emerged, the value of
resources exchanged might have led to a greater degree of inequality and
conflict than in a barter economy, highlighting the difficulty of evolving
Hamiltonian Cooperation [+, +]. Of course, an economy based upon monetary
transactions would not be expected to completely eliminate coercion, force,
exploitation, or “subterfuge.” A related quote by Crook (1971) is, also,
pertinent: “Social behaviour is a subterfuge.”</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Division-of-Labor:</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Division-of-labor is
defined as “cooperation between specialists” (Cooper & West 2018). There is
a large literature on “generalization” and “specialization,” and their evolution,
in the Ecology literature. Division-of-labor necessarily implies “complexity,”
in the sense that a whole task or other outcome is “broken down” into component
parts for which different individuals are responsible. In the social insect
literature, the “fitness” advantages to reproductive division-of-labor have
been speculated or shown to be: increased survival rates; increased
reproduction rates; increased “productivity;” more efficient developmental
timing; and/or more efficient co-ordination among breeders and “workers.”
Research on social insects has, also, shown that the benefits of reproductive division-of-labor
scale positively with group size [mean group size in naked mole-rats is
comparatively large: Bennett & Faulkes 2000]. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">I assume that specialization
must evolve before or concurrent with Cooperation, possibly, during an advanced
stage of Interdependence—precursor to the evolution of Cooperation [+, +], “gateway”
to the evolution of Complex Sociality [reproductive division-of-labor;
specialization]. There seems to be a consensus among mammalogists that bodily phenotypes
are usually generalized, though, body plan does not seem to predict, say, a
generalized diet [see, for example, many rodent species], and behavioral
flexibility, as well as, phenotypic plasticity, may compensate for or enhance a
generalized morphology. Environmental regime, in particular, moderate to extreme
heterogeneity [variability], relative to generation time, may favor generalized
body plans (see Jones 2009).<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">--</span></b><b><span style="font-family: Wingdings; font-size: 12pt;">à</span></b><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;"> </span></b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">“Helping” may be age-dependent [“temporal division-of-labor” (“age
polyethism”)]. Among Primates, adult female, as well as, adult male mantled
howler monkeys [<i>A. p. palliata</i>] exhibit “age polyethism” (Jones 1996,
2020), and, in the Asian Hanuman langurs, females are characterized by “age
polyethism.”<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #222222; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><i><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></i><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Do humans exhibit Complex Sociality? According to the Major
Transitions Approach, the criterion for the first stage of Complex Sociality—subsequent
to the evolution of Cooperation, should Cooperation evolve within groups—is reproductive
division-of-labor and, after Crespi (2014), the general consensus is that humans
do not exhibit reproductive division-of-labor; thus, by inference, humans have
not evolved, Complex Sociality, though, in future, application of the continuum
approach to the evolution of eusociality proposed by Keller & Perrin (1995)
may revise this conclusion. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">--</span></b><b><span style="font-family: Wingdings; font-size: 12pt;">à</span></b><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;"> </span></b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">Division-of-labor is conceptualized in the Social Sciences [e.g.,
Adam Smith: Economics; Lewis H. Morgan: Anthropology] to be highly
advantageous, indeed, necessary, to the appearance of “complex”
[multi-component, multi-level, modular] groups, including, organizations and nation-states.
In particular, Adam Smith’s (2003, 1776) many ideas and verbal formulations
about division-of-labor in humans can often be extrapolated to general discussions
about social evolution. In the same book, Smith suggested, for example, that division-of-labor
reduces average costs of output, increases quality of output, facilitates
large-scale production, and “reduces scarcity—the latter, in terms of
Population Ecology, possibly, meaning that division-of-labor reduces competition
among members of a group or population for limiting resources, freeing up
availability. Smith, as well, advanced the seemingly obvious idea that division-of-labor
facilitates teamwork and Cooperation [n.b., coalitions and alliances]. This
economist, known as, “the father of Economics” also, believed that division-of-labor
was linked to a higher standard of living and quality of life, possibly,
relevant for all female mammals. <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">--</span></b><b><span style="font-family: Wingdings; font-size: 12pt;">à</span></b><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;"> </span></b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">In an interesting and useful paper, Smith & Riehl (2022)
recently reviewed and assessed empirical studies of and additional evidence for
"division-of-labor," emphasizing animal taxa other than social
insects. These authors rely upon common usage in the Animal Behavior and Social
Sciences literature whereby all non-agonistic interindividual interactions are
termed, "social." As highlighted in a major transitions approach,
however, population structure may transition from one "grade" to
another, say, from "solitary" breeding to breeding in a group, if,
and only if, it benefits the reproduction of individuals in a population to do
so, presumably, in response to changing environmental regimes and
energy-savings sufficient to allocate to reproduction, moving closer to an
individual's "fitness optimum." Indeed, Smith & Riehl barely
give a nod to the ecological factors—causes, as well as, consequences—favoring
social evolution and its contingent transitions.</span><span style="font-family: "Times New Roman", serif; font-size: 13.5pt;"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><span style="font-family: "Times New Roman", serif; font-size: 12pt;">Furthermore,
transitions to group-living may, under propitious conditions, occur without the
evolution of Cooperation, in particular, as pointed out by West et al. (2015),
it must be demonstrated that Interdependence has evolved as a precursor to
Cooperation. The point is that transitions are not inevitable, and group-living
does not necessarily imply that Cooperation characterizes interindividual
interactions in groups. In mammals, for example, breeding females may cohabit
without cooperating—in the Hamiltonian sense as treated herein [+, +]. </span><span style="font-family: "Times New Roman", serif; font-size: 13.5pt;"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><span style="font-family: "Times New Roman", serif; font-size: 12pt;">Finally,
the "grade," Complex Sociality, entails the evolution of
"reproductive division-of-labor" and task, role, and/or morphological
specialization. By concentrating almost exclusively on "task"
specialization, these authors overlook the concept that the fundamental
"division"/specialization entails a division between Breeder and
Helper [in social insects, "worker"], not specialization by task per
se.</span><span style="font-family: "Times New Roman", serif; font-size: 13.5pt;"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Four<a name="_Hlk112955382"> FINE Seminar Series video-lectures [Social Insects: C.
Penick; R. Libbrecht; R. Gadaghar; K. Kapheim: available on YouTube or on the
FINE website]</a>, suggest that there are other ways, besides,
Cooperation, that may be precursors to reproductive division-of-labor [e.g.,
asymmetries in dietary profiles or physiology; dominance rank relations], as
well as, the researchers providing insights on what factors might lead some
females to become “pure” breeders, some females, workers [in mammals,
totipotent “helpers”—the differentiation being the first appearance of “caste,”
according to Hölldobler & Wilson (2009)]. If differential dominance rank is
shown to be a precursor to Cooperation in mammals as one possible feature of
the Interdependence “grade,” then, we might conclude that density-dependent
contest competition may factor in to the evolution of Cooperation. It seems to
me that differential dominance rank may be viewed as a type of rudimentary</span><span style="font-family: "Times New Roman", serif; font-size: 12pt;"> “caste” system, though systematic study is required
to evaluate the utility of this idea.</span><span style="font-family: "Times New Roman",serif; font-size: 12pt;"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">In my 2020 monograph
on [herbivorous] mantled howler monkey [<i>A. p. palliata</i>] female
life-history using a major transitions approach, a negatively age-graded
dominance hierarchy [younger individuals dominant to older individuals]
combined with differential access to limiting nutrient [food] resources [diet],
in particular, ephemeral food sources, instantiated “temporal division-of-labor”
[“age polyethism”], though, it is not clear to me how to treat “temporal
division-of-labor” in my schema proposed above. Neotropical howler monkeys [<i>Alouatta</i>],
do not exhibit reproductive division-of-labor. This topic, as well as, the possibility
that other mammals utilizing limiting, ephemeral, food resources [e.g.,the
Neotropical primates in the Family, Atelidae] or who are folivores [e.g.,
langurs among the Old World Primates (see Hrdy & Hrdy 1976 for “temporal
division-of-labor” in langurs)], deserve systematic investigation. To my
knowledge, social mole rats exhibit “temporal division-of-labor,” and age as a
main effect should be systematically investigated in all group-living mammals. </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p><p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><b>--> </b>It is important to keep in mind that "division-of-labor" is defined as "cooperation between specialists;" thus, a major impediment to the evolution of Complex Sociality in mammals, including, humans, is that, for the most part, mammalian phenotypes are generalized, as pointed out by Eisenberg (1981), making the cooperatively-breeding mammals and the social mole-rats, especially, <i>H. glaber</i>, all the more remarkable.</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Humans:</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">When considering human
evolution, it is, always, important to keep in mind “behavioral flexibility,” “phenotypic
plasticity,” and, variation writ large, in addition to, noise and error in the
sense of complexity and Information Theory.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Geographically, humans
and social insects are the most widely distributed terrestrial taxa, a
concordance that, alone, justifies comparing and contrasting mammals and social
insects using a major transitions approach. For example, both taxa are heralded
for their notable behavioral flexibility and phenotypic plasticity. As top
predators, humans dominate food chains; thus, they have less energy to extract
from available food resources compared to animals at other trophic levels. This
condition should lead to intense intraspecific, as well as, interspecific,
competition for limiting resources that might, in some regimes, favor group-living,
Interdependence, and, possibly, Cooperation. Intraspecific competition needs to
be considered as a likely context for the evolution of mammalian sociality,
including, density-dependent competition, frequency-dependent competition, and the
spectrum of possible causes and consequences of intense intraspecific
competition—pressures that are likely to have favored traits with the potential
to minimize the deleterious effects of competition for conspecifics’ “fitness”
[e.g., the evolution of dominance hierarchies; mechanisms to partition
resources; dispersal patterns; behavioral flexibility and phenotypic plasticity].
Mortality may, also, be a significant effect of intraspecific competition
influencing life-history strategies (see Stearns & Koella 1986).<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Humans’ unusual
life-history strategy combining a relatively long developmental period with a
relatively “fast” reproductive rate (see Stearns 1992 for “fast” and “slow”
life-histories) may, also, be related to competition for limiting resources, as
might be, the evolution of noteworthy behavioral flexibility and phenotypic
plasticity, the latter traits being ones that Macdonald (2001) considers
characteristic of the Class. In addition, Stearns & Koella (1986) highlight
the importance of studying mortality rates whenever life-history is
investigated. All human, and other mammalian species’, traits should be viewed
in the light of intraspecific competition and mortality, particularly, as they
may relate to evolutionary transitions to sociality, keeping in mind that
transitions depend upon the differential reproductive costs and benefits to
individuals in given environmental regimes and are not inevitable.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">According to Bernie
Crespi (2014; also, see Jones 2020, 2021), </span><a href="https://twitter.com/hashtag/Humans?src=hashtag_click"><span style="color: black; font-family: "Times New Roman", serif; font-size: 12pt; text-decoration-line: none;">humans</span></a><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> do not exhibit reproductive division-of-labor.
Some have claimed that the presence of post-reproductive “sterility” in human
females strongly suggests eusociality (Foster & Ratnieks 2005). I argue in
my 2020 and 2021 monographs that any gregarious groups that post-reproductive
females may be part of are not necessarily characterized by a Breeder-Helper
class/”caste” structure, nor are post-reproductive females in this species
obligated to perform “helping” roles or tasks at all. The wide variety of
composition of human reproductive units requires systematic investigation and
“unpacking.” A human “ethogram” would be of great assistance. Though the
potential combinations and recombinations of motor and action patterns exposed
to the environment, particularly, other individuals, would be impossible to
measure and classify, the motor and action patterns, themselves, should not be
numerically exhaustive to describe because of fundamental constraints within
and between human populations imposed by similar anatomical structures. When
attempting to unpack human “complexity,” perhaps cultural evolutionary
anthropologists should begin with the null hypothesis, Humans are <b>not</b>
“complex.”—contrary to the opposite assumption automatically assumed in the
Social Sciences. West et al. (2021) point out the importance of employing
“null” models.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Related to the topic
of human social evolution, broadly speaking, quite a few authors have concluded
that most human interactions are Selfish [Actor benefits reproductively;
Recipient, bears reproductive costs (as per Hamilton 1964: +, -)]. These papers
can be found in the earlier Social Psychology and Learning Theory [Behaviorism]
literature. Recently, two empirical papers by Burton-Chellew (e.g.,
Burton-Chellew & West 2013, Burton-Chellew & West 2021) reported that
humans demonstrate primarily “selfish” tendencies. Suggestively, West et al.
(2021) advance the opinion that Altruism will be found to be rare [or, absent?]
in humans.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #0f1419; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: "Times New Roman"; mso-bidi-font-family: "Times New Roman"; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: "Times New Roman"; mso-symbol-font-family: Wingdings;">à</span></b><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
</span></b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Kirschner &
Gerhart’s (1998) treatment of “Evolvability” provides support for the following
ideas. First, that the extreme behavioral flexibility and phenotypic plasticity
exhibited by humans, functions to enhance or maximize Evolvability [potential
for evolution], <a name="_Hlk113033800">on whole.</a> Extreme behavioral
flexibility and phenotypic plasticity, also, may function as, “tinkering,” and
Kirschner & Gerhart discuss the “exploratory” functions of evolvability in
the context of their core discussion about functions that are, at once,
flexible and robust. There is a large literature on “exploratory behavior” in
the Psychology literature which may provide some useful ideas regarding evolvability
in humans and other mammal taxa. Investigation of mammal species with a broad
array of “stereotyped” signals and displays [e.g., Neotropical howler monkeys,
Atelidae, <i>Alouatta</i>; Old World cercopithecines], should, also, be studied,
since they provide examples of “plastic” [irreversible] traits that can be
combined and recombined. Notably, human phenotypes are virtually devoid of
ritualized signals and displays, with at least one exception being the “eyebrow
flash” (Eibl-Eibesfeldt 1989). <o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Second, extreme behavioral
flexibility and phenotypic plasticity may function to conceal the intent of
Actor and/or to confuse Recipient when it is in the [reproductive] interest of
Actor to do so [“Social behaviour is a subterfuge.”, <i>op cit</i>.]. This idea
tentatively suggests that Actor may “control”—not necessarily in a conscious or
aware manner—the expression of some of the components of his/her phenotypic
arrays, genetically-correlated or not. Related to the aforementioned,
Information Theory emphasizes costs to complexity—“noise” and “error.” These
topics require systematic investigation.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">--</span></b><b><span style="font-family: Wingdings; font-size: 12pt;">à</span></b><b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">
</span></b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">When thinking about the extant
Great Apes, including, humans, a</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">s Gene Robinson's “sociogenomic” (Robinson,Grozinger, &
Whitfield 2005) work with social insects shows, every taxon has its own “toolkit,”
though convergences are often identified, even, among distantly-related taxa,
including, across Classes [see Robinson's landmark paper on convergent
evolution at the genomic level, Woodard et al. (2011), which includes data for
mammals]. Berens et al. (2015) demonstrated that, in fact, across taxa, genes
may differ—it is the pathways that are often be conserved.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><b><span face=""Arial",sans-serif" style="color: #222222; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">--</span></b><b><span style="color: #222222; font-family: Wingdings; font-size: 12pt; mso-ascii-font-family: Arial; mso-bidi-font-family: Arial; mso-char-type: symbol; mso-fareast-font-family: "Times New Roman"; mso-hansi-font-family: Arial; mso-symbol-font-family: Wingdings;">à</span></b><b><span face=""Arial",sans-serif" style="color: #222222; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b><span style="font-family: "Times New Roman", serif; font-size: 12pt;">Like all non-human apes, chimpanzee [<i>Pan troglodytes</i>] females
breed in a group but breed independently [little or no “allomothering,”
“babysitting,” or other contact or “help” by kin or non-kin of either sex].
This phenomenon [that has been noted in the Animal Behavior literature] compared
to the impressive variety of interdependent traits in humans, deserves
systematic investigation.</span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">This contrast between humans and other apes
may relate, evolutionarily, to other diagnostic human features [e.g., gestures
and language; maximum group sizes; extreme reliance upon reward mechanisms (e.g.,
associative learning; Herrnstein's "matching;" observational
learning); mimicry; "fast" reproductive rate combined with slow
development; potential to completely decouple Survival & Fecundity (only
modern Humans, given the advent of technology, can completely decouple Survival
and Fecundity after egg production and harvesting), emancipating human females
to allocate energy to other activities besides breeding and caretaking—should
they “choose;” extreme "phenotypic plasticity," as well as,
behavioral variability/flexibility—primitive traits in animals (Nijhout 2003);
intense sensitivity to and awareness of contextual stimuli (sensation and
perception); mechanisms of punishment (see Clutton-Brock & Harvey 1995) and
other types of “enforcement;” “social learning” and culture;
"complex" societies, as per, combination and re-combination of
differentiated interindividual responses; learned, rather than inherently
predisposed, task and role division-of-labor; broad geographical success of <i>Homo
sapiens</i>]. Not all of these features depend upon higher-order and/or conscious
and aware, cognitive processes, including, individual recognition.</span><span face=""Arial",sans-serif" style="color: #222222; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><o:p></o:p></span></p><p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><b>--> </b>Human social evolution probably requires significant revision. Though there seems to be general agreement that humans do not exhibit "reproductive division-of-labor," the remarkable behavioral flexibility and phenotypic plasticity of human responses permits a wide range of conformations, including, division-of-labor," possibly, subsequent to further investigation, "reproductive division of labor" in some reproductive units." Keep in mind that some human responses will be genetically-correlated.</span></p><p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"><br /></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">CONCLUSION:</span></b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">
The Major Transitions Approach to the evolution of organismal complexity is the
consensus model, including, its assumptions for and applications to
conceptualizing and investigating Social Evolution (West et al. 2015) which I
hope will be adopted by all Mammalian Social Biologists.<o:p></o:p></span></p>
<p class="MsoNormal" style="background: white; line-height: normal; mso-margin-bottom-alt: auto; mso-margin-top-alt: auto;"><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 10pt; mso-fareast-font-family: "Times New Roman";"> </span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">ACKNOWLEDGMENTS<o:p></o:p></span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><span style="color: #222222; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";">Too many people to
mention individually have contributed to my scientific career, directly and
indirectly, by providing information and feedback, as well as, encouragement
and constructive criticism. Nonetheless, without the early assistance of Mary
Jane West-Eberhard, as well as, the late Harry Levin, the late John F.
Eisenberg, and the late Masao Kawai, in addition to, the faculty of Tropical
Ecology course, OTS-‘73, my career never would have happened. I appreciate
being permitted to use the amazing photographs shared by Chris Faulkes [Fig. 1]
and Brian Wood [Fig. 2]. Much gratitude to J. Gordon Faylor for providing the
technical expertise required to create a lulu.com text based on my blogpost at
vertebratesocialbehavior.blogpost.com. My highest praise and thanks are
extended to my son, Miguel Luke Jones, who makes it all possible.</span></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><b><span style="color: #0f1419; font-family: "Times New Roman",serif; font-size: 12pt; mso-fareast-font-family: "Times New Roman";"> </span></b></p>
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<p class="MsoNormal" style="background: white; line-height: normal; margin-bottom: 0in; mso-margin-top-alt: auto;"><br /></p>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-66146190989446712142022-09-21T12:53:00.007-07:002023-12-09T13:01:25.028-08:00BIOACCUMULATION: A NEW HYPOTHESIS FOR THE EVOLUTION OF "AGE-REVERSED" DOMINANCE SYSTEMS BY CLARA B. JONES<p><br /></p><p>BIOACCUMULATION AND THE EVOLUTION OF "AGE-REVERSED" DOMINANCE SYSTEMS IN ANIMALS, WITH A FOCUS ON NON-HUMAN PRIMATES: A NEW HYPOTHESIS Clara B. Jones August 2023</p><blockquote style="background-color: white; color: #222222; font-family: Arial, Helvetica, sans-serif; font-size: small;" type="cite"><div dir="auto">I have what I consider a robust & testable hypothesis for the evolution of "age-reversed" ["age-inverse"] dominance systems [dominance systems whereby younger adults are dominant to middle-aged & older adults; MA adults dominant to old adults; old adults subordinate to all adults younger than themselves, a dominance system so far documented only in howler monkeys & langurs--both leaf-eating taxa ... I asked ChatGPT to explain its origin ... the LLM suggested a few obvious & generic answers (e.g., female choice, differential access to food) ... nonetheless, the entity "said" something that really got me thinking--that the youngest, dominant animals must be competitively superior ... after much thought, & after rejecting several ideas, I settled on the explanation that toxicity from a folivorous diet builds up in individuals over time ["Bioaccumulation"]--from weaning to old age, leaving progressively older individuals with less Energy--fewer metabolic resources--to allocate to reproduction & survival, as well as, competition with same-sex adults for access to limiting resources [e.g., food, mates] ... this idea is consistent with the hypothesis that howlers are under greater, though significant, metabolic stress than Hanuman langurs since howlers' behavioral repertoire [including little aggression & a diverse display of stereotyped & ritualized signals] is highly truncated compared to langurs ... also, howlers don't have a modified gut; unlike langurs that have a pseudo-rumen ... the suggestion, then, is that, as individuals age, the "age-reversed" ["age-inverse"] dominance system emerges as a result of the age-dependent bioaccumulation of toxic compounds with metabolic consequences constraining/inhibiting allocation of Time & Energy to reproduction & survival as ageing advances ... fwiw, cows, grazers, with a rumen, also, exhibit this dominance system [SB Hrdy, personal communication], as well as, the social insects, <i>Apis mellifera</i> [honey bees] ... furthermore, in cercopithecines characterized by matrilines [e.g., baboons], the dominance hierarchy of sisters is "age-reversed" ["age-inverse"], with youngest sisters dominant to older sisters; mothers are dominant to offspring ... furthermore, Bioaccumulation may explain "wasteful" feeding whereby primates, other mammals, &, probably, other folivorous or herbivorous vertebrates, bite into one or a few items before throwing the item away; mantled howlers, for example, consistently throw away mango fruit after biting into the toxic fruit [many of us have experienced the unpleasant effects of consuming too many mangos] ... interestingly, these monkeys visit the same trees year after year without, seemingly, learning the general rule--"Mangos are toxic." ... finally, selection for behaviors/strategies opposing Bioaccumulation may help to explain the short duration of feeding bouts in food "patches" by some folivorous or herbivorous animals [e.g., folivorous gorillas] ... whether or not Bioaccumulation can explain the aforementioned cases deserves systematic investigation, research that, probably, can be easily effected in the field ...</div></blockquote><p>Clara B. Jones</p><p>August 2023 </p>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-10019498143082747402022-03-17T07:32:00.025-07:002022-09-24T11:45:02.862-07:00SCROLL DOWN TO NEXT BLOGPOST<p>scroll down to next blogpost</p>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-10076783829793217172021-10-18T11:50:00.014-07:002023-12-26T20:43:38.195-08:00Female traits associated with Mammal Social Evolution [Cooperation (Hamilton); Complex Sociality (specialization)] ... CB Jones, 10/2023<p> </p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;"><b>Female traits associated with Mammal Social Evolution [Clara B. Jones October 2023]</b></span></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;">see 1st blogpost in this blog [link below], Mammal Social Evolution: Major Transitions Approach-- also, available at lulu dot com in hard copy or pinned to my Profile on Twitter [X], @cbjones1943 [see West et al. 2015, cited below]... see, also, cited book linked to my Twitter [X] profile concerning mantled howler monkey (<i style="font-weight: normal;">Alouatta palliata</i>) <b>female</b> life-history strategies ...</span></span></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;"><br /></span></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;">https://vertebratesocialbehavior.blogspot.com/2022/11/mammal-social-evolution-major.html</span></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;"><br /></span></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;">c.f. for an early, classic, treatment of Mammal Evolution, including, group-living mammals, see Eisenberg (1971); for an early treatment of <b>females</b> and Mammal Social Evolution, see Wittenberger (1980); among other points, Wittenberger (<i>op. cit.</i>) argued that, for all intents & purposes, mammal <b>females</b> are the "social" sex ... West et al. (2015) is a necessary introduction to "major transitions" & social evolution ...</span></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;"><br /></span></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;">Terminology & Context: </span></span><span style="color: #0f1419;">Terminology in the field of Social Biology, here, related to Social Evolution, is not standardized ... for example, WD Hamilton & Stu A. West, define "social" in different ways, the former limiting use of "social" to Cooperation & Altrusim, the two [of four] categories of behavior [Selfish, Cooperation, Altruism, Spite] in which Recipient gains "fitness" benefits ... West, on the other hand, defines "social" as any interindividual interaction where either or both interactants benefit reproductively ... recall that "interdependence" is a necessary precursor to Cooperation, while Cooperation is a necessary precursor to the evolution of Complex Sociality ... here, I am thinking of Cooperation as per Hamilton's general formulation--an interindividual interaction in which both interactants benefit in "fitness" ... related to the Major Transitions Approach, Complex Sociality [division-of-labor (cooperation between specialists); task, role, &/or morphological specialization] is, particularly, problematic, it seems to me ... if we view specialization along a continuum, say, from feeding specialization to DoL to reproductive DoL with "totipotent" [<b>female</b>] workers or "helpers" to reproductive DoL with more or less "sterile castes" [social insect workers], do we accept the typical usage whereby Advanced Eusocial taxa are the most "complex" societies [the "pinnacle" of sociality as per Wilson (1971)]? ... though many Social Scientists would claim that interindividual interactions in humans represent the "pinnacle" of Complex Sociality, expert consensus among Social Biologists (e.g., Crespi 2014) have concluded that humans cannot be classified, Eusocial, because this species does not display "reproductive division-of-labor," although, the mechanism, Learning, permits humans to maximize the benefits and minimize the costs of generalization & specialization [e.g., (learned) division-of-labor] ... it is worth pointing out that the wide variety of human reproductive structures have not been systematically described, classified, or diagnosed, including, analysis of populations in which more than one architecture is found, such as all modern nation-states [e.g., monogamy, polygamy, promiscuity, etc.] ... whether & how & why these various conformations interact over Time & Space remains to be determined ... it is assumed that 1st Principles [T, E, acquisition, consumption, allocation] relative to abiotic [e.g., climate] & biotic [e.g., food dispersion; breeding sites] environmental regimes, as well as, constraints of limiting resources, as well as, species-typical female traits, always apply ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Most Mammals are "sexually segregated"* [sometimes termed, "solitary" in the literature], with polygynous male home ranges or territories overlapping the home ranges or territories of however many <b>females</b> he can monopolize ... however, aggregations [temporary clumping of animals] may occur around clumped, limiting resources, especially, food [as, white-tailed deer may do in Spring when food is abundant & clumped]; other aggregations may form when animals migrate from one feeding &/or breeding ground to another; the term "herds" often refers to an aggregation ... the term, "group," is reserved for a reproductive unit ... Hamilton's Rule [</span><i style="color: #0f1419;">rb</i><span style="color: #0f1419;"> - </span><i style="color: #0f1419;">c</i><span style="color: #0f1419;">>0 ----> </span><i style="color: #0f1419;">rb</i><span style="color: #0f1419;"> > </span><i style="color: #0f1419;">c</i><span style="color: #0f1419;">] is widely accepted as a general formulation of Social Evolution ... according to Hamilton (1964), "social" behavior is limited to Cooperation & Altruism--the only conformations of interindividual interactions [Selfish, Cooperation, Altruism, Spite] whereby Recipient gains in "fitness" ... this definition of "social" should be derivable from Hamilton's equations ...</span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-size: small;"></span></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Mammalian phenotypes are usually generalized (see Eisenberg <i>op. cit.</i>) though there are noteworthy exceptions, & many mammal species display generalized, as well as, specialized, traits ... the evolution of Complex Sociality requires Specialization [division-of-labor; task, role, &/or morphological specialization]; Interdependence [among interacting group members] is a necessary precursor to the evolution of Cooperation ... since Cooperation is a necessary precursor to the evolution of Complex Sociality, specialization must evolve at some point during the Cooperation "grade" ... Hamilton (<i>op. cit.</i>) classifies "cooperation" as an interaction in which both Actor & Recipient benefit reproductively ... as a significant aside, in taxonomic studies, "specialization" is used to diagnose "primitive"/"ancestral" from "derived"/recent characters, where "specialization" is a criterion for "derived" classification ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Depending upon environmental regime, an advantage of a generalized phenotype is promotion of phenotypic flexibility [reversible response] &/or phenotypic plasticity [irreversible response]--responses that are usually condition-, context-, situation-dependent ... in extreme &/or highly time-varying environments, responses may be "decided" statistically or probabilistically, even under Hamilton's Rule [</span><i style="color: #0f1419;">op. cit.</i><span style="color: #0f1419;">; often termed "kin selection"] ... it is worth pointing out that a] Hamilton's Rule [</span><i style="color: #0f1419;">rb</i><span style="color: #0f1419;"> - </span><i style="color: #0f1419;">c</i><span style="color: #0f1419;"> >0 ----> </span><i style="color: #0f1419;">rb</i><span style="color: #0f1419;">></span><i style="color: #0f1419;">c</i><span style="color: #0f1419;">] does not predict that it is always beneficial to assist the reproduction of kin] the evolution of sociality does not necessarily require individual recognition, e.g., "greenbeards" may operate or "decisions" may be based on probabilities or likelihoods ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">The observation that most mammalian phenotypes are more or less generalized is usually attributed to evolution in time-varying environments ... because most mammalian phenotypes are generalized & because Complex Sociality requires the evolution of specialization, it is expected that Complex Sociality, &, possibly, Cooperation, would be uncommon in the Class & in other taxa with generalized phenotypes ... except for cooperatively-breeding mammals & the [eu]social mole rats, reproductive division-of-labor [cooperation between specialists] has not been described to date in mammals, though, "temporal division-of-labor" ["age polyethism"] has been described for mantled howler monkeys (Jones <i>op.cit.</i>) and probably exists in numerous other species, particularly, folivore/herbivores relying on "clumped" limiting resources [e.g., food] ... "bioaccumulation" has been proposed for the evolution of TDL in mantled howler monkeys [see blogpost above] task, role, & morphological specialization are described so far only for the social [eusocial] mole rats & is learned in humans ... several other species of mammals are potential candidates for specialization [see David Macdonald's edited reviews of Mammal Orders, and many species of mammals exhibit both generalized & specialized traits ... in humans, whose phenotypes are generalized in the mammalian mode, the evolution of learning mechanisms permits [learned] specializations such as division-of-labor and task, role specialization ... learning may have evolved in mammals with generalized phenotypes, in part, to minimize the costs of solitary living and maximize the benefits of group-living ... the question of whether morphological specialization has evolved in humans has not, to my knowledge, been systematically studied in humans; however, Race [ecotype] may be a candidate for morphological specialization in <i>Homo sapiens</i> ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"><b>Since most mammals' phenotypes are more or less generalized, the evolution of Complex Sociality [&, possibly, Cooperation] is expected to be severely constrained in the Class</b> ... however, it is noteworthy that some mammals are specialists, & many mammals are characterized by both generalized & specialized traits ... importantly, some mammalian characteristics [e.g., Learning; phenotypic flexibility (reversible) & phenotypic plasticity (irreversible) may have been favored by selection to minimize the costs & maximize the benefits of Generalization &/or Specialization.</span></p><div><span style="color: #0f1419;"><br /></span></div><div><span style="color: #0f1419;"><br /></span></div><div><span style="color: #0f1419;">1... <b>females</b> are expected to be "energy-maximizers," males, "time-minimizers" (Schoener 1971), a formulation based on 1st Principles ... all <b>female</b> [&/or male] traits/phenotypes should be investigated & measured relative to 1st Principles [Physics: T, E; Ecology: acquisition, consumption, allocation [to survival, &, especially, reproduction] ... see Bateman for evolution of <b>female</b> & male life-history trajectories ["fitness optima"] from initial gamete size [eggs large & costly to produce; sperm many & cheap to produce] ... the evolution of any phenotypic trait can be partitioned into G[enetic] x E[nvironmental] components ... 1st Principles influencing individual [genotype] "fitness" may result from variability of abiotic &/or biotic factors, e.g., climate; dispersion of limiting resources, especially, food, mates, where "dispersion" is the distribution & abundance of limiting resources in Time & Space; interactions with other group members [interindividual events], etc. ... interindividual interactions may be beneficial to, deleterious to, or neutral with respect to an individual's [an entity's] lifetime reproductive success ["fitness"] ... it is imperative for Behavioral Ecologists studying mammals [& other Vertebrates] to investigate the many strategies that have evolved to reduce reproductive, especially, <b>maternal</b> costs ... these strategies often take the form of <b>females</b>' reducing <b>maternal</b> care by one means or another [e.g., "parking;" reduction of fat in milk; evolution of precocial young; "bet hedging;" seasonal breeding, etc.] ... human <b>females</b> are extreme in these strategies since they can reduce maternal costs completely, including, the metabolic costs of ovulation [by culling eggs] ...</span></div><div><span style="color: #0f1419;"><br /></span></div><div><span style="color: #0f1419;"><br /></span></div><div><span style="color: #0f1419;">2... Vertebrates, social behavior [Cooperation, Altruism, as per Hamilton </span><i style="color: #0f1419;">op. cit.</i><span style="color: #0f1419;">] & Complex Sociality [specialization*], cannot evolve unless <b>females</b> transition from breeding in a solitary state to breeding in a group* [a "<i>bauplan</i>"], & it is important to define what "breeding in a group" can mean ... for example, the offspring of solitary mammal breeders may disperse when the next generation is produced--or, soon after ... this solitary state may, depending upon relative costs & benefits to <b>breeders</b>, including, species-typical traits & environmental regime [abiotic & biotic], evolve to a state whereby young from one generation overlap with the next generation, a precondition for the evolution of Complex Sociality [Specialization, as per "reproductive division of labor" characterizing Cooperative Breeding & Eusociality] ... more commonly in mammals, <b>breeders</b> may form communal units ... other conformations exist, such as, Chimpanzees (</span><i style="color: #0f1419;">Pan troglodytes</i><span style="color: #0f1419;">) for which <b>females</b> coexist more or less in a solitary state with their young in association with other <b>females</b> breeding in a more or less solitary state on overlapping home ranges [also see <b>breeders</b>' conformations in other non-human great apes] ... in this system, related, collaborative [cooperative?] males monopolize some number of <b>female</b> home ranges, & the <b>female</b> sex disperses--opposite from the norm in Mammals ... all non-human ape <b>breeders</b> exhibit a more or less solitary mode of maternal care ... the chimp, and other non-human ape sociosexual pattern may have evolved to minimize costs of breeding in a solitary state while maximizing benefits of breeding in a group, e.g., predator defense, access to mates ... a similar tradeoff may explain the <b>female</b> reproductive strategy in mantled howler monkeys (</span><i style="color: #0f1419;">Alouatta palliata</i><span style="color: #0f1419;">) whereby <b>females</b> breed in a more or less solitary state embedded within a multimale-multifemale sociosexual group structure ... each of the aforementioned examples is characterized by <b>female</b> traits minimizing <b>maternal</b> costs ... <b>female</b> reproductive strategies described for all mammalian sociosexual systems can be assessed for mechanisms to reduce <b>maternal</b> costs [primarily, gestation &, especially, lactation], e.g., "parking" young in protected place, usually, associated with reduced number of nursing bouts; reducing amount of fat in milk--in a few mammalian species, the amount of fat varies with stage of infant development; in groups, <b>mothers</b> may reduce <b>maternal</b> costs by receiving "help" from other group members [usually, other <b>female</b> offspring or <b>female</b> kin], sometimes termed, "allomothering" (Hrdy 1976); mantled howler monkeys reduce the duration of <b>maternal</b> care combined with harsh weaning of young as soon as the issue can feed on its own ... the great apes, except humans, are an interesting case because <b>females</b> breed in groups but permit little contact between offspring & other group members, thereby, assuming high <b>maternal</b> costs, an apparent strategy whereby costs and benefits of group-living are minimized, as well as, maximized ... importantly, humans may be the only animal species in which all reproductive costs can, effectively, be eliminated by <b>females</b> [and males, also]... all phenotypic traits of vertebrate, here, mammal <b>females</b> should be evaluated in terms of their ability to minimize the costs of <b>maternal</b> care ... costs & benefits to young should, also, be evaluated and whether young can ever "win" in "fitness" conflicts with the <b>mother</b> ... Trivers (1972) was of the opinion that, in certain contexts, offspring can "win;" Alexander (1974) was skeptical ... the question can only be resolved by measuring the "fitness" costs of offspring's behavior toward <b>mothers</b>, & vice versa ...</span></div><div><br /></div><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;">3... Currently, the consensus is that Humans are not characterized by "reproductive division-of-labor;" thus, the consensus is that humans are not "eusocial" (cf. Crespi 2014) ... </span><span style="color: #0f1419;">Human <b>females</b> [& human males] are unique in being able to relieve themselves of reproductive costs except for the production of eggs [ovulation (& ejaculation) can be avoided by culling of eggs (& sperm)] ... this remarkable capacity for reproductive flexibility contrasts with social insect Queens [<b>females</b>] who relieve themselves, obligately, from all tasks associated with reproduction except for insemination, egg production, & laying ... in Mammals, Eusociality may be viewed as a continuum whereby more or less exclusive [though, not, obligate] "Queens" [social mole rats; cooperative breeders] may perform most or all production of offspring and, also, perform other tasks or roles [e.g., feeding of young; defense] ... it is important to note that, given our knowledge to date, all mammalian [more or less non-reproducing] "helpers" are "totipotent," capable of breeding, & mammals exhibit a variety of mechanisms whereby reproduction is suppressed ... as David Macdonald has pointed out, Class Mammalia is characterized by phenotypic flexibility ... as an aside, solitary breeding, non-human great ape breeding mode, Cooperative Breeding, & Eusociality may be differentiated by breeders' relative degrees of task & role specialization [& investment] ...</span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;">4... Finally, recall that the evolution of groups* [from group formation to group maintenance], as well as, the evolution of/transition to sociality [Cooperation as per Hamilton <i>op. cit.</i>, &, under some environmental conditions, the possible transition (evolution) to Complex Sociality (specialization: division-of-labor--especially, Reproductive DoL; task, role, &/or morphological specialization)] is not inevitable or necessarily reproductively beneficial from the <b>female's</b> ["fitness"] perspective ... Interdependence, the precursor to the evolution of Cooperation is not inevitable, or, necessarily, beneficial, either ... the evolution of groups depends upon abiotic [e.g., climate] and biotic [especially, (clumped) food dispersion] environmental regimes, as well as, species-typical phenotypic traits ... the evolution of Complex Sociality is necessarily dependent upon the evolution of specialization which must first evolve at some point in the "grade," Cooperation ... in addition to environmental factors, the evolution of the <b>female</b> transition from the reproductive costs & benefits of breeding in a solitary state to breeding in a group depends, in part, upon evolved species-specific traits of <b>females</b> [& offspring], as well as, environmental context, such as, dispersion of breeding sites, and predation pressure] ... competition with other <b>females</b> from access to these and other limiting resources will, to a large degree, determine the relative reproductive costs & benefits of breeding in a solitary state to breeding in a group and, ultimately, <b>female</b> dispersion [distribution and abundance in Time & Space] ... a major factor determining whether or not it is reproductively beneficial for <b>females</b> to transition [evolve] from breeding in a solitary state to breeding in a group is her ability to narrow her niche [in particular, her feeding niche], becoming more specialized, in order to minimize competition for limiting resources, one of the major "fitness" costs of group life [see RD Alexander 1974; also, cf. reference cited below] ... in short, <i>ceteris paribus</i>, comparing the relative reproductive costs and benefits to a <b>female</b> from breeding in a solitary state to breeding in a group, either might be most beneficial depending upon initial conditions--species-typical <b>female</b> [& offspring] traits, as well as, environmental regime ... it is not necessarily beneficial for <b>females</b> to breed in groups or for sociality [Cooperation as per Hamilton <i>op. cit.</i>] & Complex Sociality, to, subsequently, evolve ... "decisions" about breeding in a solitary state vs. breeding in a group context depend upon differential costs & benefits to breeders' "fitness" ["relative reproductive success"] ...</span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><br /></span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;">*after group-formation, group-maintenance will not evolve unless limiting resources are "clumped" in T & S ... the evolution of group-maintenance is not necessarily beneficial to an organism's lifetime reproductive success ["fitness"] ... some Behavioral Ecologists have hypothesized that groups will not evolve in response to predation alone ...</span></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><br /></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"><b>References</b></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Alexander RD (1974) The evolution of social behavior. <i>Ann Rev Ecol Syst</i> 5: 325-383.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Eisenberg JF (1981) <i>The mammalian radiations: an analysis of trends in evolution</i>. The University of Chicago Press, IL.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Hamilton WD (1964) The genetical evolution of social behavior. <i>J Theor Biol</i> 7: 1-52.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Hrdy SB (1976) Care and exploitation of non-human primate infants by conspecifics other than the mother. <i>Adv Stud Behav</i> 6: 101-158.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Jones CB (2020) Female mantled howler monkey (<i>Alouatta palliata palliata</i>: Primates, Atelidae) life-history strategies--a "major transitions approach" to mammalian social evolution. lulu dot com</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Schoener TH (1971) Theory of feeding strategies. <i>Ann Rev Ecol Syst</i> 22: 369-404.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Sheppard CE, et al. (2018) Intragroup competition predicts individual foraging specialization in a group-living mammal. <i>Ecology Letters</i> 21: 665-673.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Trivers RL (1972) Parental investment and sexual selection. In B Campbell (ed) <i>Sexual selection and The descent of man</i> (1871-1971). Aldine, Chicago, pp 136-179.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">West SA, et al. (2015) Major evolutionary transitions in individuality. <i>PNAS</i> 112(33): 10112-10119.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Wilson EO (1971) <i>The insect societies</i>. Belknap, Harvard, Cambridge, MA.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Wittenberger JF (1980) Group size and polygamy in social mammals. <i>Am Nat</i> 115: 197-222.</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"><br /></span></p><div><span style="color: #0f1419;"><br /></span></div><p style="font-style: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;">Clara B. Jones, October 2023, Silver Spring, MD, USA</span></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">
<span style="color: #0f1419;"><span style="font-size: small;"></span></span></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><span style="color: #0f1419;"><p></p></span>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-29006812174196919092021-10-10T21:40:00.006-07:002022-09-01T18:38:01.818-07:00Negative Impacts of the Social Sciences on Behavioral Ecology & Social Biology<p> </p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">
<span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Clara
B. Jones</span></span></span></p>
<p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span style="font-style: normal;"><span style="font-weight: normal;">Email:
</span></span></span></span></span><span style="color: navy;"><span lang="zxx"><u><a href="mailto:foucault03@gmail.com"><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span style="font-style: normal;"><span style="font-weight: normal;">foucault03@gmail.com</span></span></span></span></span></a></u></span></span><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span style="font-style: normal;"><span style="font-weight: normal;">;
</span></span></span></span></span><span style="color: navy;"><span lang="zxx"><u><a href="mailto:mapcbj@gmail.com"><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span style="font-style: normal;"><span style="font-weight: normal;">mapcbj@gmail.com</span></span></span></span></span></a></u></span></span><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><span style="font-style: normal;"><span style="font-weight: normal;">
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<p style="margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419; font-family: "Times New Roman", serif;">Twitter:
@cbjones1943</span></p>
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<span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Date: 8/28/2022</span></span></span></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></span></p><p style="font-style: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><span style="color: #0f1419;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">In many ways, the Social Sciences have a disproportionate & unfortunate influence on <b>Behavioral Ecology, including, Social Biology</b>, some of which follow [in no particular order]:</span></span></span></p>
<p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><span style="font-style: normal;">
The Social Sciences Approach to Behavioral Ecology, including, Social Biology, and, Animal Behavior, in general, might be viewed as the </span><i>Scala Naturae</i><span style="font-style: normal;"> Approach since it centers </span><i>Homo sapiens</i> as the measure of and pinnacle of Social Evolution, especially, as derived from the purported complexity of human societies driven by higher-order cognitive traits, including, an obsession with "big brains." A fundamental principle of Complexity Theory is that complexity emerges from simple rules [after which it may be acted upon by Natural Selection, in the cases of evolved traits, see Duarte et al., 2011].</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">The Social Sciences Approach begins with the assumption that humans are complex [contrast with the Major Transitions Approach]. Instead of making the <i>a priori</i> assumption that humans are complex, practitioners of the Social Sciences approach should begin with the null hypothesis, Humans are not complex.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">It is common for Anthropologists to suggest that human phenotypic diversity is too complex to figure out ["irreducible complexity" as per Stephen Wolfram] or that it has no adaptive value. One of a scientist's responsibilities is to search for patterns and to "unpack" variation. Social scientists have made no headway on this score. For example, the ideas in the field, "evolutionary cultural Anthropology" are tortuously complex [sic] & obfuscating. This field is an example of investigators of human behavior starting with the assumption that humans are complex, rather than beginning with the above null hypothesis, as well as, simple hypotheses & mechanisms.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">The Social Sciences Approach [<i>Scala Naturae</i> Approach] is not based on 1st Principles [Physics; Ecology (acquisition; consumption; allocation)].</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">The Social Sciences do not define theory as mathematics, as the Sciences do. In the Social Sciences, theory is, generally, verbal, & many studies are purely descriptive Natural History, without quantitative treatment beyond Descriptive Statistics.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Social Science research is generally not hypothetico-deductive.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Social Science researchers, in Behavioral Ecology, including, Social Biology, are, generally, averse to field experiments, math modeling, & simulation modeling. It is rare to find studies in the previous fields using Individual- [Agent-] Based simulations to conduct experiments.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Social Science researchers have generally not studied Introductory Ecology & Population Ecology taught by specialists. Behavioral Ecologists, including, Social Biologists, are Autecologists & Population Ecologists. Social Scientists, in general, seem unaware of the literature in Ecology, Population Biology, & Social Biology [e.g., EO Wilson's 1971, The Insect Societies, arguably, the greatest book written to date in Animal Behavior]. Even JH Crook's 1964, classic weaver bird monograph, arguably, marking the inception of the field, Behavioral Ecology, seems unknown to the majority of Social Scientists.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Social Science researchers rarely link their work to [population-level] Evolutionary causes & outcomes. </p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Social Science researchers are fond of Group Selection.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Social Science research rarely uses terminology consistently [e.g., "aggregation" is often used synonymous to "group"].</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Terminology in Social Science research is not standardized. To be fair, terminology is not standardized or used consistently in Animal Behavior, generally, including, Behavioral Ecology & Social Biology. For an introductory discussion on "terminology" in the previous fields, see my YouTube video.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Most Social Scientists studying Behavioral Ecology, including, Social Biology, seem to misunderstand WD Hamilton's Rule, <i>rb</i> - <i>c</i> >0, by assuming that the formulation predicts that it is always in Actor's [reproductive] interest to benefit the reproduction of kin.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">To my knowledge, where social evolutionary transitions are discussed by Social Scientists [including, Evolutionary Psychologists] at all, the Major Transitions Approach, a general model, has not influenced the Social Sciences, including, Anthropology. Readers are referred to my YouTube video, Mammal Social Evolution: A Major Transitions Approach, for an introduction, as well as, to the first blogpost of this blog.. In the MTA applied to social evolution, Cooperation is the gateway to Complex Sociality [Reproductive Division-of-Labor; Specialization]. In his 2019 book, <i>Genesis</i>, Wilson broadens the definition of "eusocial," limiting it to the character trait, Reproductive Division-of-Labor, & denotes Humans, as well as, some other Mammals as "eusocial." It is important to note that Wilson creates a series of 6 transitions from simple to complex sociality that differ in significant ways from the classical formulation. Wilson's [and, some others'] pinnacle stage of social complexity is "language," consistent with a social scientific, <i>Scala Naturae</i> Approach.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><p style="margin-bottom: 0in; orphans: 2; widows: 2;"><b>References</b></p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Crook JH (1964) The evolution of social organization and visual communication in the weaver birds (Ploceinae). Behaviour Supplement #10: 1-201.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Duarte ANA, Weissing FJ, Pen I, Keller L (2011) An evolutionary perspective on self-organizing division-of-labor in social insects. Ann Rev Ecol Syst 42: 91-110.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Wilson EO (1971) The insect societies. Belknap (Harvard), Cambridge, MA.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;">Wilson EO (2019) <i>Genesis: the deep origins of societies</i>. Liveright, NY.</p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><p style="font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p><p style="font-style: normal; font-weight: normal; margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p>
<p style="margin-bottom: 0in; orphans: 2; widows: 2;"><br /></p>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-43302043682004814672020-08-26T19:20:00.003-07:002020-12-11T12:59:32.750-08:00General Schema For The Analysis Of Mammalian Inter-individual Interactions (Clara B. Jones)<p> In <b>Community Ecology</b>, there are three (3) types of species-species interactions: Competition [-, -]; Predator-Prey [+, -]; Mutualism [+, +]. </p><div><br /></div><div>For studies in <b>Population Ecology</b>, we can interpolate these three types of interactions from the Community-level to the Population-level--interactions between individuals of the same species (inter-individual interactions*). Thus, the three (3) types of inter-individual interactions become: Competition [-, -]; Selfish or Social Parasitism [+, -]; Social [Cooperation (+, +), Altruism (-, +)]. This schema can, also, be employed to analyze inter-group interactions.</div><div><br /></div><div>When quantified, these 3 types of inter-individual interactions can be employed to classify all interactions, effecting the decomposition of variability.</div><div><br /></div><div>As for inter-specific interactions, inter-individual interactions may be Facultative or Obligate.</div><div><br /></div><div><br /></div>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-11973832577945186732020-08-15T17:10:00.010-07:002022-06-03T19:49:00.526-07:002020 self-published book: mammal social organization, female mantled howler monkey life history [PDF; order hardcopy] (Clara B. Jones)<p> <b>Citation [122 page book available in hard copy at Lulu "dot" com]</b></p><p><br /></p><p>Jones CB (2020) Female mantled howler monkey (<i>Alouatta palliata</i></p><p><i>palliata</i>: Primates, Atelidae) life-history strategies—a “major transi-</p><p>tions” approach to mammalian social evolution. Lulu.com.</p><p><br /></p><p><b>Abstract</b>: Howler monkeys [<i>Alouatta</i> spp.] are wholly herbivorous. Based on earlier work by the present author [1978; 1980], the "age-reversed" dominance system is described whereby young adult mantled howler females are dominant to older females; middle-aged females are dominant to old females and are subordinate to young females; and, old females are subordinate to young and middle-aged females. The dominance system is characterized by "temporal division-of-labor" ["age polyethism"] whereby "social foraging" exhibits work [foraging, in the present case] graded by age, with old female "workers" ["helpers"] engaged in most foraging activities for variable plant resources, young females accounting for least. Adult female life-history parameters are described based on a "life table" and shown to correspond to patterns of temporal environmental cycles, in particular, the 6-month pattern of rainfall which females can "track" relative to "generation time." To my knowledge, this is the first demonstration of "temporal division-of-labor" in primates. Cooperatively-breeding primates/mammals exhibit, by definition "reproductive division-of-labor," the first stage of "complex sociality" in mammals. Reproductive division-of-labor is absent in mantled howler monkeys. Social mole-rats have been shown to exhibit both temporal and reproductive division of labor.</p><p><b>References</b></p><p>Jones CB (1978) Aspects of reproduction in the mantled howler monkey (<i>Alouatta palliata</i> Gray). Unpublished Ph.D. dissertation, Cornell University, Ithaca, NY.</p><p>Jones CB (1980) The functions of status in the mantled howler monkey (<i>Alouatta palliata</i> Gray): intraspecific competition for group membership in a folivorous Neotropical primate. Primates 21: 389-405.</p><p><br /></p><p><br /></p>Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-78185676158212755832020-04-23T08:07:00.000-07:002020-04-23T08:07:47.484-07:00Abstract: Predictors of male residence patterns in groups of black howler monkeys (Jones et al)<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB, Milanov V, Hager R (2008) Predictors of male residence patterns in groups of black howler monkeys. <i>Journal of Zoology</i> 275: 72-78.</b><br />
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<b>Abstract: </b>Males may share access to fertilizable females (polygynandry) in one environment while, under other conditions, polygynous (one-male or "harem") mating is the norm [in mammals]. However, few studies in mammals have empirically investigated the factors predicting when males will oexist in bisexual reproductive units rather than live in one-male associations with females. We examined patterns of male group membership in a population of black howler monkeys, Alouatta pigra, residing in two habitats (deciduous and riparian of a tropical moist forest environment in Belize, Central America. Using general linear and logistic regression modeling, we evaluated nine variables as possible predictors of male residence patterns (one-male groups or multimale groups). Our results suggest that adult sex ratio and group size are the best predictors of male residence patterns in both habitats. Our findings provide empirical support for theoretical expectations that male reproductive strategies will be a function of habitat-related demographic patterns and the subsequently varying potential of males to monopolize females in heterogeneous regimes. This study may have important implications for our understanding of features of mammalian societies in which males compete directly for access to females.<br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-3543799829983060902020-04-22T17:49:00.000-07:002020-04-22T17:49:57.770-07:00Summary: An exploratory analysis of developmental plasticity... [CB Jones]<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB (2005) An exploratory analysis of developmental plasticity in Costa Rican mantled howler monkeys (<i>Alouatta palliata palliata</i> Gray). In A. Estrada, PA Garber, MSM Pavelka, LeA Luecke (eds.), <i>New perspectives in the study of Mesoamerican primates: distribution, ecology, behavior, and conservation</i>. Springer, New York.</b><br />
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<b>Summary: </b>The topic of developmental plasticity is fundamentally related to life-history evolution (West-Eberhard 2003), in particular, patterns of survival and reproduction. Jones (1997b) employed matrix analysis (see Alberts & Altmann 2003) of Scott's census data with age structure for mantled howlers at Hacienda La Pacifica to estimate life-history parameters including survivorship, fecundity, and mortality. The suite of life-history traits described by this author (e.g., low survivorship in more than one age class, iteroparity, relatively small reproductive effort) is consistent with the view that mantled howlers, and, possibly other members of the genus, express tactics and strategies minimizing costs to fecundity. Since changes in CC [Chest Circumference] and/or CC:P [Chest Circumference : Pubis Width] are irreversible morphological changes, it is proposed that female mantled howlers are capable of responding to local conditions with mechanisms of developmental plasticity, a within-individual strategy compatible with the life-history strategy of mantled howlers (Meyers & Bull 2002; Table 1; see Ravosa et al. 1993). Further research is required to test alternate hypotheses for the present results (e.g., natural selection [C.P. Groves, pers. comm.; F. Nihout, pers. comm.]) and to examine the possibility that there is a threshold of response to locally stressful conditions in irrigation habitat exhibited by female howlers and manifested as developmental plasticity in CC and CC:P.<br />
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The present report is consistent with the program of Stearns et al. (2003: 311) expressed in the following statement: "Alternative explanations for characteristic male and female growth schedules, and the consequences of the patterns seen in each species...all call for investigation across the spectrum of primate social systems." The study of the functional ecology, including physiological ecology and developmental plasticity, of primates is in its early stages (Milton 1998; also see Strier 1992; Ravosa et al. 1993; Crockett 1998; Reader & Laland 2003: 20-21; Jones 2005), investigations which are likely to occupy laboratory and field investigators for many years. This body of research will have important implications on primate and other mammalian development, energetics, life history evolution, and conservation, as it involves an understanding of growth, survival, and reproduction relative to environmental regimes.<br />
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<b>References</b><br />
<b><br /></b>
Alberts SC, Altmann J (2003) Matrix models for primate life history analysis. In PM Kappeler, ME Pereira (eds.), <i>Primate life histories and socioecology</i>. University of Chicago Press, pp 66-102.<br />
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Crockett CM (1998) Conservation biology of the genus <i>Alouatta</i>. <i>Int. J. Primatol</i>. 19: 549-578.<br />
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Jones CB (2005) <i>Behavioral flexibility in primates: causes and consequences</i>. Springer, New York.<br />
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Meyers LA, Bull JJ (2002) Fighting change with change: adaptive variation in an uncertain world. <i>Trends Ecol Evol</i> 17: 551-557.<br />
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Milton K (1998) Physiological ecology of howlers (<i>Alouatta</i>): energetic and digestic considerations and comparison with the Colobinae. <i>Int J Primatol</i> 19: 513-548.<br />
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Ravosa MJ, Meyers DM, Glander KE (1993) Relative growth of the limbs and trunk in sifakas: heterochronic, ecological, and functional considerations. <i>Am J Phys Anthropol</i> 92: 499-520.<br />
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Reader SM, Laland KN (2003) <i>Animal innovation: an introduction</i>. In SA Reader and KN Laland (eds.), Animal innovation, Oxford University Press, Oxford, pp 3-35.<br />
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Stearns SC, Pereira ME, Kappeler PM (2003) Primate life histories and future research. In PM Kappeler, ME Pereira (eds.), <i>Primate life histories and socioecology</i>. University of Chicago Press, pp 301-312.<br />
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Strier KB (1992) Ateline adaptations: behavioral strategies and ecological constraints. <i>Am J Phys Anthropol</i> 88: 515-524.<br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-60847916664782039202020-04-22T16:46:00.000-07:002020-04-22T16:46:17.856-07:00Abstract: Multi-modal communication by male mantled howler monkeys [Jones & Van Cantfort]<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB, Van Cantfort TE (2007) Multimodal communication by male mantled howler monkeys ( <i>Alouatta palliata palliata</i> Gray) in sexual contexts: a descriptive analysis. Folia Primatol 78: 166-185.</b><br />
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<b>Abstract: </b>We analyzed continuously sampled focal and <i>ad libitum</i> data of male mantled howler Cmonkeys (Alouatta palliata palliata Gray) observed in random order. Males resided in two groups in a Costa Rican tropical dry forest environment (riparian habitat group: 3 adult males, 15 adult females, 402 h observation; deciduous habitat group: 2 adult males, 8 adult females, 114 h observation). Samples were limited to sexual contexts, in particular, the 60-min periods before and after each copulation observed within each group for each adult male. Time samples for each male were distributed equally before and after their own copulations. Before statistical analyses were conducted, data were corrected for differences in time sampled for males within each group. Four types of multimodal signaling were resolved: (1) audiovisual, (2) olfactory-visual, (3) olfactory-visual-tactile, and (4) tactile-gustatory. Olfactory and tactile signals were never observed in combination with auditory signals. Consistent with expectation for a Neotropical, arboreal species, audiovisual signals were the most frequently observed type of multimodal communication in both groups (riparian habitat group: n= 139; deciduous habitat group: n= 66). Our evidence strongly suggests that unimodal signals may be combined and recombined to form complex, multimodal signals. Subordinate males in each group were more likely than dominant males to emit audiovisual signals before their own copulations. Male dyads were compared to assess the relative rate of audiovisual signaling by one male before another male's copulations. On average, the subordinate male of the riparian habitat group exhibited audiovisual signals at a higher rate before his own copulations compared to the rate of audiovisual signaling by his dominant challengers. The same comparisons are not significant for males in the deciduous habitat group. The pattern of male response that we report whereby subordinates emit some complex signals at a higher rate than dominants supports the "terminal investment hypothesis" predicting that organisms should increase reproductive effort with age since, in mantled howlers, age correlates negatively with dominance rank. Additional, qualitative observations suggested that subordinates in both groups were most likely to obtain copulations when they increased rates of complex signaling and/or escalated interactions with their male challengers. Group differences were apparent, however, and we suggest factors that may account for these patterns. We assessed responses by female receivers of complex signals emitted by males in sexual contexts. In general, higher-ranking males are more attractive to females and are more successful at monopolizing them. Findings for other, less frequently displayed, multimodal signals (olfactory-visual, olfactory-visual-tactile, and tactile-gustatory) are presented and discussed. We conclude with the suggestion that howlers may be a robust model for the investigation of complex signals in Neotropical primates, including research on functionally referential communication and context-dependent syntax.</div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-41126274082245896932020-04-22T15:51:00.000-07:002020-04-22T16:11:39.149-07:00Abstract: Population structure and group productivity...female socioecology [CB Jones]<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Horwich RH, Brockett RC, James RA, Jones CB (November 2001) Population structure and group productivity of the Belizean black howling monkey (<i>Alouatta pigra</i>): implications for female socioecology. <i>Primate Report</i> 61: 47-65.</b><br />
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<b>Abstract: </b>The assembly and architecture of populations are functions of decisions made by individuals for the optimization of lifetime survival and reproductive success. We analyzed the results of 12 longitudinal surveys (209 group counts) of Belizean black howling monkeys (<i>Alouatta pigra</i>) at the Community Baboon Sanctuary (CBS) in an attempt to describe population structure and group productivity over time. Similar to previous reports of black howlers at several sites, modal group size was found to be one adult male and 2 adult females. Group size ranged from 2 to 16 and maximum female group size* was 4 as reported for other species of polygynous <i>Alouatta</i>. Population density ranged from 8.14 - 178.19 individuals per km^2, one of the highest densities ever recorded for <i>A. pigra</i>. Group size was significantly positively correlated with population density, and 52% of the variance in group size was explained by population density. Female group size and number of immatures [J+I+ sub-adults] per group were positively correlated. An analysis of the least squares regression line for female group size and the number of immatures per group found 9 of 12 surveys experiencing density-dependent conditions. Relative reproductive success [RRS], the mean number of immatures : females per female group size, decreased with increasing female group size suggesting that females in larger groups are at a disadvantage due to decreases in survivorship and/or fecundity [i.e., no Allee Effect]. Again, density-dependent conditions appeared to be operating. Our analysis of gains and losses to 19 groups from 1995-1997 suggests that the black howler population at the CBS is at equilibrium or slightly increasing, primarily as a result of recruitment of immatures (infants, juveniles, and/or sub-adults). If female black howlers at the CBS experience density-dependent conditions, they may undergo significant food competition contrary to predictions of the "ecological model" for folivorous primates. The different conditions predicted by the least squares regression analysis (density-dependence, density-independence, or an advantage to large groups [see method used on page 54: "An indirect test of female survivorship and/or fecundity]) may define the domains of selective pressures generating variations in group size as a function of decisions made by individuals optimizing inclusive fitness. These and other findings have important implications for female social relations <i>Alouatta</i>. At present we cannot distinguish between competition for limiting food resources and infanticide as the proximate mechanism [or, both?] limiting female group size in Belizean black howlers and other polygynous howlers.<br />
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*<b>Related Reference</b><br />
Jones CB, Milanov V, Hager R (2008) Predictors of male residence patterns in groups of black howler monkeys. <i>J Zool</i> 275: 72-78.</div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-76036215500116317642020-04-22T13:21:00.000-07:002020-04-22T13:21:08.630-07:00Relative Reproductive Success... [Methodology]... (CB Jones)<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB (March, 1996) Relative Reproductive Success [RRS] in the mantled howler monkey: implications for conservation. <i>Neotropical Primates</i> 4(1): 21-23.</b><br />
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<b>First Paragraph of Brief Communication: </b>The structure of primate groups is thought to result from the tendency of females to select rich patches of food and that of males to select large aggregations of females (Wittenberger 1980; Emlen & Oring 1977). Because patch richness and the consequent number and quality of females may vary, the relative reproductive success (RRS) of females may also vary over space and time. RRS is a population parameter, since it is one characteristic of demographic or life history traits describing subunits of a species within and between environmental regimes (see Vehrencamp & Bradbury 1984). RRS is important to the field of conservation biology since an increase in the variance of reproductive success in a population reduces effective population size (Primack 1993). Information about RRS facilitates viability analysis of population fluctuations required for recovery from environmental perturbations.<br />
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<b>Methods</b>: This report analyzes relative reproductive success (RRS--Method via Sandy Vehrencamp, Cornell University, ~1976) of mantled howler monkeys (<i>Alouatta palliata palliata</i> Gray) in two Central American forests as the mean number of juveniles plus infants (J+I) per female group size per site. This report uses data from several studies...at two research sites where mantled howler monkeys have been studied most intensively....Mantled howler monkeys, large cebids [n.b., now classified Atelidae]....<br />
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<b>References</b><br />
Emlen ST, Oring L (1977) Ecology, sexual selection, and the evolution of mating systems. <i>Science</i> 197: 215-223.<br />
Primack RB (1993) <i>Essentials of conservation biology</i>. Sinauer Associates, Sunderland, MA.<br />
Vehrencamp SL, Bradbury JW (1984) Mating systems and ecology. In <i>Behavioural ecology: an evolutionary approach</i>. JR Krebs, NB Davies (eds.). pp. 251-278. Sinauer Associates, Inc., Sunderland, MA.<br />
Wittenberger JF (1980) Group size and polygyny in social mammals. <i>Am Nat.</i> 115: 197-222.</div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-30543478606028286552020-04-22T12:42:00.000-07:002020-04-22T15:09:40.384-07:00Abstract: Ethology, neuroethology, and evolvability in vertebrates... (CB Jones)<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB (February, 2008) Ethology, neuroethology, and evolvability in vertebrates: a brief review and prospectus. <i>Primate Report</i> 75: 41-61.</b><br />
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<b>Abstract: </b>The implications of recent developments in cellular and developmental biology are discussed for vertebrate ethology, describing behavior as neuromuscular elements with the potential to generate non-lethal phenotypic novelty induced by environmental stimuli (<i>evolvability</i>). I present a modified schema of a recent model for the origin of adaptive phenotypic novelties. <i>Behavioral accommodation </i>is hypothesized to lead to <i>genetic accommodation</i> if recurrence of environmental effects upon biochemical pathways of novel genetically correlated neuromuscular elements enhances survival and/or reproduction, I review, discuss, and interpret findings which have been implicated in neural plasticity and subsequent reorganization of the phenotype (e.g., "trial-and-error" learning), emphasizing, in particular, the importance of <i>hypervariable exploratory systems</i>. It is suggested that hypervariable neuromuscular elements and subsequent phenotypic plasticity may be induced by long-term potentiation (LTP), potentially deconstraining conserved action patterns and exposing novel patterns of response to selection. The idea that the phenotype is a heterogeneous landscape of neuromuscular elements varying in function from selfish, including parasitic, to mutualistic is proposed, and I suggest that conflict may be ubiquitous, enhancing the potential for deconstraint. A simple theoretical treatment is applied to my proposal that semi-autonomous, antagonistic <i>transposable behavioral elements</i> (TBE) may parasitize one another within and between individuals, inducing hypervariability. I suggest topics for future research, in particular, the role of environmental stressors as inducers of hypervariability and evolutionary adaptability.<br />
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Background Reference<br />
Kirschner M, Gerhart J (1998) Evolvability. <i>Proc. Natl. Acad. Sci</i>., USA 95: 8420-8427.</div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-87640314874532274592020-04-22T12:15:00.000-07:002020-04-22T15:09:02.064-07:00Abstract: Allouatta palliata politics... [CB Jones]<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB (April, 2000) <i>Alouatta palliata</i> politics: Empirical and theoretical aspects of power. <i>Primate Report</i> 56: 3-21.</b><br />
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<b>Abstract: </b>Social scientists have studied social influence, in particular, aspects of power, for more than 50 years. Social influence in two groups of the mantled howling monkey (<i>Alouatta palliata palliata</i> Gray) was investigated. Dyadic aggressive interactions were observed 131 times in 516 h of observation. Ritualized aggression (the "branch-break" display), primarily exhibited by males, accounted for 40% of the observed aggression, while fighting and chasing comprised the remainder. Females exhibited fighting proportionately more than males, but the sexes were equally likely to chase. No aggression was observed between males in the 2-male deciduous forest group. A class (socioeconomic) effect was noted in dyadic aggressive interactions among females since aggressors and victims were usually of similar rank. Females may "disrupt" one another's sexual activities, suggesting that female-female competition is intense. There was little evidence for female bonding. Male-female aggression occurred infrequently, almost always in sexual contexts. Females often used the submissive "bared-teeth display" to rebuff males and appear to be significantly "emancipated" from male control. Aggression by adults toward immatures was rare. High-ranking individuals were observed to harass low-ranking individuals, the primary tactic of group expulsion in both sexes. Males were observed to intervene in the aggressive interactions of females, a form of "policing." Coalitions were observed in the 3-male riparian forest group within both sexes and, for females, appeared to be opportunistic. Post-conflict behavior was analyzed to test the hypothesis that submissive behaviors are expressed more frequently after conflict. No significant differences in the exhibition of submissive behavior occurred post-conflict compared with matched controls. Specific behavior patterns occurred more frequently during post-conflict or matched-control periods, however. In particular, "approach" and "vocalize" were more frequent post-conflict, the latter possibly representing "reconciliation" to repair or to stabilize relationships. The lowest-ranking male in the three-male riparian forest group was experimentally translocated to assess the effects of changes in proximity as a function of male identity and dominance rank. The presence or absence of individuals appeared to affect competitive relations among males. French & Raven's (1959) "bases of power" were identified in <i>A. palliata</i>, but mechanisms of social influence are not necessarily "cognitive-based" as they may be for chimpanzees and humans. French & Raven's paradigm may provide a useful framework for comparative studies.<br />
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"Power is a general matrix of force relations at a given time, in a given society." Dreyfus & Rabinow, 1982, p 186<br />
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References<br />
Dreyfus JL, Rabinow P (1982) <i>Michel Foucault: beyond structuralism and hermeneutics</i> (2nd edition). University of Chicago Press, Chicago, IL, USA..<br />
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French, Jr., JRP, Raven, B (1959) The bases of power. In: Cartwright D (ed.). <i>Studies in social power</i>. Institute for Social Research, Ann Arbor, MI, USA.<br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-79852878374302199732020-04-22T11:28:00.000-07:002020-04-22T13:22:50.877-07:00Abstract: The number of adult females in groups of polygynous howling monkeys... (CB Jones)<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB (January, 2004) The number of adult females in groups of polygynous howler monkeys (<i>Alouatta</i> spp.): theoretical inferences. <i>Primate Report</i> 68: 7-25.</b><br />
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<b>Abstract: </b>Several reports have documented that adult female group size in polygynous howler monkeys (<i>Alouatta</i> spp.) rarely exceeds 4. This paper evaluates three schemas for the interpretation of this phenomenon: (1) a simple game theoretical model; (2) the resource dispersion hypothesis; and, (3) certain models of reproductive skew. Similarities among these schemas are noted, and their possible utility in explaining differences between the typically polygynandrous <i>A. palliata</i> and typically polygynous species of the genus is discussed. Suggestions for future research are proposed, including, data required to test each schema. </div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-76721485666324785272020-04-22T10:36:00.003-07:002020-04-22T10:36:35.998-07:00Abstract: Life history patterns of howler monkeys in a time-varying environment. CB Jones<div dir="ltr" style="text-align: left;" trbidi="on">
<b>Jones CB (1997) Life-history patterns of howler monkeys in a time-varying environment. <i>Biol. Primatol. Lat</i>. 6(1): 1-8.</b><br />
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<b>Abstract: </b>This report examines the relationship between life-history characteristics and environmental predictability for mantled howler monkeys (Alouatta palliata palliata Gray) at Hacienda La Paccifica, Guanacaste Province, Costa Rica. A census with age structure was employed to estimate life-history parameters [calculations of life table after Wilson & Bossert, 1971] including survivorship, fecundity, and mortality, [& generation time]. A time-series analysis of yearly rainfall at La Pacifica was conducted to test inferences from life-history theory whereby variations in mortality across the lifespan [across age stages] are a function of environmental predictability. La Pacifica was found to be a relatively predictable environment, and, consistent with theory, howlers exhibit life-history traits expected for their regime. These include low survivorship during more than one age class, iteroparity, a relatively small reproductive effort, a single young per litter, relatively few young across a lifetime, and relatively long lifespan. The predictable environment of howlers at La Pacifica appears to favor adult over juvenile (including infant) survival, and howler life history is consistent with that for other large mammalian herbivores whose females may time reproductive investment to reduce the [deleterious] effects of environmental heterogeneity ("bet-hedging"). [A moving average model of the rainfall data is provided in the paper.]<br />
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Reference<br />
Wilson EO, Bossert WH (1971) <i>A primer of population biology</i>. Sinauer Associates, Stamford, CT.</div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-44525909573205162262020-01-10T10:06:00.002-08:002020-07-08T16:18:10.208-07:00Alouatta [howler monkeys] spp: unstable "temporal division-of-labor" [Clara B. Jones, 1996]<div dir="ltr" style="text-align: left;" trbidi="on">
Jones CB (June, 1996) Temporal Division-of-Labor In A Primate: Age-Dependent Foraging Behavior. <i>Neotropical Primates</i> 4(2): 50-53. [with minor edits from the original]<br />
<br />
<b>Introduction</b><br />
Division of labor based on age or size ["polyethism"] may reflect the reproductive condition of individuals in social groups. In 1967, West proposed the general hypothesis that hierarchical relations may be advantageous to both dominants and subordinates and that individuals of low rank may be inferior reproductives who benefit genetically from associations with and contributions to reproductively superior individuals. Since increasing age or size eventually entails decreasing reproductive value (Vx), several authors have noted that the display of social behavior, such as foraging behavior that benefits all members of a group, especially kin, should increase with age as the benefits from individual (selfish) reproduction decline (e.g., West-Eberhard 1975; Hrdy & Hrdy 1996). As individual reproductive value decreases, benefits (genetic or other) from assisting the reproduction of conspecifics (social behavior [cooperation, altruism as per W.D. Hamilton 1964]) may increase because costs (genetic or other [including, delayed costs or benefits] of social behavior) decrease with decreased benefits from individual reproduction. In order to test this hypothesis, I studied the relationship between adult female age, dominance rank, reproductive value, and social foraging behavior (food search and pursuit) for adult female mantled howler monkeys (<i>Alouatta palliata</i> Gray). [Author's note, 1/20/2020: "Temporal division-of-labor" (TDL) may, also, be termed, "age-polyethism" or "primitive" (totipotent) eusociality or Totipotent Eusociality (TE); see blogpost on General Mammalian Patterns, #28]<br />
<br />
<b>Subjects and Methods</b><br />
During an extended period of study at Hacienda La Pacifica, Canas, Guanacaste, Costa Rica, I studied two marked, aged groups of mantled howler monkeys in two tropical dry forest habitats [Riparian, Group 5, and Deciduous, Group 12: see Jones 1980, Table 1]. For this species [and others of the genus], age and dominance rank are negatively correlated [Jones 1978, Jones 1980].<br />
<br />
Social foraging was operationally defined as the behavioral series: feed-rest-move [at least 100 m]-feed, by a unit of more than three adults. These criteria were adopted in order to standardize measurement and to eliminate periods of food search within unusually large patches and by consort pairs. I identified which females in the primary study groups initiated foraging sequences and analyzed these observations by age.<br />
<br />
My null hypothesis held that the frequency of foraging by females of any age class would be proportional to the total number of females who foraged in an age class. Two of the 15 [adult] females in one group [both young adults--Riparian Habitat Group 5] were never observed to direct foraging sequences and are excluded from analysis. Three [adult] females were aged on the basis of physical and behavioral traits other than tooth wear, and assignment to age classes for these females was made independent of the present analysis. Two of these females were observed from sub-adult through adult growth and classified as young adults; a third [adult] female, classified as middle-aged, was the mother of a sub-adult and a juvenile offspring, a highly unlikely combination for any other age class [see Glander 1980]. In my analysis of the second group [eight adult females--Deciduous Habitat Group 12], two young adult immigrant females were never observed to forage socially and were excluded from analysis. The pattern of results reported here would remain unaffected by alternative treatments of the raw data.<br />
<br />
A monthly foraging rate for each forager was computed by dividing the frequency of foraging by the female's number of months resident in a group, a period of time varying from 10-14 months since some females emigrated during the study. These rates were compared with a female's age class, on the one hand, and dominance rank, on the other, to assess the relationship between the display of social foraging behavior and rank, and reproductive value [Vx, for her age class, (computed from) population data in Malmgren 1979, Table 23; equation after Wilson & Bossert 1971; c.f. Jones 1997] where relative contribution to future generations of an individual of a given age is quantified.<br />
<br />
<b>Results and Discussion</b><br />
Table 1 [below--scroll down] presents the results of my analysis for the first group [Group 5] of foraging frequency as a function of female age, including, expected frequencies, and Chi Square. Computing "goodness of fit" led to an unequivocal rejection of the null hypothesis [P <- 0.001, X2= 107.64, df= 3]. Thus, old age and foraging frequency are significantly related. Young adult females initiate foraging significantly less than expected on the basis of their numbers [in Group 5: P <- 0.001], suggesting that such individuals are relatively "selfish" or are conserving time [T] and energy [E], possibly for reproduction and/or competition. Table 1 also shows that the middle-aged to old female foraged more than expected by chance [P <- 0.01], and this female succeeded the oldest and lowest-ranking female as the most frequent [social] forager when the old female emigrated in 1977 [personal observation*].<br />
<br />
Additional observations support the reliability of the above patterns. The oldest female in the second group [Group 12] foraged more frequently than any other adult female [P <- 0.001, X2= 17.29, df= 2; c.f. Jones 1998]. Similarly, the relationship between foraging rate and age class [Fig. 1] yields a significant positive correlation [rs= +0.629, P <- 0.05]. Related to this, the correlation between foraging rate and dominance rank [Fig. 2] is significant but negative [i.e., the higher the foraging rate, the lower the dominance rank, rs= -0.63, P <- 0.05]. Thus, the initiation of [social] foraging is significantly associated with female age and dominance rank.<br />
<br />
It was hypothesized above that the expression of social behavior would increase with increasing age since reproductive value [Vx, Fig. 3] decreases with age and with it the benefits from selfish reproduction [i.e., adult females have less to lose and more to gain in fitness as they age]. Figure 3 shows the reproductive value curve for the population of mantled howler monkeys at Hacienda La Pacifica [after Jones 1997]. Comparing Fig. 3 with Figs. 1 and 2, consistent with expectation, a strong negative association appears to exist between reproductive value and rate of foraging. Reproductive value in the four adult age classes is negatively, and significantly correlated with social foraging rate/month [rs= -0.95, P <- 0.02]. These results support the view that increasing age or size eventually entails decreasing reproductive value and that the display of social behavior should increase with age as the benefits from individual [selfish] reproduction decline.<br />
<br />
What features of the howlers' environment might favor temporal division-of-labor? On 52 occasions, I was able to record the specific resource upon which foraging sequences terminated. Forty-four [85%] of these sequences terminated on ephemeral food [i.e., fruit, flowers, or new leaves: see Jones 1996], while eight [15%] sequences terminated with feeding on mature leaves [P <- 0.001, X2= 49, df= 1]. Thus, the initiation of social foraging sequences appears to be associated with food, the local distribution of which is temporally uncertain; new leaves, flowers, and fruit. The old female initiated 21 of the 52 [foraging] bouts, 20 of these for ephemeral food [c.f. Jones 1998].<br />
<br />
An old female's presumed experience with the mosaic of her home range might enhance her efficiency as a forager so that her foraging activity may yield an energetic and nutritional gain to other group members. Temporal uncertainty of preferred food resources [see Jones 1996] may favor individuals that are the beneficiaries of the foraging activity of others, particularly, kin, when reproductive value is low. Division-of-labor through differential social roles may be a function of relative reproductive value, and behavioral roles may be understood within the context of life history patterns. [------>across Social Mammals** & other Social Vertebrates**? across Social Animals**?].<br />
<br />
<b>Acknowledgments</b><br />
I appreciate the comments of R.C. Lewontin, E.O. Wilson, M.J. West-Eberhard, I.S. Bernstein, W.C. Dilger, and K.E. Weber on an earlier draft of this note. I thank the W. Hagnauer family for permission to work on their property, Hacienda La Pacifica, and for logistic assistance. My gratitude to Norman J. Scott, Jr. [USFWS], for expert introductions to the conduct of fieldwork and for imparting a variety of skills, is immeasurable. The work was supported by grants from the National Fellowships Fund and the National Research Council.<br />
<br />
<b>Clara B. Jones</b>, Institute of Animal Behavior, Rutgers University-Newark, 101 Warren Street, Newark, New Jersey 07102, U.S.A.<br />
<br />
<b>References</b><br />
Glander KE [1980] Reproduction and population growth in free-ranging mantled howling monkeys. <i>Am J Phys Anthropol</i> 53: 25-36.<br />
Hamilton WD [1964] The genetical theory of social behavior. <i>J Theor Biol</i> 7: 1-52.<br />
Hrdy SB, Hrdy DB [1976] Hierarchical relations mong female hanuman langurs (Primates: Colobinae, <i>Presbytis entellus</i>]. <i>Science</i> 197: 913-915.<br />
Jones CB [1978] <i>Aspects of reproduction in the mantled howler monkey</i>, Alouatta palliata Gray. Ph.D. Dissertation, Cornell University, Ithaca, NY.<br />
Jones CB [1980] The functions of status in the mantled howler monkey, <i>Alouatta palliata</i> Gray: intraspecific competition for group membershi in a folivorous Neotropical primate. <i>Primates</i> 21: 389-405.<br />
Jones CB [December, 1996] Predictability of plant food resources for mantled howler monkeys at Hacienda La Pacifica, Costa Rica: Glander's dissertation revisited. <i>Neotropical Primates</i> 4(4): 147-149.<br />
Jones CB [1997] Life history patterns of howler monkeys in a time-varying environment. <i>Bol. Primatol. Lat.</i> 6(1): 1-8.<br />
Jones CB [March-Dec, 1998] A broad-band contact call by female mantled howler monkeys: implications for heterogeneous conditions. <i>Neotropical Primates</i> 6(2): 38-40.<br />
Malmgren LA [1979] <i>Empirical population genetics of golden mantled howling monkeys</i> (Alouatta palliata) <i>in relation to population structure, social dynamics, and evolution</i>. Ph.D. Dissertation, University of Connecticut, Storrs.<br />
West MJ [1967] Foundress associations in polistine wasps: dominance hierarchies and the evolution of social behavior. <i>Science</i> 157: 1584-1585.<br />
West-Eberhard MJ [1975] The evolution of social behavior by kin selection. <i>Quart Rev Biol</i> 50: 1-33.<br />
Wilson EO, Bossert WH [1971] <i>A primer of population biology</i>. Sinauer Assoc., Stanford, CN.<br />
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* The last time I saw Group 5's D4 female, she was seated alone and immobile on a tree limb; her face impaled with quills of the prehensile-tailed porcupine [or, coendou: <i>Coendou</i>]. I never encountered this female again.<br />
**EO Wilson's 2019 book, <i>Genesis</i>, advances the idea that many Mammals, including, humans, may be "eusocial." Where "temporal division-of-labor" is demonstrated, taxa can be classified, "primitively eusocial;" this classification would apply, also, to any other Vertebrates or, indeed, to any other Animals, where "age polyethism" is identified. If "tradeoffs" [e.g., energetic, reproductive, survivval] are most likely to be observed in "poor" conditions [e.g., heterogeneous regimes where "fitness" is compromised; recurrent drought, unpredictable food or water supply], "age polyethism" may evolve to minimize energetic costs in time and space. While females are expected to be most sensitive to energetic effects, males, also, may benefit, under some conditions, from age-dependent responses. Furthermore, there may be energetic [reproductive] benefits in coordinating many maturational [age-dependent] and developmental [age-dependent] milestones or markers with one another as genetic and physiological energy-savings tactics and strategies.<br />
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<b>Table 1</b>. Age class, estimated age in years, number of
females in each age class (N), observed (O), and expected (E)
frequencies of social foraging, and chi square (<i>X</i>2) for a test of the null hypothesis. In “Age Class” column,
YA= Young Adult; M-a= Middle-aged Adult; M-a-O= Middle-age to Old
Adult; O= Old Adult.<br />
<table border="1" bordercolor="#000000" cellpadding="4" cellspacing="0" style="width: 100%px;">
<colgroup><col width="51*"></col>
<col width="51*"></col>
<col width="51*"></col>
<col width="51*"></col>
<col width="51*"></col>
</colgroup><tbody>
<tr valign="TOP">
<td width="20%"><b>Age Class</b></td>
<td width="20%"><b>N</b></td>
<td width="20%"><b>O</b></td>
<td width="20%"><b>E</b></td>
<td width="20%"><b>(O-E)2/E (<i>X</i>2)</b></td>
</tr>
<tr valign="TOP">
<td width="20%">YA (5-7)</td>
<td sdnum="1033;" sdval="5" width="20%">5</td>
<td sdnum="1033;" sdval="15" width="20%">15</td>
<td sdnum="1033;" sdval="42.4" width="20%">42.4</td>
<td sdnum="1033;" sdval="17.71" width="20%">17.71</td>
</tr>
<tr valign="TOP">
<td width="20%">M-a (7-10)</td>
<td sdnum="1033;" sdval="5" width="20%">5</td>
<td sdnum="1033;" sdval="35" width="20%">35</td>
<td sdnum="1033;" sdval="42.4" width="20%">42.4</td>
<td sdnum="1033;" sdval="1.29" width="20%">1.29</td>
</tr>
<tr valign="TOP">
<td width="20%">M-a-O (10-15)</td>
<td sdnum="1033;" sdval="1" width="20%">1</td>
<td sdnum="1033;" sdval="18" width="20%">18</td>
<td sdnum="1033;" sdval="8.1" width="20%">8.1</td>
<td sdnum="1033;" sdval="12.11" width="20%">12.11</td>
</tr>
<tr valign="TOP">
<td width="20%">O (15+)</td>
<td sdnum="1033;" sdval="1" width="20%">1</td>
<td sdnum="1033;" sdval="33" width="20%">33</td>
<td sdnum="1033;" sdval="8.1" width="20%">8.1</td>
<td sdnum="1033;" sdval="76.54" width="20%">76.54</td>
</tr>
<tr valign="TOP">
<td width="20%"><b>Total</b></td>
<td sdnum="1033;" sdval="12" width="20%">12</td>
<td sdnum="1033;" sdval="101" width="20%">101</td>
<td sdnum="1033;" sdval="101" width="20%">101</td>
<td sdnum="1033;" sdval="107.65" width="20%">107.65</td>
</tr>
</tbody></table>
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-54158851086449348062019-09-19T12:18:00.000-07:002019-09-23T14:03:49.061-07:00My amazon.com comments on Eisenberg JF (1981)...Mammal "radiations" (Clara B. Jones)<div dir="ltr" style="text-align: left;" trbidi="on">
<b>My amazon.com comments on Eisenberg JF (1981)...Mammal "radiations"...</b><br />
<br />
The mammalogist, John F. Eisenberg's, oeuvre remains one of the most highly respected and important in his field. Because most of his research investigated Neotropical mammals, rather than mammals of the Old World, his is not a household name. Because of my own specializations, in this brief review, I limit my comments to Eisenberg's treatments of mammal Behavior and Social Organization [group-living, interindividual interactions, cooperation, and altruism]. With E.O. Wilson's 1975 important chapter on patterns of sociality in Class: Mammalia ["The secret to the evolution of sociality in mammals is milk."], Eisenberg's "radiations" is the first attempt to identify PATTERNS of Behavior & Social Biology across all group-living mammals for which there was data by 1981. No subsequent treatment has attempted successfully to summarize the social biology of mammals comparable to Wilson's 1971 treatment of Social Insects [necessary reading for Behavioral Ecologists & Social Biologists] or Holldobler & Wilson's 1990 treatment of the Ants. Though Tim Clutton-Brock recently published a book [2016] titled Mammal Societies [sic], it stands as a highly selective literature review, rather than a synthesis. The limited and highly selective chapter by Smith et al. in Rubenstein & Abbott's 2017 Comparative Social Evolution omits Primates and posits bats as the Order needing research as a model for the evolution of sociality in mammals [a view, possibly, derived from Wilson, 1975's discussion of bats]. To the contrary, bats are a highly derived group not suited for a model of general patterns. Rodents are the taxon of choice, including large-bodied and small-bodied groups, sexually-segregated ["solitary"] to highly social species [including the social mole-rats], and taxa with generalized as well as specialized phenotypes--living across virtually all ecotones, and with a commensal relation to humans--that will be most helpful in the early phases of identifying common features across mammals, as well, possibly, across vertebrates. Additionally, critical to any scientific treatment, and as Eisenberg points out in "radiations," the marsupials are the only group of mammals that can serve as a "control group" for tests of hypotheses and apparent patterns [see comments on this idea in Wilson '75]. My 2014 book, The evolution of mammalian sociality in an ecological perspective, is synthetic, emphasizes Ecology, but is brief. The Class is sorely in need of a synthesis across taxa for which data are available; unfortunately, behavior & sociality of rodents are not well known [but, see, Wolff & Sherman's 2007 volume, Rodent Societies]. Wilson, 1971, 1975, are critical for the standardization of terminology, as well as, factors critical to the study of Social Biology and patterns of group life [e.g., the evolution of Communication, "polyethism"], and Robert Trivers', Social Evolution, as well as, James Costs's, The other social insects, should be consulted. Critically, as a few mammalogists have pointed out [e.g., Bob Selander], initial attempts to synthesize patterns across group-living mammals should begin with analyses that study large and small mammals separately. Finally, we want to assess the observation that the structure of mammal [vertebrate?] groups results from the tendency of females to select rich patches of food and that of males to select the largest relative aggregation[s] of females. Other patterns are identifiable in existing literature. Since the time of John Hurrell Crook [Behavior Monograph X], we know that patterns of group organization depend upon the distribution and abundance [dispersion] of limiting resources [e.g,, food, mates, sleeping sites]. Also, related to this overview, females are, cet. par., "energy maximizers;" males, "time-minimizers."</div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-85901835932494328372019-09-19T09:32:00.003-07:002023-12-16T18:03:22.497-08:00General Mammalian Patterns, esp., Patterns of Group-living Species [compiled by Clara B. Jones]<div dir="ltr" style="text-align: left;" trbidi="on">
<b>General Mammalian Patterns [?--in no particular order] with an emphasis on Group-living taxa, especially, Social Biology & Behavioral Ecology--also see Twitter @cbjones1943... Conceptual Framework... </b><br />
<b><br /></b>
<span face="" style="background-color: #f5f8fa; color: #14171a; white-space: pre-wrap;">FIRST PRINCIPLES OF BEHAVIORAL ECOLOGY:: E[nergy]: Acquisition->Consumption->Allocation====> Worker &/or Reproductive &/or Dependent...(Males, T[ime] Minimizers; Females, E[nergy] Maximizers)</span><br />
<b><br /></b>
<br />
<div style="margin-bottom: 0in;">
<span style="color: #14171a;"><span face="">Mammals: Anisogamy-->Sexually-Antagonistic Selection-->Sexual Conflict-->Differential
T-E investment, including, Life History (males, time-minimizers;
females, energy-maximizers)-->Sexual Dimorphism, inc, Body
Size...constrained by dispersion of limiting resources [especially,
food for females, mates for males]--that may or may not not favor
group-living...[apply, for example, to evolution of male Alternative Reproductive Behaviors (ARBs) & female "counterstrategies"]...</span></span></div>
<div style="margin-bottom: 0in;">
<br /></div>
<br />For analysis, and where data permit, we will separate comments for large [>100 kg] & small mammals--as suggested in literature, e.g., <a href="https://pdfs.semanticscholar.org/3281/ad106c0d6a9fdc831b7725dc5d3460222136.pdf">https://pdfs.semanticscholar.org/3281/ad106c0d6a9fdc831b7725dc5d3460222136.pdf</a><br />
<br /><p style="margin-bottom: 0in;"><span style="color: #0f1419;">Terminology & Context: Terminology in the field of Social Biology, here, related to Social Evolution, is not standardized ... for example, WD Hamilton & Stu A. West, define "social" in different ways, the former limiting use of "social" to Cooperation & Altrusim, the two [of four] categories of behavior [Selfish, Cooperation, Altruism, Spite] in which Recipient gains "fitness" benefits ... West, on the other hand, defines "social" as any interindividual interaction where either or both interactants benefit reproductively ... recall that "interdependence" is a necessary precursor to Cooperation, while Cooperation is a necessary precursor to the evolution of Complex Sociality ... here, I am thinking of Cooperation as per Hamilton--an interindividual interaction in which both interactants benefit in "fitness" ... related to the Major Transitions Approach, Complex Sociality [division-of-labor (cooperation between specialists); task, role, &/or morphological specialization] is, particularly, problematic, it seems to me ... if we view specialization along a continuum, say, from feeding specialization to DoL to reproductive DoL with "totipotent" workers or "helpers" to reproductive DoL with more or less "sterile castes" [social insect workers], do we accept the typical usage whereby Advanced Eusocial taxa are the most "complex" societies [the "pinnacle" of sociality as per Wilson (1971)]? ... though many Social Scientists would claim that interindividual interactions in humans represent the "pinnacle" of Complex Sociality, expert consensus among Social Biologists (e.g., B Crespi 2014) have concluded that humans cannot be classified, Eusocial, because this species does not display "reproductive division-of-labor" ... it is worth pointing out that the wide variety of human reproductive groups have not been systematically described, classified, or diagnosed, including, analysis of populations in which more than one architecture is found, such as all modern nation-states [e.g., monogamy, polygamy, promiscuity, etc.] ... whether & how & why these various conformations interact over Time & Space remains to be determined ... it is assumed that 1st Principles [T, E, acquisition, consumption, allocation] relative to abiotic [e.g., climate] & biotic [e.g., food dispersion; breeding sites] environmental regimes, as well as, constraints of limiting resources, as well as, species-typical female traits, always apply ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"><br /></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">1... most Mammals are "sexually segregated" [sometimes termed, "solitary" in the literature]; however, aggregations [temporary clumping of animals] may occur around clumped, limiting resources, especially, food [as, white-tailed deer may do in Spring when food is abundant & clumped]; other aggregations may form when animals migrate from one feeding &/or breeding ground to another; the term "herds" often refers to an aggregation ... the term, "group," is reserved for a reproductive unit ... Hamilton's Rule [<i>rb</i> - <i>c</i>>0 ----> <i>rb</i> > <i>c</i>] is widely accepted as a general formulation of Social Evolution ... according to Hamilton (1964), "social" behavior is limited to Cooperation & Altruism--the only conformations of interindividual interactions [Selfish, Cooperation, Altruism, Spite] whereby Recipient gains in "fitness" ... this definition of "social" should be derivable from Hamilton's equations ...</span></p><div><span style="color: #0f1419;"><br /></span></div></div><div dir="ltr" style="text-align: left;" trbidi="on"><br /></div><div dir="ltr" style="text-align: left;" trbidi="on">2... Several authors have claimed that large mammals have, <i>cet. par</i>., generalized phenotypes (e.g., Eisenberg 1981), presumably, related to evolution in heterogeneous regimes ["environmental grain" theory proposed by Richard Levins 1968. Wilson '75 posits that [1] phenotypic flexibility/plasticity is a general feature of social evolution of large mammals [n.b., the evolution of "learning" should be discussed here, including, the evolution of "social" learning requisite to Culture]; however, generalized phenotypes may represent [2] "canalization." Whether [1] or [2] is an effect of temporal variability & "grain" will depend upon patterns of variability [e.g., rainfall, hurricanes] relative to Generation Time [Is the environment trackable?] . For all of the topics in #2, you might want to take a look at my 2012 Springer Brief. Furthermore, to the extent that mammals exhibit generalized phenotypes, this trait would be expected to inhibit evolution of specialized morphologies, such as, morphologically distinct castes. See sections on generalized phenotypes in Eisenberg (1981).</div><div dir="ltr" style="text-align: left;" trbidi="on"><br /></div><div dir="ltr" style="text-align: left;" trbidi="on"><p style="margin-bottom: 0in;"><span style="color: #0f1419;">3... Mammalian phenotypes are usually generalized [see Eisenberg <i>op. cit.</i>] ... the evolution of Complex Sociality requires Specialization [division-of-labor; task, role, &/or morphological specialization]; Interdependence [among interacting group members] is a necessary precursor to the evolution of Cooperation ... since Cooperation is a necessary precursor to the evolution of Complex Sociality, specialization must evolve at some point during the Cooperation "grade" ... Hamilton (<i>op. cit.</i>) classifies "cooperation" as an interaction in which both Actor & Recipient benefit reproductively ... as a significant aside, in taxonomic studies, "specialization" is used to diagnose "primitive"/"ancestral" from "derived"/recent characters, where "specialization" is a criterion for "derived" classification ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">4... depending upon environmental regime, an advantage of a generalized phenotype is promotion of phenotypic flexibility [reversible response] &/or phenotypic plasticity [irreversible response]--responses that are usually condition-, context-, situation-dependent ... in extreme &/or highly time-varying environments, responses may be "decided" statistically or probabilistically, even under Hamilton's Rule [</span><i style="color: #0f1419;">op. cit.</i><span style="color: #0f1419;">; often termed "kin selection"] ... it is worth pointing out that a] Hamilton's Rule [</span><i style="color: #0f1419;">rb</i><span style="color: #0f1419;"> - </span><i style="color: #0f1419;">c</i><span style="color: #0f1419;"> >0 ----> </span><i style="color: #0f1419;">rb</i><span style="color: #0f1419;">></span><i style="color: #0f1419;">c</i><span style="color: #0f1419;">] does not predict that it is always beneficial to assist the reproduction of kin] the evolution of sociality does not necessarily require individual recognition, e.g., "greenbeards" may operate or "decisions" may be based on probabilities or likelihoods ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">5... the observation that most mammalian phenotypes are generalized is usually attributed to evolution in time-varying environments ... because most mammalian phenotypes are generalized & because Complex Sociality requires the evolution of specialization, it is expected that Complex Sociality, &, possibly, Cooperation, would be uncommon in the Class & in other taxa with generalized phenotypes ... except for cooperatively-breeding mammals & the [eu]social mole rats, [reproductive] division-of-labor [cooperation between specialists] has not been described to date in mammals; task, role, & morphological specialization are described so far only for the social [eusocial] mole rats ... several other species of mammals are potential candidates for specialization [see David Macdonald's edited reviews of Mammal Orders ... in humans, whose phenotypes are generalized in the mammalian mode, the evolution of learning mechanisms permits [learned] specializations such as division-of-labor and task, role specialization ... learning may have evolved in mammals with generalized phenotypes, in part, to minimize the costs of solitary living and maximize the benefits of group-living ... the question of whether morphological specialization has evolved in humans has not, to my knowledge, been systematically studied in humans; however, Race [ecotype] may be a candidate for morphological specialization in <i>Homo sapiens</i> ...</span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;"></span></p><p style="margin-bottom: 0in;"><span style="color: #0f1419;">6... <b>since most mammals' phenotypes are generalized, the evolution of Complex Sociality [&, possibly, Cooperation] is expected to be severely constrained in the Class</b> ... on the other hand, some mammal species display specialized phenotypes, & many species display both generalized and specialized traits ... selection may favor some characters [e.g., Learning; phenotypic flexibility (reversible) & phenotypic plasticity (irreversible)] because they minimize the costs &/or maximize the benefits of generalization &/or specialization ... thinking in terms of differential costs & benefits [to "fitness"], research has led to the conclusion that specialists are good competitors but vulnerable to extinction while generalists are poor competitors but comparatively less vulnerable to extinction ...</span></p><div><span style="color: #0f1419;"><br /></span></div><div><span style="color: #0f1419;"><br /></span></div></div><div dir="ltr" style="text-align: left;" trbidi="on">7... Selander & Kaufman (1973) and others [see paper linked in #1 above] argue that large mammals have higher tolerance for genetic monomorphism. <br />
<br />8... The basis for Social Evolution in mammals is "milk", according to EO Wilson [1975, <i>Sociobiology</i>, Belknap: HUP] n.b. a number of authors claim that the basis for Social Evolution in mammals is the primitively [non-cooperative, non-altruistic] communal female group leading to polygyny [c.f. Wittenberger 1980]. The basis of Wilson's point, however, is that Mammal females are obligate caretakers of young, predisposing them to certain life-history trajectories, including, probably, receiving help, all limited by energetics. Recall the litany that females are "energy-maximizers," males, "time-minimizers." Some female mammals, extend maternal care [&, often, lifespan], a trait that will retard the evolution of "complex" sociality [specialization, eusociality] because extended maternal care prevents the evolution of reduced maternal care and the evolution of the de-coupling of Survival and Fecundity. Importantly, in humans, social insects, & "primitively" eusocial mole-rats [& "cooperatively-breeding" mammals?], an apparently rare life-history strategy is found [among females] whereby extended lifespan [a "slow" life-history trait] is combined with a high reproductive rate [a "fast" life-history trait]. In humans, however, extended lifespan is, apparently, combined with increased investment in maternal care; while, in social insects, social mole-rats, & "cooperatively-breeding" mammals [?], extended lifespan in associated with reduced maternal care--a necessary, though, not sufficient, condition for the evolution of higher "grades" of sociality [specialization, eusociality, genotypic altruism]. Further, reduced maternal care is a necessary trait associated with decoupling of Survival & Fecundity.<br />
<br />9... According to Wilson (1975, p 379), the four "pinnacles" of social evolution are: the colonial invertebrates [e.g., corals, bryozoans], the social insects, the non-human mammals, & humans]. n.b. reflective of a "major transitions" approach to the Tree of Life<br /><br /></div><div dir="ltr" style="text-align: left;" trbidi="on">10... Traits of the non-human mammals: aggressiveness & discord carried further in vertebrate, including, mammal, societies than in social insects...selfishness rules inter-individual interactions...no sterile castes [though reproduction may be suppressed chemically or by "choice"]...acts of altruism infrequent and usually directed toward kin, especially, offspring...cooperation usually rudimentary; though inequity & hierarchy virtually ubiquitous [wherever there is division of labor there will be inequity]...by human standards, some acts "brutal" [e.g., treatment of infirm & dying]...mostly after Wilson '75, p 380<br />
<br />11... Human social traits: humans remain "essentially vertebrate" in their social structure, but more "complex," not by reduction of selfishness but by evolution of intelligence and language [relate to phenotypic plasticity; learning, including, social learning (cultural mechanism); social parasitism; persuasion; manipulation; exploitation, etc]...elaboration of kin relations [cost-benefit]...human societies "approach" levels of cooperation in insect societies & far exceed social insect societies in communication...mostly after Wilson '75, p 380<br />
<br />12... Most "interesting" mammal orders: rodents, artiodactyls, primates, marsupials--"control group" for Class after Wilson '75 & Eisenberg '81 [social biology of largest mammal orders--bats, rodents--virtually unknown as of 1975]...mostly after Wilson '75, p 468 x<br />
<br />13... Traits labile [e.g., mode of intrasexual cooperation, degree of cohesion, openness of societies (c.f. "open" vs. "closed" groups)]. Many forms of interaction change seasonally, & patterns differ at species level. [after Wilson '75]<br />
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14... Phylogenetically, more primitive living marsupials and insectivores tend to be solitary. Species that forage nocturnally or underground are, also, mainly solitary. Eisenberg [1981] & Wilson [1975] posit that marsupials are the only "control group" we have for Class Mammalia as a whole.<br />
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15... As a rule, the most complex social systems within each Order occur in the physically largest numbers...true of marsupials, rodents, ungulates, carnivores, and primates [Why? maybe because diurnal and because of large brains (intelligence?)? Wilson '75 Also, refer to J.H. Crook's work with weaver birds and other work in Behavioral Ecology: need to evaluate dispersion (distribution and abundance in time and space) of limiting resources relative to behavior & social biology].<br />
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16... Species adapted to life in open environments more likely to be social. Rodents known to form coteries of mixed sexes are all inhabitants of grasslands. Ungulates--great herds predominantly found in grasslands and on savannas, though herds mostly loosely structured--but see, horses, mountain sheep, elephants, and a few others [zebras?] Wilson '75<br />
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17... The structure of mammal [vertebrate?] groups is thought to result from the tendency of females to select rich patches of food and that of males to select large aggregations of females. Because patch richness and the consequent number [and quality] of females [female group size] is expected to vary, the relative reproductive success [RRS] of females may, also, vary over time and space. See the following paper, for example:<br />
<br />
<a href="http://eprints.uberibz.org/1173/1/jones_2008.pdf">http://eprints.uberibz.org/1173/1/jones_2008.pdf</a><br />
<br />
Also, see:<br />
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Jones, CB (March, 1996) Relative reproductive success [RRS] in the mantled howler monkey: implications for conservation. <i>Neotropical Primates</i> <b>4</b>(1), pp 21-23.<br />
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18... Chapters on Mammals in Rubenstein & Abbott (2017), may not be very useful for treatments of general patterns of Social Biology in Mammals; however, see stimulating final, summary chapter in this ambitious edited volume.<br />
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19. IMO, the chapters in Wilson (1975) on non-human primates and on humans are not very useful for identifying patterns across the Primates, in <i>Homo sapiens</i>, across Mammals [or, across vertebrates].<br />
<br />20... Tim Clutton-Brock's text, <i>Mammal Societies</i>, not very useful for identifying general patterns of Social Biology in Mammals. This book is not a Synthesis for the Mammals in the way that Wilson's [1971], <i>Insect Societies</i>, is a Synthesis for the Social Insects--even though TC-B titles his book, <i>Mammal Societies</i>. This book is a highly selective literature review. See, however, the useful table of contents & #generateideas. re: sub-eusocial insects--their traits & trends--see James Costa's book, <i>The other insect societies </i>which includes many ideas for research.<br />
<br />21... Neotropical social mammals require thorough treatment, review, and integration into the broader literature on mammals which is dominated by research in the Paleotropics. In South America, see, especially, caviomorph rodents [infraorder, Caviomorpha], uniting all So. American hystricognaths. n.b. all or most of these spp. are small-bodied, & they are related to Old World [hystricognath] social mole rats.<br />
<br />22... General Patterns...Gene Evolution...Convergence...Gene E. Robinson Lab [see Jenny Tung's preliminary, though, cutting-edge, work on mammals @jtung5]:<br />
<br />
<a href="https://www.pnas.org/content/108/18/7472">https://www.pnas.org/content/108/18/7472</a><br />
<br />23... Temporal division of labor [age (-graded) polyethism: Wilson 1971] is ubiquitous in social insects; how common is it in Mammals & other vertebrates? See [especially, tables]: Jones CB (June, 1996) Temporal division of labor in a primate: age-dependent foraging behavior. <i>Neotropical Primates</i> 4(2): 50-53.<br />
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also, Wilson EO (1971) <i>The Insect Societies</i>. Belknap [HUP], Cambridge, MA.<br />
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24. The following book by Bernie Crespi discusses correlates of sociality [p 66, pp 73-74]...note, especially, significance of "high value" of limiting resources [abode, mates, food, etc.]...again, shows need to study limiting resource dispersion [distribution & abundance in time & space]...<br />
<br />
<div style="margin-bottom: 0in;">
<span face="">Crespi BJ, Morris DC, Mound LA (2004) <i>Evolution of ecological and behavioural diversity: Australian</i> Acacia <i>thrips as model organisms</i>. Australian Biological Resources Study, Canberra.</span></div>
<div style="margin-bottom: 0in;">
<span face=""><br /></span><span face=""><span style="font-size: small;"><a href="https://www.environment.gov.au/science/abrs/publications/thrips">https://www.environment.gov.au/science/abrs/publications/thrips</a></span></span><br />
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<span face="">25... How much of the variation in mammal behavior & social biology is explained by body size?</span><br />
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<span face="">26. Nursing [mammary glands] predisposes mammal females to caretaking, dependency, subordinate [e.g., "helper"] status, & favoritism of kin &/or other females. But how are these adaptations affected by evolution in suboptimal ["poor"] conditions or conditions of intense competition [for limiting resources], or, say, drought or collapse of prey population?</span><br />
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<span face="">27. See Part 4 of Eisenberg [1981] for discussion [also, tables] of mammalian behavior, including mating systems & social biology--see, especially, Chapters 31-33.. Eisenberg [1981] classifies mammal "mating systems" into 10 categories, reduced to 3 basic types: polygyny [1 male, >1 females]; polyandry [multiple-male : multiple female], in which females mate with >1 male [not necessarily "promiscuous", i.e., females may demonstrate selectivity or preferences; also, there may be "favoritism" in which an adult female & an adult male are likely to mate exclusively or commonly with each other]; monogamy [uncommon] </span><br />
<span face=""><br /></span>
<span face="">See Eisenberg's outdated but still interesting and classic paper, Eisenberg JF (1966) The social organizations of mammals, <i>Handbuch der Zoologie</i>, Band 8. Lieferung 39: 10(7): 1-92.</span><br />
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<span face="">28... Feldhamer GA, Drickamer LC, Vessey SH, Merritt JF, Krajewski C (2007) <i>Mammalogy</i>. Johns Hopkins UP, MD: "typical mammalian pattern of preferring unrelated distant kin as mates;" Emlen's "ecological constraints" model + Trivers' reciprocity [+ "pay to stay"?] explain "helping" in mammals?--especially, patterns in social mole rats [do naked mole rats have castes since there is a specialized "disperser form" in this species?]...see <b>Jarman [1974] for classification of mammalian social organization x feeding selectivity: least selective most gregarious, large, polygamous, & sexually dimorphic</b>--Jarmon PJ [1974] The social organization of antelope in relation to their ecology. Behaviour 48: 215-267...<b>need to integrate Jarmon's findings with those of Crook [1964, <i>Behaviour Monograph X</i>] on weaver birds, i.e. relationship between food dispersion [distribution & abundance] and social structure/organization</b>...</span><br />
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<span face="">29... Ricklefs RE, Miller GL [2000] Ecology, 4th Edition. W.H. Freeman & Co., NY: Part 4, Population Ecology, Section 4.2, The dispersion of individuals reflects habitat heterogeneity and social interactions--<b>"The dispersion of individuals within a population describes their spacing with respect to one another."--clumped [aggregation (maybe poor choice of word)]--clumped [limiting] resources may be most common determinant of these patterns, also, tendency of offspring to remain in natal group --see #13 above; spaced [even spacing= hyperdispersion]; random ["In the absence of social antagonism, which results in a spaced dispersion pattern or mutual attraction, which leads to clumping, individuals may distribute themselves at random"</b> Can use Poisson methods to evaluate dispersion patterns...also, see Index of Dispersion [Clark & Evans, 1954]...also, can evaluate "nearest-neighbor distance"--"interindividual distance" [IID] in primatology literature...again, see Crook, 1964, <i>Behavior Monograph X</i>...</span><br />
<span face=""><br /></span><span face="">30... When discussing patterns of grouping or "social" organization [in populations], I have the strong impression that Mammalogists tend to attribute the patterns to phylogeny, philopatry [patterns of dispersal x sex], and/or predation...a Crookian [Behavioral Ecology] framework seems not to influenced Mammalogy very much where a "Crookian framework" would entail the mapping of animals in a population on to the dispersion of limiting resources [distribution and abundance of limiting resources in time and space--<i>ceteris paribus</i> & on average relative to differential engergetics x sex--males, at least, adult males, expected to be "time-minimizers," females, at least, adult females, "energy-maximizers"]...</span><br />
<span face=""><br /></span><span face="">31... The (2007) edited volume on Rodents by Wolff & Sherman [link] should be read in association with John F. Eisenberg's 1981 book on mammalian "radiations" [op. cit.]...Eisenberg provides broad context for many topics [e.g., parental behavior, scent marking, infanticide]...</span><br />
<span face=""><br /></span>
<span face=""><a href="https://www.amazon.com/s?k=wolff+rodents&i=stripbooks&ref=nb_sb_noss">https://www.amazon.com/s?k=wolff+rodents&i=stripbooks&ref=nb_sb_noss</a></span><br />
<br />32... General patterns in Social Mammals [Social Vertebrates? Social Animals?]: EO Wilson [<i>Insect Societies</i>, Belknap (Harvard), 1971] discussed "age polyethism" ["temporal division-of-labor": TDL; "primitive" (totipotent) eusociality] in social insects as a feature of eusociality, particularly, "primitive" eusociality [taxa displaying "totipotency," the capacity to switch from a reproductive to a non-reproductive state & vice versa, as opposed to "advanced" eusociality, taxa characterized by sterile castes]...this article* provides the first empirical evidence of temporal division-of-labor in a mammal [1996], the mantled howler monkey, <i>Alouatta palliata</i> Gray, studied by the present author in Canas, Guanacaste, Costa Rica. Temporal division-of-labor ["age polyethism," <b>"primitive" (totipotent) eusociality</b>] may occur widely among social mammals, social vertebrates, and, possibly, all social animals, where Cooperation and/or Altruism [including, reciprocity] have evolved in group-structured populations...<b>i would speculate that TDL ["age polyethism;" "primitive" (totipotent) eusociality] is more energetically efficient than more opportunistic, and, perhaps, more flexible or plastic, modes of [social] behavioral expression & might be expected to evolve/occur in contexts favoring conservation of energy [e.g., in heterogeneous regimes]</b>...since females are expected to be "energy maximizers" [metabolically, reproductively], TDL may be more likely to be observed in females compared to males ["time-minimizers"]...TDL is expected to be developmentally programmed [within some (statistically normal) species-typical, population-typical, range (x sex)], & conscious & aware processes may not be required for its generation...in ////?, Jones reported age-dependent expressions of conflict-resolution among adult males of the same <i>Alouatta</i> species in the same location & group; these results require intense investigation, lab & field experiments, & modeling, including, for humans...it is important to investigate those conditions under which different energy-conservation strategies evolve x sex...since males are expected to be more robust under energy constraints, it seems likely that thresholds will be identified differentially x sex for the same endogenous and exogenous, including, social, stimuli/factors/conditions.<br />
<br />
EO Wilson's 2019 book, <i>Genesis</i>, advances the idea that many [social] Mammals, including, humans [ see**], may be "eusocial." <b>Where "temporal division-of-labor" ["age polyethism"] is demonstrated, taxa can be classified, "primitively eusocial" and/or Totipotent Eusocial [TE] this classification would apply, also, to any other Social Vertebrates or, indeed, to any other Social Animals, where "age polyethism" [TDL; "primitive" eusocial; TE] is identified.</b> If "tradeoffs" [e.g., energetic, reproductive, survival] are most likely to be observed in "poor" conditions [e.g., heterogeneous regimes where "fitness" is compromised; recurrent drought, unpredictable food or water supply], "age polyethism" may evolve to minimize energetic costs in time and space. While females are expected to be most sensitive to energetic effects, males, also, may benefit, under some conditions, from age-dependent responses. Furthermore, there may be energetic [reproductive, survival] benefits in coordinating many maturational [age-dependent] and developmental [age-dependent] milestones or markers with one another as genetic and physiological energy-savings tactics and strategies.<br />
<br />33... Wolff & Sherman (2007, link) includes some interesting & useful chapters, though, terminology is not standardized [& is often confusing (e.g., "single-breeding," "multiple-breeding"--particularly, as they are discussed in relation to philopatry...see, for example, Chapter 21 by Lacey & Sherman, who needed to make Emlen's work on Ecological Constraints fundamental to their discussion of "philopatry")], and "species" is often employed where "population" is, rather, the correct term. Each chapter in this book has a Summary at the end that can be read before digging into the whole chapter. Some chapters in Sections, "Introduction," "Social Behavior," & "Comparative Sociality" are of particular interest to students of Behavioral Ecology & Social Biology. There are many allusions to the importance of environmental [exogenous] factors, including, dispersion of resources; however, there is no explicit integration of [John Hurrel] Crookian socioecology [Behavior Monograph X] nor is there integration of Emlen's Ecological Constraints model which is fundamental when combined with Crookian socioecology. For context & references (especially, Emlen, Crespi), see blogpost, this blog, of EO Wilson's, <i>Genesis</i>.<br />
<br />
Wolff & Sherman [2007], amazon.com:<br />
<br />
<a href="https://www.amazon.com/Rodent-Societies-Ecological-Evolutionary-Perspective/dp/0226905365/ref=sr_1_1?keywords=jerry+wolff+rodents&qid=1580071530&s=books&sr=1-1">https://www.amazon.com/Rodent-Societies-Ecological-Evolutionary-Perspective/dp/0226905365/ref=sr_1_1?keywords=jerry+wolff+rodents&qid=1580071530&s=books&sr=1-1</a><br />
<br />
EO Wilson <i>Genesis</i> review [blogpost, this blog]:***<br />
<br />
<a href="http://vertebratesocialbehavior.blogspot.com/2019/04/review-of-eo-wilsons-new-book-genesis.html">http://vertebratesocialbehavior.blogspot.com/2019/04/review-of-eo-wilsons-new-book-genesis.html</a><br />
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34... Fundamental questions in Behavioral Ecology pertain to Population Structure relative to dispersion [distribution & abundance] of limiting resources in T(ime) & S(pace), also to Population Structure relative to Basic Laws of Ecology--Acquisition, Consumption, Allocation. Ultimately, we should be able to reduce "fundamental questions" to Thermal Biology [all Biology reduces to Heat], & all material phenomena reduceable to Physics----->expressible in mathematical terms.<br />
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31. T<span face="" style="background-color: #f5f8fa; color: #14171a; white-space: pre-wrap;">owards general principles [GUTs] via studies of Biological Assembly...for group-living---->sociality, see literature in Physics on "queuing"...</span><br />
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<span class="css-901oao css-16my406 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" face="" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; color: #14171a; display: inline; font-stretch: inherit; line-height: 1.3125; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: pre-wrap;"> ... </span><a class="css-4rbku5 css-18t94o4 css-901oao css-16my406 r-1n1174f r-1loqt21 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" data-focusable="true" dir="ltr" href="https://t.co/mqFuz8k2Py?amp=1" rel="noopener noreferrer" role="link" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; color: #1b95e0; cursor: pointer; display: inline; font-size: 15px; font-stretch: inherit; font-variant-east-asian: inherit; font-variant-numeric: inherit; line-height: 1.3125; list-style: none; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; text-decoration-line: none; white-space: pre-wrap;" target="_blank" title="https://reintegratingbiology.org/wp-content/uploads/2019/12/Universal-Forces-That-Result-in-Emergent-Properties-Across-Different-Levels-of-Biological-Organization-Dustin-Rubenstein.pdf"><span aria-hidden="true" class="css-901oao css-16my406 r-1qd0xha r-hiw28u r-ad9z0x r-bcqeeo r-qvutc0" color="" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; cursor: pointer; display: inline; font-stretch: inherit; font-style: inherit; font-variant: inherit; font-weight: inherit; line-height: 1.3125; list-style: none; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: inherit;">https://</span><span style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; color: #1b95e0; cursor: pointer; display: inline; font-stretch: inherit; line-height: 1.3125; list-style: none; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: pre-wrap;"><span style="background-color: #f5f8fa; border-color: black; border-image: initial; border-style: solid; box-sizing: border-box; cursor: pointer; font-stretch: inherit; line-height: 1.3125; list-style: none; white-space: pre-wrap;">reintegratingbiology.org/wp-content/upl</span></span><span aria-hidden="true" class="css-901oao css-16my406 r-1qd0xha r-hiw28u r-ad9z0x r-bcqeeo r-qvutc0" color="" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; cursor: pointer; display: inline; font-stretch: inherit; font-style: inherit; font-variant: inherit; font-weight: inherit; line-height: 1.3125; list-style: none; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: inherit;">oads/2019/12/Universal-Forces-That-Result-in-Emergent-Properties-Across-Different-Levels-of-Biological-Organization-Dustin-Rubenstein.pdf</span></a><br />
<br />
...as "assembly" patterns & rules are investigated and identified, it is important to address the Physics literature on "queuing" [see my 2014 Springer Brief on mammalian social evolution, Chapter 4]...<br />
<br />
...as "assembly" patterns & rules are investigated and identified, it is important not to ignore Thermal Biology pertaining to Social Biology...<br />
<br />
35... Needs lots of research: Resource Partitioning in Group-living vertebrates--search for patterns search for rules [within & between trophic levels, including, differential focus on "frugivores," "carnivores," "omnivores," etc.]<br />
<br />
36. <span class="css-901oao css-16my406 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" face="" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; color: #14171a; display: inline; font-stretch: inherit; line-height: 1.3125; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: pre-wrap;">Mammalian Patterns [here, Rodents]: Chapter 3 in </span><span class="r-18u37iz" face="" style="background-color: #f5f8fa; color: #14171a; white-space: pre-wrap;"><a class="css-4rbku5 css-18t94o4 css-901oao css-16my406 r-1n1174f r-1loqt21 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" data-focusable="true" dir="ltr" href="https://twitter.com/hashtag/Rodent?src=hashtag_click" role="link" style="background-color: rgba(0, 0, 0, 0); border: 0px solid black; box-sizing: border-box; color: #1b95e0; cursor: pointer; display: inline; font-size: inherit; font-stretch: inherit; font-style: inherit; font-variant: inherit; font-weight: inherit; line-height: 1.3125; list-style: none; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; text-align: inherit; text-decoration-line: none; white-space: inherit;">#Rodent</a></span><span class="css-901oao css-16my406 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" face="" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; color: #14171a; display: inline; font-stretch: inherit; line-height: 1.3125; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: pre-wrap;"> Societies [Jerry Wolff & Paul Sherman, Eds, U of Chicago Press, 2007] concludes that most rodents are polygynous or promiscuous [the author prefers the latter]...the litany for </span><span class="r-18u37iz" face="" style="background-color: #f5f8fa; color: #14171a; white-space: pre-wrap;"><a class="css-4rbku5 css-18t94o4 css-901oao css-16my406 r-1n1174f r-1loqt21 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" data-focusable="true" dir="ltr" href="https://twitter.com/hashtag/Mammals?src=hashtag_click" role="link" style="background-color: rgba(0, 0, 0, 0); border: 0px solid black; box-sizing: border-box; color: #1b95e0; cursor: pointer; display: inline; font-size: inherit; font-stretch: inherit; font-style: inherit; font-variant: inherit; font-weight: inherit; line-height: 1.3125; list-style: none; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; text-align: inherit; text-decoration-line: none; white-space: inherit;">#Mammals</a></span><span class="css-901oao css-16my406 r-1qd0xha r-ad9z0x r-bcqeeo r-qvutc0" face="" style="background-color: #f5f8fa; border: 0px solid black; box-sizing: border-box; color: #14171a; display: inline; font-stretch: inherit; line-height: 1.3125; margin: 0px; min-width: 0px; overflow-wrap: break-word; padding: 0px; white-space: pre-wrap;"> is that most mammals are polygynous...</span><br />
<br />
37... Mammalian Patterns, Ch. 4 on female reproductive strategies in Wolff & Sherman, op. cit. does not identify general patterns but exposes several problems inherent to the literature on mammalian sexual selection in this book & in many other treatments [e.g., <i>Mammal Societies</i> by TCB, op. cit.]...for example: 1...the reason treatments of male reproductive strategies dominate the literature is that females are a LIMITING resource for males; males are not a limiting resource for females [------>selection]; 2...many treatments, such as this chapter, do not integrate optimality language and logic into their analyses; 3...many/most treatments are not derived from Crookian model of Behavioral Ecology/Social Biology [dispersion [distribution & abundance] of limiting resources [food, mates, sleeping sites, etc.] in Time & Space; 4...many treatments do not integrate Hamilton's Rule & logic into their analyses; 5...many treatments, especially, in Primatology/Anthropology, do not consider the various hypotheses advanced in an attempt to explain Multiple-Mating by Females [in many reports, only Hrdy's idea is advanced and/or the hypothesis that females mate multiply in order to diversify the genetic composition of litters]; 6...Trivers' 1972 chapter in Campbell's volume seems not to have been digested in many treatments of female reproductive strategies [e.g., mammal females, after parturition and after some threshold investment in "current" offspring, are understood to begin investment in future offspring (e.g., repair, mate choice, fertilization, gestation, etc.)]; this generalization may need to be and has been modified for older/old females; 7...females are, cet. par., "energy-maximizers"; males, "time-minimizers"...<br />
<br />
38... this is a classic chapter by Jerry Wolff on alternative reproductive strategies in non-primate mammals...<br />
<br />
<a href="https://www.cambridge.org/core/books/alternative-reproductive-tactics/alternative-reproductive-tactics-in-nonprimate-male-mammals/88AB2B7352642314032233C426A4A231">https://www.cambridge.org/core/books/alternative-reproductive-tactics/alternative-reproductive-tactics-in-nonprimate-male-mammals/88AB2B7352642314032233C426A4A231</a><br />
<br />
39.... following Chapter 7, by John L. Koprowski, in Wolff & Sherman...: "Conflict, both intersexual and intrasexual, has been remarkably influential in the evolution of mating systems of tree squirrels. Conflict among males appears to have been particularly important in promoting evasive behavior in females, led to reduced copulatory duration, promoted alternative reproductive tactics, and influenced postcopulatory tactics for minimizing multiple paternity. Conflict among the sexes likely acts to diminish reproductive success of any dominant individual male, as females mate with multiple males and often avoid active pursuit males. The reproductive skew among males, however, is substantial, and active pursuit males clearly are most successful. Intersexual conflict appears to increase the number of males that are able to mate due directly to the evasive behavior of females, a behavior that maintains the satellite tactic among males. The benefits to females of manipulating the types of competition remain unclear. Future research must quantify the costs and benefits of male and female tactics under different environmental and social conditions...." [95]<br />
<br />40... Gene E. Robinson and other researchers have pointed out that, in social insects, "coordination and control" is decentralized, while, in [most?] human societies, "coordination and control" is centralized...is C & C in [most?] Mammal societies [i.e., in most group-living Mammals], centralized? ... my initial impression based upon my knowledge of all mammalian Orders is that centralized group structure is not common ... dominance hierarchies, for example, would be characterized as a type of "Interdependence," a necessary precursor to social behavior [Cooperation] ... thus, perhaps, the evolution of centralized group architecture is evolutionarily "derived" ...<br />
<br />41... In Wolff & Sherman [op. cit.], Chapter Eight by Dobson & Oli, [using PCA] body mass & phylogeny explain most of the variation in life-history traits [age at maturity; age at last reproduction; juvenile & adult survival; fecundity]...these traits predict a species' position on the "fast-slow [life-history] continuum...should this dataset, also, be analyzed by body size separately--"small" Mammals vs "large" Mammals [unless i am mistaken, a "small" Mammal is considered one < 100 lbs]...<br />
<br />42... in the final analysis, is the most important reason to study Primates the fact that such a large proportion of species is group-living ["social"]--structured around philopatric females, often kin?<br />
<br />
43... In Wolff & Sherman [op. cit.], Chapter 13 by Scott Nunes, "dispersal and philopatry" is discussed: "The dispersal process can be categorized as either natal or post-breeding.... In mammals, natal dispersal is more common than breeding dispersal, occurring in nearly all species. Natal dispersal has a strong sex-bias in mammals, with males typically emigrating in new home areas at higher rates or over greater distances than females." [150] "A variety of hypotheses have [sic] been proposes to explain the prevalence of female philopatry in rodents and other mammals.... The evolution of complex social systems among rodents and mammals in general has been suggested to be predicated upon philopatry and cooperation among kin living near each other [especially, philopatric females, often kin]." [162]<br />
<br />
44... though authors may mention environmental or ecological factors, in general, few, if any, of the chapters in Wolff & Sherman integrate a Crookian model into their analyses [dispersion (distribution and abundance) of limiting resources in Time & Space]--but see Section, Comparative Socioecology, Chapters 29-37...these <b>excellent and important chapters</b> address Families, Genera, & Species that could reasonably become model systems for the investigation of general mammalian patterns, Behavioral Ecology, & Social Biology [<i>Rattus</i> & <i>Mus</i> are already model systems in research Biology and Medicine]...taxa covered are: ground squirrels; marmots; semifossorial desert rodents; <i>Rattus</i> & <i>Mus</i>; Capybaras and Maras; Octodontid & Ctenomyid rodents; rock-dwelling rodents; African mole-rats; black-tailed gunnison's & Utah prairie dogs...<br />
<br />
See Emlen's "ecological constraints" model:<br />
<br />
<a href="http://max2.ese.u-psud.fr/epc/conservation/PDFs/HIPE/Emlen1982.pdf">http://max2.ese.u-psud.fr/epc/conservation/PDFs/HIPE/Emlen1982.pdf</a><br />
<br />
See Greenwood's classic 1980 paper on dispersal in birds & mammals:<br />
<br />
<a href="https://psycnet.apa.org/record/1981-29692-001">https://psycnet.apa.org/record/1981-29692-001</a><br />
<br />
45... Wolff & Sherman Chapter 14: Gene dynamics and social behavior [F. Stephen Dobson] uses "F" statistics [Population Genetics] to investigate population structure, finding that, in many mammals, population structure is a function of dispersion of female kin groups...<b><u>n.b. "social breeding groups may slow the loss of genetic diversity from populations," a process adding to the list of benefits to group-living and, possibly, sociality [cooperation, altruism]</u></b>...indeed, cooperation & altruism may evolve to insure or enhance these benefits [but what are the tradeoffs, the costs?]...though Anthropologists generally maintain that human sociality evolves from an ancestral state of monogamy, the evidence from Mammals summarized in this blogpost strongly suggests that the ancestral state is the female group, especially, the female kin group, responding to clumped, limiting resources [in heterogeneous regimes?]--as most treatments by mammalogists hold...polygyny [and/or promiscuity?] appear to be the most common mammalian sociosexual system, &, perhaps, as suggested by many researchers, males, in part, monopolize female groups to defend females & their limiting resources [saving energy for female "energy-maximizers"]...as an aside, what role dies Sexual Selection play in all of this...<br />
<br />
46... is eusociality much more common than we generally think? Bernie Crespi, WD Hamilton, Nancy Moran quoted in this <i>NY Times</i> article linked below...note importance of protected, reliable food source[s] and of protective, reliable housing [& protection from predators--see Bernie Crespi article linked below]...how about humans?; i have suggested that humans are "facultatively eusocial" in my brief communication, "Are humans cooperative breeders" in <i>Archives of Sexual Behavior</i> discussed on this blog and linked below...<br />
<br />
<a href="https://www.nytimes.com/1993/02/16/science/social-castes-found-to-be-not-so-rare-in-nature.html">https://www.nytimes.com/1993/02/16/science/social-castes-found-to-be-not-so-rare-in-nature.html</a><br />
<br />
"Are humans cooperative breeders: a call for research" (2011), CB Jones:<br />
<br />
<a href="https://link.springer.com/article/10.1007/s10508-011-9741-5?shared-article-renderer">https://link.springer.com/article/10.1007/s10508-011-9741-5?shared-article-renderer</a><br />
<br />
...thre conditions for the evolution of eusociality, Bernie Crespi [Are these general?]:<br />
<br />
<a href="https://www.semanticscholar.org/paper/Three-conditions-for-the-evolution-of-eusociality%3A-Crespi/feb94b7e861ac0e14b8d575f39e33c0a28338419">https://www.semanticscholar.org/paper/Three-conditions-for-the-evolution-of-eusociality%3A-Crespi/feb94b7e861ac0e14b8d575f39e33c0a28338419</a><br />
<br />
47... when we speak of "generality" we are often speaking of Convergence--whatever our level[s] of analysis from gene/allele to whole organism...here is link to a now classic paper from Gene E. Robinson's lab; note generalizeability, convergent evolution, & "toolkit" concept...see Table 3.1 in my 2014 Springer brief--phylogenetic & convergent social features of "prehistoric" mammals...<br />
<br />
<a href="https://www.pnas.org/content/108/18/7472">https://www.pnas.org/content/108/18/7472</a><br />
<br />
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----------------------------------<br />
*<br />
<a href="http://vertebratesocialbehavior.blogspot.com/2020/01/age-polyethism-temporal-division-of.html">http://vertebratesocialbehavior.blogspot.com/2020/01/age-polyethism-temporal-division-of.html</a><br />
<br />
**<br />
<a href="https://link.springer.com/article/10.1007/s10508-011-9741-5?shared-article-renderer">https://link.springer.com/article/10.1007/s10508-011-9741-5?shared-article-renderer</a><br />
<br />
***<br />
<a href="http://vertebratesocialbehavior.blogspot.com/2019/04/review-of-eo-wilsons-new-book-genesis.html">http://vertebratesocialbehavior.blogspot.com/2019/04/review-of-eo-wilsons-new-book-genesis.html</a><br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-19963968992942968182019-06-25T16:44:00.001-07:002019-06-26T20:26:26.717-07:00Research questions based on Ratnieks et al. 2006 (Clara B. Jones)<div dir="ltr" style="text-align: left;" trbidi="on">
<u>Research questions for group-living vertebrates based on Ratnieks et al. 2006</u>*, **<br />
<br />
<i>Introduction</i><br />
<br />
<i>Inclusive fitness theory explanations for both cooperation and conflict</i><br />
<br />
<i>What do we mean by conflict resolution</i>?<br />
1. "Potential conflict is any difference in the reproductive optima of individuals or groups within a society."<br />
2. Individuals have three options...to be workers, reproductives, or dependents (e.g., "social parasites") or some combination of these serially or concurrently. Categories can be analyzed by class or rank, by sex, by age, by race or ethnicity, etc.<br />
3. For vertebrates, including, humans, how do we estimate "fitness optima?"<br />
<br />
<i>Kinship, coercion, and constraint</i><br />
"...low relatedness makes wasteful conflicts more likely because the cost of conflict falls upon more distant kin."<br />
<br />
<i>Sex allocation</i><br />
...differential investment in males and/or females & differential costs/benefits...<br />
<br />
<i>Conflict-resolution</i><br />
1. "Coercion and constraint may help to resolve conflict if one party has complete power over sex allocation." ...etc., e.g., over access to limiting resources...<br />
2. "Mixed power seems likely in many but not all situations." ...e.g., "division of task"..."division of labor"...n.b. DoL more efficient than DoT; also note, male & female interests may not coincide since males are expected to be time-minimizers, females, energy-maximizers...<br />
3. Under some conditions, is it more beneficial for an individual to assume the costs of conflict rather than to resolve conflict? ...i.e., When are costs of conflict [over reproduction &/or limiting resources (convertible to offspring] tolerated or preferred relative to benefits of conflict-resolution?<br />
4. Importance of information, especially, competition for information [see EO Wilson 1971 on "communication" in societies]...Humans: language, who shares information with whom, etc...<br />
5. Investigate "policing"*** as a form of conflict-resolution [mitigation of selfishness], mitigation of conflict [e.g., force, coercion, constraint, persuasion, etc. (see French & Raven's "bases of power")]...<br />
6. What factors lower incentives for individual selfishness [& subsequent "tragedy of the commons"]<br />
7. High "<i>r</i>" lowers the proportion of reproductives in a population...<br />
See "reproductive skew" literature, e.g., R Hager & CB Jones 2009 CUP...<br />
<br />
<i>Conflicts among totipotent individuals</i><br />
1. "Totipotency greatly increases the potential for conflict...."<br />
2. If optimal family and/or group size can be estimated, excess individuals are likely to increase potential for conflict...<br />
<br />
<i>Discussion & Conclusions</i><br />
1. Class conflicts relatively easy to resolve [because dominants "police" subordinates?]...<br />
2. Conflict easily resolved when one party "powerless"...<br />
3. Should the incorporation of steriles or other non-reproductives in groups reduce conflict?...relative to what [e.g., "<i>r</i>"]...<br />
4. How do humans escape control in groups? [see French & Raven's "bases of power"]<br />
5. What role does genetics play in conflict-resolution [e.g., genetic conflict]...<br />
<br />
*Ratnieks FIW, Foster KR, Wenseleers T (2006) Conflict resolution in insect societies. <i>Annu. Rev. Entomol.</i> 51: 581-608.****<br />
**Sentences in quotation marks are quotations from paper; other statements, comments, or questions are my interpolations relative to group-living vertebrates, including, humans.<br />
***New paper: "'Enforcement' is central to evolution of cooperation"...i.e., &, also, to evolution of repression of selfishness... <a href="https://www.nature.com/articles/s41559-019-0907-1">https://www.nature.com/articles/s41559-019-0907-1</a><br />
<br />
****Topics/questions based on titles of literature cited:<br />
--evolution of male and female traits<br />
--causes & consequences of informational constraint<br />
--conflict over class/SES determination<br />
--status-allocation to offspring<br />
--Does it "pay' kin to favor relatives who are "losers" or inferior reproductives and/or competitors?<br />
--n.b. Charnov EL (1978) Evolution of eusocial behavior: offspring choice or parental parasitism? <i>JTB</i> 75: 451-465.<br />
--n.b. Crespi BJ, Ragsdale JE (2000) A skew model for the evolution of sociality via manipulation: why it is better to be feared than loved. <i>Proc R. Soc. London B Biol. Sci.</i> 267: 821-828.<br />
--facultative policing<br />
--conflict over paternity [n.b. causes & consequences of multiple-mating by females]<br />
--n.b. Keller L (1997) Indiscriminate altruism: unduly nice parents and siblings. <i>TREE</i> 12: 99-103.<br />
--genes regulating complex social behavior<br />
--ancestral states of complex sociality<br />
--sex-ratio determination in groups & populations [e.g., conflict over sex-ratio determination]<br />
--causes & consequences of sexual deception [e.g., in some mammal species, males are indistinguishable from adult females for some time before expressing secondary sexual characteristics (see, e.g., mantled howler monkeys, <i>Alouatta palliata</i>)]<br />
--selfish tactics & strategies that promote sociality [e.g., "social cohesion"]<br />
--implications: "heirs & spares"<br />
--mechanisms of conflict-management & conflict-resolution in eusocial taxa<br />
--n.b. Ratnieks FLW, Wenseleers T (2005) Policing in insect societies. <i>Science</i> 307: 54-56.<br />
--alternative reproductive strategies and conflict-resolution/conflict-management<br />
--n.b. Sachs JL, Mueller UG, Wilcox TP, Bull JJ (2004) The evolution of cooperation. <i>Q. Rev. Biol.</i> 79: 135-160.<br />
--policing x age, sex, class/SES, race, etc.<br />
--Is multiple-mating by females rare in vertebrates?<br />
--causes & effects of totipotency; ubiquitous in vertebrates?<br />
--genetic basis of sterility<br />
--mechanisms of kin discrimination<br />
--What traits differentiate members of classes [within & between]?<br />
--relevance of behavioral flexibility & phenotypic plasticity to conflict-management & conflict-resolution [also, flexibility of group structures]<br />
<br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-86413418010627444852019-04-11T20:44:00.004-07:002023-12-30T16:30:16.108-08:00Review of Tim Clutton-Brock's, Mammal Societies (by Clara B. Jones, 2016)<div dir="ltr" style="text-align: left;" trbidi="on">
<div style="margin-bottom: 0in;">
<i>Mammal Societies</i></div>
<div style="margin-bottom: 0in;">
Tim Clutton-Brock</div>
<div style="margin-bottom: 0in;">
2016</div>
<div style="margin-bottom: 0in;">
Wiley-Blackwell (Oxford, UK)</div>
<div style="margin-bottom: 0in;">
744 pp</div>
<div style="margin-bottom: 0in;">
ISBN 97811119095323<br />
<br /></div><div style="margin-bottom: 0in;"><br /></div>
<div style="margin-bottom: 0in;">
“The key to the sociobiology of mammals is milk.” E.O. Wilson (1975)<br />
<br />
Reviewed by Clara B. Jones (2016; revised 2020)*</div>
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Knowledge about group-living mammals may contribute to an understanding of vertebrate social evolution and the evolution of gregariousness in animals with generalized phenotypes [specialization being a signature of high "grades" of social evolution (specialization, reproductive division-of-labor]. Compared to social insects and birds, the social biology of mammals is poorly known with the exception of ungulates, carnivores, and primates (3 of ~25 Orders). Among many similar papers in the mammalian literature on Social Paleontology, in 2011, Ladev<span style="font-family: "times new roman" , serif;">è</span>ze et al. reported fossil evidence appearing to document mammalian gregariousness and its associated ecology from the basal Tertiary of Bolivia. These findings suggested that extinct, marsupial-like <i>Pucadelphys andinus</i> were group-living, probably exhibiting frequent interactions, strong sexual dimorphism, and male-male competition, as well as, polygyny. Based on the spatial and ecological settings of their specimens, as well as, the climate, in addition to, physical and situational associations and patterning of adult, sub-adult, and juvenile remains in their sample, these authors speculated that the species may have been "social" [gregarious]. In 2012, employing phylogenetic analyses, Briga et al. showed that relatedness and allomaternal<span style="font-family: "times new roman" , serif;">¹</span> care are positively correlated in Class Mammalia. These papers indicate that, though the population dispersion of most extant mammals is sexually segregated (“solitary”) and though fossil remains cannot definitively preserve Behavior, group-living may have a long history in this Class (also see Jones 2014, Table 3.1, pp 19-25).</div>
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Tim Clutton-Brock (henceforth, “TC-B”) is a highly-regarded empiricist at the University of Cambridge (UK), recognized, particularly, for his field studies on primates, red deer, and meerkats. He is a prolific scientist with a knack for asking good questions and choosing animal models that have yielded flagship research. The author will be familiar to most animal behaviorists and behavioral ecologists as a specialist of cooperative breeding and evolutionary aspects of reproduction (e.g., female mating strategies, sexual selection). In the book under review, TC-B notes that his undergraduate training was in Anthropology and that he completed his doctorate under Robert Hinde, an animal behaviorist that Psychology typically claims as one of its own. I have been familiar with TC-B's work since the 1970s, and my personal favorites among his copious publications are his 1995 paper with Geoff Parker and the 2003 volume edited with R.M. Sibley & J. Hone. I am pleased to have the opportunity to review <i>Mammal Societies</i>. I have interacted with TC-B on several occasions, once face-to-face, and, more than once, <i>via</i> e-mail. He has always been generous and courteous to me. </div>
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Previous books by JH Crook, “Griff” Ewer, J Eisenberg, EO Wilson, R Estes, D MacDonald, CB Jones, and others, have treated mammalian social biology to one degree or another. <i>Mammal Societies</i>, however, is the first attempt to provide a comprehensive literature review of the topic. The publisher's description of the volume states that it is intended for “behavioral ecologists, ecologists, and anthropologists,” and TC-B self-identifies as a “behavioral ecologist.” The book is, to all purposes, a literature review emphasizing publications on Old World taxa [a tradition attributable, in particular, to Primatology]. While the Table of Contents presents a detailed outline of topics of interest to social biologists, the book is not organized using ecological [e.g., JH Crook] or evolutionary models [e.g., the "major transitions"]. To provide context, professors using <i>Mammal Societies</i> as a course textbook or reference work are strongly advised to acquaint their students early on with Wilson's (1975) treatment (pp 456-574) presenting an explicitly articulated conceptual framework for mammalian social biology, including, trends, conventional terminology, general and comparative features in the Class, an extensive glossary, as well as, case studies and summary tables, figures, and diagrams. John Eisenberg's [1981], "mammalian radiations," is, also, an invaluable source of information on mammalian patterns, including, behavior, as well as, mating & group architectures.</div>
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Chapter 1, “Social evolution,” omits definitions of terms the first time they appear in the book (e.g., “aggregation,” “social”, “cooperation”), leading to obfuscation throughout, particularly, since there is no discussion of how to measure social traits (cooperation, altruism) and to discuss their pertinence to reproductive success. In this chapter, the author might have defined “Mammal” and should tell the reader why mammalian social biology is of import. The reader will want to understand possible trajectories to cooperation and altruism from aggregations to group-formation to group-maintenance and how the (spatial and temporal) distribution of limiting resources favor or disfavor the evolution of mammalian sociality. Chapter 1 is, in great part, a selective account of the history of Animal Behavior combined with some mention of theoretical issues (e.g., Darwinism, competition, reciprocity, game theory). However, for rigorous discussions of verbal and quantitative theory in Behavioral Ecology, as well as, overviews of Methods and G x E interactions, readers are referred to Davies et al. (2012) and Westneat & Fox (2010).</div>
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Chapters 2-9 address topics related to features of female behavior, particularly, as they pertain to mating, maternal tendencies, and gregariousness. Focusing on females, their strategies, and their energetic requirements as the primary driver of group-living and patterns of male behavior and dispersion is fundamental to an understanding of mammal societies because fertilizable females are a limiting resource for males and, subsequently, an ultimate determinant of male “fitness." Though these and other important concepts are implicit in some of TC-B's discussions, explicit use of many principles inherent to Behavioral Ecology are unclear or lacking (e.g., integration of Hamilton's rule [<i>rb</i> > <i>c</i>] throughout chapters, acknowledgment of the many competing hypotheses in Ecology pertaining to dispersal or multiple-mating by females, use of optimality formulations). As an example from Chapter 5 (“Maternal care”), TC-B's treatment asserts, accurately, that mammalian females invest heavily in current offspring, but theory holds that, after parturition, female resources, above some critical minimum, are channeled into future reproduction and lifetime reproductive success--“fitness.”</div>
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Chapters 10-16 pertain to males, especially, mating strategies, relations with females, and paternal care. Characteristic of <i>Mammal Societies</i> as a whole, these chapters are literature reviews of mostly descriptive publications from the Animal Behavior literature. Life history evolution is addressed in this chapter without mentioning the importance of tradeoffs, the distinction between semelparity and iteroparity (“fast” and “slow” life history trajectories, respectively), the importance of life-tables and the role of mortality as a driver of life-history evolution (Stearns 2000). Chapter 17 reviews “Cooperative breeding,” one of TC-B's specializations, and Chapter 18 presents a discussion of “Sex differences." Throughout the book, the author impresses the reader with the centrality of sex, sexual competition, and mating—topics of import in TC-B's career, though one is surprised that more attention is not given to Sexual Selection, per se. Chapters 19 and 20 address hominoids and hominids, including, modern humans, topics often missing or skimmed in other Animal Behavior texts.</div>
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TC-B presents at least one controversial formulation in <i>Mammal Societies </i><span style="font-style: normal;">b</span>y asserting, with no supporting evidence or logical arguments, that no mammals are “eusocial”<span style="font-family: "times new roman" , serif;">²</span>—that the highest grade of sociality in mammals is “cooperative breeding.” This view is orthogonal to standard practice in Mammalian Social Biology whereby the social mole rats are typically classified as “primitively” eusocial. Technically, according to common usage, “cooperative breeders” might, as well, be classified “primitively” eusocial because of the presence of reproductive division of labor [cooperation between specialists] in the form of totipotent “helpers” (see Jones 2014, p 48-52). <i>Mammal Societies</i> exemplifies the need for practitioners of Natural History, Animal Behavior, and Behavioral Ecology to revisit topics such as standardization of terminology, advancement of the Hamiltonian Project, the roles of quantitative theory and modeling (in particular, agent-based modeling), field experiments, as well as, hypothesis-testing [including, the role of null-hypotheses] based on 1<sup>st</sup> principles. <span style="font-style: normal;">The text </span>will appeal to professors wanting a Natural History, mostly, non-quantitative, review allowing supplementary reading to be incorporated into a syllabus. Future syntheses of Mammalian Social Biology will rely on mainstream schemas from Ecology & Evolution, in particular, employing a Major Transitions Approach (cf. West et al. 2015), in addition to, Population Ecology, of which Behavioral Ecology is a sub-field.</div>
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<span style="font-family: "times new roman" , serif;">¹</span><span style="font-size: x-small;">Care of offspring by conspecifics other than the mother</span></div>
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<span style="font-family: "times new roman" , serif;">²</span><span style="font-size: x-small;">”The evolution of eusociality, here defined as the emergence of societies with reproductive division of labour and cooperative brood care, has occurred under specific ecological, genetic, and life history conditions. Although sophisticated levels of cooperation have evolved in the largest and more complex societies, conflicts among individuals are still common because, in contrast to cells of an organism, they are not genetically identical,”--i.e., not "clones" (Keller & Chapuisat, 2010)</span></div>
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<span style="font-size: small;"><b>References</b></span></div>
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<span style="font-size: small;">Bradbury JW (1981) The evolution of leks. In </span><span style="font-size: small;"><i>Natural selection and social behavior</i></span><span style="font-size: small;">. (RD Alexander, DW Tinkle, eds). Chiron Press, New York, pp 138-169.</span></div>
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<span style="font-size: small;">Briga M, Pen I, Wright J (2012) Care for kin: within-group relatedness and allomaternal care are positively correlated and conserved throughout the mammalian phylogeny. </span><span style="font-size: small;"><i>Biology Letters</i></span><span style="font-size: small;">:</span><span style="font-size: small;"><i><span style="font-weight: normal;"> </span></i></span><span style="font-size: small;">p.rsbl20120159</span></div><div style="margin-bottom: 0in;"><span style="font-size: small;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-size: small;">Clutton-Brock TH (2021) Social evolution in mammals. <i>Science</i> 373(6561): doi:10.1126/science.abc9699. </span></div>
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<span style="font-size: small;">Clutton-Brock TH, Parker GA (1995) Punishment in animal societies. </span><span style="font-size: small;"><i>Nature </i></span><span style="font-size: small;">373: 209-216.</span></div>
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<span style="font-size: small;">Davies NB, Krebs JR, West SA (2012) </span><span style="font-size: small;"><i>Introduction to behavioral ecology</i></span><span style="font-size: small;">. Wiley-Blackwell, 4</span><sup><span style="font-size: small;"><span style="font-style: normal;"><span style="font-weight: normal;">th</span></span></span></sup><span style="font-size: small;"> edition. Oxford, UK.</span></div><div style="margin-bottom: 0in;"><span style="font-size: small;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-size: small;">Eisenberg JF (1981) <i>Mammalian radiations</i>. U Chicago Press.</span></div>
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<span style="font-size: small;">Jones CB (2014) </span><span style="font-size: small;"><i>Evolution of mammalian sociality in an ecological perspective</i></span><span style="font-size: small;">. Springer, New York.</span></div>
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<span style="font-size: small;">Keller L, Chapuisat M (2010) Eusociality and cooperation. In </span><span style="font-size: small;"><i>Encyclopedia of life sciences</i></span><span style="font-size: small;">. Macmillan, published online: DOI: 10.1002/9780470015902.a0003670.pub</span></div>
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<span style="font-size: small;">Ladev</span><span style="font-family: "times new roman" , serif;">è</span><span style="font-size: small;">ze S, de Muizon C, Beck RMD, Germain D, Cespedes-Paz R (2011) Earliest evidence of mammalian social behaviour in the basal Tertiary of Bolivia. </span><span style="font-size: small;"><i>Nature </i></span><span style="font-size: small;">474: 83-86.</span></div>
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<span style="font-size: small;">Sibley RM, Hone J, Clutton-Brock TH (eds) (2003) </span><span style="font-size: small;"><i>Wildlife population growth rates.</i></span><span style="font-size: small;"> The Royal Society: Cambridge University Press, Cambridge University Press, UK.</span></div>
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<span style="font-size: small;">Stearns SC (2000) Life history evolution: successes, limitations, and prospects. </span><span style="font-size: small;"><i>Naturwissenschaften</i></span><span style="font-size: small;"> 87: 476-486.</span></div><div style="margin-bottom: 0in;"><span style="font-size: small;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-size: small;">West SA, Fisher RM, Gardner RA, Kiels ET (2015) Major evolutionary transitions in individuality. <i>PNAS</i> 112(33): 10112-10119.</span></div>
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<span style="font-size: small;">Westneat D, Fox C (eds) (2010) </span><span style="font-size: small;"><i>Evolutionary behavioral ecology</i></span><span style="font-size: small;">. Oxford University Press, Oxford University Press, UK.</span></div><div style="margin-bottom: 0in;"><span style="font-size: small;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-size: small;">Wilson EO (1971) <i>The insect societies</i>. Belknap (Harvard), Cambridge, MA.</span></div>
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<span style="font-size: small;">Wilson EO (1975) </span><span style="font-size: small;"><i>Sociobiology: the new synthesis</i></span><span style="font-size: small;">. Belknap (Harvard), Cambridge, MA.</span></div>
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*Originally published in <i>International Society for Behavioral Ecology Newsletter</i>, 2016.<br />
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Addendum</b>: I want to apologize to Tim Clutton-Brock for this review which expresses, as much as anything else, my disappointment in his text--of which I had great expectations. My expectations were so high--I expected a synthetic masterpiece in the context of evolution and the Major Transitions Approach--something on the order of EO Wilson's, <i>The Insect Societies</i> (1971), but, for Mammals. Even a book like JF Eisenberg's, mammalian "radiations" (1981), but about mammalian social evolution--extending Eisenberg's relevant chapters--would have been a major advance for the field. Instead, Clutton-Brock's book, whose title is a misnomer, is little more than a review of many highly selected publications without a conceptual framework [e.g., The Hamiltonian Project (the General Law of Social Evolution); Major Transitions, including, Sociality (Cooperation, after Hamilton 11964), as well as, Complex Sociality: SPECIALIZATION: division-of-labor (Cooperation between specialists); reproductive DoL, task, role, &/or morphological specialization] without any treatment of general patterns, without standardization of terminology, with only a nod to, systematic, quantitative theory--including, Hamilton's rule, and all in the context of a colonial historical-academic framework [e.g., every photograph of people of African descent is of primitive groups save one, and that one of an African guide, apparently, on a hunting safari with white male tourists. Unless I am mistaken, only a very few women are highlighted. Surely, TC-B might have provided a nod to changing worldviews by highlighting one or more of his African & female colleagues who must exist since TC-B has been conducting research in Africa for decades; if TC-B has not had African collaborators, volumes are silently spoken [admittedly, I know nothing about his personal or academic or research affiliations or his views on race, ethnicity, class or gender; and, I "get" it--the "greats" in Animal Behavior are, mostly, white men--even, until recently, in Anthropology, & I defer to the historical record ... nonetheless, it would have been generous to include, say, "Griff" Ewer, as deserving of recognition, for example--if only for the sake of appearances, but, then, my own biases are showing; in the final event, it would not detract from C-B's legacy for him to have been more generous in his historical acknowledgements].In short, as one who has admired TC-B's exhaustively rendered empirical work for decades, my expectations and anticipations, as well as, intellectual curiosity, were totally deflated by the long literature review treated herein. A synthesis of mammalian Ecology & Evolutionary Biology, in other words, a text on Mammal Societies, awaits future treatment. John F. Eisenberg's 1981 book, "radiations," is the closest thing we have so far to a synthesis of mammalian social biology. 5/9/2020; slight edit, 12/30/2023 ...</div><div style="margin-bottom: 0in;">
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<a href="http://vertebratesocialbehavior.blogspot.com/2019/09/my-comments-on-eisenberg-jf-1981mammal.html">http://vertebratesocialbehavior.blogspot.com/2019/09/my-comments-on-eisenberg-jf-1981mammal.html</a><br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-41763560985040814182019-04-11T20:36:00.005-07:002022-06-20T22:38:36.945-07:00Review of Robert L. Trivers' Memoir, Wild Life (by Clara B. Jones, 2016)<div dir="ltr" style="text-align: left;" trbidi="on">
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<i>Wild Life: Adventures of an Evolutionary Biologist</i></div>
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Robert L. Trivers</div>
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Biosocial Research</div>
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New Brunswick, NJ</div>
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2015</div>
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225 pages</div>
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$9.58 (Paperback and Kindle)</div>
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Reviewed by Clara B. Jones, Asheville, NC, USA (February, 2016; slightly revised, 4/11/2019)</div>
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To dispense with preliminaries, I have interacted with Robert L. “Bob” Trivers intermittently since I was in graduate school but have never been in his “inner circle.” I requested a complimentary copy of <i>Wild Life</i> from Robert, telling him that I intended to review it for a journal that was, at the time, undetermined. Robert was enthusiastic about the idea and sent the memoir. I have reviewed a number of books for <i>ISBE Newsletter</i> so it was logical for me to consider it as the venue for publication, particularly, since I speculated that many readers would be interested. I appreciate Andreas Svensson for giving me this opportunity.</div>
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In addition to several other biologists (e.g., Louise Emmons, Dan Janzen, Russ Lande, Nancy Moran, Steve Stearns, Mary Jane West-Eberhard, Don E. Wilson, EO Wilson), I consider Trivers to be a National Treasure. Against numerous odds, he has risen to the top of his field, having published papers that are fundamental contributions to Social Biology, generators of immeasurable bodies of systematic and informal research and publications, as well as, continuing streams of productive thought—among his peers, other scientists, educators, students, journalists, and the general public. Many persons in and outside the scientific community will purchase a copy of <i>Wild Life</i> hoping, even, expecting, to find inside their names or mention of one of their vivid memories with Robert. Most of these men and women will be crushed, and I can think of four or five who will never recover. Robert's new book should have included a detailed index, not only for reference purposes, but, also, to prevent time wasted by some who would otherwise rifle through the book in search of evidence that Trivers considers them worthy of mention.</div>
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In the literary world, a memoir belongs to the genre, Creative Nonfiction. <i>Wild Life</i>, however, for the most part, appears to be uncontrived, a convincing example of a brilliant, though, controversial, figure telling it as he truly sees it. As the saying goes, “What you see is what you get.”, though Robert is clear to point out that he is unable to measure the full extent to which deception and self-deception influenced this project. After reading <i>Wild Life</i><span style="font-style: normal;">,</span> I was left with several messages, one being that its author has written a memoir hoping to determine the arc of his legacy. With the exception of Chapters 14 and 15, he crafts his story, as he has presented all of his ideas, with clinical, even, scapular, precision. There is a sense in which the dominant theme of the book can be summarized by a single observation expressed on page 186 when speaking of W.D. Hamilton: “I thought of Bill as perhaps the greatest evolutionary theorist since Darwin. Certainly, where social theory based on natural selection is concerned, he was our deepest and most original thinker.” Many would place E.O. Wilson in second place. However, in <i>Wild Life</i>, as well as, elsewhere, Robert seems of the opinion that Wilson is greatly overrated (an opinion with which I disagree). Though he does not say so explicitly, Trivers likely considers himself to hold second rank after Hamilton, a self-assessment that many would challenge if only because of the author's non-normative record of behavior. Nonetheless, as Robert told a reporter at Rutgers University in 2014, “<span style="font-style: normal;">I don’t want to sound immodest, but I am one of the greatest social theorists in evolutionary biology alive, period.”</span></div>
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<i>Wild Life</i><span style="font-style: normal;">, dedicated to Robert's teacher, William H. Drury, Jr., comprises fifteen chapters, seven of which, in addition to numerous additional remembrances, concern his life and research in Jamaica. He reports, with obvious pride, that all of his five children are “American/Jamaican,” referring to himself as an “out-breeder” created by his peripatetic experiences as the son of a career diplomat. For decades, I was an obsessive reader of the print-version of </span><i>The New York Times</i><span style="font-style: normal;">. One day in 1987, scanning the obituary page, I noticed the name, Howard Trivers [</span><i>sic</i><span style="font-style: normal;">, no middle initial], and, because, at that time, I had received second-hand reports of Robert's life from one of his close friends, I determined from relevant details that the deceased was the evolutionary biologist's father. Among other prominent roles in the U.S. Department of State, Howard Trivers had been Director of its Office of Research and Analysis. It was impossible not to note that the diplomat's obituary included no personal information—no mention of parents, a wife, or other family members. I had been told that Robert had a very contentious relationship with his father, in part, because of the latter's profession. Robert has only mentioned his father to me once, in passing; thus, I was not totally surprised to find virtually no reference to his family, including his mother, in </span><i>Wild Life</i><span style="font-style: normal;">. For this reason, combined with the, sometimes shocking, even, disturbing, particulars of his life, Robert would provide a Freudian scholar with copious material and room for speculation (e.g., Oedipal failure?). In addition to passing mention of his “family of origin” and his children, the Preface expresses the author's opinions about the conventional lives of scientists (“This kind of life never appealed to me.”), and the remaining few paragraphs introduce the reader to themes that follow.</span></div>
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Chapter 1 is revealing to all students of Trivers' work. He documents his early precocity in mathematics, particularly, The Calculus. I consider myself an amateur student of the thinking strategies of famous scientists. Many population ecologists, for example, think like physicists, characterizing genes and other events as mass flowing in space and time. Ecosystem ecologists, on the other hand, are likely to think, not only spatially, but at multiple scales at once. Robert's mind is that of a quantitative modeler expertly identifying and manipulating decision rules. A limitation but, even, more, a strength, of his mostly verbal theories is that they bypass complexity to identify fundamental principles of natural selection for mechanisms and functions of general import. Trivers reveals that Drury taught him to begin with interesting questions about human behavior, and Robert has exhibited a remarkable ability to choose topics basic to non-human, as well as, human, Social Biology, always cognizant of intra- and inter-individual conflict (-of-interest) and of genetics operating at the level of individuals, constrained by Hamilton's Rule. While I imagine there is intuition and art involved in Trivers' generative processes, I think his successes can be attributed, primarily, to his cogent choice of topics and his ability to write with laser clarity, as close as verbal models can come to mathematical ones. In my humble opinion, Robert is a better verbal modeler than Charles Darwin.</div>
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Chapters 2 and 3 describe early experiences conducting fieldwork and the impact of Drury and Ernst Mayr, respectively, on the development of the memoirist's early thinking, publications, and career. Robert convincingly communicates the extent to which he honors these men, and he shares with candor the pivotal role they played in the formation of some of his most important ideas. Trivers has reason to be confident that his acknowledgments of others' inputs will not reduce his reputation and that, though some critics may attempt to diminish his project, particularly, posthumously, Robert's theories are not subject to claims that they are derivative. Chapter 4 is titled, “I Become a Lizard Man in Jamaica,” describing how he became “a green lizard freak” after accompanying Ernest Williams there as a research assistant. Some readers will be offended by both men's evaluation of primatologists and, by implication, Anthropology as a field (but, see Chapter 6). In another chapter he denigrates Psychology, and it is clear throughout the memoir that Robert's opinion about what constitutes a Science is a narrow one.</div>
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<i>Wild Life</i><span style="font-style: normal;"> is peppered with interesting facts about lizards and other animals, as well as, snapshots of geography and human nature. Chapters 5, 7, 8, 9, and 10 recount a variety of experiences in Jamaica, some of them life-threatening, as well as, significant friendships, mostly with men. The only women who receive a lingering nod in the memoir are “mother-in-law,” “Miss Nini,” and her daughter, Robert's “wife,” Lorna, mother of four of his children. Even though these chapters, and a few other accounts, document Robert's capacity for deep, sincere, and reciprocated feelings, he sometimes refers to friends using clinical, seemingly detached, language.</span></div>
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Chapter 11 is an interesting one in which the author reminisces about his relationship to Huey Newton and the Black Panther Party, and Chapter 12 continues descriptions of his entanglements with intra-specific conflict, including, incarceration. Robert seems not to have learned a litany from Behavioral Ecology that the costs of aggression (or, spite) usually outweigh its benefits (see the self-analysis in Chapter 15, and Parker 1974). Chapter 13, titled, “Vignettes of Famous Evolutionary Biologists,” suggests that, in addition to Drury, Mayr, and Williams, Richard Dawkins and W.D. Hamilton, effectively, complete Trivers' list of illustrious figures in the field who have influenced him and his ideas, and in this chapter, Robert includes a few sarcastic paragraphs about Stephen J. Gould that I consider gratuitous and unnecessary.</div>
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The final two chapters are intimate in nature, Chapter 14 is titled, “Ambivalence About Jamaica,” describing the unsettling and increasing rates of violence there. Some readers will find the final chapter haunting since Robert candidly assesses personal failings and outlines his burial plans. I was surprised that he had nothing to say about politics or the state of the world (e.g., climate change, income inequality, racism, biodiversity loss, terrorism), if only to inform the reader about how his opinions and values might have changed since his Black Panther days (“One man's terrorist is another man's freedom fighter.”, as the CIA used to say). His life is not as dark as it may appear, however, since Robert has recently outlined several proposals for future research, specifically, speciation processes (with Koos Boomsma); evolutionary heterogamety; natural selection of honor killings; evolutionary dynamics of homosexuality; and, human evolutionary genetics.</div>
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<span style="font-style: normal;">It would be remiss and unrealistic not to provide some academic critique of Trivers' work, and I consider it appropriate to suggest a couple of limitations*. Since I was introduced to his publications in the 1970s by Behavioral Ecologists at Cornell, it has concerned me that, though "inclusive-fitness maximizing" and sexual selection are fundamental to Robert's writings, his body of work fails to reflect the importance of Evolutionary Ecology, particularly, life-history evolution (e.g., Stearns 1976), as well as, evolution in changing environments about which there is a significant literature pre-dating Robert's first publication (e.g., Levins 1968, Lewontin 1957). Though Evolutionary Ecology is relatively recent as a systematic discipline, Robert is aware of Population Genetics and G x E interactions, an operation receiving limited treatment in his publications (e.g., How do “selfish” genes and social traits </span><i>behave</i><span style="font-style: normal;"> when conditions vary or along gradients? When and under what conditions is social behavior situation-dependent? ...flexible?). As important as his neglect of evolution in heterogeneous regimes [Population Ecology] & of life-history evolution, Robert, in addition, fails to address Behavioral Ecology, especially, the fundamental & "classic" work of John Hurrell Crook (1964: group structure in response to limiting resource dispersion [distribution and abundance] in Time & Space). All of these concerns relate to the "condition-dependence"/"situation-dependence" of organismic responses & adaptation, or, lack thereof, in the context of "local adaptation."</span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;">On the other hand, Robert's “feel” for statistical thinking is that of an expert, and I would expect more treatment of variation and deviations from central tendencies in his canon. Of particular import, is that, regardless of having produced seminal papers in his early career, as well as, an interesting book on sociality (Trivers 1985), Robert has shown little interest in the EVOLUTION of sociality [as, Major Transitions Approach: West et al. 2015], per se, as per transitions from solitary breeding, to breeding in groups [including, group formation & group maintenance], to Presocial state, to Subsocial state, to Cooperation [as per Hamilton '64], & to Complex Sociality (reproductive division-of-labor; specialization as per Eusociality variously defined); see, Wilson 1971's schema]. Furthermore, treatment of social evolution in comparative perspective [within & between genera, families, orders, classes] is lacking in Robert's oeuvre.</span></div>
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<span style="font-style: normal;">On the other hand, one way that Trivers' work achieves elegance, is by simplifying complex phenomena; thus, environmentally-focused, realistic theories might not have been as successful, productive, or fundamental. In a sense, the author's insights are </span><i>primitive</i><span style="font-style: normal;"> in the deep, honorific sense that the word is employed to describe some treatments in pure mathematics. Nonetheless, all theoretical work of import is subject to vetting by subsequent theory, models, experiments and other empirical tests. Already, a few researchers have modified certain details of Trivers' theory of sex ratio selection, and his ideas about parental investment and sexual selection have been challenged by some feminist biologists. Related to any discussion of Robert's legacy, in </span><i>Wild Life</i><span style="font-style: normal;">, the memoirist states how important the appreciation of </span><i>conflict</i><span style="font-style: normal;"> (genetic and whole organism, intra- and inter-individual) has been to his success. In my opinion, this observation confirms Trivers' understanding that asymmetries produce differential “fitness optima” and that differential (asymmetric) phenotypes are exposed to environments upon which selection may act. I strongly recommend </span><i>Wild Life</i><span style="font-style: normal;"> to all who are interested in Ecology and Evolutionary Biology, not only, for its explication of “wild” experiences, but, also, for insights into how a stunning mind works.</span></div>
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<b>References</b></div>
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<div style="margin-bottom: 0in;"><span style="font-style: normal;">Crook JH (1964) The evolution of social organization and visual communication in the weaver birds (Ploceinae). </span><span><i>Behaviour</i></span><span style="font-style: normal;"> Supplement #10: 1-201.</span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;"><br /></span></div><div style="margin-bottom: 0in;">
<span style="font-style: normal;">Levins R (1968) </span><i>Evolution in changing environments: some theoretical explorations</i><span style="font-style: normal;"> (No. 2). Princeton University Press.</span></div>
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<span style="font-style: normal;">Lewontin RC (1957) The adaptation of populations to varying environments. </span><i>Cold Spring Harbor Symp Quant Biol</i><span style="font-style: normal;"> 22: 395-408.</span></div>
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<span style="font-style: normal;">Parker GA (1974) Assessment strategy and the evolution of fighting behaviour. </span><i>J Theor Biol</i><span style="font-style: normal;"> 224: 115-126,</span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;">Stearns SC (1976) Life-history tactics: a review of the ideas. </span><span><i>Quart Rev Biol</i></span><span style="font-style: normal;"> 51: 3-47.</span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;">Trivers RL (1985) </span><span><i>Social evolution</i></span><span style="font-style: normal;">. Benjamin-Cummings.</span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;">West SA, Fisher RM, Gardner RA, Kiels ET (2015) Major evolutionary transitions in individuality. </span><span><i>PNAS</i></span><span style="font-style: normal;"> 112(33): 10112-10119.</span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;"><br /></span></div><div style="margin-bottom: 0in;"><span style="font-style: normal;">Wilson EO (1971) </span><span><i>The insect societies</i></span><span style="font-style: normal;">. Belknap (Harvard), Cambridge, MA.</span></div>
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*<span style="font-family: "times new roman" , serif;">First, no American social biologist can compete with Wilson's expertise as a student of a single social group, ants, in Wilson's case. Second, IMO,</span><span style="font-family: "times new roman" , serif;"> </span><i style="font-family: "times new roman", serif;">The Insect Societies</i><span style="font-family: "times new roman" , serif;"> </span><span style="font-family: "times new roman" , serif;">(1971) is the greatest book ever written in Social Biology, indeed, in Animal Behavior and Ethology as a whole.</span><br />
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Third, unlike E.O. Wilson, Trivers is not a synthesizer, though his most heralded papers have general import. Trivers has not communicated much interest in a search for general patterns--within, between, and across taxa. Also, again, in contrast to E.O. Wilson, Trivers' canon pays scant attention to Genotype<----->Phenotype<----->Environment<-----> causes and effects. One seeks, for the most part, in vain, to locate Ecology--abiotic & biotic Environments [however conceptualized]--in Trivers' writings. Furthermore, to my knowledge, Trivers has not emphasized the topics--group-formation, group maintenance, and the environmental conditions facilitating the emergence of sociality [which may or may not follow group-formation]. Especially pertaining to the latter are the topics, competition and differential access to limiting resources, as well as, limiting resource dispersion [distribution and abundance in time and space].<br />
<br />
Trivers may be one of the last remaining extreme genetic thinkers. He typically asks a question, then, considers what consequences would obtain given alternate, pairwise combinations of related individuals [parents, full sibs: 1/2; first cousins: .1/8, etc.]. This approach has yielded several fundamental papers; however, Trivers' work does not satisfactorily address variations in inter-individual interactions nor evolution in heterogeneous regimes nor phenotypic plasticity nor the principle that behavior is condition-dependent. In other words, an actor & a recipient, whatever their "<i>r</i>" [coefficient of relationship], will respond relative to "<i>b</i>" [benefits to recipient] and "<i>c</i>" [costs to actor ("donor"--of reproductive units)], rather than, strictly, "<i>r</i>" [Hamilton's Rule: <i>rb</i> - <i>c</i> >0].<br />
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Trivers' literal logic based, apparently, on "<i>r</i>" alone, may reveal one unfortunate consequence of the term, "kin selection" that leads many to assume that it is always in <i>ego</i>'s favor to exhibit social behavior towards kin. The latter assumption may be an assumption behind Trivers' [very successful and justifiably heralded] publications. Furthermore, as I [and several others before me (thanks to James Marshall and Andrew Bourke for making me aware of this literature)] have suggested, it may be useful to consider the role of competition influencing behavior between actor [donor] and recipient and to question whether Hamilton's Rule adequately incorporates the consequences for actor and recipient and for the expression or non-expression of cooperation or altruism [i.e., "social behavior"] of interindividual competition [for limiting resources, e.g., food, mates, space].<br />
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Thus, sometimes, kin may be ego's "worst enemy," a litany of Behavioral Ecology [it may not be beneficial for ego to assist the reproduction of kin; it <b>may</b> be in the interest of ego to assist the reproduction--depending upon environmental regime] where predation is non-random by genotype [where cooperation or altruism toward a relative would increase ego's chances of becoming prey]. But, complicating the matter, in certain conditions, death, however, defined [e.g., self-induced, other-induced], can benefit kin. Clearly, systematic empirical and theoretical studies, in addition to modeling, are needed.<br />
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In the final analysis, however, the impressive success of Trivers' verbal models based on "<i>r</i>" may demonstrate the power of Hamilton's Rule to predict a very broad array of the social acts [cooperation, altruism] observed in Nature, including, Human Nature. However, we should not only ask, "What is "<i>r</i>"?, but, also (or rather?), "'<i>r</i>' relative to what?" According to Hamilton's Rule, the effects of "<i>r</i>" are expected to be constrained by the components that comprise "<i>b</i>" & "<i>c</i>".</div>
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<span style="font-style: normal;"><br /></span><span style="font-style: normal;">Clara B. Jones (</span><span style="color: navy;"><span lang="zxx"><u><a href="mailto:foucault03@gmail.com">foucault03@gmail.com</a></u></span></span><span style="font-style: normal;">)</span></div>
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Asheville, NC, USA (now, Silver Spring, MD, USA)</div>
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<span style="font-style: normal;">February, 2016 In </span><i>International Society of Behavioral Ecology Newsletter</i></div>
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-23362495767233537522019-04-10T12:11:00.001-07:002020-08-31T19:42:20.692-07:00Review of E.O. Wilson's new book, Genesis (Clara B. Jones, 2019)<div dir="ltr" style="text-align: left;" trbidi="on">
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Genesis:
the deep origin of societies</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Edward
O. Wilson</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">2019</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Liveright
Pub. Co. (W.W. Norton & Co.)</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">153
pp</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">$15.88</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Reviewed
by Clara B. Jones, Ph.D. (2019)</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><br /></span></span>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">"The key to the sociobiology of mammals is milk." EO Wilson (1975)</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Social
evolution is an important topic of investigation by behavioral
ecologists and evolutionary biologists. The two categories of
sociality, cooperation and altruism (Hamilton 1964), have arisen
infrequently across vertebrate taxa because, <b>in propitious
environmental regimes</b>, group-level coordination and control is
usually derailed by “cheaters” who fail to comply with group
norms. As Wilson pointed out in 1971, groups of cooperators and
altruists characterize the most “successful” (i.e., widely
distributed) extant terrestrial taxa—social insects and humans. In
his new book, </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Genesis
</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><span style="font-style: normal;">[</span></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>sic</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><span style="font-style: normal;">]</span></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">,
the entomologist, E.O. Wilson, winner of the Crafoord Prize in 1990 and America's premier social biologist*, assesses the
emergence of eusociality, the highest social “grade” (Wilson
1971). </span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><br /></span></span>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Perhaps the primary contribution of this brief book is that
Wilson classifies humans as eusocial, a system characterized by
overlap of generations, cooperative brood-care, and non-reproductive
"helpers." If Wilson is correct, humans would be
classified, </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>"primitively"
eusocial</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
(Wilson, 1971) or Totipotent Eusocial [TE; see "General Mammalian Patterns" blogpost, this blog, #28, 9/19/19], since most human "helpers" (except
post-menopausal females or other sterile persons) are expected to be
"totipotent"—"helpers" capable of independent
reproduction, able to reverse their non-reproductive status [TE]. Members
of permanently sterile “castes,” are labeled, </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>“advanced”
eusocial</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
(Wilson, 1971), and Wilson's treatments in this book suggest to me
that he might be inclined to label permanently non-reproductive human
groups as “caste”-like, associations that should be investigated, as well, for the possible presence of "temporal division-of-labor" ("age polyethism": see Wilson, 1971).. </span></span>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">The
first five chapters of </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Genesis</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
include limited explications of some topics (e.g., “multi-level”
selection, “phenotypic plasticity”). Wilson clearly explains that
the conceptual frameworks of </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Genesis</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
are Maynard Smith & Szathm</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">á</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">ry's
(1995) classic treatment of “major transitions of evolution,” as
well as, “multi-level” and “group selection,” terms used
interchangeably. </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">In
Chapter 6, Wilson appears to be primarily interested in proffering a
defense for Charles Darwin's explanation for the evolution of sterile
castes—an argument based on group selection which Wilson defines as
follows: "...within groups, selfish individuals win against
altruists, but groups of altruists beat groups of selfish
individuals" [attributed to David Sloan Wilson]. Among impediments to "proofs" of "group selection," defenders need to show how "cheaters" are controlled within groups, demonstrations that will require empirical studies.</span></span><br />
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Here
and throughout the book, Wilson fails to incorporate the ecological
literature showing, for example, that intragroup competition is
generally stronger than intergroup competition or that behavioral
ecologists have, since the early 1980s, advanced general criteria for
the evolution of cooperative groups (e.g., Emlen 1982) and of eusociality
(e.g., Crespi 1994; also see, Choe & Crespi 1997, Crespi et al. 2004 [see, especially, p 66 & pp 73-74], Bourke & Franks 1995, Bourke 2011); West-Eberhard 1975 on p 169, col. 3, par. 2 in Queller & Strassmann 1998 pp 169-170). More
specifically, Wilson fails to cite other researchers who have
advanced the idea that humans are eusocial (e.g., Foster &
Ratnieks 2005, Jones 2011, Crespi 2014).<br />
<br />
Nonetheless, combined with
related studies (e.g., Emlen 1982, Emlen 1984, Hrdy 2011), there
seems to be an expanding literature justifying systematic and
quantitative investigation of eusociality in humans, in particular,
and in vertebrates, broadly, including, standardization of
terminology, experiments, and modeling (e.g., “agent-based”
modeling). For an early, published paper that might generate ideas
for these future projects, Lotka (1928) is suggested.</div>
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The final chapter (7), titled, “The
human story,” reviews “transitions” to eusociality across apes,
from chimpanzees (<i>Pan troglodytes</i>), as well as, bonobos (<i>P.
paniscus</i>) continuing to <i>Australopithecus</i> and the <i><span style="text-decoration: none;">Homo</span></i>
line. Unlike other chapters, this one emphasizes the importance of
ecological factors (habitat) for the evolution of social mechanisms
among hominids and their ancestors, and Wilson endorses the “social
brain hypothesis” as well as the importance of fire for the “rapid
evolution” of large brains and the facilitation of group-life,
respectively—as well as, their consequent adaptations.
Interestingly, in this chapter (p 114), the author compares human
eusociality to other social mammals, in particular, African wild
dogs, demonstrating that he is prepared to classify “other mammal
species,” eusocial, in addition to the social mole rats (see Jones 2014, pp 48-52)****. For another interesting example, see Dwarf Mongoose.</div>
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Wilson does not dismiss “kin
selection;” but, he holds that “multi-level” or “group”
selection is the primary driver of the route to eusociality, behind
which kin effects may follow [he may be right; see this lecture, "Ecology Of Societies," by Simon Levin of Princeton:<br />
<br />
<a href="https://www.youtube.com/watch?v=rQUsApf3RHs">https://www.youtube.com/watch?v=rQUsApf3RHs</a> ].<br />
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Most social biologists are certain to
be surprised to read Wilson's claims that "Hamilton's Rule"
suffers "fatal weaknesses" and is no longer "useful (see Bourke 2011, Bourke & Franks 1995)."
Wilson does not support these flippant statements with mainstream
literature about which there is wide consensus in favor of Hamilton's
Rule, and nowhere in his text does he assess assumptions underlying
considerations of differential benefits to recipients of social
behavior (cooperation or altruism) or differential costs to "donors,"
terms subsumed in Hamilton's Rule (see, for example, Bourke 2011,
Marshall 2015). Related to this, Wilson all but completely avoids
optimality [cost-benefit] thinking, and social biologists will, I
think, find his explication of group selection obfuscating when
applied to genetics, including the assertion that population
geneticists have shown the verity of group selection.<br />
<br />
Nonetheless,
researchers, including, evolutionary psychologists, human biologists,
and anthropologists, will derive many testable hypotheses from
Wilson's claims, among the more provocative of them, the statement
that division of labor by human professional categories is evidence
of eusociality and group selection (that they are "caste-like)." I am led to wonder if some human
guilds might be characterized by high <i>r</i> (coefficient of
relationship), a possibility that would be easy to test. Likewise, guilds, and other formal human groups (e.g., fraternities and sororities, political groups, and the like), should be investigated for evidence of "temporal division-of-labor" ('age polyethism": see Wilson, 1971).</div>
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<div style="margin-bottom: 0in;">
In service to economy, organization,
and clarity, <span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Genesis</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
might have been more wisely presented as a "tight"
technical paper rather than a manifesto in book form, though Wilson
deserves to be applauded for advancing bold ideas, for insisting that
human social behavior be subjected to the same analyses that we apply
to non-human animals, and that, ultimately, evolutionary explanations
will need to be "gene-centered," a long-standing hallmark
of Wilson's approach (e.g., Wilson 1975) and that of the heralded
evolutionary biologist, Robert Trivers (see, e.g., Trivers 1985 and the remarkable Trivers & Hare, 1976**; also see citations by Bernie Crespi). I
recommend this creative and controversial text to specialists,
students, and the general audience. It will raise many questions,
stimulate thought, and, hopefully, generate conversations*** and
research** about variations in human socio-sexual units, as well as,
the origins and evolution, the causes and consequences, of group life
across all vertebrates.</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><b>References</b></span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Bourke
AFG (2011) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Principles
of social evolution</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Oxford University Press, Oxford.</span></span><br />
<span style="font-family: "times new roman" , serif;">----, Franks NR (1995) <i>Social evolution in ants</i>. Princeton [NJ] University Press.</span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Choe
JC, Crespi BJ (1997) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>The
evolution of social behavior in insects and arachnids</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Cambridge University Press, London.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Crespi
BJ (1994) Three conditions for the evolution of eusociality: are they
sufficient? </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Insectes
Sociaux</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
41(4): 395-400.</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Crespi
BJ (2014) The insectan apes. </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Human
Nature</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
25(1): 6-27.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Crespi BJ, Morris DC, Mound LA (2004) <i>Evolution of ecological and behavioural diversity: Australian</i> Acacia <i>thrips as model organisms</i>. Australian Biological Resources Study, Canberra.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><br /></span></span>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><a href="https://www.environment.gov.au/science/abrs/publications/thrips">https://www.environment.gov.au/science/abrs/publications/thrips</a></span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><br /></span></span>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Emlen
ST (1982) The evolution of helping I: an ecological constraints
model. </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>American
Naturalist</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
119: 29-39.</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Emlen
ST (1984) Cooperative breeding in birds and mammals. Pp 305-339 in
</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Behavioral
ecology an evolutionary approach</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">,
2</span></span><sup><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">nd</span></span></sup><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
ed. (JR Krebs, NB Davies, eds.). Sinauer, Sunderland, MA.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Emlen ST, Oring L (1977) Ecology, sexual selection, and the evolution of mating systems. <i>Science</i> 197: 215-223.</span></span></div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Foster
KR, Ratnieks FLW (2005) A new eusocial vertebrate? </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Trends
in Ecology and Evolution</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
20(7): 363-364. </span></span>
<br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Frank SA (1995) Mutual policing and repression of competition in the evolution of cooperative groups. <i>Nature</i> 377: 520-522.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">----(2003) Repression of competition and the evolution of competition. <i>Evolution</i> 57(4): 693-705.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Hamilton
WD (1964) The genetical evolution of social behavior. </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Journal
of Theoretical Biology</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
7: 1-52. </span></span>
<br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Holldobler B, Wilson EO (1990) <i>The ants</i>. Belknap (HUP). Cambridge.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Hrdy
SB (2011) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Mothers
and others</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Belknap-Harvard.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Jones
CB (2011). Are humans cooperative breeders? A call for research.
</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Archives
of Sexual Behavior</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
40(3): 479-481.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">----(2014) <i>The evolution of mammalian sociality in an ecological perspective</i>. Springer, NY.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Lotka
AJ (1928) Sterility in American marriages. </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>PNAS</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">
14(1): 99-108.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Marshall
JAR (2015) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Social
evolution and inclusive fitness theory: an introduction</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Princeton University Press, Princeton, NJ.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Maynard
Smith J, Szathm</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">á</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">ry
E (1995) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i><span style="font-weight: normal;">The
major transitions of evolution</span></i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
W.H. Freeman Spektrum, New York.</span></span></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Queller DC, Strassmann JE (1998) Kin selection and social insects. <i>BioScience</i> 48(3): 165-175.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Trivers
RL (1985) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Social
evolution</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Benjamin-Cummings Pub. Co., San Francisco.</span></span><br />
<span style="font-family: "times new roman" , serif;">Trivers RL (2015) </span><i style="font-family: "times new roman", serif;">Wild Life</i><span style="font-family: "times new roman" , serif;">. Biosocial Research. New Brunswick, NJ. [& e-book, amazon.com] </span><br />
<br />
<span style="font-family: "times new roman" , serif;"><a href="http://vertebratesocialbehavior.blogspot.com/2019/04/review-of-robert-l-trivers-memoir-wild.html">http://vertebratesocialbehavior.blogspot.com/2019/04/review-of-robert-l-trivers-memoir-wild.html</a></span><br />
<br /></div>
<div style="margin-bottom: 0in;">
<span style="font-family: "times new roman" , serif;">----, Hare H (1976) Haplodiploidy and the evolution of the social insects. <i>Science</i> 191(4224): 249-263.</span><br />
<div>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">West-Eberhard MJ (1975) The evolution of social behavior by kin selection. <i>Quarterly Review of Biology</i> 50: 1-33.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Wilson
EO (1971) </span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>The
insect societies</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Belknap/Harvard, Cambridge, MA.</span></span></div>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">----(1975)
</span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><i>Sociobiology</i></span></span><span style="font-family: "times new roman" , serif;"><span style="font-size: small;">.
Belknap/Harvard, Cambridge, MA.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Wittenberger JF (1980) Group size and polygamy in social mammals. <i>Am. Nat</i>. 115: 197-222.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Yamamura N, Higashi M (1992) An evolutionary theory of conflict resolution between relatives: altruism, manipulation, compromise. <i>Evolution</i> 46: 1236-1239.</span></span><br />
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;"><br /></span></span>
<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">*First, no American social biologist can compete with Wilson's expertise as a student of a single social group, ants, in Wilson's case. Second, IMO, <i>The Insect Societies</i> (1971) is the greatest technical book ever written to date (and <i>The Ants</i> [Holldobler & Wilson, 1990] the greatest popular book ever written to date) in Animal Behavior and Ethology.</span></span><br />
<br />
Third, unlike E.O. Wilson, Robert L. Trivers, also a renowned social biologist, winner of the Crafoord Prize in 2007, is not a synthesizer, though Trivers' most heralded papers have broad import. Trivers has not communicated much interest in a search for general patterns--within, between, and across taxa. Also, again, in contrast to E.O. Wilson, Trivers' canon pays scant attention to Genotype<----->Phenotype<----->Environment<-----> causes and effects. One seeks, for the most part, in vain, to locate Ecology--abiotic & biotic Environments (however conceptualized)--in Trivers' writings. Furthermore, to my knowledge,<br />
<br />
Trivers has not emphasized the topics--group-formation, group maintenance, and the environmental conditions facilitating the emergence of sociality (which may or may not follow group-formation). Especially pertaining to the latter are the topics, competition and differential access to limiting resources, as well as, limiting resource dispersion (distribution and abundance in time and space). A litany of Behavioral Ecology is that, in some environmental regimes, kin may be ego's worst enemy. Such conditions may occur, for example, where there is intense local competition for limiting resources [above some critical threshold of decreasing reproductive gains].<br />
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Trivers may be one of the last remaining extreme genetic thinkers among living social biologists (see my review of his memoir, <i>Wild Life</i> [2015] linked above). He typically asks a question, then, considers what consequences would obtain given alternate, pairwise combinations of related individuals [parents, full sibs: 1/2; first cousins: 1/8, etc.]. Trivers' canon is about mechanisms, not, about causes. His approach has yielded several fundamental papers; however, Trivers' work does not satisfactorily address variations in inter-individual interactions nor evolution in heterogeneous regimes nor phenotypic plasticity nor the principle that behavior is condition-dependent nor the litany of Behavioral Ecology that patterns of group-living will "map" onto dispersion [distribution and abundance in time & space and abiotic phenomena (e.g., climate)] of limiting resources nor the ideas that females are energy maximizers--males, time minimizers. Trivers does not seem to be sensitive to Hamilton's "<i>b</i>" & "<i>c</i>" whereby an actor & a recipient, whatever their "<i>r</i>" [coefficient of relationship], will respond relative to "<i>b</i>" [benefits to recipient] and "<i>c</i>" [costs to actor ("donor"--of reproductive units)], rather than, strictly, "<i>r</i>" [Hamilton's Rule: <i>rb</i> - <i>c</i> >0].<br />
<br />
Trivers' literal logic based, apparently, on "<i>r</i>" alone, may reveal one unfortunate consequence of the term, "kin selection" that leads many to assume that it is always in <i>ego</i>'s favor to exhibit social behavior towards kin. The latter assumption may be an assumption behind Trivers' (very successful and justifiably heralded) publications. Furthermore, as I [and several others before me (thanks to James Marshall and Andrew Bourke for making me aware of this literature)] have suggested, it may be useful to consider the role of competition influencing behavior between actor [donor] and recipient (see Yamamura & Higashi, 1992) and to question whether Hamilton's Rule adequately incorporates the consequences for actor and recipient and for the expression or non-expression of cooperation or altruism (i.e., "social behavior") of interindividual competition for limiting resources (e.g., food, mates, space, etc.).<br />
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<span style="font-weight: 400;">Thus, sometimes, kin may be ego's "worst enemy" [it may not be beneficial for ego to assist the reproduction of kin; it </span><b>may</b> be in the interest of ego to assist the reproduction--depending upon environmental regime] where predation is non-random by genotype [where cooperation or altruism toward a relative would increase ego's chances of becoming prey]. But, complicating the matter, in certain conditions, death, however, defined [e.g., self-induced, other-induced], can benefit kin. Clearly, systematic empirical and theoretical studies, in addition to modeling, are needed.</div>
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In the final analysis, however, the impressive success of Trivers' verbal models based on "<i>r</i>" may demonstrate the power of Hamilton's Rule to predict a very broad array of the social acts (cooperation, altruism) observed in Nature, including, Human Nature. <span style="font-family: "times new roman";">However, we should not only ask, "What is "</span><i style="font-family: "times new roman";">r</i><span style="font-family: "times new roman";">"?, but, also (or rather?), "'</span><i style="font-family: "times new roman";">r</i><span style="font-family: "times new roman";">' relative to what?" According to Hamilton's Rule, the effects of "</span><i style="font-family: "times new roman";">r</i><span style="font-family: "times new roman";">" are expected to be constrained by the factors comprising "</span><i style="font-family: "times new roman";">b</i><span style="font-family: "times new roman";">" & "</span><i style="font-family: "times new roman";">c</i><span style="font-family: "times new roman";">". For example, when and under what conditions are kin, enemies?</span><br />
<span style="font-family: "times new roman";"><br /></span>
<span style="font-family: "times new roman";">**See comments on this paper in Holldobler & Wilson (1990, p 184)</span></div>
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***For example, cooperation and altruism might occur by way of "mutual policing," coercion, force, or persuasion, in addition to, "self-restraint" (see, especially, Steve Frank's work, in particular. 1995 and 2003).<br />
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****Addendum 4/26/2019: Regarding the ideas that humans are "cooperative brreeders" (e.g., Hrdy 2011) and that "cooperatively-breeding" mammals should be classified, "eusocial" (Jones 2014), I am going to go out on a limb to suggest that, based upon my reading to date on mammal, including, human, group-living patterns, mammalian "cooperative breeding" evolved from groups of communal females, and their offspring, that may or may not, then, evolve to share "economic" tasks, including care of young [see epigraph]...since females, <i>ceteris paribus</i>, are expected to be "energy-maximizers," thermal efficiency, broadly defined, will be paramount as a selective factor and in determining optimal "inclusive fitness maximizing" (see Jones 2014, Chapter 3). Anthropologists specializing in human "cooperative breeding" hold that the system evolved via monogamy, as it is thought to have done in insects and birds. However, mammalian sociosexual systems generally exhibit "sexual segregation" ("solitary" dispersion between the sexes) and, in mammals, monogamy is derived. Related, Mammal sociosexual systems and group structure are thought to result from the tendency of females to select rich patches of [limiting] food and that of males to select the largest possible number, sometimes, an aggregation, of females (Wittenberger 1980, Emlen & Oring 1977).<br />
<br />
In summary, <span face="" style="background-color: white; color: #222222; font-size: x-small;">it seems to me that the more likely "route" to eusociaity in humans would have been via communally nesting mammalian ancestors, particularly, since monogamy is "derived," mammals are very likely to exhibit a "sexually-segregated" ["solitary"] socio-sexual organization, monogamy is relatively rare, and, in humans, at least, monogamy is often imposed by some authority. Routes to eusociality that might be applied to humans have, also, been proposed by West-Eberhard (1975) and by Queller & Strassmann (1998: "fortress defenders" and "life insurers") [see Queller & Strassmann (1998) for a brief summary of these ideas on pp 169-170]. </span>These issues need to be unpacked.</div>
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<span style="font-family: "times new roman" , serif;"><span style="font-size: small;">Clara
B. Jones is a retired behavioral ecologist living in Silver Spring,
MD (USA). </span></span></div>
<br /></div>
Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-50448314142312543542019-03-13T18:56:00.002-07:002019-07-20T13:43:49.548-07:00When to exhibit Social Behavior [Cooperation, Altruism]? Clara B. Jones [Graph, Legend]<div dir="ltr" style="text-align: left;" trbidi="on">
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjoUFNLWGdz0QOubbS0mAPaH_RPNmkwbXxI2mo46l06tTR2uwHgXM_w_n0G1p5xr1Fq75fsukB1TX3LLo7AmBfEWJfL3IExNW2boQd-PJctzyx4tgMGSP0EUSxxwN7sR5sX2jnOSf53dhuD/s1600/CB-IF-TE-Graph2.jpg" imageanchor="1" style="margin-left: 1em; margin-right: 1em;"><img border="0" data-original-height="1086" data-original-width="1324" height="262" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjoUFNLWGdz0QOubbS0mAPaH_RPNmkwbXxI2mo46l06tTR2uwHgXM_w_n0G1p5xr1Fq75fsukB1TX3LLo7AmBfEWJfL3IExNW2boQd-PJctzyx4tgMGSP0EUSxxwN7sR5sX2jnOSf53dhuD/s320/CB-IF-TE-Graph2.jpg" width="320" /></a></div>
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<b>Fig. 1. </b>When should an Actor or a Recipient exhibit Social* Behavior [Cooperation, Altruism]? Clara B. Jones, 3/13/2019<br />
<br />
<b>1.</b> This [idealized] graph**** depicts & predicts when an [idealized] organism--an Actor or a Recipient--should give/exhibit Cooperation or Altruism, defined, formally, as those interactions during which the Recipient benefits [in relative reproductive units].<br />
<b>2.</b> X-axis: Time [T], Energy [E] investment in Social Behavior [reflects organism's T/E budget for a given interaction].<br />
<b>3.</b> Y-axis: Costs [C, in reproductive units to Actor or Recipient] or Benefits [B, in reproductive units to Actor or Recipient] to Inclusive Fitness** [IF= selfish reproductive units (usually, & clasically, offspring, though, offspring might be delayed, such as when transaction in wealth or other goods--see "Altruism" below) + reproductive units of kin]. <i>r</i>, coefficient of relatedness, is incorporated in measure/estimate of IF [how to measure?].<br />
<b>4.</b> C rise or reach asymptote over T. <b>Costs, for Actor or Recipient, or both, may prevent Cooperation or Altruism between kin when, for example, Cooperation or Altruism increases Competition [for limiting resources such as food, mates, burrows], increasing [for <i>ego</i> or <i>ego</i>'s offspring] likelihoods of dispersal, "competitive exclusion" [demand for ~identical limiting resources], death, etc. Cooperation and Altruism are expected, by definition, to be responses for the avoidance or neutralization of Competition.</b><br />
<b>5.</b> B rise, then decrease over T.<br />
<b>6.</b> "x" represents the optimal point at which an Actor or a Recipient should exhibit Cooperation [both Actor and Recipient gain reproductive units] or Altruism. [Actor loses, Recipient gains reproductive units]. "x" is the extension to the X-axis of the vertical, black line connecting the points with the greatest separation between the C and B curves--where B > C. "x" is the idealized point maximizing inclusive fitness benefits.<br />
<b>7.</b> The shaded area represents the zone around "x" for which Actor's benefits from Social Behavior are highest. Ideally, the broken vertical lines extend vertically & downward from those points on either side of "x" on the B curve at which the B curve first begins to descend. The shaded area represents the zone of maximal benefit to Inclusive Fitness of A or B, in this case, maximum benefit from an act of Cooperation or Altruism ["IF maximizing"].<br />
<b>8.</b> Actor & Recipient curves and values will usually be asymmetric, by definition [possibly, even, for clones (e.g., identical twins)]. In other words, zones of maximal benefit from social behavior should differ for Actor and Recipient. Overlaying Actor's & Recipient's curves will determine whether there is overlap between Actor & Recipient zones of greatest benefit to <i>ego </i>from Cooperation or Altruism. If there is overlap, the zone of mutual benefit is defined, so that Actor & Recipient should exhibit Cooperation [+, +] within these parameters [of T, E]. If there is no overlap, then conflict arises [between Actor and Recipient &, possibly, kin of one or both].<br />
<b>9.</b> Similarly, an Actor should exhibit Altruism [-, +] to a Recipient when the long-term B of such action[s] outweigh the short-term C [determined by hard-wired Hamiltonian algorithm?].<br />
<b>10.</b> If B from Social Behavior are minimal or non-existent [e.g., low T/E "budget;" little or no overlap between Actor-Recipient zones of greatest benefit], then interactants should adopt an alternate behavior [Alternative Reproductive Behavior--tactic or strategy], such as fight***, avoidance, flight, force, coercion, persuasion, manipulation, exploitation, coexistence--or some combination of these.<br />
<b>11.</b> Group-living organisms experience relatively ongoing "decisions" about how to behave relative to conspecifics; however, it is important to keep in mind that "fitness budgets" change over T and that behavior is condition- [context-] dependent.<br />
<b>12.</b> <i>Ceteris paribus</i>, and on average, curves, B & C, as well as, shapes of zones of maximum benefits for Males [time-minimizers] and Females [energy-maximizers] should differ. Curves, also, are expected to be Age-, Class-, & Role-dependent.<br />
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*After Hamilton (1964), "Social Behavior" is defined as Cooperation [both Actor and Recipient gain reproductive units] or Altruism [Actor forfeits reproductive units; Recipient gains reproductive units]<br />
**The term "inclusive fitness" can be confusing since, technically, "fitness" is the frequency of an allele in a population averaged across the reproductive success of all individuals in the population bearing the allele, leading one to inquire how important it is to measure individual variation of traits [since individual variation will be subsumed in the measure of an allele's "fitness"].<br />
***As pointed out by Geoff Parker [1974], aggression should not usually be a first resort because the C of damaging behavior generally outweigh the B.<br />
****The same graphical/conceptual framework could be used for Coexistence if axes modified so that X-axis= savings in T, E to Mean fitness of allele 1 [or 2]; Y-axis= B, C from T, E savings to fitness [Mean frequency] of allele 1 [or 2] in a population. In the case of Coexistence, a Community Ecology process, the population would be escaping Competitive Exclusion. [See, also, my 2014 Springer Brief, Ch 2, for discussion of Coexistence in relation to Hamilton's Rule.]<br />
<br />
Acknowledgments: Thanks to my son, Luke [M. Luke Jones] for stimulating discussion and for drawing the figure. Further, via e-mail, I asked Andrew Bourke [Univ. of E. Anglia] to comment on this blogpost. On 4/11/2019, I received the following reply [n.b.: my original blogpost above, including, the graph, have NOT been revised subsequent to Bourke's comments]:<br />
<br />
<div class="m_5680186934002065603m_-3377720628254791425MsoListParagraph" style="background-color: white; color: #222222; font-family: Arial, Helvetica, sans-serif; font-size: small;">
<span style="color: black; font-family: "arial" , sans-serif; font-size: 10pt;">1.<span style="font-family: "times new roman"; font-size: 7pt; font-stretch: normal; line-height: normal;"> </span></span><u></u><span style="color: black; font-family: "arial" , sans-serif; font-size: 10pt;">Perhaps the X axis needs clarifying a little – after all, time and energy seem to have most relevance in this context insofar as they are proxies for fitness, so then there would be some relationship between the metrics on the X axis and on the Y axis, and how these interact might then require consideration.<u></u><u></u></span></div>
<div class="m_5680186934002065603m_-3377720628254791425MsoListParagraph" style="background-color: white; color: #222222; font-family: Arial, Helvetica, sans-serif; font-size: small;">
<u></u><span style="color: black; font-family: "arial" , sans-serif; font-size: 10pt;">2.<span style="font-family: "times new roman"; font-size: 7pt; font-stretch: normal; line-height: normal;"> </span></span><u></u><span style="color: black; font-family: "arial" , sans-serif; font-size: 10pt;">It would be good to have concrete, empirical examples of when the scheme in the figure needs invoking, i.e. showing how considering the time/energy investment dimension in the manner advocated in the figure enlarges our understanding of the occurrence of social behaviours. Personally I find it hard to think of things in the abstract all the time, so even some hypothetical biological examples/applications might be useful.<u></u><u></u></span></div>
<div class="m_5680186934002065603m_-3377720628254791425MsoListParagraph" style="background-color: white; color: #222222; font-family: Arial, Helvetica, sans-serif; font-size: small;">
<u></u><span style="color: black; font-family: "arial" , sans-serif; font-size: 10pt;">3.<span style="font-family: "times new roman"; font-size: 7pt; font-stretch: normal; line-height: normal;"> </span></span><u></u><span style="color: black; font-family: "arial" , sans-serif; font-size: 10pt;">I think your footnote 8 makes a good point, i.e. that actor and recipient interests need not overlap even for altruistic and cooperative interactions. This point was explored theoretically in a fairly old paper by Yamamura and Higashi (1992)*****. Their work does not seem to have been followed up empirically to any great extent, perhaps because the usual assumption that one of the parties has most control over the interaction (in which case conflict is minimised – e.g. an insect larva may often have little choice but to accept the level of care it’s given) holds.<u></u><u></u></span></div>
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<b>References</b><br />
Hamilton WD (1964) The genetical evolution of social behaviour. <i>J Theor Biol</i> 7: 1-52.<br />
Parker, G (1974) Assessment strategy and the evolution of fighting behavior. <i>J Theor Biol</i> 47: 223-243.<br />
*****Yamamura N, Higashi M (1992) An evolutionary theory of conflict-resolution between relatives: altruism, manipulation, compromise. <i>Evolution</i> 46: 1236-1239.<br />
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<a href="http://www.kurims.kyoto-u.ac.jp/~kyodo/kokyuroku/contents/pdf/0827-14.pdf">http://www.kurims.kyoto-u.ac.jp/~kyodo/kokyuroku/contents/pdf/0827-14.pdf</a><br />
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.comtag:blogger.com,1999:blog-4604556790862046588.post-34332881772820565562019-01-01T11:06:00.000-08:002019-01-01T11:06:05.273-08:00Fragment: Inferences from West-Eberhard (1975) pertinent to Mammals (Clara B. Jones)<div dir="ltr" style="text-align: left;" trbidi="on">
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><b>2.4
Inferences Pertinent To Mammalian Sociality Can Be Drawn From
West-Eberhard (1975)</b></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">West-Eberhard’s
(1975) summary of general models of social evolution reducible to
mechanisms dependent upon inclusive-fitness maximizing is useful as a
reminder that Hamilton’s rule is manifested in several ways,
dependent upon local condition, sex, role (e.g., reproductive or
helper), and lineage. It is noteworthy that each of the six
strategies (1a – 3b) is, in one manner or another, applicable to
social mammals and to mammalian females living mutualistically or
cooperatively in groups. In insects and most mammals, olfaction is
the primary mechanism of communication. Thus, coordination and
control of conspecifics is expected to be constrained by the
spatiotemporal dynamics of chemical properties (rapid delivery,
relatively rapid decay, pheromonal repression of selfishness).
Despite interspecific similarities, it is possible to derive
inferences from West-Eberhard’s (2005) outline that may pertain,
especially, to social mammals. </span></span>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
1: Because the variance on reproductive success is lower among
females than among males, </span></span><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><i>ceteris
paribus</i></span></span><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">
(Trivers 1972), mammalian females will be more closely related, on
average, to other females in their group or population than will
males be, on average, to each other, to group or population females,
and to each other’s offspring. For the same reasons, a mammalian
population is likely to include more females than males, and females
are more likely to exhibit sociality.</span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
2: Trivers’ (1972) model, fundamentally, concerns differential
energetic investments by males and females and the life-history
trajectories deriving from polymorphic allocation patterns. Thus,
because female “fitness budgets” are more constrained
energetically that those of males, the reproductive female component
of a group or population is expected to be more stable,
spatiotemporally, than the dispersion of males in the same
population. For the same reason, on average, female turnover is
likely to be lower, female survivorship higher, female emigration
rates lower than for males. </span></span>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><b>Verbal
Model I</b> was advanced by Trivers (1972) and was not, originally,
presented as a general model of social evolution. However, this
author predicts that social evolution will be biased by initial
reproductive allocations or energetic investments (Schoener, 1971).
This verbal model is not limited to evolution by sex, encompasses all
conditions in which organisms make (“Hebbian”) “decisions”,
including, decisions to join (group-formation), or, remain in
(group-maintenance), groups, and has the potential to be developed,
qualitatively, and, expressed, quantitatively, as a synthetic
formulation.</span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">//////////////////////////////////</span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
3: On average, since females are “energy-maximizers”,
“inclusive-fitness maximizing” is expected to mitigate energy
losses, and, on average, females should benefit from transferring
some component “fitness budget” to others. Thus, females and
their female offspring are likely to be the major donors and
beneficiaries of the benefits of cooperation and altruism, and,
relative to males in her population, an adult female has more to gain
in her reproductive lifetime from facilitation.</span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
4: Since males, “time-minimizers”, are expected to favor direct
over indirect reproduction, “inclusive-fitness maximizing”, a
relatively time-intense strategy, is unlikely to characterize
reproductive males, on average. </span></span>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
4: </span></span><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><i>Ceteris
paribus</i></span></span><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">,
and, depending upon threshold reproductive effects relative to
ecological conditions, where males cannot discriminate their own
young, they should be indiscriminately selfish, concentrating on
mating allocation strategies rather than facilitation of
conspecifics. </span></span>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
5: It follows from “parental manipulation” and “maternal
control” models that females can discriminate their mothers and,
there own offspring, where females are not “promiscuous” and do
not express “favoritism” of a single male during an estrus cycle.
On the other hand, cortical circuitry may be favored, permitting
females to make “decisions” based on likelihoods of paternity.
In order for this neurophysiological strategy to be favored by
selection, losses from error must not, on average, compromise
relative reproductive success (of the pertinent genotype). It is
important to note that “decisions” based on kinship may yield
lower group sizes than those based on selfish strategies (Hamilton
1964); thus, the Malthusian optimum for a reproductive female may not
concord with idealized mean fitness of her population.</span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Inference
6: Following from previous inferences, a reproductive female, on
average, is expected to lose less from probabilistic tactics and
strategies than will a reproductive male of the same population.
Related to the previous inference, “inclusive-fitness maximizing”
is expected to mitigate the maternal investment : ageing tradeoff for
tactical and strategic females, on average, possibly explaining the
reproductive advantage of extended post-reproductive lifespan in
humans and killer whales: ////.</span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><br /></span></span></div>
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<span style="font-family: Times New Roman, serif;"><span style="font-size: small;">The
previous discussion of West-Eberhard’s (1975) classification
(1a-3b) suggests that “inclusive-fitness maximizing” should be
more characteristic of reproductive females, on average, than of
reproductive males in the same conditions. It is hypothesized that
this condition obtains since, theoretically, females are expected to
be “energy-maximizers”. The latter life-history strategy should
privilege time-intense (“non-damaging”) rather than
energy-intense (“damaging”) strategies, theoretically,
characteristic of males. Should the aforementioned allocation
(thermal) trajectories withstand quantitative, including,
experimental, testing, each of them will yield information pertinent
to the evolution of mammalian sociality. In particular, on average,
each reproductive morph in the same population will respond
differentially to density effects that are expected to impact
“energy-maximizers” to a greater degree than “time-minimizers”
since an increase in population density should correlate positively,
</span></span><span style="font-family: Times New Roman, serif;"><span style="font-size: small;"><i>ceteris
paribus</i></span></span><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">,
with increased intensities of competition. Under these regimes,
reproductive females should “switch” to or increase dependence
upon, time-intense, energy-saving, helper tactics and strategies,
including, in some regimes, self- (suicide, “give-up” points) or
offspring-elimination (foetal resorbtion, abortion).</span></span></div>
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Clara B. Joneshttp://www.blogger.com/profile/09089578792549394529noreply@blogger.com