Social Selection Examples Turtles Salamanders Anurans

On the evolution of empathy based reward sensations in Reptiles and Amphibians

(salamanders)Chemical application > (frogs and toads)Physical atomic mass/sound motion -> (turtles) motor pattern mediated sound communication > (snakes) tactile exchange SALAMANDERS Salamanders posses pheromones that are either wafted in the water or applied directly to the nares of others. Co-evolution of the receptors and the pheromones may found the basis for motivational state communication in urodels and provide a reward sensation to pairs forming groups that are attempting to affect the number of offspring surviving into the next generation.

ANURANS Ryan suggests that neuroanatomy affects speciation rates in frogs and toads (PNAS 1986). He does recognize that other traits which show a trend in line with the AP may confound the influence of the inner ear on the number of species in a lineage but can not think how any other known trends could affect the speciation process like call type frequency change in his technological analog (signaler receiver) applied to behavioral ecology. The basic correlation that Ryan is attempting to explain as a causality is the increased number of species with more complex (AP) Amphibian Papilla.

Furthermore, Ryan makes the claim that proposed explanations of observed evolutionary changes in AP structure which may represent adaptive modifications has no bearing on the relationship between neuroanatomy and speciation ostensibly because the he asserts only a proposal of a species-selection argument could reverse the cause and effect scenario Ryan writes to establish.

BUT as Roughgarden has said that although previous work describes evolutionary mechanisms that promote or stabilize different social behaviors, we still have little understanding of the factors that drive animal behavior proximately. Robert Wright balked at Rougardens project of social evolution citing that complex adaptations are too hard to explain clearly. Ryans thinks that only a species selection argument can distrupt his cause for the correlation because he utilizes a simple technological message carrier analog for how the anuran ear trembles amongst individuals. He reduces the species specific character of call type to frequency and does not recognize that behavior may modify preference for the extremes of the call type rather than the mean (Scaphiopus species will mate with other species since females prefer the extremes when more species are present but the mean when only one is present) This is not necessarily a species-selection rebuttal but a social selection hypothesis and the mutations may be polymorphic under plastic behavioral traits rather than simplistically cognized by Ryan to simply to increase the domain of frequencies available (probability of frequency change) for neuroanatomical synthesis and processing. So if the anuran chorus involves timbre as well as frequency and duration as well one-off call-mating cause and effect which can be affected by the sounds of others in calling for the evening and there is a difference in the affect on offspring survival depending on calling and mating early vs call throughout the season and mating at different times a species-selection alternative is not the only thing that can overturn Ryans explanation of the correlation. Instead, use of the AP to affect Nash bargaining can explain the correlation causally as well. The very large difference in the number of species which use AP types A,B,C vs D suggest that the linear increase in frequency rather than a more complex (possibly two dimensional) electric mechanical communication continuum under Nash Bargaining is not the cause of the observed correlation. TURTLES Reward Sensations Determine Cooperation in a Repeated (behavioral) Game. Reward sensations have been proposed as the (driving force/ physiological instantiation) behind learnably affectable motivational states of individual organisms that can be maximized in social interactions through modeled objective functions. This theoretical framework embeds a cooperative behavioral dynamic within an evolutionary dynamic and leads to the/an idea that empathy-based altruism may be more common in nature than otherwise recognized. Here we propose that turtle foreclaw display is such a behavior. We suggest a detailed physiological manifestation underlying the motivation and show that the physics of the proximal behavioral factor entails no essential difference between a preference or strategy/action in the repeated games played out in any behavioral time so embedded. A cooperative behavioral dynamics embedded in an evolutionary dynamic has been proposed (7) as necessary to understand the evolution of social behavior

It has been a presumption that fore-claw display behaviors objective is courtship fore-play causally prior to mating. This has been canonized/ aggregated through the use of the term titillation we still have little understanding of the factors that drive animal behavior proximately. we study goal-seeking behavior in the context of a social interaction by developing a model of a pair of interacting animals whose motivations derive from their internal reward sensations, which they aim to maximize. These reward sensations, which could be encoded in specific neural circuits such as dopamine pathways involved in learning (11), are represented in our model as objective functions. detailing a possible motivational kinematics grounded in the physical phenomenon of resonance within the behavioral actions of fore-claw display as the reward sensation physiology. The physiological mechanism sets limits to the extent that repeated games can be played to effect (only a certain quantity of resonance induced reward sensations are possible for each behavioral bout or series of bouts (resonance is not infinite)). Within single depolarization there is a limit that resonance reaches as it achieves peak depolarization and maximal deflection. There is spontaneous deviations that resonanat with incoming fore claw forced motions to reach maximal peak depolarizations. Thus even small reward enticements (few nanometers) if resoanant can affect actual depolarizations. By combined cell effects other higher uses of resonance is possible. Our model rests on motivations to achieve objectives This paper suggests how some turtles may motivate each other. the dynamics of a social interaction at the behavioral time scale This is composed of the fore-claw display in its various forms and relations. other-regarding motivations In this case are the ablity of the displayer to motivate the other through physical resonance in the inner ear. The proximal mechanism of resonance repeated (game wise)(different motor patterns and individuals)

previous work describes evolutionary mechanisms that promote or stabilize different social behaviors, we still have little understanding of the factors that drive animal behavior proximately.

Most directly, field observations of an other-regarding individual would see it spontaneously undertaking actions that could be costly to itself but benefit another individual with whom it interacts. This spontaneous helping should occur even among unrelated individuals and in situations in which immediate reciprocity is unlikely

Fore-claw display occurs between genera of turtles and thus is spontaneous in this sense. If the fore-claw display indeed supplies the physical force (as here proposed) that causes sensations in the other then the metabolic cost of the motor pattern is born by the purveyor in behavioral time and accruing to the benefit of the other but evolutionary stable if the result is a cooperative bargain. If the visual cues (posturing, eye blinking) determine reciprocity, the actual action to vibrate the fore-claws is independent of any immediate reciprocity. Posturing and eye-blinking (direct assay of motivation) thus would neurologically reveal actual care for the pay-off of the other. Change of behavior from fore-claw display to biting (in melanistic individuals) may represent transitions to direct pay-off expression. Responsiveness is not a global description of the behavioral dynamics as this involved both gene changes in the hair cells themselves as well as costs and benefits of different motor patterns/ posturing and because the result of a behavioral interaction is to increase the resonance and thus the reward there is no distinction between a strategy and behavioral rule both lead to the same but not in global descriptor but rather are driven by payoff-based motivations, and is an index summarizing the feedback brought about by the individuals' motivations in the vicinity of the behavioral equilibrium. In general, can describe a wide variety of evolved behavioral feedbacks; thus, our behavioral model represents a less-restrictive approach to how animals make decisions. It is a double feedback system (within the nervous system and between the individuals). SN AKES Snakes have evolved the ability to use their muscles to constrict prey. The muscles have also been supposed to provide heat to eggs in some Pythons. I have witnessed a curious behavior in a pair of Corn snakes that have grown up together since young. It appears that the snakes help each other out and cooperate on finding prey (dead mice). They have never been fed live mice and thus they must use their olfactory sense principally to locate the food when placed in the cage. One snake prefers to grab the prey as soon as it is placed in the cage and the other to locate it after touching it repeatedly. Although the snakes have been observed to bite at the prey in the mouth of the other (they tear the mice to pieces) and to bite the other, both snakes perform an odd muscle flexing of the entire body (only when prey is present) which causes one snake to jerk alongside the other. This appears to help the snakes differentiate the other snake from the rest of the cage and to possibly signal where the prey is not? It may be that through touch snakes can communicate socially. It has long been observed that male rattle snakes will rub against each other (to get the upper side? Or just to show where the female is not?) and the dens of garter snakes are well known. This may be a gimbal based neurobiology of mutual pleasure and not a war of any kind. That fangs are not used for injury is irrelevant and a part of the older sexual selectionist perspective view.

The information is developed out of the 7 1-D freize patterns in which sound is decomposed and analyzed depending on the goal. The differences in the material substrates (different amount of electrical or mechanical affects) lead to back and forthing , individual threats, and teamworks in pairs or larger firms.(contra Hulswit 1996)

The proximate reward sensation is the efficient cause of the ultimate acquisition of the trait. This trait may contain a double quaternion use where individual threat points are operative asymmetrically thus providing a purely computational interval to friably separate material teamworks regardless of the physical waveforms (depending on how pleasure is neurobiologically instaintiated in turtles). Social selection thus can provide a kind of dynamics that is not reducible to the optimization framework suggested for universal biological dynamics.
An optimization framework of biological dynamical systems Ryota Horie Transcendetals categorize the countably infinite recovered in the social selection evo tier but remain logically within a superset of the noncountably infintly divisible sound types.

The difference between Williams and Mayr use of Pittenridgh depend on how infinity is found in the analysis.

What is the origin of that information (1-D symmetries in force impacting space creating sounds and pressures and waves) and the development of canals to separate double quaternions into single quaterions (lateral lines from lancelets to morphologically made torus quaterions. With recapituatlions.) . It is possible to discuss explanatory and descriptive teleonomy within social selection. Teleonomic description depends on just what 1-D symmetries are in the environments past and present of the teleonomy so being explained. These symmetries may arise as by products of the behavior computer as somatic program recapitulated, through teleomatic necessities, or due to otherwise adapted features. The evolutionary tier is not preprogrammed but historical and depends on many things other than just the behavior computers churning while changing. This churning and changing is in this case followable as the entropy measure of the teleonomic program from juvenile to adult behavior as the different quaternionic monomial become determinant for the entropy as the belt-trick is algorthmitized out of various homogneous polynomial functions of differing degress. It happens to coincide with development of melanin. The proximate reward sensation is the efficient cause of the ultimate acquisition of the trait. The proximate causes (quantum mechanical 4pie rotations in chemical bonds) mirroring this ultimate social selection have yet to be identified but are likely to be parts of the mechancanical electrical transduction means in the hair cells.

Social selection begins with the offspring and works back to the mating from there. Thomas has observed that female turtles will obtain displays from young males but not older males which concurrently display rather to the younger. Here I suggest that social evolution in Trachymes occurs as coalitions founded by melanistic individuals of displaying turtles form. A turtle displayed to form a team

A turtle displayed and formed a team What is the goal of turtle organization (to untwist sound using a particular kind of belt (horizontal vs anterior /posterior) This approach may be supportive of the ecological theory of orientation which does not support the notion of gravinertial coordination. Head Tilt signals may provide quaternionic information to divide the effects of double quaternions from left and right regular function differentials of single ones.

These coalitions form because they end up producing more offspring than simply mating more. The coincident self interest in the coalitions arise as teamworks that avoid recapitulated threat point deviations from perfect teamwork in locked vestibular pleasure circulate in a population. One can have a focused attention on natural selection through past selections as recapitulations against threat points in pleasured based teamworks as coalitions format the phenotypic trait. Thus there is no theoretical reason to deviate from Roughgarden as Potchnick thinks. It is an empirical case if turtles actually do follow this or not.
Berry and Shine (1980) set out a general viewpoint on sexual selection in turtles. They characterize the foreclaw display behavior as purely precoital functioning to stimulate the female to accept intromission. This category is compared to forced intromission and male-male combat. This division is then divided as indicating female choice in the first but male sexual selection in the 2nd and third. The idea follows Darwin purely and simply that the females choose the males with longer finger nails and smaller body sizes but modifies this on Ghiselns idea of contact rates with varying population density suggesting the that aquatic turtles with the display resulted from highly mobile turtles under Ghiselns view where females could get away from males and males had to be highly mobile to get the girls and mate. The display is part of coerceing the female into mating and somehow the females choose to be coerced by long nailed displayers. This is the kind of scenario that social selection was devised to provide an alternative for. Gibbons and Lovich (1990) questioned the notion of relying on Ghiselns additive when speaking of turtle mating systems but continued to view turtle nail traits as a part of sexual selection differing in how sexual selection may operate in turtles.

I then look at what the social selection view vs sexual selection view presents for geographic differences in Trachymes and discuss the evolution of the aquatic emydids.

The notion of natural selection causing a turtle to enjoy working as team is applied here. Synergy is accomplished by synchronizing vestibular outputs to pleasure centers (dopamine regions) where each turtle can choose to increase synchrony or individual threaten (biting, moving away) to revert to individualism. The coalitions that form are in part external (visible groups of turtles) but also internal (effecting similar operation of somatic programs behavior computers). If the turtle only displays(self-regarding) Show picture but does not vibrate or pulsate (show picture) then though it may receive pleasure and obtain an NCE (same with other conspecific that only accepts the first pulsation or whatever it pleases rather than allowing its quaternions to lock). The NBS can however be realized if the sound input is allowed to direct the locking of quaternions (other regarding) and the mulitiplcation (exponentially weighted product of the fitness incremenets) by combining the vestibular and sound outputs after untwisting. That is the possibility of perfect teamwork and is case where the causality of the somatic program is materially determining. Imperfect teamwork may occur if the programs are not exactly the same (the math formalism whereby the symmetries are recorded does not contain all of the information on the social evolution as then understood) and some kinds of chance are involved. Binmore insistence to follow Rubestein and insist on individual attainment of the NBS can tested in this model if its physical instantiation is as presented since both the individual and the team works can be distinguished in the molecular chain of events physiologically. The molecular chains of events that enable an individual hair cell to fire or not is not evolutionary biology nor is it social evolution but insofar as it can format the topology of the proximate reward sensation of selectable social infrastructure it bears on the evolution quite directly. I use the social infrastructure selection as a means to decide why some ontogenetic developments need somatic programs and why others do not (difference between places of double and single quaternions). Irreversible ontogenetic development of melanin does not need a somatic program but growth of nails that occurs within the same temporarlity does. Here the vestibular system foreclaw physical impacts provide the somatic program for sound detection. Sound apperception requires the recapitulation of the lateral line cilia vestibularly. Recapitulation can have a renewed understanding if the evolutionary tier can be dissected out of the behavioral tier. When we consider social selection in a ll emydid turtles for instance or how sound is used by turtles in general will follow.

Here we show how social selection allows one to unconfound the immediate physiological mechanism from the evolutionary history in only some turtles. Application to sound use will later be compassed. The description of the goal-direction is clear, it is to untwist the sound into translations and reflections pleasurably and the untwist can occur differently depending on which quaterion axes are directed towards.

How we come back and know what is maintained by natural selection is to find how social infrastructures are selected and how 1-D symmetries are dense thoughought the environments that they are evolved from. It is Lewontins creature constructs environment constructs creature scenario. We cash in the different countably infinite counts for infinite divisions of volumes restricted per density and we do worry about Aristotle (there is a minor confusion in how intelligent man made design relates to purposes invetable). Natural selection is NOT a program in Mayrs sense (contra Thompson), social selection through somatic programs are! The NBS determines the location of the somatic programs goal direction. Social selection can occur without direct ontogenetic irreversibilities however. Social selection is an explanatory system that enables direct definition of descriptive teleonomy. Social selection sensu WestEberhard can not do this. We describe the 1-D environment but explain the particular programs use of particular 1-D symmetries. It is not circular. This places anti-Gould heterochornic space inbetween any velocities (accelerations retardations) and thus make a heterogenous set of polynomials but restricted in dynamics they can record given past recapitulations. The open vestibular-sound program produces the adapted feature of long claws as a property of the somatic program socially selected.

Salamanders
Involuted coalitional model for social selection in salamanders. Payoffs are associated with pheromones and compensations are additionally provided by the coalition that forms creating demes that maximize rescources avaible to offspring through parental investment and extra-pair/within coalition paternity.

It is postuatled that following Iyer and Rougarden that tail ornaments are badges for entry into rescourse controlling cliques which with the

evolution of other families the biding becomes too high such that supplemental stratigies develop different coalitions for access to different rescources and continue by alternation of complement and supplements and intra and extra coalitional social selection.

They state, one rationale is that ornaments serve to avoid further conflicts once the stable coalitional structure is attained, for instance by acting as signals or badges that an individual belongs to the more powerful clique. Such a trait would be beneficial if the cost of producing the ornament is less than the cost incurred in the conflict that the display of the ornament avoids.

The crest of newts may be thought of avoiding the conflict of matching pheromone and receptors (the evolution of other regard) and signaling possession of offspring beneficial pheromone mutation and reception which would serve to visually enhance the reward sensations and obviate further syncing of the receptor and pheromone. This sets up the cost of the pheromone receptor dance to be higher than the cost of the crest. The condition with respect to offspring is dependent on the coalitional payment of compensation through paternity and the payoff due pheromone-receptor syncing on behavior and not represented by the crest. This correlation (between condition and crest) can develop during the course of the year and may be responsible for the differences between species with and without crests. Admission into larger coalitions is wholly due to the payoffs of receptor-pheromone co-evoltuion as it affects extra pair paternity in increasing access to

rescources of the offspring and or ameliorating issues due to gametic contact rates.

The ornaments function under this as bids for entry into dominant rescource controlling cliques but because the crest ornaments do not indicate condition there may be crest variations that correlate with bidding and those that correlate secondarily with condition while being associated with removing the need for one on one conflicts

Pleurodeles waltl<iframe width="560" height="315" src="http://www.youtube.com/embed/6PwNkB7E7ZA" frameborder="0" allowfullscreen></iframe> seems to use ribs to keep female moving in circle with male by providing equal and opposite force to the forward circling thus keeping the female inside rather than outside a spiral. The ribbed newt can swing ribs forward 50degrees http://brainrobot.wordpress.com/2009/08/22/snikt-amphibian-stabpredator-with-sharp-ribs/ when agitated and if the ribs are also used to keep the female from going forward then the male clasp (under arms) could be the perception by the male that the female had gone forward and so he moves his forlegs forward to catch her before she gets an angle away from him.

is it possible that all salamanders move their ribs during courtship? Is the tail straddle walk associated with a straightening of the spine??

The circular tail-straddling walk of the clouded salamander, Aneides ferreus: a deviation from the highly conserved linear tail-straddling walk of the Plethodontidae Authors: Sapp, Jerod R.1; Kiemnec-Tyburczy, Karen M.2 Source: Amphibia-Reptilia, Volume 32, Number 2, 2011 , pp. 235-243(9) http://www.ingentaconnect.com/content/brill/amre/2011/00000032/00000002/art00009

It could be that Aneides circles first to get the straight spine that other plethondids already use.

Can/DO SALAMANDER Cooperative Games supplant the sexual selection scenario below?

Selection on pheromones may be the result of natural or sexual selection (Arnold and Houck 1982), and our salamander data cannot distinguish between these possibilities. Stabilizing selection on the delivery system seems to argue against sexual conflict, in which pheromone delivery or costs of mating reduce female fitness while increasing male fitness (Parker and Partridge 1998; Chapman et al. 2003). In this scenario, female resistance to males drives signal diversification, leading to perpetual coevolution of signals and receptors and to high levels of polymorphism within populations (Gavrilets 2000; Gavrilets and Waxman 2002). Sexual selection can produce an evolutionary pattern in which female receptors constantly change as a correlated response to the evolution of male signals (Lande 1981) or as a result of male exploitation of a female bias towards a complex signal. Natural selection, for example arising from virally encoded cytokine mimics (Moore et al. 1996), or drift acting in females might result in an ever-changing population of receptors that males must track (Lofstedt 1993). Additional observations are needed to distinguish between these selection scenarios http://mbe.oxfordjournals.org/content/21/6/1032.full

Sequence homology places PRF in the same cytokine family as interleukin-6 (IL-6). These cytokines act as soluble ligands that sequentially bind extracellular domains of at least two transmembrane receptors and so bring together cytoplasmic domains capable of signal transduction (Bravo and Heath 2000). A four-helix bundle forms a structural core of the proteins to which receptors bind at conserved positions (Kallen et al. 1999; Bravo and Heath 2000; Hill, Morea, and Chothia 2002).

The act of vaccination/scratching behaviorally induces extracellular domain mixing (by skin cutting) which may result in repetitive spurious signal transduction during courtship. If this serves as a reward into the nervous system then a behavior Nash bargaining equilibrium could be set up under the premaxillary cutting behavior associated with the sodeferin molecule of older cloacal tail wafting behaviors (>50-100mya) and other behavioral /evolutionary equilibria . If soluble cytokines are present in the saliva it is possible that the non-mental gland behavior head sliding in Gyrinophilus is a behavioral tier activity that plays into the ESS of mental gland pheromone establishment. Thus social selection shows that this behavior is not a precursor nor is independent of the molecular evolution but part and parcel of a two tier system of social evolution. The individual scratching (independent of the older pheromone vaccination affect) with spurious/accidental reward signaling may function to reinforce male dorsal tail chemical uptake (also independent of mental gland action) and thus the behavioral bargain of straddling to achieve internal

fertilization may become even more cooperative.

Thus increased payoffs would accrue to behavioral interactions that had more cytokine variability exposed and lead to the result that the behavioral tier activity effects an evolutionary tier outcome.

The physical act of scratching (thus depending on the tooth size and quality, the depth and kind of scratches and the actual micro-behaviors) works as a sidepayments for an older pheromone reception and nasolabial groove uptake of tail chemicals bargaining agreement. In this scenario the transition to olfactory delivery of the cytokine in 19mya Plethodon clade is not a decoupling of the behavioral and the evolutionary molecular tier kinematics under positive selection but the dynamical outcome of an even better equilibrium unifying behavioral and molecular evolution together (physical damage to the skin is eliminated and presentation to the nose which sense the tail anyway gets to the same place nervously than the less direct skin route).

Changes in the female receptors to this positive selection may show relation in to changes in tail uptake chemicals. This would not be predicted sexual selection male-female Lande 1981 exponential correlational model which only would operate in the restricted binding site spaces of the PRF and be independent of micro behavior ethograms.

Description of the individual (keep the older bargain, limit physical injury, general cytokine variation) and team fitness functions (increased side payment, more direct alignment of straddle). It may be that cooperative salamander games that synergistically associate ciliary neurotrophic factor (CNTF) to cytokines during skin cutting by the maxilla and hence increase motor neuron survivability and increase motored behavior thusly drives the transformation from vaccination to olfactory pheromone delivery in salamanders rather than sexual selection (assumed on the molecular level) epsiodes.

Two different papers have sought to justify the suspicion that there is phenotypic-molecular

evolutionary decoupling in pheromone trait associations that was masked Stabilizing Selection on Behavior and Morphology Masks Positive Selection on the Signal in a Salamander Pheromone Signaling Complex Molecular Biology and Evolution Volume21, Issue6 Pp. 1032-1041 2004 Evolutionary replacement of components in a salamander pheromone signaling complex: more evidence for phenotypic-molecular decoupling. Evolution. 2007 Jan;61(1):202-15.

Instead this may simply be the that any strict ESS found in a restricted strategy space will only be neutrally stable against mutants from a larger strategy space that lead to the same behavioral equilibrium which representing a less-restrictive approach to how animals make decisions than those models where responsiveness is as a global description of the behavioral dynamics.Here rather than those where thereare both the local and global dynamics in the model driven by payoff-based motivations there is an index summarizing the feedback brought about by the individuals' motivations in the vicinity of the behavioral equilibrium. This can be driven by positive selection in there pheromone receptor relations where complementarity syngerstically reinforces both the behavior and the point mutations and need not occur by sexual selection per say.

Proc Natl Acad Sci U S A. 2009 November 10; 106(45): 1906119066. A theory for the evolution of other-regard integrating proximate and ultimate perspectives Erol Akay,1,2 Jeremy Van Cleve,1,3 Marcus W. Feldman, and Joan Roughgarden

In an attempt to understand how different pheromone components evolve Kiemnec et al 2009 create a model that focuses on mating (The probability of a male inseminating a female will not only be a function of her reproductive status and how attractive she finds the male, but will also be affected by her need to forage and her need to engage in predation avoidance.)

Evolutionary shifts in courtship pheromone composition revealed by EST analysis of plethodontid salamander mental glands Gene 2009

It is proposed that over the phylogeny plethodontids from salamandrids that Sodefrin pheromone transitioned with others through PMF and other chemicals to PRF in a recent clade of Plethodon using a different behavior. It was noted that while Sodefrin and PRF tend to decrease courtship time PMF taken alone increases it. What if instead the social behavior of salamanders was dependent both on the individual advantages but also on a kind of team play not merely to mate but to accomplish the most number of offspring into the next generation. Thus the invidivuals could have a threat point where they depart from team play and revert to individual actions where the team play is to elongate the entire breeding courtship season (much like frog breeding choruses are of different lengths). Salamandrids may be thought to use cleaved sodeferin as an attractant to a place while PMF may have functioned to increase the breeding season. Thus the simple model focusing on probability of insemination fails to take into effect the behavioral feedback of prolonged courtship and failure to inseminate but ability to return to be inseminated at another time in the extended season. would be possible for Plethodontids but not Salamandrids or Ambystomatids where the straddling behavior and spermatophore uptake outside the water is the behavioral equilbrium being targeted. This could lead to new behaviors , the establishment of a new pheromone such as PRF and the raised PMF in the newer situation as the NBS transitions to a different ESS while individual actions find a cooperative solution of team play.

Here we present a salamander courtship objective (motivated by evidence suggested in recent studies of pheromone exchange behavior and molecular

biology) (function) to the effect that ultimate molecular changes interrelate with the proximate behavioral goal to evolve a restricted ES strategy space subject to syngerism and complementarity that expresses pareto- efficient equilibira in a two tiered structure.

The goal is terrestrial spermatophore uptake (internal fertilization as an adaptation) and the behavior is the male-female straddle walk. The possible female-receptor male-pheromone co-evoulution is thus the result of complementary investments of the two individuals and not due to sexual selection (choosy receptor females and promiscuous pheromone broadcasting males).

Mechanical to Chemical cooperative NBS (some lack of chemical transfer in NOtphlamus) to investment pheromone-recpetpor point mutation *(minus the cost to invest (dependence on mental gland route rather than choice to use genial gland or tail wafting) positive se3lections rather than sexual conflict based receptor-pheromone affect through runaway processes The mechanical processes use enlarged sperm size in the spermatophore location depenednet on the breeding pond substrate while chemical pheromone straddle walk locates the placemenent of the re-enlarged sperm size capsule at the place of the bodies (adult locations) of the species rather than the habitat of the breeding pond(young locations) Spermataphore gelatin is a contact property. Internal fertilization in salamanders from broadcast fertilization may have resulted in situations where gamete contact rates could possibly fall below that which gave rise the difference in size of egg and sperm to begin with. Spermatophore gleatinazations were cases where chemical response to this effectively increased the size of the sperm. In a geographic to bodily positioned evolution the index of assortativitiy may be expected in geographic situation but not necessaritly in the bodily position position. Thus by a detailed understanding of the molecular evolution of the pheromone-trait positive selections we may be able to differentiate runaway sexual selection, strategy evolution which limits the behavioral input, and preference or social selection evolution. It may turn out to be a case that the evolution of plethodon olfactory transmittal behavior arose from social evolution through a bottleneck of strategy evolution as the sexes began to invest in the straddle walk and its chemical foundations otherwise simply directed in the mechanically forced internal fertilization (to keep the number of gamete encounters high enough). This will explain how individual proteins can have both highly conserved and highly variable

regions. So under strategy evolution where it is theoretic al supposed that a realistic feature of many social dilemmas is the net benefit of contributing becomes smaller as others contributions increase will result in some parts of he pheromone being conserved as the other contribhtuion becomes smaller but if it variable if the contribution of the males itself is smaller (provided within the same dilemma/game/strategy space)) The conserved region also applies to the conservation of the pheromone itself. So the transition from Soderfin through PMF to PRF with different variances of positive selections and conservagtions of regions will be due to actual strategy evolution (if a common explanation can be made for all mutant changes (on both scales) relative to scratching to olfacgtory behviro change suppoing it is in the same social dilemma that transformed soderfin from the clocal to the mental gland OR preference evolution if behaviors are more strongly correlated with synergismtic conditional regard. Alternatively the behavior to trait extablishment/expression may be even more losslely connected by Fishers runaway sexual selection. Thus by looking at the positive selection mutantant loci in the various pheromones we may be able to evaluate both how acute the angle is back towards an incumbent (male-female receptor-phermone prior amino acid placement) trait and the general format of the evolution gradient. This works because the geographic localiazation in addition to setting up restricted strategy space of the behavioral equilrbium also establishes the altruism/spite baseline from which strategy evolution may have been conjectured while both are independent of runaway sexual selection where males are cut of the process (as so far modeled by Lande 81) into the breeding seasons which is increased by a straddle walk regardless of whether this is due to strategy or preference.

Contrarily is the view of Reis and Houck

In contrast to the sexual selection view consider the possibility PRF as a resource founded in Eastern Plethodon and that he development of olfactory delivery occurred through coalitional formation of gland ornamentation and attendant behavior. Alternative PRF morphs would not arise as special cases of male and female templates but rather from coordinated intraspecific competition that excluded the cinerus group. Thus Darwin may be wrong to suppose that newt tail ornamentation is caused by sexual selection if it is a badge of coalitional structure for sodferin chemicals that can only be maximally affected by wafting the water that dilutes the coalitional signal and thus causes overly costly development of the ornament when compared to developing a more direct delivery method as in Plethodon.

So rather than being badges of fighting or competitive exclusion of other males it may signal a better ability to attract females chemically (Darwin seems mistaken to think of the male finding the female when the ornament may be part of the communicating the attractant to the female) and thus to have in its directly back lineges created more offspring. It may work to attract males to the area where reproduction is occurring and upbraids the importance of a particular location as having brought forth generations in the past. Thus like the possible use of frog calls the chemical system in salamanders may give individuals differential access to good breeding resource areas through the formation of coalitions of ornamenting individuals (that produce more offspring) he number of matings themselves.

Salamanders provide an interesting group to work out the distinction of natural selection, sexual selection and social selection. The concept of social selection has been critically inspected by Milam to the effect that the concept may not be able to be distinguished from some intuitions on sexual selection. Is social selection a strict alternative to sexual selection? What would the practical implications of strategy evolution and preference evolution amount to? to the extent that (whether reproduction is an exaptation for evolvability) Salamanders may a fairly unique group where cognitive abilities ( a particular system of reward sensations) to enter into an agreement to care for offspring arose fairly de novo. Nusbaaum has divided salamander parental care into three classes (lentic , lotic and terrestrial) and note that parental care is lacking in lentic populations/species. Thus if the cognitive ability to care for young which exists in some salamanders and is lacking in others may be investigating social evolution. Miliam offered critically that for Darwin, Sexual selection was his theory to explain all stable intraspecific differences, including why males and females differed in their appearance and behavior, yes, but also differences between races of a single species Miliam imputes that the concept of social selection conflates the sexual division within intra-specific differences that are supposedly implied by sexual selection as distinct from natural selection. Both male and female salamanders provide parental care. Some salamanders provide none. The evolution of parental care in salamanders may be cognized as a transition of intra-specific differences into sexual choice of care. Thus we may be able to distinguish the making of the concept of social selection distinct from making a distinct concept of social selection. This will help to differentiate the larger project of re-envisioning the evolution of social behavior from claims/assertions about intuitions of sexual selection. One of the striking features and benefits of Roughgarden social

evolution through social selection is the symmetry breaking perspective and clarity it provides to tabulating the effects of sex correlatable differences. Intuitional tendencies often reside at this level of consideration. Furthermore, in salamanders, the provisioning of parental care appears to be associated with changes the development of the young themselves. The enlarged tail structure that Darwin described as an ornament surely is at least, an adaptation to more efficient wafting of pheromones and exists as a geographic/racial variant in some specie. Is this trait due to natural selection, sexual selection of social selection? Does theoretical and empirical research into salamanders verify social selection in this case? Are behaviors projected towards rearing the most number of offspring in the next generation or towards genetic quality? With the new adaptation of spermatophore production and the separation in space and time of sperm uptake and oviposition/hatchling site prevention of oophagy would be needed under sexual selection Nussbaum notes that parental care has been argued to function to (1) prevent intra- and interspecific oophagy, (2) apply antibiotic skin secretion to the embryos, (3) agitate embryos to prevent yolk sedimentation, adhesions and developmental anomalies, and (4) provide moisture via maternal skin and urinary balder and to retard desiccation by bodily covering of the embryos. Nussbaum presents the thesis that parental care results from changed developmental needs of the hatchlings to available food size distributions (that salamander laravae had to be larger to eat larger prey items and that the larger eggs to achieve these size differences in the larvae needed to be attended to (to reduce the death rate of embryos that take longer to develop)). On the view of social evolutionthen, courtship may be a job interview to ensure that larger larvae will be raised. Male pheromone delivery would not be a signal of better genetic quality of past reproductions as in sexual selection but of future commitments for increase developmental size of offspring. Correlation of increased pheromone production and enlarged ornamented morphology to deliver such is incidentally correlated with developmental pathways that tend to increase physiological structure formation and hence promote increased size development in general and is not thus due to males simply attracting females in cases where there is no parental care, as Darwin observed. This increase is independent of the process suggested by Darwin where the non-muscle structure (not for getting males near females) arises by sexual selection through either the male or female. Instead this particular ornament indicates a particular developmental subset of possible embyrologies and offers by bevahiorcal choice entrance of salamander populations into a coalition of different embryonic strategy space not into different visual sexual selection space of better genes. The embryological classes are dependent on the choice required not simply that there is choice.

This view is different than Clutton-Brock who seems to relate a sex-conflict or different male and

female pobjectives view for the same info rather than cooperation and unified goalable view.

The choice of the mating place becomes less relevant as the bargain becomes more canalized. In those with external fertilization (Hynobius) both sexes have been reported to care for the eggs.

http://salamandersofchina.lifedesks.org/pages/4029

Breeding season is between late winter and early spring. Adults mate in the ponds next to streams/creeks. Parental care includes guarding the eggs sac and larvae by both sexes (see refs in Hou et al., 2010). Egg sacs are attached to the bottom of stones in water. Each egg sac contains only 4-8 eggs. Larvae hatch after two months and reportedly feed on algae and decomposed plant residue (refs in Hou et al., 2010). One month later metamorphosis takes place, when snout-vent length is about 14.5 mm (Hou et al., 2010). Metamorphosed juveniles have been found in March (Chen & Lue, 1986), suggesting a reproductive pattern similar to that of the other Taiwanese Hynobius species.

Females do not need to be "choosing" the sites to begin with, but salamanders seem to have found a way to increase offspring production by internal fertilization and choice of where the young are not the adults.

A bargain may be the use of active pheromone- receptor co-evolution to probe the extent of development to larger egg sizes by syncing it with frozen to random networks as indicative of larger larval sizes. The Active to Frozen nature of the receptor-pheromone mutations may be a small scale (few loci) correlate of larger general developmental macrothermodynamic extent measureable by random vs active gene networking and if correlated to egg size can facilitate a behaviorally triggered bargain to increase larval sizes to place of young where more food is. This could occur by breaking through a check Darwin would have considered.

In analogy with ACKAY and Roughgarden, I consider a salamander side-payment to be a consequence of a phenotype of the sequence of the pheromone gene or its receptor. Males provide a pheromone sequence and females a receptor sequence. This sequence is fixed for a given social/behaviroal interaction of males and females. The consequence of the differences in the sequences among the individuals can not be withdrawn. Whatever the effect of a difference in the sequence is, its effect in the behavioral interaction socially can not be ethologically substituted So if PMF is sensed by the female VNS and the female picks up a spermatophore sooner than otherwise the requirement of AKAY and Roughgarden that individuals can discern the actual payoffs and react quickly by adjusting their actions. If a male for instance refuses to deliver pheromone, the female could quickly break off the follow up straddle walk. This is what ensures that receptor- pheromone co-evolve. Alternatively one may suppose that sexual selection causes the back and forth play of receptor-pheromone evolution. Going from SPF to PMF to PRF makes it possible to construct a fairly specific proximate mechanism for the straddle walk behavior engaged during salamander courtship sociality. During a transition from SPF to PRF the relation of PMF in transitional sub sociality behaviors may highlight conditions where fitness interests are not aligned completely. Also the common receptor-phermone situation across the threepheromones enables a generalized model of other-regarding motiviations to be modeled into the mechanism. In the process we will be able to investigate how the payoffs themselves change. Here we will be able to see the behavioral game as the paypoffs interms of fertilization and coalitional formation vs evolutioanary co-evoltuion of the receptor-and pheromones. We will be able to see how different the same straddle walk can give rise to different evolutionary games of different pheromones and delievery mode transitions (mechanical chemical (wafting cutting - touch)) and how repetition of the straddle walk (or its precursor) affects the ability to sense wafting payoffs, cutting payoffs, and touch payoffs)) under a common reward sensation development. We thus fix the decision making rule and find that a switch to a new pheromone or pheromone gene is equivalent to the recovery of a new Pareto boundary (where both players (male-female) can not improve both payoffs any futher both players payoffs. This supplants the imagery of the moving dance floor of P. The conflicts of interest thus are due to the simultaneous interest to change the receptor or pheromone sequence but still have the highest payoffs and other-regard behavorially. So if a male has a new pheromone sequence that binds tighter to a new female receptor and the female does not respond to move away then a point will be reached as to how much the dance can change the sequences before the behavior results in a much lesser payoff due to poor response to the bindings. Thus crucial in this application of the model will be to elucidate the reward sensations that associated with the payoffs. If the stronger bindings can lead to either higher payoffs or lower payoffs it may be possible to empirical discern whether multiple Nash Eequilibria extant and which are preferred. The establishment of stable polymorphisms may explain how SPF continues (possibly with a lower behavioral individual fitness but is polymorphic with P MF or how PRF split into two genes) The small number of pheromones and modes of delivery as phenotypic plasticity and genetic polymorphism may be the result of the small number loci that can support such a situation. This could be compared with the general social

selection scenarios in frogs and caecilians, especially the later where the move has been from broadcast int frogs through sperm uptake by the female to male intromission. It would be extremely exciting if behavioral differences can be correlated with sequence polymorphisms both within and between modes of delivery such that though constrained within modes by different overall fintesses ranges per mode different patterns of positive selections can emerge with relation to reward sensations. The incentives to cooperation may thus be in the location of the mutant sequence changes whether a mutation needs to be at the binding site or if can be less costly father away but still be effective in getting a tango on the dance floor with multiple plays of the game. Is it possible that the prisoners dilemma can be used with salamanders? Both players playing SPF but each prefers the other to play PMF but with a mutant one plays SPF and the other PMF which leads to a change in where the binding site cost per mutation place lay in sequence space such that more mutations can occur in PMF if none occur in SPf and both prefer this?

Between mode pheromone divergence may be caused by matainence of genetic polymorphism of behavioral influence on the evolution of the pay off matrix. The lack of positive selection in the scratching and slapping modes in SodeferinSPF may be due to the higher level of mechanical-behavioral association of the traits.

The internal insemination begun in salamanders may be thought of an adaptive transition to care more for future offspring by offering more physiological constraint than broadcast methods.

Milstein criticism of Roughgarden can be inspected in the case of Darwins citation of the Newt Ornamanet in the case of males that do not have the pheromone associated expression (unknown to Darwin) vs those that establish polymorphisms due to evolution of the payoff matrix. These males without the pheromone would not be able to form coalitionts and would not be able to move to the chemical from the mechanical as the trait evolved but WOULD show positive selection as the salamanders evolved new delivery modes of the class of receptorpheromone coevolutions whether by sex differences or behavioral polymorphisms in the payoff matrix. Thus salamander data indicates that milsteins criticsm is factually mistaken. Darwin in truth implicated the moral perspective but with Tetration and the Ackerman FUNCTION applied to Fishers runaway process the juncture in evo-devo will show that words can not substitute where data exists. Acutal infinity was developed at the same time of Darwin.

Sexual selection can amount to infinite drift relations (since uses aesthetic) in the process and thus is different than natural selection (Fisher vs Wright generally) but social selection amount

of drift depends on the evolutionary existence of the behaviroral development (hence why successes rather than ability) variation in the most general symmetric case. In this case if preference occurs in these cases over strategy then cooperation does replace selfish altruism and spite.

Mechanical to Chemical cooperative NBS (some lack of chemical transfer in NOtphlamus) to investment pheromone-recpetpor point mutation *(minus the cost to invest (dependence on mental gland route rather than choice to use genial gland or tail wafting) positive se3lections rather than sexual conflict based receptor-pheromone affect through runaway processes The mechanical processes use enlarged sperm size in the spermatophore location depenednet on the breeding pond substrate while chemical pheromone straddle walk locates the placemenent of the re-enlarged sperm size capsule at the place of the bodies (adult locations) of the species rather than the habitat of the breeding pond(young locations) Spermataphore gelatin is a contact property.

Internal fertilization in salamanders from broadcast fertilization may have resulted in situations where gamete contact rates could possibly fall below that which gave rise the difference in size of egg and sperm to begin with. Thus behavirors in external fertilzers could only go so far in increaseing the size of the egg before losing the gains in gametic contant which seperated sperm and egg size in prior time.

Spermatophore gleatinazations were cases where chemical response to this effectively increased the size of the sperm. (increasing size just like increasing "size" of eggs with gelatin. Soderferin pheromone attraction may have been a simple way to attempt to aggregate reproduction to bring back larger gamate contact rates.

In a geographic to bodily positioned evolution the index of assortativitiy may be expected in geographic situation but not necessaritly in the bodily position position. Thus by a detailed understanding of the molecular evolution of the pheromone-trait positive selections we may be able to differentiate runaway sexual selection, strategy evolution which limits the behavioral input, and preference or social selection evolution. It may turn out to be a case that the evolution of plethodon

olfactory transmittal behavior arose from social evolution through a bottleneck of strategy evolution as the sexes began to invest in the straddle walk and its chemical foundations otherwise simply directed in the mechanically forced internal fertilization (to keep the number of gamete encounters high enough). This will explain how individual proteins can have both highly conserved and highly variable regions. So under strategy evolution where it is theoretic al supposed that a realistic feature of many social dilemmas is the net benefit of contributing becomes smaller as others contributions increase will result in some parts of he pheromone being conserved as the other contribhtuion becomes smaller but if it variable if the contribution of the males itself is smaller (provided within the same dilemma/game/strategy space)) The conserved region also applies to the conservation of the pheromone itself. So the transition from Soderfin through PMF to PRF with different variances of positive selections and conservagtions of regions will be due to actual strategy evolution (if a common explanation can be made for all mutant changes (on both scales) relative to scratching to olfacgtory behviro change suppoing it is in the same social dilemma that transformed soderfin from the clocal to the mental gland OR preference evolution if behaviors are more strongly correlated with synergismtic conditional regard. Alternatively the behavior to trait extablishment/expression may be even more losslely connected by Fishers runaway sexual selection. Thus by looking at the positive selection mutantant loci in the various pheromones we may be able to evaluate both how acute the angle is back towards an incumbent (male-female receptor-phermone prior amino acid placement) trait and the general format of the evolution gradient. This works because the geographic localiazation in addition to setting up restricted strategy space of the behavioral equilrbium also establishes the altruism/spite baseline from which strategy evolution may have been conjectured while both are independent of runaway sexual selection where males are cut of the process (as so far modeled by Lande 81) into the breeding seasons which is increased by a straddle walk regardless of whether this is due to strategy or preference.

Negotiating Larval Size : Social Selection in Salamanders If it is true that bindings of pheromones and receptors are correlated with better general developmental alignments and stronger and stronger bindings (within molecules and across to different ones) are associated with longer development times then salamanders may negotiate for larger larval sizes by controlling the delivery of pheromones rather than relying on diffusion at the location of mating. This correlation may kill two birds with one stone since it could permit both decreases in the number of sperms per successful offspring (spermatophore vs small gametes diffusing cloud) and larger zygote size and also be a rebalancing of asexual-sexual modes expressed interms of contacts per geography (suitable breeding locations) towards negotiations that revolve the contrainsts on egg survival with larger size vs numerous small male gametes. Since egg survival can be further enhanced by parental care increased alignments may the cause of behavioral modifications in many Plethodon (thus allowing the developmental correlation to be expressed as behavioral diversification). This hypothesis may also enable one to get a general numerical ration between cost of meiosis (under

assumption of no relation of gamete contacts and geography) and the asexual survival in Ambystoma from a geography (indicative of node to track relations) of A. barbouri to that where social selection is effective. Female disruption of Courting may be a case where pheromone diffusion and abundance of glacial ponds offer a difference of opinion than is possible by developmental tracking of pheromone receptor coevolution, resulting in pheromones only used to attract and distrupt (behavior based) malefemale when the ponds are stable enough and plentiful enough. The larger size of Ambystoma ensures large enough zygote size for those cases where not enough blooms of plankton occur? The projection of Eurycea and the ringed salamander track leads to the node that is indicative at once for the unisexual ambystomas and the other plethodon.

We begin by considering that chemical communication between males and female externally fertilizing salamanders follows the model of a trust game that evolves into a cooperative situation of team play of internal fertilization. The neontological species of external fertilizers continue to practice run ups to determinations of enough information to determine who is trustworthy with the alternative to resort to individual play continues. Instead in internal fertilizers the side payments. Side-payments are the amount of wafting by the males or the amount of approach by the female in response. The behavioral ethograms of external fertilizers are split into categories of sequential decisions In which different amounts side-payments accumulate to the overall payoff but without explicit team play the cooperation remains restricted to the genetic polymorphism effects rather than the behavioral plasticity. Trust results in changes in the pheromone itself and the receptors as other-regarding behavior appears. The specific amino acid changes and binding changes indicate what physiological correlations can result in team play resulting beyond simple other-regard as expressed in the behavioral cycles of catogrized decisions making in later generations. This permits with small amounts of loci involved the possibility of new pheromones to appear in the same function as into internal fertilzsers. And this appears in the internal fertilizers as the synergy between the social functionality (reproduction) and the behavioral mechanism (amplexus and sperm and egg combination). Team internal fertilization only arises from polymorphic gene behaviors where the trust and other regard even when resorting to individual play (synergy outweighs indivdiaul selfishness).

If this application is supported in facts then it may help to differentiate(Okasha 2009) "commoninterest" interactions supposed to be a part of group selection vs the genetics of the social selection while the common-interest" acquires enough synergy to over ride prior reversion of indivual behaviors in systems of threat points past. This team play of for spermataphore pick up through cycles of decisions trusted and evolved can be further compared temporarily to the 2-3 hour internal insemination time of caecilians. Thus as reported on-line

http://www.amnation.com/vfr/archives/009763.html "However, is not this difficulty overcome via a "transitional form," namely the salamander? As the Encyclopedia of Earth article tells us, the female salamander picks up the male sperm packet and puts it inside her cloacum and has internal fertilization. This extraordinary behavior would seem to reduce significantly the number of simultaneous random mutations that must occur in order for the male and female to mate successfully. Instead of there having to be both the simultaneous development of the complementary mating organs and behavior in both sexes and the development of the female's internal reproductive functions, the latter would already be in place, so that only the sexual contact would have to be added to the picture in order to create the new species behavior" we can understand the "complementary mating orgs and behavior in both sexes" such that internal reproductive functions could be futher transited. The threat points are dependent on the correlation of the receptor-pheromone variability onto behavior. Thus when a newt pheromone is changed by one amino acid it can attract a specific geographic population but not another. Plethodontid salamanders have some positive selections in the pheromones and the bahviroaer coorelated with these will help to describe the behaviroarl dynamic threat points. Signal decoupling (of multiple pheromones in a common straddle walk) is thus understood as variation in the genetic polymorphic loci that support behavioral actions through an abilty to change the threat points towards more cooperative states and is not separated as Picard thesis expressed. Functinoal integration is thus a matter of individual play threat points vs synergy of cooperative team play WITHIN the loci variation permissible to keep the basic behaviors of coalitiational subscription (either geographic or within body). The idea that there must be a coupling which is in truth better described as a decoupling results from the false premise that sexual selection would tend couple the signal elements.(Picard page 4 integrate all parts of this signaling complex). The threat is simply that since both males and female plethodon mate multiple times in a season they can simply withdrawl from mating but they can not withdrawl from pheromones released or reward sensations nervously conducted. The details of the correlations between chemical-receptor variation and behavior is the effect of one spermataphore uptake or failure to take up on the next courtship uptake or not. It is important to witness correlations between multiple uptakes of spermataphores in an ethogram so as to more accurately describe threat points. It is possible that series form with various structures of density and transitivity.

Contra the idea that shorter courtships are selectively beneficial because they decrease predation possibility and conflict with conspecifics is the idea that shorter courtship time enables more intricate serial decision making between series of a multiple courtships and sperm uptakes in the single breeding season.

Do salamander avoid strict application of Hamiltons rule? The biology of how the benefits are produced determines whether mutual regard will evolve or not. The biology of pheromone use during courtship in salamanders has been analyzed utilizing sexual selection without an alternative. Here we present a proposed case of social selection to the same data set and indicate a situation where mutal regard evolves as new pheromones become ESS. It is suggested that primitive salamanders had payoffs that could conflict between males and females but that through synergism new mutant pheromones (and their attendant receptors) lead to other regard in a particular benefit and cost scenario.

This application will permit an alternative to be compared to that narrative of salamander evolution given by sexual selection but also compare so-called strategy vs preference evolution. It may be that salamander avoid the issue of thought of by Hamiltons rule in that they utilize both substitute strategies (new pheromones) and complement strategies (positive selection on binding) to osscillate across clades. This could falsify the class of models that seek to find preference evolution cooperativity in the structure of relations to altruism/spite under assortativity rather than in the population genetics of social selection. The modeling also suggests an alternative to sexual selection.

There is no test here only the fleshing out of the narrative. Higher frequency of persuasion behaviors with historical success may simply be a result of an NBS for more offspring not more matings. Having larger variance in more matings may

also be for not permtting loss of a breeding season per male (number of offspring produced) rather than simple number of matings no matter how many breeding seasons the males undergo. Thus if other regard behavioral interactions increase (accelerate) (by continual production of pheromone) the nervous system reward sensations (quicker behavioral responses) over the life o the salamanders there may be cases where more matings occur and thus the local strains genes increase faster but over the long run and across a larger region those that have higher payoffs of he others (by variation in the receptor-pheromone indepnednet of the amount produced). Thus social selection may lead to differential intra-inter deme selection of the individuals overall (no group selection).

Abstract
An important function of the courtship displays of male animals is to stimulate, or persuade, females to mate. While many studies of courtship success have shown that females choose stimulating males as partners, fewer have demonstrated that the most persuasive males in a population also enjoy the highest mating success (i.e. mate with the greatest numbers of partners). Such a relationship provides a stronger test of the hypothesis that male displays evolve by sexual selection via female choice. We tested this hypothesis in the laboratory using the North American plethodontid salamander,Desmognathus ocoee, in which females only elicit spermatophore deposition if rendered sexually responsive by male courtship. First, we determined variance among individual males in the numbers of females with which they mated across 35 encounters, in the absence of differences in partner encounter rates and direct intermale interactions. Second, we demonstrated that males with high historical scores of mating success performed significantly higher frequencies of persuasive courtship displays, which provide tactile, chemical and visual stimulation to females. We conclude that sexual selection via female choice favours persuasive displays because they confer high mating success on the males that perform them at the highest frequencies. *1 D. H. TaylorS. I. Guttman
f1

Correspondence: P. Verrell, Department of Zoology and Center for Reproductive Biology, Washington State University, Pullman, WA 99164-4236, U.S.A. Animal Behaviour Volume 56, Issue 2, August 1998, Pages 443-448

Picard reports that (empirical) studies have indicated that the delivery of each pheromone component (PRF, PMF and SPF) alone, elicits a change in female sexual receptivity as measured by courtship duration (page 14) and he reasons that sexual selection is expected thus to play an important role in shaping the evolution of these signal components and in formatting (affecting) the evolution of the behavioral and morphological traits associated with the delivery. Here we present a model that inverts this reasoning and shows how behavior itself through social evolution effectively enables other regard to arise with respect to signal component physiology wherein shorter duration courtship effects smaller effective variances by density increases across a breeding season resulting in more offspring raised regardless of the choice by males or females or receptor-pheromone signaling binding strength as multiple signal components become incorporated in an ESS manner. Vinneage and Verrell ( Animal Behaviour Volume 56, Issue 2, August 1998, Pages 443-44)draw out the general sexual selection argument for plethodontid salamander courtship by arguing from data on Desmognathsu ocoee, that sexual selection via female choice favours persuasive displays because they confer high mating success on the males that perform them at the highest frequencies but the model we present here shows how the behavioral action of high frequency of pheromone delivery in the persuasion display can be a pay off indicative of more offspring production regardless of the amount of matings that occur. With the two alternatives social selection and sexual selection we investigate what studies are needed to decide between the two possibilities. Rollmann Houck and Feldhof f(Animal Behaviour 2003 66, 857861)suggest that pheromones are not used to discriminate species (conspecifics vs heterospecifc pheromones with PRF experimentally applied) but nonetheless do decrease courtship times which in the model here appears as the global objective function wherein team play vs individual play can occur. But contrary to the implication in the work of RHand F pheromones are not to be thought of a candidate for behavioral isolation between species but rather as a team play signal after the individual threat points have been passed for as they noted there is variance in the rate and amount of deliveries individual males may make. The decrease in courtship time on application of the pheromone itself expresses the synergism already achieved. This synergism maintained by behavioral diversity in the small number of pheromone signal components results in heterozygote advantages that explain the persistence of the delivery modes rather than stabilizing selection(canalization). This fact that the hypothesis thatartifical conspecific pheromone application was not preferential (when measured by diminsihgn courtshiptime) to heterospecific pheromones may indicate that pheromone difference could be a first step to team play that involves all of the other behaviors associated being complentary

but not fully conditional in syngergism.

These six perspectives become one in the model and we offer what had not been heterfore possible without social selection

The six perspectives and their predictions are: (1) A sexual selection perspective on communication systems predicts either stasis or rapid evolution, depending on whether selection acts on the receiver. OTHER REGARD -COOPERATION (2) An empirical perspective based on observations of morphological and behavioral evolution in Plethodon salamanders predicts either stasis or rapid evolution, depending on which branch of the phylogeny is under scrutiny. DIFFERENT THREAT POINTS TO TEAM PLAY (3) A neurophysiological perspective based on the pathways mediating female behavioral response to the male signal suggests that radical change in both the signal and receiver should co-occur in one particular branch of the Plethodon phylogeny. REWARD SENSATIONS (4) A molecular perspective on the cytokine protein family to which one of the pheromone signals belongs suggests that some parts of the pheromone signal should be static (e.g., binding sites), but other parts of the molecule should be capable of rapid evolution. LINKAGE of multiple LOCI sustaining heterozgote conditions giving rise to multiple behaviors The particular binding sites being conserved may simply reflect that individual advantage occurs to increase ones partners payoff only when one is sufficiently certain that the partner will also do the same. This is not part of the strategy but functions as hard wired with respect to conditional regard itself. It specifies for any given complemtarity the total amount of syngergism the given payoff matrix can produce without its evolution.

(5) A molecular perspective based on recent case studies of proteins suggests that our pheromone signals are candidates for rapid molecular coevolution.

SYNERGISM AND COMPLEMENTARY STRATGIES IN OTHER-REGARD (6) Preliminary analysis of pheromone gene sequences from four Plethodon species reveals rapid, selection-driven evolution of some parts of one pheromone molecule. Taken together, these perspectives motivate a research strategy in which we focus tests on particular branches of the Plethodon phylogeny. We ask whether the same kinds of selection (stabilizing or diversifying) and same evolutionary mode prevail at all levels of the signaling system. DIRECT PAY-OFF DIFFERENCES There is no one unifying construct that makes theory-based predictions at all levels (e.g., from proteins to neural response). Sexual selection theory, for example, can address expectations concerning a male signal and the female response, but offers little insight into structural changes of a three-dimensional protein signal. We therefore used a variety of perspectives to make predictions about evolutionary stasis or change. These perspectives make contrasting predictions. Our goal was to assess which perspective(s) best predict the evolutionary mode each level of our signaling system. Also, if we discover that stasis, for example, characterizes signal evolution among species having an olfactory delivery mode, does stasis also characterize female behavioral and neural response to this signal?

Thus while supplying a unifying social evolution construct to unite these perspectives it is also possible to resolve the discrepancy in a proposed difference of strategy and prereference evolution by showing under a small number of loci involved (three pheromones) that these three enable macro evolution to oscillate with changing complements and supplements as given by the three pheromones through their connections to behavior as the change in sequence space in this preference both below and above that suggested by Hamiltons rule for the entire sequence space changes in the entire phylogeny.

Does the biting behavior during courtship of 3 Desmogs, the excretion of mental gland pheromone through the mouth side of the gland (teeth) and the proglucogon peptitdes that are controlled by Homeobox genes suggest how wafting behavior to biting was able to construct a bargain of Plethodontids from Salamandrids for a negotiation to raise more offspring by agreeing on having more

development time (to be able to have larvae eat larger prey and not worry about ineffiencies in gametic contact rates with external fertiliaton)?

Here we attempt to use salamander pheromones associated with courtship to determine that, if neither cooperation nor competition is to be assumed, which makes more sense for these amphibians? Both a reproductive conflict view of sexual selection and reproductive cooperation view of social selection suggest that there may be co-evolution of the pheromones and the receptors used to sense the chemical s(signal).

We have referred to the process producing this evolutionary pattern in male courtship pheromones as a molecular tango and have hypothesized that it is driven by coevolution with receptors in the female VNO (Watts et al. 2004, Palmer et al. 2005). To test this coevolutionary hypothesis we prepared cDNA from female VNO tissue and used RT-PCR and 5-RACE PCR to amplify large segments of pheromone receptors. This strategy revealed 67 distinct V2R isoforms, belonging to at least 6 families, in P. shermani. Ongoing discovery of new isoforms shows that the V2R family is as diverse in Plethodon as it is in rodents and other vertebrates. Our preliminary studies in Plethodon have also documented a type of receptor not previously reported from the VNO. These gp130 and related receptors may mediate PRF reception in the VNO because these receptors are universally utilized by those cytokines most similar to PRF (Watts et al. 2004, Bravo & Heath 2000). Both gp130 and CNTF-r from Plethodon VNO show unusually high levels of sequence diversity compared to other animals. This departure from the usual purifying selection mode for gp130 receptors and cytokines is consistent with a coevolutionary process driving diversification of PRF and these gp130 receptors. In social evolution this molecular tango may be thought of as the set of outcomes upon which it is not possible to improve each players payoff simultaneously as expressed in a genetic polymorphism that gives rise to many different behaviors. So when a mutant pheromone occurs behaviors associated with the mutant can be side-payment (biting in Desmognathus with glucogon like peptides) with a cost and one can decide if the mutant can invade the same game the payoffs are expressed in. Let us assume that Plethodontid salamanders play a rock-scissors-paper game with SPF, PMF and PRF and see what we can learn about the kinematics.
By embedding the difference of preference and strategy evolution in quaterionic space it seems to me that preference evolution need not be thought to conflict with Hamiltons rule since even the the acute angle back to the resident with mutant in the splitting of a biquaterion it seems that the general asymmetric two player game can ossilate below and above the necessary index of assortativity in three

part game and perhaps salamanders have done this SALAMANDER FAMILIES are there any? Statement Nussbaum (1985 The Evolution of Parental Care in Salamanders), Either the female or the male cares; there are no cases of biparental care in salamanders. It is argued that the sex that chooses the oviposition site will be the care-giving sex among salamanders 2012, Roughgarden, J. and Song, Z. Incentives in the family I: The family firm, an evolutionary/economic theory for parent-offspring Relations. In: Narvaez, D., Panksepp, J., Schore, A., and Gleason, T. (Eds.) Human Nature, Early Experience and the Environment of Evolutionary Adaptedness. New York: Oxford University Press, in press The idea that social evolution can occur within families through incentives has been developed for birds by Roughgarden, Song and others http://www.situsci.ca/event/incentives-family-firm I ask whether the theoretical modeling of Groves 1973 which underlies this familial notion may be applied to discussions of, and research in, parental care and/or parental investment in salamanders. Analogically, one may attempt to see if the parent as manager and the offspring as divisions applies. More generally one would like to know what kind of incentive structure may be divided amongst those salamanders giving care or receiving benefits from the same. T. H. Clutton-Brock has observed that Nussbaums extended reasoning leads to a removal of any advantage to females of choosing particular mating sites to begin with. The logic of this understanding is that given there is no biparental care and there appears to be a correlation/association between the care giving parent-sex and the mode of fertilization (males with external fertilization and females with internal) and that in the internal case there is a tendency to separate mating location from oviposition location, if the choice of the location of the oviposition site (by males in external fertilizers and by females where spermatophores arepicked up) is determinative, regardless of sex, and that there is also a benefit to separate in space the mating from the oviposition selection pressure is removed from mating location components and since this happens mostly in the internal cases females have no hypothetical pressure in salamanders to choose mating sites. This kind of thinking however is dependent on a sexual conflict or asymmetry view of sex in relation to reproduction rather than a harmonious or cooperative view. It insists that the correlation between fertilization mode and sex giving benefits (parental investment) is dual and polar. This may not be. Fertilization mode may be related to sex through gametic contact rates rather than maximum number

of matings and the separation of mating location from oviposition location may simply be in the internal case a secondary effect increased contact rates (compared to external fertilization), regardless of the number of matings. Females may still have an advantage to choose mating locations and could do so when it increases the incentives to parental care or benefits to the offspring even under fluctuating amounts of matings per care.

So can the parent (male or female) be the manager and the offspring the divisions? Do Salamander familes exist?? Nussbaum had suggested that http://www.springerlink.com/content/t635783883707510/ food availability differences between lentic (smaller items) and lotic (larger itmes) causes selective differences in parental investment in stream salamanders than pond salamanders. Nussbaum suggests that plankton rich environments have less parental investment selection levels. He analyzes the payoff evolution in terms of the size of the offspring ( more payoff for many small individuals when plankton is readily available) but fewer larger individuals (and more mortality certiaus paribus) where offspring prey are less dense and more numerous. Maternal care is thus a compensatory mechanism to reduce mortality after larger sized individuals are developed. This view does not symmetrically account for Fraternal care in external fertilizers. So if there is any beneficial skin to skin chemical interchange in either sex to the offspring ( offspring contact with mothers has been demonstrated http://www.bioone.org/doi/abs/10.1670/09-181.1?journalCode=hpet ) and/or if the care can directly effect larger size individuals (shown experimentally with Plethodon http://deepblue.lib.umich.edu/bitstream/2027.42/47699/1/442_2003_Article_1410.pdf ) and this behavioral/developmental dynamics drives egg size parental care may be constitutive component of selection for larger sized offspring and part of a Nash bargain amonst species living under lotic correlated pressures rather than a compensation for changes in mortality in the different environments, more on the r than the K selection side but rather yet again understood by social evolution. http://philosophy.ucdavis.edu/millstein/papers/SexAndSensibility.pdf By applying the incentive view of Groves 73 http://jmvidal.cse.sc.edu/library/groves73a.pdf to salamander families both the mating and oviposition and post oviposition behaviors and development become integrated with parental care symmetrically with respect to sex and differences

of egg size vs spermatophore size and number are dissected in terms of gametic contact rates rather than number of matings or specific selection pressures for larger individuals. Thus if salamander parenting struggle is on the r side of K selection of Nussbaum then social selection could be compared with r and K causally in salamanders.

(said Roughgarden in Sex and Sensibility)

Can male midwifeing in Hynobids be biomechanically related to female straddle walking in Plethodontids? Can salamanders rather be cooperating than subject to sexual selection during external fertilization Do glands in the larger size of the male head secrete chemicals or can they be used to move eggs around rather than exclude other males? http://www5d.biglobe.ne.jp/~hasumi/doc1/study_e.html

One of the more popular criticisms of Joan Roughgardens attempt to replace sexual selection with a discipline of social evolution via social selection is to say that Darwin himself was not mistaken at all. In Sex and Sensibility Milam insists that Roughgarden should not have the option of casting todays research into sexual selection as essentially the same as Darwins. While this may seem to be of consequence for historical interpretation only, this is not really so, not surely if cooperation and the late 50s notion of the animal family through social evolution are to replace strict sociobiological directums in more recent research and is significant when it comes to displacing Ghiselins attempt parse Darwin to the advantage of an earlier generation of biologists. http://research.calacademy.org/redirect?url=http://researcharchive.calacademy.org/calwild/ While it is true that vigorous and well-armed to the left and and in other respects more attractive pertain to different parts in the sequence of Darwins presentation, it is the entire view of sexual selection as moderns and Darwin have it that is in question. The principle question is, Does social selection theory

offer new range of hypothesis testing not available in the thought process on sexual selection and do the hypotheses constructible stand up to evidence to the exclusion of sexual selection scenarios? I will show how both sexual selection and social selection theories can be designed for the social reproductive behavior of Ambystoma texanum and we will await results of tests (observation and experiment) rather than rhetoric to determine the fate of the alternatives. The texas mole salamanders have a relatively simple reproductive behavior when compared to salamanders in general ranging from may be considered male male competition incidents and the existence of polyandrous aggregations, thus spanning systematically the range Gieshlen attempted to use language rather than data to constrain judgment of whether Roughgarden or Darwin was mistaken. Thus through long observation and experiment rather than vitriol and innuendo will we decide on whether any use of quotations from Darwin are fake or not.
Sasha R.X. Dall1*, John M. McNamara2, Nina Wedell1, & David J. Hosken1

1. Sexual Selection Cannot be Replaced by Cooperative Game Theory (and It Doesn't Need Replacing)

On the other hand are not splitting hairs as Ghislen had tried to do linguistically over the modes of selection determining specific agents (man vs environment vs other member of the population) but once again using position that A. texanum been asserted to maintain behavior that is a competitive game theory ESS, both competitive and cooperative alternatives will be postulated theoretically and we will let the data decide rather than apriori think we can judge animal and plant populations only with our human based understandings of economics/theory applied. There is a difference between a distinct concept (using cooperative as opposed to non-cooperative game theory) and making a concept distinct (allowing the data obtained to drive the discrimination/distinction/difference). It has been claimed that the behavior of spermataphore capping as exists throughout the genus Ambystoma is a Maynard Smith (1974) ESS since once capping has begun and a reverse mutant that does not cap evolves it cannot invade the capping clan. This is a clear example of competitive game theory in sense Rouggharden et al challenge (as resulting in ad hoc consequences, .) Harriss experiments appear to remove experimentally the idea that perhaps capping is a cooperative behavior since the data seem to indicate that females do not have an easier time finding spermatophores that capped vs those not capped and appears to put an end thus to the selfish vs cooperative change social evolution was designed to adjudicate. Alger has further theoretically distinguished strategy from preference evolution but in an expanded Prisoners dilemma applied to A. Texanum which alternative complementary vs substitionary strategies of males (whether fighting or making peace) with females (on the diagonal in the game when both fight or both make peace) symmetrically under the only other major behavior (more nudging/fighting of males to males than males to females and none of females to males per polyandrous mating territory) it is possible to construct a specific genetic polymorphic situation in which cooperative behavior forms a new game theory equilibrium when invaded by a

mutant that still caps but does not fight as much. Capping is thus a cooperative behavior that sustains mating site location fidelity under nudging/physical contact that both moves males away and draws females attention to the place to accept a spermatophore preferentially for males that do some pushing but less than the average OVER generations where the pushing is either in complement or subsititute for female choice of Darwins supposed in sex selection. We then look to the examination of futher experimental designs to decide if capping is a competative adaptation to avoid sperm competiation or a cooperative behavior evolutionarily selected to ensure across generation mating/oviposition site fidelity. This is thus tested within the species, explained throughout the genus and expanded to other genera. It seems that without the possibility of social selection the notion of males and females would remain within that decided by the causal modes of selection divided without this difference. A replacement of older generations of evolutionary theorists in order and is an incentive of Wright.

Anurans
The use of a tympanum may have been socially selected away from Leiopelmids and Ascaphids which utilized internal fertilization developed into internal egg development in caecilinans and internal fertilization (spermatophore for the intromittant organ) in salamanders. Late jump recovery is unique in Leiopelmatidae. When Leiopelmatid spp. jump, they land in a "belly flop" fashion repositioning their limbs for takeoff for the next jump only after hitting the ground with the ventral surface of the torso. After landing, Ascaphus skids to a halt before recovering. The appearance of early jump recovery in more advanced taxa Jump recovery may be nothing advanced but simply the adapation in forms which have tight muscels needed to hold ear bones in place so as not to have too loud noises affect nervous input.

Assume that anuran parents make decisions in behavioral time. Are there such things a frog and toad parents able to effect decisions concerning their offspring?

Well if you were this mommy or daddy it seems that behavioral decisions made (jump into dense foliage that may scrape of eggs to escape a predator vs jump into deep clear water where both eggs and parent are easily seen ) by her can quite dramatically affect the future of her offspring.

It may be that anurans got to carry eggs after camaflouging them with colors to their back, go under some and found it better camaflogue. Also harder to get eaten.

If anurans socially evolved sound transfers as means to negotiate a behavioral bargain then either the more primitive frogs that lack such means to negotiate must utilize a different mechanism to effect cooperation and this (or the lack thereof) must be able to physically indicate a different mode preexisted such use of sound in terms of the offspring themselves.

The Anuran Chorus Competition or Cooperation?

Communication across an Electrical-Mechanical Continuum After publication of Joan Rougarden et.al. 2006 Science paper there was an exchange of opinion between McNammara etal and Rougarden etal on whether competition or cooperation or neither need be assumed. Obviously, it should be neither, but just what are the possiblites?

We can look at this from the narrowed issues of theoretical application of game theory or the general issue of sexual selection and genders. I will show that it is possible to cognize the Frog and Toad chorus as model of cooperation in the sense of Rougharden et al and not that attributed to sexual selection of Darwin. It will be possible to show how game theory directly applies in the two tier model and that chorus calling ( as opposed to lone calling) inititates what is otherwise a binding bargaining contract. The Nash bargaining equilibrium is directly indicated in the ossilations within and across cells in the anuran ear. The extent to which the NBS can be measured a priori in the ear will be developed. Charles Darwin wrote, sexual selection. This depends on the advantage which certain individuals have over others of the same sex and species solely in respect of reproduction. If this is read to indicate advantages not due to growth and development or behavior other than reproductive (where natural selection may act during habit activity in life) then it may still be possible to recognize cooperative reproductive activites under sexual selection if one sex and an individual of another sex cooperate to the disadvantage of other individuals of another gender. But because Darwin included same sex and species it is unclear how cooperation of one sex and one individual of another sex is also at the advantage of the species unless it is the same phenomenon in which case Darwin would not have need to have written same sex and species. Rather what it appears Darwin means is individual advantage in one sex over others of the same sex or the species as a whole regardless of sex. With the development of the modern synthesis it becomes problematic how single gender structure directed difference in the habits of life inherited do not mix with sexually selected differences. A mutation for locomotive behavior may lead to sexually selected competition or cooperation and thus though selected naturally is fixed sexually or a niche may selected naturally and then offer feed forwards for otherwise sexually selected dynamics. Of course a solution to this is to deny the existence of sexual selection altogether. With respect to beetles Darwin makes plain that the ticking sound is a sexual call that facilitates the finding locomotion of the male and the female (0832). He concludes this from the observation that bettles respond to each others ticks and then are found together. The anuran chorus he likewise attributes to an appropriate expression but actually saw the alternative when he noted that bull-frogs are inppropriate to our taste or ornamentation. He decides emphatically that the structure of vocal organs (in all cases) is attributed to sexual selection. It may not it may be androgen driven instead. Darwin describes sexual selection as of two kinds (pager1051) one between individuals of the same sex generally the males to drive away or kill their rivals (compete) the other is a struggle of competition to excite or charm the opposite sex and are selected by this sex. Some others have thought female frogs may be doing this with respect to mating calls. I will provide a different scenario where the male frogs are cooperating to produce a chorus that triggers a bargain with females when the cooperation reaches an NBS. Thus if cooberated the Darwinin narrative with respect to anuran chorus and vocal structure can be demonstrated as false. Darwin thought the effect was due to differences in timing of breeding but in a comparison of tropical and temperate chorus where such differences are evident and necessary it will be demonstrated tha t the same cooperation to using a choral series to put a mechanical electrical continuum into sync is the effient cause the difference in vocal structure.

The basic logical apparatus that establishes Dawins sexual selection is that because females are unarmed and unadorned and yet survive that vocal music strucutures allureing and exting the female (age 764) are the result of sexual and not natural selection. This will be shown suspect in cases wher females call instead of males and the reasons why. Macnamara etal wrote: Nevertheless, it is fair to say that cooperative games typically involve
bargaining tactics, which require some form of communication (including extended interaction) between players.However, this does not mean that any form of communication between players necessitates a cooperative game and the corresponding solution concepts - so-called 'unimprovable' outcomes such as Nash bargaining solutions (NBS).

We will see that any form of auditory anuran communication affects the electro-mechanical continuum of the ear necessitating unimprovable outcomes given the state of inheritance of monophyly underconsideration.
So how much can cooperative game theory, with its intrinsically binding 'contracts', help us to understand sexual interactions in biology, which is the domain of sexual selection? On the one hand, sexual reproduction, by definition, must involve interactions between organisms, however remote, to exchange genetic material. However, this does not require that the individuals involved are bound to any 'contract' they form in the process.

It will be shown that the nature of the chorus sounds produced by individual callers creates binding contracts serially that are negotiated with respect to density and length of the series which differentiates the serial chorus from some other one possibly useable by another species under the same syncing system of the genders.
Such stricture implies some form of enforcement that is external to the interaction, which may make sense in many human interactions occurring within legal and moral bounds. However, if commitments are not implicitly enforceable then games are non-cooperative. Furthermore, to presume that individuals will behave so as to avoid discommodating other interacting individuals, which is central to the 'unimprovable' solution concepts in cooperative games, is unrealistic for organisms that have evolved by natural selection.

The commitments are implicitly enforceable because of the random Brownian nature of the motion both in the electrical and the mechanical motions. So while a contract may be altered by speeding up a call or spacing it out differently in the average (the number of the next generation callers) the Brownina motion will enable enforcement of the behavioral tier in evolutionary time. This solves a common problem of relating ecological time to evolutionary time. Silence could be a threat and vocalization a promise. Silence would operate in behavorial time by affecting the choral ouput. Promising to exert metabolic energy to create audio sounds to feed in to to change the electrical continuum that only operates otherwise in silence is also behavorial . Amplexus thus silences when non reproductive (precalling silence duration) silence value is no longer effective in silenceing within the chorus itself. Thus the so-called limiting rescourse of the audio bandwith is only limiting when the series can not be any denser or shortend or lengthened. Competition will thus arise at

the species or levelsa beyond if ecosystem can not support multiple kinds of populations rather than invidiual population sizes. Maximization of payoffs are thus within a bandwith that can be reached by a given series dependent on its ordinality not cardinality. The cross eletro-mechanical continuum communications depend on kinds of series of choral singers not on any choral sound whatsoever. That is what Darwin had in mind when relating the taste in music to beauty and female choice. This is not what is happening physiologically even though the female may silence the male and the offspring spring forth.

The males (below)may be cooperating through electrical oscillations(from muscles to metabolism and nerves) affecting the mechanical oscillations generally (sound etc) no matter the biomechanics and not fighting nor under sexual selection at all.

Attenborough tries to make the case that hand waving and soft calling which can not be heard well in the rushing stream environment of the golden frog is a case of males fighting for a female but this could well be the case that for some physiological reason that ultra sound is not developed in these frogs as in China and Borneo that rather instead they utilize (the black and )gold motion through the eyes to stimulate the electrical oscillations in the ear making the call (in evolutionary time) only once /after directly encountering the balck and gold male that does not move unlike the female that will jump to the water. So the hand motion is a moving of something gold to see if the other male will jump and sound is used less. This is a cooperative behavior with the hand motion, and lack of motion on the bottom and the call on the top (two males) as the promise. The threat point is the ability to clasp hard on the female when other males also find the same motion if the female is jumping towards water. Instead, we suggest that animals cooperate to rear the largest number of offspring possible, because offspring are investments held in common. Generally, extant amphibians categorically are divisible into the rearing of offspring by external broadcast fertilization (anurans), internal sperm retention (urodeles), and internal egg development(caecilians). Social selection should be applicable in all three kinds of cases if sexual selection actually is not true. We introduce a notion of allocating time into various relationships to maximize cooperative or team fitness. In this theory, we can observe that diverse social organizations emerge from how individuals accure direct benefits from the relationships they develop with one another within diverse ecological contexts.

We suggest an approach that relies on the exchange of direct ecological benefits among cooperating animals without reference to genetic benefits In frogs and toads the development of eye-locomotion ability (which salamanders and caecilians in part lack) and the mechanical to electrical bidirectionality of the anuran auditory system provides the communication and coordination system necessary to/for teamwork as modeled with cooperative game theory independent of actual genetic benefits. The anuran chorus is the standard game that frogs and toads play to increase the number of the offspring in the next generation. Thus with these amphibians there is a literal not an analogical relationship as expressed by Rouhgarden et al with A social system develops from the interaction of individuals just as body parts develop from the interactions of tissues. The development time differentiation of the auditory system provides a structural morphological-cognitive variation of the behavioral tier that doubles in the social interaction of individuals thanks to the dual random motion in the ear with electrical oscillations and mechanical ones. This provides a special case wherein arguments against social selection can be tested. unimprovable enforceable contracts physiological continuum and behavioral variation vs discrete individual behaviors and actual correlated speciation patterns. & Doubts about how to define team play where mates may switch from season to season (social selection vs mutation and migration and drift). In our view of selection in relation to sex, the number of eggs produced by the female is of direct significance, and both males and females should have evolved to maximize this quantity by exchanging ecological benefits. Frogs and toads operate with a multiplayer bargaining structure where males cooperate to increase the choral volume so as to have common access in a given beneficial ecological niche that maximizes the number of eggs produced. If this explanation is correct then a non-cooperative signal network (expressed in bandwidth distribution amongst multiple species in the same ecological environment)game (for the same data set) will be shown to lead to malformed and inefficient behaviors and failure to maximize the number of offspring developing in the next generation. The issue of communication may end up being sufficiently defined by the ability to send signals from the electrical to the mechanical continuum or from the mechanical to the electrical by any means (proximate or ultimate etc) across the randomization effected by Brownian motion on either side. Individual strategies will thus be decided by the ability to effect particular influences (of frequency or amplitude or serial density or serial elongation or serial ordinality) through a transitive asymmetrical relation with respect to the entire trajectory of the behavioral and evolutionary tiers. Perception of the team fitness function thus became a convergent mathematizable sequence of

series when applied to amphibians. The applicability of ESSs will depend the nature of these maths regardless of the philosophy implicated. When Schiotz explains choral sound volume (Evolution of Anuran Mating Calls Ecological Aspects p 312) by, It may be explained as a result of a selection by females of males with the loudest voice, the voice that is heard best in the chorus he exemplifies the difference responded to by Roughgarden et al , that, The difference between our theory and sexual selection is that we claim selection in relation to sex operates through direct ecological benefits and exchange of these through social interactions. On the other hand, sexual selection theory stipulates selection occurs for genetic benefits and says almost nothing on sociality, and what purpose it should serve. Thus contrary to sexual selection narrative exemplified by Schoitz who used of female choice to explain the unexplained amplitude of the chorus, in a cooperative interpretation the males increase the volume because of constraint the particular timbre benefits ecologically and the female is not picking out males from the chorus but is offering a payoff once the volume gets loud enough(has had enough influence in behavioral time) and the randomness in the communication channel is no longer able to convey the message. The payoff timing thus may not be a result of only heritable variation but instead includes a developmental constraint in the behavioral tier due to the mathematics of serial ordination of sounds that must vibrate across two systems of physical force surfaces. The physiology of the anuran social selection system is such that these constraints are related directly to number of offspring produced because egg laying decreases the physical amplitude of the chorus ipso facto in terms of life history. Social selection thus allows one to differentiate the developmental constraints effect behaviorally on form-making from heritable trait variability in the form of the phenotype. This division of structural shape into two kinds of mathematical components seems to be missing from multiple level selection theories and is due to the novel theoretical formation of the two tier modeling that has expanded the range of hypotheses that can be formulated.
Male green frogs lower the pitch of acoustic signals in defense of territories: a possible dishonest signal of size? has been published but this may not be a form competition but instead a function of cooperation if

the lower pitch for instance preferentially stimulates the electrical oscillations so as to make either the individual ear more sensitive to pitches in general or offers a broader background sound in which more inclusion of callers is possible in the continuum (that may accumulate in behavioral time over years but is built up in its barganing solution (in the nervous system (years) or on the tectum membranes (miniutes) year by year (more so for the smaller frogs)). Thus the switch to lower frequencies may indicate conversion from individual threat points to the attrainment onto the team fitness function.

If this is applicable for frog chours frequencies in general than the difference in the calls of Acris

indicate a further development of the team fitness function of the noththern cricket frog over the sothern. These frogs do not move as much as others and the sothern hops more/further than the northern. This may be how the Southerns can get by with a less developed team fitness function.

Non-vocal ripples caused by the green frog below may mechanically trigger electrical oscilations and permit movement from individual fitness gradient inclinations to team fitness attainment as calls are produced.

In rejecting a standard sexual selection narrative interpretation the proposed difference in energy ( as measured by call number and duration for those who mate and those that do not (mating individuals call less and for shorter periods of time) may reflect a variable timbre character/characteristic of the dual continuum, dependent on total length and density and not be reflective of pure whole body energetic. Cooperative callers may be able to mate more by being properly situated in the density of the chorus so situated for the longest mechanical auditory pressure and this may be the bargain (NBS)established to keep the frogs at the breeding site which under an evolutionary stable strategy is the best for the local territory (average disrespecting establishment and variation).It would occur through social selection thus rather than sexual selection dependent on explanations of the first males at the site and females choosing males based only on calls. (frequency etc) evolutionarily.

to develop a discipline of social selection study in cooperative chours characteristics sustained by the physical interrelations of a mechanical and an electrical continuum in the anuran ear.

The study of anuran aural communication has largely been investigated in terms of signaling theory but given the dual structure of the ear organ it is possible that two way communication is not between

individuals but between individual effects on both components of the EAR. Cooperative bargains between individuals can result from coordinated two-way paths socially selected through the components to the exclusion of those routes used by others. Honest signaling theory (energetic constraints on call output) becomes much more involved (lots of calling for short time, little calling for long time) with the details of physiology within the frog than with the nature of the signal channel use cost between them. Complex serial effects of the chorus sound structure informs the strategic equilibrium acquired by structured cooperations. This offers an alternative to sex selection in the explanation of differences in sound use. http://neurocomputing.org/AuditoryInnerEar.aspx A dual mode auditory discrimination strategy is needed when a female frog approaches a pond surrounded by a multitude of calling males. The calls all merge together such that any temporal characteristics of individual calls are hidden until the female gets suffeciently close to a calling male. TRANSITION FROM MECHANICAL TO ELECTRICAL AFFECTS DOMINATING . Consequently, only the frequency and intensity components of their calls can initially be characterized. The frequency and intensity parameters are characterized by the dorsal medullary nucleus (DMN) and the superior olivary nucleus (SON). The torus semicircularis is where temporal pattern information is initially characterized just as it is in the tactile system. Background context of the temporal pattern information is accomplished by the thalamus.

SINCE THE MUSCLES CAN FREEZE THE BONES TO PREVENT LOUD NOISES FROM AFFECTING THE CONTINUUM, ENERGETIC CONSTRAINTS ON METABOLISM EFFECTS THE AMPLITUDES/ENERGY POSSIBLE CALL CHARACTERISTICS DEPEND NOT ON DETECTION AND TRANSMISSAL BUT ALSO ON THE STATE OF THE MUSCLES (AREOBIC ANAEROPBIC). TIGHT MUSCLE SPECIES LIKELY HAVE LOWER VARIANCE IN CALL PARAMETERS AND LOWER USE AMPLITUDE.

Page 292 The peripheral substrates and central mechanisms for processing airborne sounds develop in a sequential fashion throughout metamorphosis (B-Horowitz and Simmons 97), but the extent to which this developmental sequence is genetically mediated or environmentally modifiable is unknown.

Is sexual selection the main driving force in the evolution of anuran acoustic communication? Is the notion of communication able to trace out the work (forceXdistance) of sound transfer between individuals? Can biogeography assist in the analysis of the features of anuran calls? Can the anuran call rather than be bandwidth to increase individual matings rather function to increase the number of offspring that survive into the next generation? Can not call intensity, call rate, call duration call pitch and temporal pattern represent characteristics of a social bargain cooperatively reached with different parameters in different species rather than be traits shaped by sexual selection alone. To assert that all of the morphological, physiological and biochemical machinery involved in call production is molded by sexual selection remands an alternative by which such claims can be chosen. Social evolution provides an alternative perspective to test this claim. The energetic costs of calling work up differently into the characteristics for sexual selection and social selection. Under sexual selection the costs result in callers that increase efficiency by calling a lot right from the beginning of the season to those that call a little (conserve energy) over a long time. The energetic constraint yields call trait characteristics that are largely independent as the transmission itself is maximized per energy available. Social selection implicates the effect of energy onto the strucuture of the call such that strategically intriacate cooperative equilibriums are behaviorally sustained which yield info on offspring survival. The dual electric-mechanical nature of the ear plays no special role in sexual selection except insofar as it can increase transmission effiencey while in social selection it enables internal and external factors create large behavioral possibilities missed so far in the ethology of frogs and toads. Does hand waving of arrow posion frogs have something to do with the call? Muscles can keep loud sounds out so energetic constraints affect the loudest calls a cooperative bargain may achieve and not simply be a competition for bandwidth space.

The two patch difference is needed for the TWO continua socially adapted.

It is not about one continuum with different frequency sensitivities it is two. That is what one thinks is consequent with the following reasoning If anurans socially evolved sound transfers as means to negotiate a behavioral bargain then either the more primitive frogs that lack such means to negotiate must utilize a different mechanism to effect cooperation and this (or the lack thereof) must be able to physically indicate a different mode preexisted such use of sound in terms of the offspring themselves.

The different mode is single patches in the papilliae shared with salamanders.

The use of a tympanum may have been socially selected away from Leiopelmids and Ascaphids which utilized internal fertilization developed into internal egg development in caecilinans and internal fertilization (spermatophore for the intromittant organ) in salamanders. Late jump recovery is unique in Leiopelmatidae. When Leiopelmatid spp. jump, they land in a "belly flop" fashion repositioning their limbs for takeoff for the next jump only after hitting the ground with the ventral surface of the torso. After landing, Ascaphus skids to a halt before recovering. The appearance of early jump recovery in more advanced taxa Jump recovery may be nothing advanced but simply the adapation in forms which have tight muscels needed to hold ear bones in place so as not to have too loud noises affect nervous input.

Assume that anuran parents make decisions in behavioral time. Are there such things a frog and toad parents able to effect decisions concerning their offspring?

Well if you were this mommy or daddy it seems that behavioral decisions made (jump into dense foliage that may scrape of eggs to escape a predator vs jump into deep clear water where both eggs and parent are easily seen ) by her can quite dramatically affect the future of her offspring

The auditory periphery (and bulbing of the kinocilia on the inside with double patching) of evolved anurans arose as part of amphibian social evolution to permit sound nash bargain negotiations through mechanical adaptations. There was no large north and south split geographically (Darlington 1957) here but rather a two tier process of social selection on the developing electrical connection. This may indicate that electical continuum is cardinally double the equipollent equivalent of the mechanical one. Transfinte numbers may directly be used to distinguish brain divisions between the basal frogs and salamanders (caecilians?) with the other frogs. Transifintes may be used to bridge/construct the applied physics and evolutionary biology of cooperation involved. Ryan supports a view of the relation between structure and function that denies any particular hypothesis of adaptive significance of the amphibian papilla alters his position on the relation between

neuroanatomical constraint and speciation. He contends that B and C forms can not speciate as much because of the lack of a middle waveform peak caused by higher frequency sensitivites regardless of whether this cardinality be examined as ordinally adapted to a traveling wave rather than a resonance model. If the adaptation is a social evolution of nash equilibrium communications afforded by dual electrical-mechanical continuua the cardinal-ordinal ordertype could speciate B and C forms as much as D if the behavioral preferences are differnetly locomoted in the pay offs. The fact that there is a higher probablity of a mutation causing a change in the call frequency with increased sensitivities to higher frequencies depends in social selection on the polymorphism the cooperative behavior sustains in the species population. Only if decreased variance (less movement around the pay off matrix possiblities) increases the polymorphic mutation probablity will the speciation necessarily be concomittant with the strucuctral differences. Function depends differently in volume than in mass in game theory. "When high-quality conspecifics resemble heterospecifics, females may be unable to engage effectively in both species recognition (identification of conspecifics) and mate-quality recognition (identification of high-quality mates). Consequently, females that engage primarily in mate-quality recognition may risk heterospecific matings, and females that engage primarily in species recognition may risk mating with low-quality mates. I examined the evolutionary consequences of this conflict between species and matequality recognition in spadefoot toads, Spea multiplicata. I compared mate preferences and the fitness consequences of these preferences in spadefoot toad populations that did and did not overlap with congeners. In non-overlapping populations, S. multiplicata females preferred an extreme call character resembling that of heterospecifics, and they had more eggs fertilized. In overlapping populations, S. multiplicata females preferred those call characteristics that were closest to the norm for their population, and they did not receive the benefit of enhanced fertilization success. Thus, S. multiplicata females appear to trade off species and mate-quality recognition, such that those co-occurring with heterospecifics forgo the benefits of high-quality matings to ensure conspecific matings. These results suggest that the interaction between species and mate-quality recognition may influence mate choice decisions in important and nonintuitive ways." This may mean that behavior guided social selection may bias a polymorphic mutation maintence either for the mode or for the extreme and show that Ryan's hypothesis that simple increase in frequency range and probablity of mutation is wrong. It does not require a species-selection alternative but simply a two tier behavior - evolution social selection of cooperation for more young in the next generation. For mammals and some birds -- "Thus, vibrations within the organ of Corti are sensed and then force is generated in synchrony to increase the vibrations. In this review, the role of force production within the cochlea to produce cochlear amplification and the micromechanical mechanisms that underlie it are discussed." This force is wholly in the structure of the periferal ear morphogeny not as force in the air by moving tissue. It is how amphbians have different papilla. Spontaneous oscillations of sterocilliary bundles in low frequency may be postive displacements to test

in the laterla line system and for use with muscles that alter the displacements themselves. "Spontaneous oscillations persisted, becoming very narrow-band, when the hair bundle was loaded with a tectorial membrane mass." the use of spontaneous oscillations in coopeative group call bargain may be how and why there is a difference between B, C and D forms. OP CIT Besides being a sensory structure, the hair bundle can also exhibit motile activity exemplified by the spontaneous periodic motion in its position, with the tip of the bundle oscillating also along the E-I axis. Spontaneous bundle oscillations have been reported at frequencies from 5 to 100 Hz in hair cell epithelia of lower vertebrates such as the turtle (4) and frog (57). Water rippling in green frogs may be a means to trigger spontaneous bundlge ossiclations by setting up resonances. The multiple ripples of many frogs in chorus as the intensity is dispersred to low frequencies entropically may also contribute to spontaneous effects. Also behavioral muscle mediated motions may interact with this (turtle courtship in Trachchmeys may be an example). Joan Roughgarden has challenged the notion of Darwin's sexual selection with a two tier (behaviorevolution) use of Nash equilibria which may manifest metaphysically the difference of penetrative surface and bodily surface suggested by Kant. I suggest that guanosine effects through microtubules mediates the different information cycle suggested by the math of social evolution and can manifest a monohierarchy.

Richard Dawkins said in 2006

"Would be similar to a scientist of... the future saying something like this: Well of course we dont literally believe in the double helix anymore But, lets think- what does the double helix have to tell us today" Whats wrong with the body of evolutionary theory? Generally there is no concept that enables a homogeneity towards a decision to know when one is observing a level of organization or a level of selection nor a common intuition of the same. This is because there is no way to know what body is. Dakwins thinks that Roughgardens notion of body of St. Paul is not one that one scientifically need bother with. He only thought this within a difference of emotional religious thought rather than

between metaphysics and transcendental philosophy. No one seems to know of the difference between flesh and bodies. Is this merely due to living things having been found to all contain cells? It cannot be. Kant had a detailed vision of the difference of fluids into strands , laminae and blocks which are segmented into solidity. This is missing from the notion of body discussed by Roughgarden and Dakwins. Roughgarden is not only wooing people of religion into the body of evolutionary thought but all people. DawkinsWhy bother to express truthsif they are truths? Why bother to make up a parable?, to make up this, tenuous allegory when you can just tell people the truth. The truth is perfectly simple. Why bother with going back to the Bible? and seaking out some sort of of analogy Why not just teach natural selection , teach evolution . It he way it is It is perfectly simple. You do not need to ..this strained , unnatural , seizing of a possible analogy from the Bible. Evolution is not perfectly simple. It may be that simple to a genic selectionist but if it is more than bookkeeping it is not simply this simple. If the fibers , laminae and blocks are biological in this sense of simplicity then every one, Bible believers and those without religion, all , have a subjective connection to the bodies under evolution that express the truth of change of the bodies. This is not strictly symbolic as Dawkins presumed but formal but beyond the logical of A and not A but is rather of A and A of Kant. So though one may think you can just tell them the truth it is not so simple since the statements themselves can inhibit to collection of a common intuition under different concepts. Roughgarden is developing a different concept of cooperation rather than conflict. We do not literally believe in the belief of DNA of Watson and Crick as expressed by Richard but instead we think of it as an elementary part of a body under repulsion and attraction that leads to bodies that can contain multiple sexes, regardless of the fact that all living bodies are made up of cells. Dawkins notion of truth told is within a particularly vested interest in a simplistic matheis directum. He inappropriately projected onto religion what is a difference between metaphysics and transcendental philosophy under the guise of politics. There is a good reason beyond the science for returning to seizing towards something different while one moves in a circle around from unity to plurality through multiplicity no matter the diversity.

That the double helix is not true anymore is thus not due to religion in the particular but due to phoronomy being applied in the opposite motions that two sexes can mechanically/dynamically instantiate in the body of evolutionary thought. There is a service here because Darwin had sought to make common locker room bravado comments for different kinds of life from ants to primates in the same mathematical philosophy without having thought out the possibilities otherwise first. Dawinks did not do this either.

Charlotte McKechnie and David Shunker of the University of Endiburgh challenge the truth of Roughgardens depreciation of sexual selection theory. They assert that the social context of Darwin does not a priori undermine the basic empirical truthof examples and interpretations from them. To state this requires some metaphysical sense, what is it? This hidden metaphysics comes out when the authors claim, there is no rationalization for equating sexual selection with a particular gender construct. The authors obtain their opinion by a general metaphysics unnamed than that which has rather specific implications as in Rougardens instead. Darwin equally tried to imagine jealously and other unusual traits for insects so asking how insects may friend each other on the facebook of a different theory is the same question as if there can be cooperation in plant sexes/genders in the body of Roughgarden. Should macrothermodynamic substance stability of aggregations as Dawkins belittled the DNA love of the future enable a form of cooperation within Roughgardens two tier theory that so leads to a better and/or newer intuition than that of the Darwinian stepping stone suggested by the Endinburgh authors, the metaphysical differences will be moot. Thus we would have rationalizing judgment applied as if true. Working this out may lead to extremely detailed understanding of plant behavior that we currently do not possess. That such is not just pie in the sky imaginePlants seeding seed are falling either to the Earth or the Sun. Evergreen trees may be doing this outside the branch and deciduous trees within the branch. Plant behavior depends on utilizing clinamatic deviations from a gravity described trajectory across generations, whether sexually or not. Sexual conflict may be defined as separations and discord of the deviations and cooperation as combinations and harmony of the deviations with respect to the particular developmental behavioral variations that arise during the growth and development of the tree relayed onto the evolutionary tier. The amount of behavioral variation in plants may even eventually appear to be larger than that phenotypically in animals and not simply dependent on squash shape variation per say. With in an increased range of hypotheses and space of plauisbly true ones the mental relations that Darwin supposed for all kinds of animals may be supplanted through a larger domain of understanding of behaviorally structured functionalities' variation (coming from plants as well) and the examples and interpretations othewise concurrent will lack the then on-going rationalized unity and thus be

depreciated to zero itself.

Social Evolution in Plants If a two tier behavioral variation plus social evolution is to replace sexual selection then cooperation and competition should be definable for plant sex difference manifestations even though the physical contact through a nervous system Is lacking.

Why are unisexual reproductive units of plants called diclinous, imperfect, or incomplete.

A local mating environment with an excess of males or females could bring about severe local competition for mates of either sex. An individual that changes to the minority sex will be favoured in such an environment. http://www.ias.ac.in/resonance/Nov1998/pdf/Nov1998p30-39.pdf

Is this really a now or never replacement of maleness for femaleness but could it no rather be cognized as a cooperation between male and female plant genders under social evolution?

1. Sexual Selection Cannot be Replaced by Cooperative Game Theory (and It Doesn't Need Replacing)
o Sasha R. X. Dall o John m. McNamara, Nina Wedell, David J. Hosken
Centre for Ecology & Conservation, University of Exeter, Cornwall Campus, Termough, Penryn, UK

Sasha R.X. Dall1*, John M. McNamara2, Nina Wedell1, & David J. Hosken1 1Centre for Ecology & Conservation, University of Exeter, Cornwall Campus, Tremough, Penryn, TR10 9EZ, UK. 2School of Mathematics, University of Bristol, University Walk, Clifton, Bristol BS8 1TW, UK Another week and another attack on Darwin, but as usual, rumours of problems with Darwinian theory are grossly exaggerated, or in this particular instance, plain wrong. In this latest instalment Roughgarden et al. [1] suggested: 1) the Theory of Sexual Selection is inadequate, 2) that it needs replacing and 3) that cooperative game theory should be the replacement. We will not go into claims 1 and 2 (they are being addressed by others) other than to note they are incorrect, are a misrepresentation of the facts, and we additionally remind readers that, as Darwin himself stated in his great sexual selection treatise [2], "False facts are highly injurious to the progress of science". Instead we will deal with claim 3, that cooperative game theory is an ideal replacement for current sexual selection theory paying particular attention to the arguments presented by Roughgarden et al. [1]

(referred to as R henceforth) to make their case. Our main contention is that R misrepresent the philosophical basis of cooperative game theory, which obscures its fundamental unsuitability for analysing evolved systems in general, including the vast majority of sexual interactions. They introduce cooperative games as alternative to sexual selection by distinguishing them from non-cooperative games. However, in doing so they deviate significantly from classical economic theory textbook distinctions between the two types of game. To quote from R: "In competitive [non-cooperative] games, the players do not communicate. ... In cooperative games, players make threats, promises, and side payments to each other; play together as teams; and form and dissolve coalitions." (p965; text in brackets added). This contrasts with a quote from the preface of a popular reference book on economic game theory [3]: "A game is cooperative if commitments --agreements, promises, threats -- - are fully binding and enforcing. It is non-cooperative if commitments are not enforceable (note that pre-play communication between players does not imply that any agreements that may have been reached are enforceable)." Thus contrary to R, the distinction between cooperative and non-cooperative games lies in the assumption of implicitly (a priori) binding 'contracts' between players; in this sense: "...the phenomenon of cooperation is to a certain extent assumed and thus not subject to a complete analysis" [4]. Nevertheless, it is fair to say that cooperative games typically involve bargaining tactics, which require some form of communication (including extended interaction) between players.

Can the tuning of a frog ear and call sound be understood as a behavioral (developmental) bargaining? Via bidirectional transduction from metabolic energy reservors through electrical matching to aduitory pressures?

However, this does not mean that any form of communication between players necessitates a cooperative game and the corresponding solution concepts - so-called 'unimprovable' outcomes such as Nash bargaining solutions (NBS). In fact, a whole branch of evolutionary game theory, which is fundamentally non-cooperative [4], is devoted to the study of communication between animals: signalling theory [5]. Furthermore, since the early 1980s economists have devoted substantial effort to analysing bargaining tactics in non-cooperative games [6]. So how much can cooperative game theory, with its intrinsically binding 'contracts', help us to understand sexual interactions in biology, which is the domain of sexual selection? On the one hand, sexual reproduction, by definition, must involve interactions between organisms, however remote, to exchange genetic material. However, this does not require that the individuals involved are bound to any 'contract' they form in the process.

The frog contract is have equal energy resonances between hair cells electrical resonances and mechanical resonances. The social evolution of a chorus provides the mechanical resonances while the behavioral interactions in the chorus may be directed by metabolic matching through the electrical resonances. Thus the two tiers are

directly associated.

Such stricture implies some form of enforcement that is external to the interaction, which may make sense in many human interactions occurring within legal and moral bounds. However, if commitments are not implicitly enforceable then games are non-cooperative. Furthermore, to presume that individuals will behave so as to avoid discommodating other interacting individuals, which is central to the 'unimprovable' solution concepts in cooperative games, is unrealistic for organisms that have evolved by natural selection.

But this may be the case of multiple species choruses dividing up the auditory spectrum where by development under metabolic control avoided discommodating and thus divides with speciation. The prediction would be that macrothermodynamic side payments as individuals team up into a chorus accrue during differential expression of otherwise socially evolved populations genes.

Indeed, it is precisely the Darwinian notion of populations of reproductive lineages competing for limited resources that has led researchers to reject cooperative game theory as irrelevant to evolutionary biology [4]. R's juxtaposed logic is illustrated clearly in the analysis of the hypothetical asymmetric game detailed under the "Logic of bargaining and side payments" section (p965- 966). In fact, the hypothetical payoff matrix presented on p965 seems a strange one to illustrate the heuristic value of cooperative games in biology; since the best payoff (in arbitrary units) Player 2 can achieve from bargaining is 7.2 (Fig 1 in R) compared to 8 if it opted out and played non-cooperatively, it is hard to imagine why it should ever agree to bargain in the first place if it is acting to maximise its payoffs (as will be favoured by natural selection). Moreover, the hypothetical sequence of threats and promises outlined in R, which is used to demonstrate how the bargaining solution can be reached, is equally unconvincing from a biological perspective. Why should Player 1's threat to play A (even if it were made credible) ever induce Player 2 to change its behaviour, given that it would gain 6 if it kept on playing A and 5 if it switched to B? The only rational reason that such threats should be effective is if Player 1's well-being was important to Player 2, which is difficult to justify for unrelated players without including additional assumptions (e.g. reciprocal interactions). In fact, most organisms are expected to minimise their relatedness to sexual partners except under very limited circumstances. Furthermore, R strongly argue that sexual reproduction is a social activity involving repeated interactions between participating individuals ensuring negotiation and threats can take place. This underpins the cooperative game theoretic approach they propose. Indeed, R admit that "the difficult issue for the cooperative game theory approach is whether the animals can carry out team play and can discern the team fitness function whose gradient they should climb" (p966).

The sexually dimorphic ear size may be explained by

repeating interactions
However, for the majority of animals (not to mention plants, many of which reproduce sexually), mating is a once in a lifetime encounter between individuals and does not involve repeated interactions, let alone opportunities to 'discern the team fitness function'. For example, external fertilizers such as broadcast spawners simply shed their gametes without ever encountering one another, and the majority of arthropods will only encounter and mate with the same individual once. R's justification for their perspective focuses exclusively on behavior that seems largely restricted to hyper-socialized vertebrates,

Macrothermodynamics however applies more so in the less socialized species

which only represent a tiny minority of the vast number of species, extant or otherwise. All sexually reproducing organisms need to be encompassed in any general theory of sexual interactions. This is not the case for cooperative game theory.

It may very well be.

Frog chorus bargaining may award side payments for movement of whrilin by MyosinXVA. The Chorus itself is a series of sounds impacting across the Brownian electrical-mechanical continuum. The chorus itself by the structure of series rewards in evolutionary time the amounts of MyosinxVA and whrilin(or equivalents) of those current injectable electrical oscillations that most probably are associated with calling behavior that increases the amplitude of the chorus. Thus a frog could hop into and out of positions in the chorus or call more on its own once it has found the correct temporality wherein its intracellular noise feeds into the Brownian motion of all its developed hair cells but thusly puts itself in sync with the continued production of intracellular electrical noise in individually macrothermodyanmically thermostated cells . Continued calling could maintain this balance and trigger amplexus in those females where the sync is overwhelmed by the chorus and the intracellular noise no longer feeds through metabolically. The males and females bargain over what sync is the best for the most number of offspring (feeback of evoltutionary onto behavoral tier). The silent pre-chorus period could function as setting up of this state where the sync will serially be consequent by hopping (muscular)behavior (muscular behahvior without sound). Muscular metabolite overwhemlming electro-mechanical sync continuity. Thus a Nash bargaining solution is obtained amongst the mutiple equilibria possible in the hop behavioral chorus spatial position, timbre of the caller, the chorus seriality, withing that side payment system of the genes that elongate the hairs and influence the metabolic to electrical noise relation independently of the sound environment.

Winfried Denk, a, b and Watt W. Webba, b

School of Applied and Engineering Physics, Cornell University, Ithaca, New York, USA Department of Physics, Cornell University, Ithaca, New York, USA

Received 8 November 1991; accepted 3 February 1992. Available online 17 March 2003. Hearing Research Volume 60, Issue 1, June 1992, Pages 89-102

Metaphysical analysis suggests that gender initiation in an organized body occurs when given a prior ponderosity a new level of cohesion is achieved sexually by coercing otherwise imponderable matter in the exhaustions of locomotion. This supports the idea that sexual/gender teams cooperate rather than conflict.

This analysis provides the "super theory" Hurd contended is lacking in Rougarden's "Genial Gene"

Hurd PL, 2009. Pitting the boys against the girls (Book review: Roughgarden J, The Genial Gene:
Deconstructing Darwinian Fitness), Trends in Ecology and Evolution. 24: 646-647. (doi:10.1016/j.tree.2009.08.010, supplemental notes).

Homosexuality would be adaptive for both sides in the prior pondersosity but because the new level of cohesion is the target of the gender origination the team's cooperation is a function of both coersion and cohesion as a whole. The failure to cognize this does give justice to Roughgarden 's claim of "homophobic" responses to the 2006 Science article.

Hurd contends that Roughgarden's use of the phrase "is it true?" "loaded language" but on listening to Dawkins (above) it is clear that this is the question that needs to be answered.

Kant's Opus Postuum also provides the reason why social evolution can be congruent with social justice but this goes through the particular mathematics of threat points in the NBS (and ESS vs NCE) and the relation to philosophy not to the particular social issues of humans

themselves. Roughgarden has not said how morality may improve science but this is also possible but is in the relation of scientists IN congregations, not philosophical differences of ID vs EVO etc. http://www.psych.ualberta.ca/~phurd/cruft/Roughgarden-notes.pdf

"Discern" may be quite definitely understood as explored in the page "Anuran Chorus" as a frog changes or does not change signal frequency in the context of background sound.

The key is in differentiating the penetrative surface genetically from the bodily one in general. Thus the four categories of social evolution directums (goal achieved by team play with proximity more important, goal achieved by side payments with proximity less important, non-goal cooperation exhausted to outcome spatially, pure competition) depend on the relations of the empty spaces not the voids in the constructability of the games as the moving forces are enclosed.

Following in the path of Gibbs where he forsook complete adequacy of all manifestations that accompany the union of atoms if we restrict attention to equal probabilities as he did and note as commented on by Plank with respect to finite numbers of particles relative to infinite divisibility and supposing equal probabilities for each gender then under these phase considerations it is possible to construct with the Macrothermodynamic thermostat levels of organization biologically as additional physical phases ((plasma, gas, liquid, solid)that are viewed permanently))to those permanent traits specifically called ornaments and armaments as resulting from expulsion of water under monoheirarchy constructions of social evolution and aggregation of higher energy chemicals by thermodynamic activity in the lower level. This can be integrated into the two tier theory of Roughgarden and explain an actual evolutionary result of phenomenological thermodynamics.

A matter cannot be penetrated by the compression of another. Thus genders which coerece more than the former ponderability cannot penetrated the surface any further than that biophysically possible through histogenic and morphogenic compression. Sexes arise on ponderosity by the new empty ponderable space created with moved imponderables through dynamically filling of impenetrability not absolute impenetrability physically. Impact of Fisher genetics and pressure of Wright affect this filling dynamically. There is a difference in the maths used that affect the philosophy of the relation to infinite divisibility of space. Nowak attempted to relate cooperation to levels of organization but mistakes the aggregations of

atoms (or naively perceptions) with a moving force of pressures (mutation, selection, immigration) under a difference of penetration and compression populationally extending reproductively. The view fails on the difference of math and philosophy. That is what is done to him (or me). He fails to apply Kant's understanding of Leibniz.

Ackermans function and infinite exponentials may help to model and work out the differences in opinions about runaway processes. How does runaway interact with entropy as understood by Wright? This may be approached with game theory differentiated into 17 2-case linearly independent translation wall paper groups with 3-D symmetry groups rolling on the surface.

If the runaway process exists the traits created thusly may not house as much energy in their structure. The importance of an overall structure is to distinguish the amounts of energy in various traits. If however a DNA computer could be interfaced with the dynamic process that gave rise to the exponential increase it may be able to get more kinetic energy out for the lower potential energy inherenet.

Can a theory enable one to predict a class of outcomes of the runaway process episodic spurt? As compared with general adaptive evolution? How does this episode fit into the entropy of wrights process and the adaaptive landscape changes in general;??

By noting that after faster runaway increase than under natural selection (with males and females) and the diminishing of the males due being over conspicuous in the exponential expression of the genes not making it into breeding group and thus not The runaway process is dependent on a individual (male female)dyad, synergisitically proceeding beyond the point possible in with natural selection (and could be made useful in general behavioral influences in a two tier model) This defines a general definition of acceleration of behavior onto entropy directed gene fixation process and produces un limited velocity increases in the embrological possibility of energy related to form-making in particluars of a lineage.

Thus depending on how the relation of the first cause of some advantage not related to preference and the particulars of the trait development are physically arranged the rapidity in the later stages Fisher said could be modeled by tetration or higher order Ackerman functions relative to generalized entropy timed gene fixations in the particular enthalpies involved (with and without certain amounts of

Gladsyhev reverse thermo flow) in the symmetric game in question per mutants.

As this value reaches its check, the relative standstill stability accrues when the change in ordinality (not cardinality since this post check stage is affected by sexual selection or behavioral selection so called) does not affect any change cardinality but if the prevention of males reaching the breeding structure is cut at "roots", a change in cardinality may permit a different ordinal series able to achieve a higher cardinality through a different ordinal increase path. Transfinite exponentials mapped back onto finite exponentials can show this? This can happen because the ethaply effects are just as important as the entropy ones in this situation(attraction and repulsions). This is not modeled in the general notion of equilibrium so far. It should be possible where preference evolution can go beyond strategy evolution.

These brief episodes ( such changes do not go on in most existing species) are examples of actual infinity dissolved or diffused in nature and can happen by behavioral team preference under perturbations of payoffs in addition to strict sexual female choice preference provided complement stratagies exist.

Wright thought that recessiveness was best thought of as inhibited (chemical) activity but it is more properly thought of as penetration without compression while dominance is penetration with compression (nothing to do with modifiers). The evolution of dominance thus depends on relative impenetrability. Philosophical concern for purpose did not consider the difference in density formed by empty spaces man-made bodies have which are not found in nature with any thought of absolute impenetrability (Fisher violated this when he thought only about modifiers changing dominance).

The 17 wallpapers 2-D symmetries with 3d symmetry orbi-spheres rolling applied to Fisher runaway scenarios generalized between two tiers across strategy and preference evolution is a case of fluidities expansive elasticity framed (original or derivative). Thus chemical (attraction -repulsions in the difference of recessive/ dominate) turn original elasticity into derived ones dependent on virial assignments tagged with thermostats.

New Ideas on Community Ecology- Implications for Evolutionary Ecology Until now, ecologists have attempted to understand a community by examining it itself, dissecting the processes that occur within it in ever greater detail The history over geological time is not considered. Heads Molecular Panbigeography of the Tropics page 452

A two tier process of social selection permits re-integration of ecology with evolution. This would mean Gould was mistaken about the intersection of evolution and ecology. Attempts to link via biomass ecosystems to ecology then needs a consideration of how the divisions of life and Earth evolving together can construct local communities given contingent past divisions/ seperations. This can be done if the community evolution has the theoretical potential to utilize tetrations rather than exponentiations but it is not essentialistic or typological as Mayr would have argued since it is the contingent vicariances that simply constrain the space structure not the space itself that the community can function in .

Social selection can be elaborated to distribute regional distributions of species within exponential combinations per tetrations for the whole ecosystem differences affecting differently the local components of biomass changes.

Social Selection: To cooperate or to compete, that is the question. Will the higher tier please stand up, that is the answer. Let us look at the case of Ambystomatid salamander spermatophore capping behavior. Males will tend

to put their spermatophores on top of the spermatophores of other males. This activity is extant in only one salamander family so far known. Is this a cooperative or a competitive behavior? Are males making it easier for females to find sperm or are they trying to keep other males from inseminating females? How can we know? Does this behavior send a signal to other salamanders at all? What is the process whereby this behavioral action is chosen and what is the outcome (choice/context) of this activity? According to Dall et al if there is any game theory application at all, it is in the negotiation rules rather than the unconditional choice of action because as they claim, the level of application is higher than the behavioral preference at the time. Alger has set up a mathematical system where this difference can retained even in the face of Roughgarden et als response that they have simply reapplied the Axelrod fundamental form contextually for all choices unconditional or otherwise. But if it is agreed that the outcomes must result from some game kinematics where the players can communicate only through the sequence of their own behavior then it does matter if the communication serialized by different behavior concussions is a signal or not when it comes to the dynamics wherein a specific action is chosen in a sequence that may be extended under negotiation (cooperative vs competitive). It appears that social selection is providing behavioral ecology with an end run around the tissue specific properties of the nervous system but when behavioral activity can format sequentially team work in reality nervous system competition (with itself and others it is communicating with) is found to have an even greater role in explaining evolution of scociality than is possible when competition is negotiated by human constructs under legal analogy alone. Creature communication has more stability evolutionarily when considered as a two tier process with behavior the lower and evolutionary ESS the higher than the reverse where human signaling theory (communicating cost) generates the distinction between the behavior rule as the higher and the act as the lower. Creatures are not consumers. They creep rather than cash out. How are the mathematical elements of game theory to apply to biology then? Can simple definitions be used to make inform an intuition towards the analog. It is clear that game theory is analogous to actions in nature. How exactly the logic and formations apply/ies depends on empirical data. But given that could organisms posses binding a priori contracts. Clearly organisms other than human can not exert the same kind of purposivity imagined by man and often made legal in contracts, but is there a metaphysics that enables one to account to living things the equivalent (for actions in dynamics) in the way that creatures communicate body English and colors etc? Does this not indicate that signaling theory itself based on human communication is itself flawed when applied to creature communication behind the difference of cooperative and competitive game? If the communication entails a bound cooperative plan can not the media of two-way cross sexual communicas be bound within the common development of the sexes independent of the formation of sex tissue? Thus since cellular development and cell memory is both within the body of the individual but beyond the reach of the behavior proximately the enforcement of the cooperative plan may be physical, chemical or

biological but be external to the interaction which is dependent on particular cell activities only. Epigenetic foundations for contract enforcement need to return to an external enforcement mechanism as seems to be the result of using human analogized signaling theory. Signaling theory depends on a receiver and transmitter, like a radio station and radio receiver, a telephone caller and a telephone receiver, Do non-human living things actually signal each other? There is not doubt that the creatures affect the senses of other creatures and even plants may be thought to react to various kinds of stimulus but are these action and reaction of living things signals in our human sense? Path analysis through this divided nervous system shows that Hamiltons rule of kn fitness is not needed to determine what the kin would do. Kin depend on the formation of the sex tissue while paths through lines of hereditity that only use a divided nervous system do not. Costly acts or costly signals are not the norm for nervous system divides but rather are expressions of deviations (spec ific action/reactions) from the series into a different sequence of concussions (mutations as fluctuation/variations). What we take for signals may simply be 1-D symmetry encodings of various moving forces and or forced directions. Thus there may be little difference between the signal and carrier for creatures unlike our invented technology.

Social Evolution

Social selection has been discussed lengthily with respect to sexual dyads but if the Nash bargaining solution, in the face of Pareto-inefficiencies , exists for differences between the sexes, a question remains as to whether there is some class or contingency of traits (that may not be sex-linked) that can increase during population increases and be ESS within soocial evolution generally. As genes becomes fixed it is possible that the irreversibility of the process itself may feedback, through entropy itself, simultaneously- to - the individual and the equilibrium the game model is targeting. Entropy and evolution has been discussed before but in the two-tier set up of social evolution it is possible to give an extended meaning to Wrights use of orthogenesis as instances of a kind of entropic orthoselection combining the two tiers. This shows that there is no real difference of preference and strategy evolution and that Fishers analogy to kinetic theory of gases can be modified in models where Nash competitive equilibria migrate into Nash bargaining solutions, even while Hamiltons rule is operating. The reproductive transactions so defined enable a new intuition of how individual bodies can be conceptually dissected. This will assist

taxonomists as well as evolutionists when dealing with overlapping phenotypic trait classes of current collections where formal differences still remain. Social evolution is beginning to assist in a new vision of linkage group correleation and epistasis.

Genetic Code Metaphysics and Dynamical Natural Philosophy Applied to Social Evolution