Monogamy within the Termite World: Mate Choice and Colonial Structure
|
|
- Mervin Allen
- 5 years ago
- Views:
Transcription
1 Becca Knox Term Paper 4/10/2013 Monogamy within the Termite World: Mate Choice and Colonial Structure Abstract Serial monogamy found in social insects, as a contrast to the promiscuity displayed by many organisms, contributes to profound advantages within a colony. For termites, mate choices often occur before copulation; the colony is first founded and the nest is secured before sexual reproduction transpires. In addition, the lifetime pairings are formed early in life. Bonds of this type are as strong of a commitment as the joining of two cells to create a zygote it is an extreme sense of commitment. Most often found in eusocial insects, the reproductive division of labor from sterile castes, and decreased mating variation as a result of this monogamy imply that lifetime commitment is a major factor in contributing to higher levels of organization within colonies. In addition, the lack of conflict over parental dominance has led to more harmonious relationships between offspring and parents. From an evolutionary standpoint, eusociality and monogamy may have stemmed from such characteristics as the overlap of generations, the advantages of group defense and colony structure, as well as the possibility to inherit the nest by offspring. The factors involved with mate choice and colony structure vary between different monogamous insects. However, recent studies on the mating biology of eusocial living have shown that termites make fascinating study species to decipher the relationship between eusociality and monogamous systems of mating. In fact, researchers have shown that even after long separations from colonies, termites are still able to recognize their colony mates. Though some antagonistic behavior was displayed, no mortality ensued. Cooperation among termites from the same colony are ways in which the species ensure that ecological competition and pressure does not promote internal aggression (Korb and Foster 2010). Kin recognition among
2 termites may be influenced by cues that are encoded genetically such as chemical olfactory cues, as well as differences in greeting behavior. These aspects of recognition raise interesting questions about how lifetime mates are chosen and how recognition of the mate becomes established. Introduction Eusociality and monogamy are often closely associated with one another within the insect world. Multiple theories about the evolutionary relationship between lifetime partnership and eusociality are important areas of study, especially as the benefits of monogamy and kin selection come to the forefront. There are several hypotheses for the evolution of this monogamous structure in termites, many of which emphasize the role of the male as crucial to the colony in terms of defense and sperm production (Nalepa and Jones, 1990). The biology of termite queens and kings as well as the establishment of different castes are fundamental aspects of the way the colony functions and becomes successful. The system of haplodiploidy and kin selection are also factors involved with the evolution of monogamy and eusociality, which contribute to the stability of a termite colony (Fromhage and Kokko, 2011). The benefits of monogamy for eusocial insects are multitudinous and include amicable cooperation among offspring and parents where conflict would lead to their demise, possible nest inheritance, and promotion of stability within the colony (Gadau and Fewell 2009). Researchers have discovered that termites make for intriguing study species to explore the relationship between eusociality and monogamous behavior, especially since mate choice is not random, but rather carefully selected before copulation occurs. In addition, termites are the second largest animal taxon to express monogamous behavior in which both parents are involved with the care Knox, 2
3 of the colony; termites are therefore a crucial component of serial monogamy research (Shellman-Reeve 1999). Though this model of monogamy does not apply to all species of termites, nor all insects that have evolved eusocially, the differences in mating patterns among other termite species and eusocial hymenoptera make for exciting opportunities for study and comparison. In many cases, the polyandry discovered within other colonies emphasizes the peaceful interactions and other advantages found within monogamous super organisms. Mate Choice and the Advantages of Monogamy Because the commitment of termite queens and kings is so substantial, they choose their partner fastidiously. Evidence for non-random mating has been discovered within populations of Coptotermes lacteus in which the different colonies were significantly deficient in heterozygotes, indicating that mate choice was selective (Thompson et al. 2006). However, it is difficult to tell whether the inbreeding of these termites resulted from ecological constraints or actual genetic preferences (Thompson et al. 2006). Females and males use different physiological characteristics of the other in order to make a mate choice. For instance, females often use large head size variance as a determinate of mate pairing, while males focus on fat and body mass when making a choice (Shellman-Reeve 1999). This investment made within a partnership choice stems from the intensity of the commitment as well as works as a way to determine and secure the quality of the mate choice. In a recent study, termites that had previously been rejected as a mate choice were more likely to end up with a partner that had also been rejected (Shellman- Reeve 1999). In this way, termites with smaller heads often paired with females of smaller body mass. These differences within the criteria used by each gender may be explained by the Knox, 3
4 indicators of each characteristic. Head width, for instance, displays whether a male can defend a nest and would be a good protector, and female body mass may be an indicator of future fertility (Shellman-Reeve 1999). Once making a mate choice, both partners are heavily involved with the construction and establishment of the colony, including involvement with care of the young, which suggests the extensive investment made by each committed partner (Shellman-Reeve 1999). It is interesting to note that mate choice occurs before consummation of the relationship which places more pressure upon the physical attributes of individuals as determinants for partners (Boomsma 2013). There are a myriad of advantages to monogamous behavior, not least of all, the harmonious cooperation of individuals which results from the lack of conflicts involving parental commitment (Boomsma 2013). Within a eusocial society, the mother conducts care of the young in cooperation with the labor performed by sterile castes. These established roles promote harmonious interactions because there is no uncertainty or conflict over behavioral roles (Boomsma 2013). In fact, these roles are often irreversible, and established early in life (Hartke and Baer 2011). Another advantage of monogamy includes the potential for a worker s ability to inherit the nest (Gadau and Fewell 2009). Because some workers still maintain their reproductive capabilities, they are able to take over the colony in the event of the death of the king and queen. For instance, in the species Reticulitermes virginicus, kings and queens may be succeeded by juveniles when sexually mature. When the king and queen decease, as opposed to being fatal for the colony, in this species, the neotenics of the colony replace the king and queen. The neotenics are developed as a result of not having access to a male. Some of the colonies of this species consist of parthenogenetic females, or those which have the ability to reproduce without Knox, 4
5 fertilization (Vargo et al. 2012).The death of the king and queen makes way for the development of neotenics, which therefore establishes the advantages of staying within the nest (Gadau and Fewell 2009). The nest, and the role of reproductive can be inherited. Additionally, advantages of group living within a monogamous colony include the safety provided by the nest itself. The risks of leaving and establishing a new colony are tremendous, and thus also influence the termite worker s desire to stay with the nest (Gadau and Fewell 2009). Though a few species of termites have the ability to develop replacement reproductives, the loss of the queen and king within a colony can be detrimental to the nest. A main reproductive pair is crucial to the long-term sustainability of the colony. A recent study shows that the survival rate of the colony is vastly superior while a colony maintains its king and queen reproductives opposed to more negative results that occur from worker developed reproductives. This biparental method of care and establishment of the colony, therefore, is vital to the health, efficiency, and stability of the nest (Hartke and Baer 2011). Mating Biology of Termites and Caste Determination In termites, the males are able to be long-lived, similar to the queens, and produce a continual supply of sperm, often for up to a couple decades at a time (Hartke and Baer 2011). In contrast to more promiscuous species, the sperm cannot be tested by the female before mating occurs. Rather, mate choice comes after the nuptial flights, and before copulation. In this way, the termite king as a sperm producer is a key concern to females (Boomsma 2013). Despite the continual presence of the king termite, the queen also stores sperm within the spermathecae. This behavior suggests that minimalization of sperm loss or waste is lessened by the storage structure (Boomsma 2013). The size of the colony is determined by the amount of Knox, 5
6 eggs the queen is able to fertilize and produce. In addition, neither the female nor the male have external genitalia, pointing to the idea that sexual selection pressures are not key factors in their evolutionary development (Hartke and Baer 2011). What is termed a royal pheromone is a substance which impedes the reproductive behavior of worker castes. Though the chemical compounds are unknown, the presence of the compounds corresponds with a reproductive s status and age, which therefore regulates the termite caste system (Hanus et al. 2010). It should not be said that termite workers are helpless dupes which suffer under the repressive reign of a king and queen. In reality, recent studies have shown that cuticular hydrocarbons act as important chemicals in insect communication and workers are decision-making individuals involved with reproductive potency coordination and modifications made within the caste system (Hanus et al. 2010). Another factor involved with caste regulation includes the differentiation of termite soldiers. The caste of a particular termite is not determined from an individual basis, but rather stems from the termite queen and king stimulation. When the primary reproductives were removed from the colony, no soldiers were developed, suggesting the necessity of the king and queen. Soldiers were seen to develop when given more proctodeal, or anus-to-mouth, feeding materials than other third instar larvae (Maekawa et al. 2012). An increased level of JH is also needed to promote the development of soldier termites, which is thought to be procured through additional proctodeal methods. For insect worker castes other than soldiers, it has been suggested that sterile castes within eusocial insect groups necessarily arose from the monogamous behavior of the parents, which then points to the idea of worker altruism and kin selection (Boomsma 2009). Knox, 6
7 Termite Worker Altruism and Kin Selection Eusocial colonies rely upon sibling behavior which consists of altruistically helping raise siblings rather than reproducing themselves (Hughes et al. 2008). These sterile individuals help take care of their siblings at the cost of personal reproduction of their own progeny (Boomsma 2009). This kin selection strategy of termites that care for their siblings rather than their own offspring, often at the possible cost of their own survival, comes from the amount of relatedness that each sibling shares with one another. This similarity continues through genes, and so the survival of offspring necessarily also coincides with the survival of similar genes, albeit within a sibling rather than offspring. Additionally, termite workers benefit from their altruistic behavior through protection and sustenance. The assistance of brood care of siblings can be viewed as a sort of surrogate, or replacement for the worker s own offspring (Boomsma 2009). Because of this relatedness of siblings, termite workers have been shown to recognize kin mates even after long periods of separation. When termites were paired with those from a different colony, significant mortality and aggression ensued; in contrast, termites paired with those they had been merely separated from but were of the same colony, featured mild levels of offensive behavior and no mortality (Olugbemi 2012). The ability to recognize mates is suggested to come from similarities of olfactory cues which may have been genetically encrypted, as well as similar receptors and greeting behavior (Olugbemi 2012). Rather than initiating a battle between related termites that had been separated for a long period, these termites were shown to begin building tunnels together. Relatedness of siblings therefore is a key element to the importance of kin recognition as well as harmonious cooperation among individuals and the promotion of altruism. Knox, 7
8 Though altruistic behavior was previously thought to have been a cause of the evolution of eusocial living, recent research shows that the behavior is actually an effect of eusociality (Hughes et al. 2008). For instance, group living is safer and has a higher survival rate than it would be for a solitary termite. Thus, though a termite colony may be composed entirely of kin, this relatedness is a result of eusociality and is beneficial toward individual workers. This can be contrasted to polyandry activity in which the offspring is increasingly less related if there are multiple breeding females within a colony (Hughes et al. 2008). A high level of relatedness among workers is an essential factor in concordant eusocial living. Evolution of Monogamy in Connection with Eusociality There are many characteristics that are involved with the evolution of eusociality including the overlap of generations, monogamy, and the benefits of group defense and protection. Individuals who help within the colony provide it with a higher probability of survival, which is beneficial to all. Eusociality may have also been derived from the ability of some workers to maintain their reproductive capabilities and thus promote the adaptation and development of a colony (Thorne 1997). A definitive conclusion about the evolution of eusociality and monogamy is impractical, but many hypotheses combine to contribute to a more complete understanding of these questions. For example, advanced eusocial insects are predominantly monogamous, which suggests that the evolution of the division of labor with sterile workers came from lifetime monogamy (Boomsma 2007). The two concepts appear indissolubly linked. Though there are multiple hypotheses for the derived nature of termite monogamy, one hypothesis states that in order to have a lifetime supply of sperm production, the continual presence of a male is needed, and another is the idea of mate guarding (Nalepa and Jones 1990). Knox, 8
9 A third hypothesis incorporates both hypotheses as well as emphasizes that ecological constraints contributed to this evolutionary phenomenon as well. The first hypothesis involving sperm production was derived from the understanding that queen termites are extremely fertile, and long lived. Therefore, without the presence of a male, the female would have to have a complex sperm storage area that would keep the sperm useful even after long periods of time which is not the case (Nalepa and Jones 1990). Additionally, within the second hypothesis, the limited availability of females at the same time as all the other females forces males to make a choice directly after the nuptial flights, and this mate guarding leads to the monogamous result. The male termite s success rate at reproduction is improved by its continual engagement with one female individual, especially since the energy and time spent finding a mate is so costly (Nalepa and Jones 1990). However, the ecological limitations of sparse food resources, the difficulties involved with being a lone termite without a colony, and the difficulty of finding a mate should also be taken into account when the evolution of monogamy is concerned (Nalepa and Jones 1990). In addition to monogamous behavior, haplodiploidy is also a factor often connected with the evolution of eusocial insects because it increases the probability for eusocial behavior to develop and become stable (Fromhage and Kokko 2011). Species that are haplodiploidy, are thought to be more conditioned to the system of eusociality because the individuals are more closely related than to their would-be offspring (Fromhage and Kokko 2011). However, a recent study furthers the importance placed upon this haplodiploid system. The study shows that the expected percentage of termite sisters that have a rare phenotype is larger in haplodiploid than in diplodipoid, suggesting that the stability of a eusocial colony relies upon haplodiploidy to guard Knox, 9
10 against rare individual alleles (Fromhage and Kokko 2011). Monogamy as well as haplodiploidy are both crucial factors for the evolution of eusociality. Polyandry within Eusocial Insect Groups As a contrast to the stability promoted within a colony by monogamy, promiscuous behavior adversely affects the benefits otherwise gained from parental commitment and kin altruism (Boomsma 2007). Because females mate multiple times, the relatedness of individuals to one another is decreased, and sexual competition over paternity is increased (Jaffe et al. 2012). However, some promiscuity in other eusocial insects seems to have some beneficial effects. Though the honey bee queen is promiscuous, this behavior seems to promote worker productivity as well as allows workers to shift their own fertility as the mating rate of the queen decreases. When the queen bee is only inseminated by one male, workers selfishly take resources from the colony to promote their own reproductive capabilities. In this way, the mating activity of honey bee queens influences the reproductive response of workers to alter the movements of individuals and results in reduced activity and performance (Mattila et al. 2012). Inversely to the monogamy found in most termites, as the honey bee colony workers become more closely related from a decrease in queen polyandry, reproductively active honey bee workers are often met with hostility by other workers. Promiscuous activity by the queen therefore establishes more peaceful cooperation among individuals (Mattila et al. 2012). Similarly, the males of the hymenopteran species Ropalidia marginata also engage in promiscuous behavior, but females do not (Shilpa et al. 2010). The male wasp of this species mates multiple times in order to secure that its sperm continues, however, the female wasp is reluctant to mate multiple times which a recent study notes contributes to increasing the harmony Knox, 10
11 of more related individuals within the colony (Shilpa et al. 2010). The idea of monogamy as enhancing cooperation among colony members coincides with the monogamy found within most monogamous termites; however, it is interesting that there is a discrepancy between the level of commitment of the female wasps versus the males (Shilpa et al. 2010). It is fascinating to note that the eusociality of bees, wasps, and termites evolved separately from one another, and yet are somewhat similar in terms of mating patterns (Boomsma 2013). Conclusion This review reveals that though there are exceptions to the evolutionary link typically made between monogamy and eusociality, the benefits of monogamous behavior within the superorganism are multitudinous. In termites as well as eusocial hymenoptera, mate-choice is made outside of the colony the event occurs after the reproductives have left their natal nest and search for their own partners, rather than being a part of societal living (Boomsma 2013). The monogamy of the primary reproductives in termites has been shown to create a more harmonious and stable living environment for their offspring, especially since kin selection is a key aspect of fostering this stability. Further research needs to be done in terms of the chemicals found within the royal pheromone to enhance understanding about caste determination of soldiers, as well as the evolutionary relationships among eusocial insect groups. Termites and the development of lifetime partnerships provide incredible insights into the benefits of monogamous behavior, especially as it relates to worker altruism and the cooperation among termite workers which decrease internal conflicts. Knox, 11
12 References Boomsma, J. J. (2013). Beyond Promiscuity: Mate-Choice Commitments in Social Breeding Phil. Trans. R. Soc B 368: Boomsma, J. J. (2009). Lifetime Monogamy and the Evolution of Eusociality Phil. Trans. R. Soc B 364: Boomsma, J. J. (2007). Kin Selection versus Sexual Selection: Why the Ends Do Not Meet Current Biology 17: Fromhage, L. and Kokko, H. (2011). Monogamy and Haplodiploidy Act in Synergy to Promote the Evolution of Eusociality Nature Communications 2: 1-5. Gadau, J. and Fewell, J. (2009). Organization of Insect Societies: from Genome to Sociocomplexity. Harvard University Press, Cambridge, 617 p. Hanus, R. (2009). Beyond Cuticular Hydrocarbons: Evidence of Proteinaceous Secretion Specific to Termite Kings and Queens Proc. R. Soc. B 277: Hartke, T. R. and Baer, B. (2011). The Mating Biology of Termites: a Comparative Review Animal Behavior 82: Hughes, W. O. H., Oldroyd, B. P., Beekman, M., and Ratnieks, F. L. W. (2008). Ancestral Monogamy Shows Kin Selection is Key to the Evolution of Eusociality Science 320: Jaffe, R., Garcia-Gonzalez, F., den Boer, S. P. A., Simmons, L. W., and Baer, B. (2012). Patterns of Paternity Skew Among Polyandrous Social Insects: What Can They Tell Us About the Potential for Sexual Selection? Evolution 66: Korb, J. and Foster, K. R. (2010). Ecological Competition Favours Cooperation in Termite Societies Ecology Letters 13: Maekawa, K., Nakamura, S., and Watanabe, D. (2012). Termite Soldier Differentiation in Incipient Colonies is Related to Parental Proctodeal Trophallactic Behavior Bio One 29: Mattila, H. R., Reeve, H. K., and Smith, M. L. (2012). Promiscuous Honey Bee Queens Increase Colony Productivity by Suppressing Worker Selfishness Current Biology 22: Nalepa, C. A., and Jones, S. C. (1990). Evolution of Monogamy in Termites Biological Reviews of the Cambridge Philosophical Society 66: Knox, 12
13 Olugbemi, B.O. (2012). Intra and Inter colonial Agonistic Behavior in the Termite, Microcerotermes fuscotibialis Sjostedt (Isoptera: Termitidae: Termitinae) Journal of Insect Behavior 26: Shellman-Reeve, J. S. (1999). Courting Strategies and Conflicts in a Monogamous, Biparental Termite Proceedings: Biological Sciences 266: Shilpa, M. C., Sen, R., Samudre, S., and Gadagkar, R. (2010). Males, but Not Females, Mate with Multiple Partners: a Laboratory Study of a Primitively Eusocial Wasp Ropalidia marginata Insectes Sociaux 59: Thompson, G. J., Lenz, M., Crozier, R. H., and Crespi, B. J. (2007). Molecular-genetic Analyses of Dispersal and Breeding Behaviour in the Australian Termite Coptotermes lacteus: Evidence for Non-random Mating in a Swarm-dispersal Mating System Australian Journal of Zoology 55: Thorne, B. L. (1997). Evolution of Eusociality in Termites Annual Review of Ecology and Systematics 28: Vargo, E. L., Labadie, P. E., and Matsuura, K. (2011). Asexual Queen Succession in the Subterranean Termite Reticulitermes virginicus Proc. R. Soc. B 279: Knox, 13
Eusocial species. Eusociality. Phylogeny showing only eusociality Eusocial insects. Eusociality: Cooperation to the extreme
Eusociality: Cooperation to the extreme Groups form colonies with reproductive and worker castes. Eusociality has evolved most often in insects: Ants Eusocial species Honeybees Termites Wasps Phylogeny
More information12. Social insects. Is it better to be social? Is it better to be social? What is social? Some costs of being social
Is it better to be social? 12. Social insects Cost and benefit viewpoint Social behavior is not always adaptive (costs exceed benefits) What are some costs of being social? What are some benefits of being
More informationChapter 14 The Evolution of Social Behavior (1 st lecture)
Chapter 14 The Evolution of Social Behavior (1 st lecture) Society A group of individuals of the same species that is organized in a cooperative manner, extending beyond sexual and parental care Colonial
More informationCooperation. Main points for today. How can altruism evolve? Group living vs. cooperation. Sociality-nocooperation. and cooperationno-sociality
Cooperation Why is it surprising and how does it evolve Cooperation Main points for today Sociality, cooperation, mutualism, altruism - definitions Kin selection Hamilton s rule, how to calculate r Group
More informationQuestions About Social Behavior
April 17: Altruism: Questions Questions About Social Behavior 1. Why live in groups? Costs: disease, competition, cannibalism, visibility to predators Benefits: more efficient foraging; defenses against
More informationWhat is altruism? Benefit another at a cost to yourself. Fitness is lost!
Altruism What is altruism? Benefit another at a cost to yourself. Fitness is lost! Does altruism exist? Best examples come from eusocial insects Bees, termites, ants Suicide in bees, etc. Non-breeding
More informationSocial Insects. Social Insects. Subsocial. Social Insects 4/9/15. Insect Ecology
Social Insects Social Insects Insect Ecology Sociality evolved multiple times in insects Much of Earth s fauna consists of social insects They play major roles in entire ecosystems Proliferation of ants
More informationSocial Insects. Insect Ecology
Social Insects Insect Ecology Social Insects Sociality evolved multiple times in insects Much of Earth s fauna consists of social insects They play major roles in entire ecosystems Proliferation of ants
More informationEvolution of Social Behavior: Kin Selection & Sociobiology. Goal: Why does altruism exist in nature?
"One lifetime, nine lifetimes are not long enough for the task of blocking every cranny through which calamity may enter A life spent, however victoriously, in securing the necessities of life is no more
More informationIntracolonial nepotism during colony fissioning in honey bees?
Intracolonial nepotism during colony fissioning in honey bees? Juliana Rangel Co-authors: Heather Mattila, Thomas Seeley Department of Neurobiology and Behavior Cornell University Apimondia Conference,
More informationThursday, September 26, 13
Helpful behaviors Alarm calls (e.g., Belding ground squirrel) Sentinel behavior (e.g., meerkats) Nest helping Eusocial behavior Actor performs some action that benefits another (the recipient). How do
More informationWhy such altruism? Why are these nymphs sacrificing themselves to protect other aphids?
12: Social Insects Some aphids in the subfamilies Pemphiginae and Hormaphidinae (Hemiptera: Aphididae) have a sacrificial soldier caste. Some first and secondinstar nymphs exhibit aggressive behavior and
More informationSOCIAL ANIMALS. -Spectacular -Photographed -Studied -Appreciated. The PINNACLE of evolution???
SOCIAL ANIMALS -Spectacular -Photographed -Studied -Appreciated The PINNACLE of evolution??? QUALITIES Social animals are aggregations of conspecifics that may have enhanced communication abilities some
More information9.916 Ingroups and Outgroups
9.916 Ingroups and Outgroups Today s Lecture One mind thinking about another: - stable - universal Social relationships: - dynamic - co-dependent - context-bound Social relationships depend on a continuous
More informationAnimal Behaviour. Mark Elgar. Eusociality II
Animal Behaviour Mark Elgar Eusociality II Outline Evolution of sociality in insects How much care to give Conflicts of interest re-visited Social doesn t always mean complex These social insects have
More informationBiology of the Colony. Dr. Deborah Delaney
Biology of the Colony Dr. Deborah Delaney Eusociality Photograph Alex Wild 2004 Photograph Alex Wild 2003 Eusocial: True Sociality Found mainly in two orders: Hymenoptera (some bees and wasps, all ants)
More informationChapter 44. Table of Contents. Section 1 Development of Behavior. Section 2 Types of Animal Behavior. Animal Behavior
Animal Behavior Table of Contents Section 1 Development of Behavior Section 2 Types of Animal Behavior Section 1 Development of Behavior Objectives Identify four questions asked by biologists who study
More information4 Questions relating to Behavior
Chapter 51: Animal Behavior 1. Stimulus & Response 2. Learned Behavior 3. Connecting Behavior to Survival & Reproduction 4 Questions relating to Behavior The Dutch behavioral scientist Niko Tinbergen proposed
More informationLocal resource competition. Sex allocation Is the differential allocation of investment in sons vs. daughters to increase RS. Local mate competition
Sex allocation Is the differential allocation of investment in sons vs. daughters to increase RS Local resource competition Biased against the competing sex Fisher: Genetic model predicts 1:1 sex ratio
More informationChapter 13 Opener: Weaver ants form superbly cooperative societies. Chapter 9. The Evolution of Social Behavior
Chapter 13 Opener: Weaver ants form superbly cooperative societies Chapter 9. The Evolution of Social Behavior Social living for gain benefits Social living is not always beneficial 13.1 The energy budget
More informationMammalogy Lecture 15 - Social Behavior II: Evolution
Mammalogy Lecture 15 - Social Behavior II: Evolution I. Evolution of Social Behavior In order to understand the evolution & maintenance of social behavior, we need to examine costs & benefits of group
More informationPATTERNS OF PATERNITY SKEW AMONG POLYANDROUS SOCIAL INSECTS: WHAT CAN THEY TELL US ABOUT THE POTENTIAL FOR SEXUAL SELECTION?
ORIGINAL ARTICLE doi:10.1111/j.1558-5646.2012.01721.x PATTERNS OF PATERNITY SKEW AMONG POLYANDROUS SOCIAL INSECTS: WHAT CAN THEY TELL US ABOUT THE POTENTIAL FOR SEXUAL SELECTION? Rodolfo Jaffé, 1,2,3 Francisco
More informationAuthor's personal copy
Kin Selection Andy Gardner, University of St Andrews, St Andrews, UK Ó 2015 Elsevier Ltd. All rights reserved. Abstract The theory of kin selection is the foundation for the study of social evolution.
More informationPolyphenic Insects. genotype X environment = phenotype POLYPHENISM. genetic polymorphism vs polyphenism. the peppered moth.
What makes for differences between individuals? Polyphenic Insects genes environment genotype X environment = phenotype POLYPHENISM poly many (more than one anyway) phen - form genetic polymorphism vs
More informationExploited superorganisms how life history shapes the reproductive strategies of honeybees
Essay Introduction to Advanced Biology M.Sc. Biodiversity, Ecology & Evolution von Jana Bundschuh Exploited superorganisms how life history shapes the reproductive strategies of honeybees Not all honeybees
More informationLevels of Selection: Multilevel Selection Theory
Levels of Selection: Multilevel Selection Theory Natural Selection higher levels lower levels Lineage Species Trait Group Kin Group Individual Cell Organelle Gene Natural Selection Basics Entities (individuals)
More informationAssessment Schedule 2016 Biology: Demonstrate understanding of the responses of plants and animals to their external environment (91603)
NCEA Level 3 Biology (91603) 2016 page 1 of 6 Assessment Schedule 2016 Biology: Demonstrate understanding of the responses of plants and animals to their external environment (91603) Evidence Statement
More informationSC741 W12: Division of Labor Part I: Fixed- and Variable- Threshold Algorithms
SC741 W12: Division of Labor Part I: Fixed- and Variable- Threshold Algorithms Outline Division of labor in natural systems Ants Bees, wasps Models and mechanisms Fixed-threshold mechanisms Variable-threshold
More informationStable eusociality via maternal manipulation when. resistance is costless
1 2 Stable eusociality via maternal manipulation when resistance is costless 3 Mauricio González-Forero 4 Monday 25 th May, 2015 5 6 7 8 Department of Ecology and Evolutionary Biology, University of Tennessee,
More informationBee Communica,on A Hive Mind
Bee Communica,on A Hive Mind Hive Mind Hive Mind One for all and all for one Kill self to protect the hive Fatal S,ng WHY? Kin Selec)on: So closely related The hive occupants: One queen a fer,le female
More informationThe basics of kin selection theory
The basics of kin selection theory Kin selection theory has its origins in attempt to unlock the puzzle of why some organisms have evolved to help other organisms of the same species. Such helping behavior
More informationThe Common Ground Curriculum. Science: Biology
The Common Ground Curriculum Science: Biology CGC Science : Biology Defining Biology: Biology is the study of living things in their environment. This is not a static, snapshot of the living world but
More informationLiving in groups 1. What are three costs and three benefits of living in groups?
Study questions Living in groups 1. What are three costs and three benefits of living in groups? 2. What is the dilution effect? What is a key assumption of the dilution effect hypothesis? What is a biological
More informationSociobiological Approaches. What We Will Cover in This Section. Overview. Overview Evolutionary Theory Sociobiology Summary
Sociobiological Approaches What We Will Cover in This Section Overview Evolutionary Theory Sociobiology Summary 7/22/2007 Sociobiological Approaches.ppt 2 Overview Total focus on internal aspects of personality
More informationSocial interaction. Kin and Group selection. Social interaction. Social interaction. Social interaction. Social interaction
Kin and Group selection Social interaction Social interactions between organisms present the opportunity for conflict and cooperation Interaction between individuals can have 4 possible outcomes on the
More informationAppendix A from G. Wild and S. A. West, Genomic Imprinting and Sex Allocation
009 by The University of Chicago. All rights reserved.doi: 0.086/593305 Appendix A from G. Wild and S. A. West, Genomic Imprinting and Sex Allocation (Am. Nat., vol. 73, no., p. E) Kin selection analysis
More informationThe Evolution of Eusociality in Bees
Nissa Osheim 4/4/2007 The Evolution of Eusociality in Bees Evolution has been called the survival of the fittest. It is a competition between individuals to see which genes will get to replicate. However,
More informationMating frequency and genetic relatedness of workers in the hornet Vespa analis (Hymenoptera: Vespidae)
Entomological Science (003) 6, 119 13 ORIGINAL ARTICLE Mating frequency and genetic relatedness of workers in the hornet Vespa analis (Hymenoptera: Vespidae) Jun-ichi TAKAHASHI, 1 Shin ichi AKIMOTO, 1
More informationLesson 1 Sexual Reproduction and Meiosis
Lesson 1 Name Date Class A. What is sexual reproduction? 1. produces an offspring when genetic materials from two different sex cells combine. a. The female sex cell, a(n), forms in an ovary. b. The male
More informationImprinting and kin recognition
Imprinting and kin recognition Imprinting Konrad Lorenz Filial imprinting Critical period Sensitive period Experimental approaches Hours after hatching precocial altricial Filial imprinting Multiple cues
More information5/7/2009. Copyright The McGraw Hill Companies, Inc. Permission required for reproduction or display.
36 1 CHAPTER 36 Animal Behavior Ethology 36 2 Science of Animal Behavior Study of animal behavior as a science had its roots in the 1872 work of Charles Darwin Ethology Science of animal behavior in its
More information1. Referring to the cladogram on page 1 and with regard to mono- / polyphyly, vertebrates are a monophyletic group; invertebrates are parayphyletc.
Answers III.4. Animals-I. 1. Referring to the cladogram on page 1 and with regard to mono- / polyphyly, vertebrates are a monophyletic group; invertebrates are parayphyletc. 2. Referring to the cladogram
More informationReproduction in primitively eusocial wasps
SOCIAL INSECTS Advanced eusocial: - morphologically sterile helpers Lecture Reproductive queueing in primitively eusocial species: predictions and tests PRIMITIVELY EUSOCIAL Polistes paper wasp hover wasp
More informationBrief history of The Prisoner s Dilemma (From Harman s The Price of Altruism)
Brief history of The Prisoner s Dilemma (From Harman s The Price of Altruism) 1948 The RAND Corporation. a civilian nonprofit think tank of an elite cadre of physicists, mathematicians, economists, and
More informationBee Behavior. Summary of an article by. Stephen Taber III from Beekeeping in the United States
Bee Behavior Summary of an article by Stephen Taber III from Beekeeping in the United States Bees Sense Organs: Vision Each compound eye is spherical in shape and comprised of some 6,300 cone-shaped facets
More informationSubmitted to Biology Letters. Patterns of split sex ratio in ants have multiple evolutionary causes based on different within-colony conflicts
Patterns of split sex ratio in ants have multiple evolutionary causes based on different within-colony conflicts Journal: Biology Letters Manuscript ID: draft Article Type: Research Date Submitted by the
More informationBinary fission occurs in prokaryotes. parent cell. DNA duplicates. cell begins to divide. daughter cells
Chapter 11 Chapter 11 Some eukaryotes reproduce through mitosis. Binary fission is similar in function to mitosis. Asexual reproduction is the creation of offspring from a single parent. Binary fission
More information28 3 Insects Slide 1 of 44
1 of 44 Class Insecta contains more species than any other group of animals. 2 of 44 What Is an Insect? What Is an Insect? Insects have a body divided into three parts head, thorax, and abdomen. Three
More informationName: Date Block A Trail of Termites
LAB Name: Date Block A Trail of Termites BACKGROUND INFORMATION: Termites play an important role in the environment. They feed primarily on wood and are important in recycling nutrients in ecosystems.
More informationAvailable online at ScienceDirect. Procedia Computer Science 20 (2013 ) 90 95
Available online at www.sciencedirect.com ScienceDirect Procedia Computer Science 20 (2013 ) 90 95 Complex Adaptive Systems, Publication 3 Cihan H. Dagli, Editor in Chief Conference Organized by Missouri
More informationThe evolution of queen control over worker reproduction in the social Hymenoptera
The evolution of queen control over worker reproduction in the social Hymenoptera The Harvard community has made this article openly available. Please share how this access benefits you. Your story matters
More informationSYMPOSIUM Student Journal of Science & Math. Volume 2 Issue 1
SYMPOSIUM Student Journal of Science & Math Volume 2 Issue 1 biology 117 B82.731 OBSERVATIONAL LEARNING IN EUSOCIAL INSECTS Background A RESEARCH PROPOSAL by Avity Norman Ants (order Hymenoptera, family
More informationPractical 5 SOCIAL BEHAVIOUR OF HONEY BEES
Practical 5 SOCIAL BEHAVIOUR OF HONEY BEES Aim: To understand communication system of a honey bee colony. Introduction Among different insect orders, only 8 have been recognized by insect taxonomists which
More informationMCDB 1111 corebio 2017 Midterm I
MCDB 1111 corebio 2017 Midterm I Name: There are 18 questions, each worth a maximum of 6 points. Multiple choice questions are worth 3 points, while explanations are worth 3 points. If you need to, use
More informationSexual Reproduction. Page by: OpenStax
Sexual Reproduction Page by: OpenStax Summary Sexual reproduction was an early evolutionary innovation after the appearance of eukaryotic cells. The fact that most eukaryotes reproduce sexually is evidence
More information2. The development of revolutionized the of life.
Science 10 Unit 7 Worksheet Chapter 15, Part 1. 1. Briefly describe the three main parts of cell theory: 2. The development of revolutionized the of life. 3. Individual cells need to take in to build and
More informationSex Ratio Conflict and Worker Production in Eusocial Hymenoptera
vol. 158, no. 2 the american naturalist august 2001 Sex Ratio Conflict and Worker Production in Eusocial Hymenoptera Max Reuter * and Laurent Keller Institute of Ecology, University of Lausanne, 1015 Lausanne,
More informationAN EVOLUTIONARY RESOLUTION OF MANIPULATION CONFLICT
ORIGINAL ARTICLE doi:10.1111/evo.12420 AN EVOLUTIONARY RESOLUTION OF MANIPULATION CONFLICT Mauricio González-Forero 1,2,3 1 Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville,
More informationSNC2D BIOLOGY 3/17/2013. TISSUES, ORGANS & SYSTEMS OF L The Importance of Cell Division (P.26-27) The Importance of Cell Division
SNC2D BIOLOGY TISSUES, ORGANS & SYSTEMS OF L The Importance of Cell Division (P.26-27) The Importance of Cell Division You began life as a single fertilized cell. Now your body is made up of trillions
More informationScience Unit Learning Summary
Learning Summary Inheritance, variation and evolution Content Sexual and asexual reproduction. Meiosis leads to non-identical cells being formed while mitosis leads to identical cells being formed. In
More informationEVOLUTION ALGEBRA. Freshman Seminar University of California, Irvine. Bernard Russo. University of California, Irvine. Winter 2015
EVOLUTION ALGEBRA Freshman Seminar University of California, Irvine Bernard Russo University of California, Irvine Winter 2015 Bernard Russo (UCI) EVOLUTION ALGEBRA 1 / 15 Understanding Genetics The study
More informationAssociation between caste and genotype in the termite Mastotermes darwiniensis Froggatt (Isoptera: Mastotermitidae)
Australian Journal of Entomology (2003) 42, 1 5 Association between caste and genotype in the termite Mastotermes darwiniensis Froggatt (Isoptera: Mastotermitidae) Michael A D Goodisman and Ross H Crozier*
More information9/6/2012. Point #1. Natural selection is purposeless and not acting for the good of anything.
Sample statements in exams and term papers... territoriality evolved for the good of the species. warning coloration helps to perpetuate the species. without echolocation, the bat species would die out.
More informationTESTS OF REPRODUCTIVE-SKEW MODELS
Annu. Rev. Entomol. 2001. 46:347 85 Copyright c 2001 by Annual Reviews. All rights reserved TESTS OF REPRODUCTIVE-SKEW MODELS IN SOCIAL INSECTS H. Kern Reeve Section of Neurobiology and Behavior, Cornell
More informationConceptually, we define species as evolutionary units :
Bio 1M: Speciation 1 How are species defined? S24.1 (2ndEd S26.1) Conceptually, we define species as evolutionary units : Individuals within a species are evolving together Individuals of different species
More informationStudying Life. Lesson Overview. Lesson Overview. 1.3 Studying Life
Lesson Overview 1.3 Characteristics of Living Things What characteristics do all living things share? Living things are made up of basic units called cells, are based on a universal genetic code, obtain
More informationSpeciation and Patterns of Evolution
Speciation and Patterns of Evolution What is a species? Biologically, a species is defined as members of a population that can interbreed under natural conditions Different species are considered reproductively
More informationChapter 13 Meiosis and Sexual Reproduction
Biology 110 Sec. 11 J. Greg Doheny Chapter 13 Meiosis and Sexual Reproduction Quiz Questions: 1. What word do you use to describe a chromosome or gene allele that we inherit from our Mother? From our Father?
More informationRegents Review Assignment #8-A08 Living Environment: Comet Part A Questions
Part A Questions 1. A student notices that fruit flies with the curlywing trait develop straight wings if kept at a temperature of 16 C, but develop curly wings if kept at 25 C. The best explanation for
More informationGenetic & Evolutionary Roots of Behavior Gleitman et al. (2011), Chapter 2
Genetic & Evolutionary Roots of Behavior Gleitman et al. (2011), Chapter 2 Mike D Zmura Department of Cognitive Sciences, UCI Psych 9A / Psy Beh 11A January 16, 2014 T. M. D'Zmura 1 Chapter Topics Genetics
More informationHONORS PSYCHOLOGY REVIEW QUESTIONS
HONORS PSYCHOLOGY REVIEW QUESTIONS The purpose of these review questions is to help you assess your grasp of the facts and definitions covered in your textbook. Knowing facts and definitions is necessary
More informationUse evidence of characteristics of life to differentiate between living and nonliving things.
Grade Big Idea Essential Questions Concepts Competencies Vocabulary 2002 Standards All living things have a common set characteristic needs and functions that separate them from nonliving things such as:
More informationOhio Tutorials are designed specifically for the Ohio Learning Standards to prepare students for the Ohio State Tests and end-ofcourse
Tutorial Outline Ohio Tutorials are designed specifically for the Ohio Learning Standards to prepare students for the Ohio State Tests and end-ofcourse exams. Biology Tutorials offer targeted instruction,
More informationThe Frequency of Multi-queen Colonies Increases with Altitude in a Nearctic Ant
The Frequency of Multi-queen Colonies Increases with Altitude in a Nearctic Ant By: Jürgen Heinze, Olav Rueppell This is the accepted version of the following article: Heinze J., Rueppell O. (2014) The
More informationMicroevolutionary changes show us how populations change over time. When do we know that distinctly new species have evolved?
Microevolutionary changes show us how populations change over time. When do we know that distinctly new species have evolved? Critical to determining the limits of a species is understanding if two populations
More informationName Class Date. KEY CONCEPT Gametes have half the number of chromosomes that body cells have.
Section 1: Chromosomes and Meiosis KEY CONCEPT Gametes have half the number of chromosomes that body cells have. VOCABULARY somatic cell autosome fertilization gamete sex chromosome diploid homologous
More informationSTUDY GUIDE SECTION 16-1 Genetic Equilibrium
STUDY GUIDE SECTION 16-1 Genetic Equilibrium Name Period Date Multiple Choice-Write the correct letter in the blank. 1. The smallest unit in which evolution occurs is a. an individual organism. c. a species
More informationTitle: WS CH 18.1 (see p ) Unit: Heredity (7.4.1) 18.1 Reading Outline p Sexual Reproduction and Meiosis
Title: WS CH 18.1 (see p.612-625) Unit: Heredity (7.4.1) 18.1 Reading Outline p. 612-625 NPD A. What is sexual reproduction? (p615) 1. _ produces an offspring when genetic materials from two different
More information4/25/12. Mutualism. Mutualism. Dominance Hierarchy. Mutualism. Selfish Behavior Spiteful Behavior
4/25/12 Dominance Hierarchy Shared gain of direct fitness Example: Prey capture by lion pride Despo0c (a.k.a. steep) Linear (a.k.a. shallow or egalitarian) Alpha Beta Gamma Helping Behavior Delayed gain
More informationPeter Dutton. 28 August Halesworth& District
Peter Dutton 28 August 2014 Halesworth& District Bees Hymenoptera - 150,000+ species Sawflies, Wasps, Bees, Ants, Bees - 20,000+ species Most solitary, a few social Honey Bee - The only one which produces
More informationPheromones by Ellen Miller November 2015
Pheromones by Ellen Miller November 2015 Greek word pherein to carry Greek word hormone to stimulate A released chemical that triggers a social response in members of the same species The chemical is transmitted
More informationKin recognition in social insects and other animals-a review of recent findings and a consideration of their relevance for the theory of kin selection
Proc. Indian Acad. Sci. (Anim. Sci.), Vol. 94, No.6, December 1985, pp. 587-621. @ Printed in India. Kin recognition in social insects and other animals-a review of recent findings and a consideration
More information(Write your name on every page. One point will be deducted for every page without your name!)
POPULATION GENETICS AND MICROEVOLUTIONARY THEORY FINAL EXAMINATION (Write your name on every page. One point will be deducted for every page without your name!) 1. Briefly define (5 points each): a) Average
More informationUNIT V. Chapter 11 Evolution of Populations. Pre-AP Biology
UNIT V Chapter 11 Evolution of Populations UNIT 4: EVOLUTION Chapter 11: The Evolution of Populations I. Genetic Variation Within Populations (11.1) A. Genetic variation in a population increases the chance
More informationSEXUAL REPRODUCTION & MEIOSIS
SEXUAL REPRODUCTION & MEIOSIS Living organisms are distinguished by their ability to reproduce their own kind. Offspring resemble their parents more than they do less closely related individuals of the
More informationBiology. Chapter 12. Meiosis and Sexual Reproduction. Concepts and Applications 9e Starr Evers Starr. Cengage Learning 2015
Biology Concepts and Applications 9e Starr Evers Starr Chapter 12 Meiosis and Sexual Reproduction 12.1 Why Sex? In asexual reproduction, a single individual gives rise to offspring that are identical to
More information2. Overproduction: More species are produced than can possibly survive
Name: Date: What to study? Class notes Graphic organizers with group notes Review sheets What to expect on the TEST? Multiple choice Short answers Graph Reading comprehension STRATEGIES Circle key words
More informationChapter 53 Animal Behavior
Chapter 53 Animal Behavior meerkats What is behavior? Why study it? Behavior everything an animal does & how it does it response to stimuli in its environment innate = inherited or developmentally fixed
More informationWhat do plants compete for? What do animals compete for? What is a gamete and what do they carry? What is a gene?
How are these animals adapted to their surroundings: - a) Polar bear b) Camel c) Cactus What do plants compete for? What do animals compete for? What is a gamete and what do they carry? What is a gene?
More informationMicrogynous queens in ants: social parasites or dispersal morphs?
Microgynous queens in ants: social parasites or dispersal morphs? Lenoir Alain 1, Séverine Devers 1, Philippe Marchand 1, Christine Errard 1, Christophe Bressac 1 and Riitta Savolainen 2 1 University of
More informationAmy Ant. Formica Mica Grant. Dr. Sheila Grant (Mica s mom)
Dr. Sheila Grant (Mica s mom) Formica Mica Grant Amy Ant By Elisabeth Schlegel, Jessica Louton, Natasha Mehdiabadi, and Ted Schultz Illustrated by Katherine Arisumi At the Smithsonian Institution s National
More informationarxiv:physics/ v1 [physics.bio-ph] 8 Feb 2000 Formalizing the gene centered view of evolution
arxiv:physics/0002016v1 [physics.bio-ph] 8 Feb 2000 Formalizing the gene centered view of evolution Y. Bar-Yam New England Complex Systems Institute 24 Mt. Auburn St., Cambridge MA Department of Molecular
More informationBee Colony Activities Throughout The Year
Bee Colony Activities Throughout The Year Written by Khalil Hamdan Apeldoorn The Netherlands A honeybee gathering nectar from a flower. Photo source: forestwander.com Bee collecting pollen. Photo source:
More informationBiology Unit Overview and Pacing Guide
This document provides teachers with an overview of each unit in the Biology curriculum. The Curriculum Engine provides additional information including knowledge and performance learning targets, key
More informationThe evolution of queen control over worker reproduction in the social Hymenoptera
The evolution of queen control over worker reproduction in the social Hymenoptera Jason Olejarz a, Carl Veller a,b, and Martin A. Nowak a,b,c arxiv:70.04873v [q-bio.pe] 6 Feb 07 a Program for Evolutionary
More informationCh 5. Evolution, Biodiversity, and Population Ecology. Part 1: Foundations of Environmental Science
Ch 5 Evolution, Biodiversity, and Population Ecology Part 1: Foundations of Environmental Science PowerPoint Slides prepared by Jay Withgott and Heidi Marcum Copyright 2006 Pearson Education, Inc., publishing
More informationAnimal Behavior (Ch. 51)
Animal Behavior (Ch. 51) Behavioral Ecology Two types of questions: Proximate questions: Focus on environmental stimuli that trigger behavior and physiology behind response How? Ultimate questions: Focus
More informationThe Science of Biology. Chapter 1
The Science of Biology Chapter 1 Properties of Life Living organisms: are composed of cells are complex and ordered respond to their environment can grow and reproduce obtain and use energy maintain internal
More informationEastern Subterranean Termite
Eastern Subterranean Termite Introduction Subterranean termites are the single greatest economic pest in the United States. These termites cause billions of dollars in damage each year to homes, historical
More informationBiology of Reproduction Spring 2007
Biology of Reproduction Spring 2007 Louis Guillette Office: 528 Bartram Hall Office Hours: Tuesday/Thursday period 4 (10:40-11:25) Phone: 392-1098; Email: ljg@zoo.ufl.edu http://www.zoo.ufl.edu/ljg/courses/index.htm
More information