vol. 191, no. 1 the american naturalist january 2018
E-Article
Thermoregulatory Behavior Simultaneously Promotes
and Forestalls Evolution in a Tropical Lizard
Martha M. Muñoz1,* and Jonathan B. Losos2
1. Department of Biological Sciences, Virginia Tech, Blacksburg, Virginia 24060; 2. Department of Organismic and Evolutionary Biology,
Harvard University, Cambridge, Massachusetts 02138
Submitted January 11, 2017; Accepted July 21, 2017; Electronically published October 25, 2017
Online enhancements: appendix.
abstract: The role of behavior in evolution has long been discussed,
with some arguing that behavior promotes evolution by exposing
organisms to selection (behavioral drive) and others proposing that
it inhibits evolution by shielding organisms from environmental var-
iation (behavioral inertia). However, this discussion has generally fo-
cused on the effects of behavior along a single axis without considering
that behavior simultaneously influences selection in various niche
dimensions. By examining evolutionary change along two distinct
niche axes—structural and thermal—we propose that behavior si-
multaneously drives and impedes evolution in a group of Anolis
lizards from the Caribbean island of Hispaniola. Specifically, a behav-
ioral shift in microhabitat to boulders at high altitude enables thermo-
regulation, thus forestalling physiological evolution in spite of colder
environments. This same behavioral shift drives skull and limb evolu-
tion to boulder use. Our results emphasize the multidimensional ef-
fects of behavior in evolution. These findings reveal how, rather than
being diametrically opposed, niche conservatism and niche lability
can occur simultaneously. Furthermore, patterns of niche evolution
may vary at different geographic scales: because of thermoregulatory
behavior, lizards at high and low elevation share similar microclimatic
niches (consistent with niche conservatism) while inhabiting distinct
macroclimatic environments (consistent with niche divergence). To-
gether, our results suggest that behavior can connect patterns of niche
divergence and conservatism at different geographic scales and among
traits.
Keywords: niche evolution, behavior, thermoregulation, Caribbean,
lizard, Bogert effect.
Introduction
For nearly a century, behavior has been recognized as a
key pacemaker for evolution (discussed in Huey et al. 2003;
Duckworth 2009). On the one hand, behavior can facilitate
evolutionary change; as organisms explore novel environ-
ments, they experience new selective pressures that can lead
* Corresponding author; e-mail: [email protected]
Am. Nat. 2018. Vol. 191, pp. E000–E000. q 2017 by The University of Chicago.
0003-0147/2018/19101-57488$15.00. All rights reserved.
DOI: 10.1086/694779
This content downloaded from 130.0
All use subject to University of Chicago Press Term
to phenotypic adaptations and speciation (Miller 1956;
Lande 1981; Wyles et al. 1983; Bateson 1988; West-Eberhard
1989; Sol et al. 20 ...
Article 4 Apes in a changing world - the effects of global warmin.docxfredharris32
Article 4: Apes in a changing world - the effects of global warming on the behaviour and distribution of African apes J. Lehmann et al. Global warming and ape biogeography.
Sourse: Lehmann, Julia, Amanda H. Korstjens, and Robin I. M. Dunbar. "Apes In A Changing World - The Effects Of Global Warming On The Behaviour And Distribution Of African Apes J. Lehmann Et Al. Global Warming And Ape Biogeography." Journal Of Biogeography 37.12 (2010): 2217-2231. Academic Search Premier. Web. 7 Feb. 2015.
O R I G I N A L
A R T I C L E
Apes in a changing world – the effects
of global warming on the behaviour
and distribution of African apes
Julia Lehmann1,2*, Amanda H. Korstjens1,3 and Robin I. M. Dunbar1,4
1British Academy Centenary Research Project,
School of Biological Sciences, Crown Street,
University of Liverpool, Liverpool L69 7ZB,
UK,
2
Department of Life Sciences, Roehampton
University, London SW15 4JD, UK,
3
Conservation Sciences, Bournemouth
University, Poole BH12 5BB, UK, 4Institute of
Cognitive and Evolutionary Anthropology,
University of Oxford, Oxford OX2 6PE, UK
*Correspondence: Julia Lehmann, Life Science
Department, Holybourne Avenue, Roehampton
University, London SW15 4JD, UK.
E-mail: [email protected]
A B S T R A C T
Aim In this study we use a modelling approach to identify: (1) the factors
responsible for the differences in ape biogeography, (2) the effects that global
warming might have on distribution patterns of African apes, (3) the underlying
mechanisms for these effects, and (4) the implications that behavioural flexibility
might be expected to have for ape survival. All African apes are highly
endangered, and the need for efficient conservation methods is a top priority. The
expected changes in world climate are likely to further exacerbate the difficulties
they face. Our study aims to further understand the mechanisms that link climatic
conditions to the behaviour and biogeography of ape species.
Location Africa.
Method We use an existing validated time budgets model, derived from data on
20 natural populations of gorillas (Gorilla beringei and Gorilla gorilla) and
chimpanzees (Pan troglodytes and Pan paniscus), which specifies the relationship
between climate, group size, body weight and time available for various activities,
to predict ape distribution across Africa under a uniform worst-case climate
change scenario.
Results We demonstrate that a worst-case global warming scenario is likely to
alter the delicate balance between different time budget components. Our model
points to the importance of annual temperature variation, which was found to
have the strongest impact on ape biogeography. Our simulation indicates that
rising temperatures and changes in rainfall patterns are likely to have strong
effects on ape survival and distribution, particularly for gorillas. Even if they
behaved with maximum flexibility, gorillas may not be able to survive in most of
their present habitat ...
The document provides an introduction to zoology, discussing several key topics:
- Theories of evolution from scientists like Lamarck and Cuvier are summarized, with Darwin's theory of natural selection identified as the most accurate.
- The complex life cycle of the monarch butterfly is used as an example to illustrate different life cycle stages like egg, larva, pupa, and adult.
- Adaptation strategies animals use to survive harsh environments like the desert and polar regions are outlined.
- The process of mitosis and key differences in replication between prokaryotes and eukaryotes are summarized.
- Homeostatic mechanisms that allow animals to maintain stable body conditions are briefly
This study examines how the presence of an endophytic fungus affects decomposition rates and fungal diversity in the grass Festuca thurberi along elevation gradients. In the first experiment, litter bags containing either endophyte-infected or endophyte-free F. thurberi litter were placed along three mountain transects to measure decomposition rates over time. The second experiment investigates how the endophyte affects host plant survival, growth, and biomass. Preliminary results show endophyte presence reduces plant biomass and survival. Fungal cultures were also taken from leaf tissues to analyze fungal community composition differences. The goal is to better understand how endophyte symbioses and climate interact to impact ecosystem processes
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Thinking of getting a Samsung flagship? We just spotted a sale for the Samsung Galaxy S23 Ultra on Shopee, with a huge RM 1,500 discount. For a 2023 flagship, this is one of the biggest, it not the biggest,
Shifts in phenology due to global climate change: The need for a yardstickSimoneBoccuccia
This document reviews examples of shifts in phenology (timing of seasonal activities) of species due to climate change. It argues that to properly interpret observed shifts in a species' phenology, a "yardstick" is needed - a measure of how much the species' environment and conditions have shifted due to climate change. The paper examines a few examples where both a species' phenology and aspects of its environment/food sources have been measured over long periods. In most examples, the species has shifted its phenology either too little or too much compared to shifts in its ecological conditions, suggesting many species are becoming mistimed due to climate change. The authors urge more research linking phenological data on species to potential yardsticks to better
This document discusses the problem of pattern and scale in ecology over the past 20 years. Some key points:
- Major advances have helped connect ecological patterns to theory and vice versa, and recognize the importance of scale in predictions.
- Technological revolutions in computing, molecular biology, sensing, and information sharing have driven radical changes in ecology.
- Research is increasingly focusing on coupling ecological and evolutionary timescales, the influence of evolution on ecosystems, integrating organism biology and ecology through networks, and understanding spatial patterns across scales.
- Studies on rotifers, algae, Darwin's finches and guppies show evolution can impact populations and ecosystems on ecological timescales. This challenges the traditional view that
This document summarizes an experiment on the evolutionary adaptation of E. coli to repeated freeze-thaw cycles. 15 populations of E. coli were subjected to 150 cycles of freezing at -80°C without cryoprotectants, followed by thawing and growth at 37°C. Competition experiments showed that the populations adapted to have increased survival during freezing and thawing as well as faster recovery after thawing, with fitness gains of around 60-90% relative to their ancestors. Future work aims to identify the genetic mutations responsible for this adaptive evolution to the freeze-thaw environment.
This document describes an experiment where 15 populations of E. coli were subjected to 150 cycles of freeze-thaw-growth (FTG) to study evolutionary adaptation. Three populations were founded from the ancestor of a long-term evolution experiment, while 12 were founded from clones that previously evolved at 37C for 20,000 generations. Competition experiments showed both groups adapted to better survive freezing and thawing, and recover growth more rapidly afterwards, with 60-90% increased fitness relative to their progenitors. Future work aims to identify genetic mutations responsible.
Article 4 Apes in a changing world - the effects of global warmin.docxfredharris32
Article 4: Apes in a changing world - the effects of global warming on the behaviour and distribution of African apes J. Lehmann et al. Global warming and ape biogeography.
Sourse: Lehmann, Julia, Amanda H. Korstjens, and Robin I. M. Dunbar. "Apes In A Changing World - The Effects Of Global Warming On The Behaviour And Distribution Of African Apes J. Lehmann Et Al. Global Warming And Ape Biogeography." Journal Of Biogeography 37.12 (2010): 2217-2231. Academic Search Premier. Web. 7 Feb. 2015.
O R I G I N A L
A R T I C L E
Apes in a changing world – the effects
of global warming on the behaviour
and distribution of African apes
Julia Lehmann1,2*, Amanda H. Korstjens1,3 and Robin I. M. Dunbar1,4
1British Academy Centenary Research Project,
School of Biological Sciences, Crown Street,
University of Liverpool, Liverpool L69 7ZB,
UK,
2
Department of Life Sciences, Roehampton
University, London SW15 4JD, UK,
3
Conservation Sciences, Bournemouth
University, Poole BH12 5BB, UK, 4Institute of
Cognitive and Evolutionary Anthropology,
University of Oxford, Oxford OX2 6PE, UK
*Correspondence: Julia Lehmann, Life Science
Department, Holybourne Avenue, Roehampton
University, London SW15 4JD, UK.
E-mail: [email protected]
A B S T R A C T
Aim In this study we use a modelling approach to identify: (1) the factors
responsible for the differences in ape biogeography, (2) the effects that global
warming might have on distribution patterns of African apes, (3) the underlying
mechanisms for these effects, and (4) the implications that behavioural flexibility
might be expected to have for ape survival. All African apes are highly
endangered, and the need for efficient conservation methods is a top priority. The
expected changes in world climate are likely to further exacerbate the difficulties
they face. Our study aims to further understand the mechanisms that link climatic
conditions to the behaviour and biogeography of ape species.
Location Africa.
Method We use an existing validated time budgets model, derived from data on
20 natural populations of gorillas (Gorilla beringei and Gorilla gorilla) and
chimpanzees (Pan troglodytes and Pan paniscus), which specifies the relationship
between climate, group size, body weight and time available for various activities,
to predict ape distribution across Africa under a uniform worst-case climate
change scenario.
Results We demonstrate that a worst-case global warming scenario is likely to
alter the delicate balance between different time budget components. Our model
points to the importance of annual temperature variation, which was found to
have the strongest impact on ape biogeography. Our simulation indicates that
rising temperatures and changes in rainfall patterns are likely to have strong
effects on ape survival and distribution, particularly for gorillas. Even if they
behaved with maximum flexibility, gorillas may not be able to survive in most of
their present habitat ...
The document provides an introduction to zoology, discussing several key topics:
- Theories of evolution from scientists like Lamarck and Cuvier are summarized, with Darwin's theory of natural selection identified as the most accurate.
- The complex life cycle of the monarch butterfly is used as an example to illustrate different life cycle stages like egg, larva, pupa, and adult.
- Adaptation strategies animals use to survive harsh environments like the desert and polar regions are outlined.
- The process of mitosis and key differences in replication between prokaryotes and eukaryotes are summarized.
- Homeostatic mechanisms that allow animals to maintain stable body conditions are briefly
This study examines how the presence of an endophytic fungus affects decomposition rates and fungal diversity in the grass Festuca thurberi along elevation gradients. In the first experiment, litter bags containing either endophyte-infected or endophyte-free F. thurberi litter were placed along three mountain transects to measure decomposition rates over time. The second experiment investigates how the endophyte affects host plant survival, growth, and biomass. Preliminary results show endophyte presence reduces plant biomass and survival. Fungal cultures were also taken from leaf tissues to analyze fungal community composition differences. The goal is to better understand how endophyte symbioses and climate interact to impact ecosystem processes
Deal: Samsung Galaxy S23 Ultra can be yours with up to RM1,500 discount today
BY Sharil Abdul Rahman
25 September 2023
12:05 pm
Comment
Thinking of getting a Samsung flagship? We just spotted a sale for the Samsung Galaxy S23 Ultra on Shopee, with a huge RM 1,500 discount. For a 2023 flagship, this is one of the biggest, it not the biggest,
https://youtube.com/shorts/3Vqu8Ui7Ros?si=8Crxj4OYqWBjWC5J
https://www.youtube.com/channel/UCyTZZP6faGeWWmWHHYzB4iQ
https://youtube.com/shorts/QKbAu7JcL_E?si=bi24Q9yYoDVih5tD
https://youtube.com/shorts/3Vqu8Ui7Ros?si=8Crxj4OYqWBjWC5J
https://rekonise.com/apk-o3cam
Deal: Samsung Galaxy S23 Ultra can be yours with up to RM1,500 discount today
BY Sharil Abdul Rahman
25 September 2023
12:05 pm
Comment
Thinking of getting a Samsung flagship? We just spotted a sale for the Samsung Galaxy S23 Ultra on Shopee, with a huge RM 1,500 discount. For a 2023 flagship, this is one of the biggest, it not the biggest,
Shifts in phenology due to global climate change: The need for a yardstickSimoneBoccuccia
This document reviews examples of shifts in phenology (timing of seasonal activities) of species due to climate change. It argues that to properly interpret observed shifts in a species' phenology, a "yardstick" is needed - a measure of how much the species' environment and conditions have shifted due to climate change. The paper examines a few examples where both a species' phenology and aspects of its environment/food sources have been measured over long periods. In most examples, the species has shifted its phenology either too little or too much compared to shifts in its ecological conditions, suggesting many species are becoming mistimed due to climate change. The authors urge more research linking phenological data on species to potential yardsticks to better
This document discusses the problem of pattern and scale in ecology over the past 20 years. Some key points:
- Major advances have helped connect ecological patterns to theory and vice versa, and recognize the importance of scale in predictions.
- Technological revolutions in computing, molecular biology, sensing, and information sharing have driven radical changes in ecology.
- Research is increasingly focusing on coupling ecological and evolutionary timescales, the influence of evolution on ecosystems, integrating organism biology and ecology through networks, and understanding spatial patterns across scales.
- Studies on rotifers, algae, Darwin's finches and guppies show evolution can impact populations and ecosystems on ecological timescales. This challenges the traditional view that
This document summarizes an experiment on the evolutionary adaptation of E. coli to repeated freeze-thaw cycles. 15 populations of E. coli were subjected to 150 cycles of freezing at -80°C without cryoprotectants, followed by thawing and growth at 37°C. Competition experiments showed that the populations adapted to have increased survival during freezing and thawing as well as faster recovery after thawing, with fitness gains of around 60-90% relative to their ancestors. Future work aims to identify the genetic mutations responsible for this adaptive evolution to the freeze-thaw environment.
This document describes an experiment where 15 populations of E. coli were subjected to 150 cycles of freeze-thaw-growth (FTG) to study evolutionary adaptation. Three populations were founded from the ancestor of a long-term evolution experiment, while 12 were founded from clones that previously evolved at 37C for 20,000 generations. Competition experiments showed both groups adapted to better survive freezing and thawing, and recover growth more rapidly afterwards, with 60-90% increased fitness relative to their progenitors. Future work aims to identify genetic mutations responsible.
1) The success of assisted colonization and gene flow conservation strategies depends on how plant species respond to changes in temperature and photoperiod across their life cycles.
2) A study exposed seeds from northern and southern populations of an annual plant, Chamaecrista fasciculata, to ambient and elevated temperatures in a potential future colonization site north of its current range to examine responses.
3) They found warming advanced development and compressed life cycles, with patterns of selection on traits changing. Performance depended on population of origin, with the northern population faring best under current conditions but the southern population potentially adapting best to future warming. However, photoperiod mismatches may limit long-term persistence, especially between
Macroevolution examines evolution over long time periods of thousands to millions of years. It studies changes above the species level, including the emergence of new species through speciation and loss of species through extinction. Key areas of study in macroevolution include building phylogenetic trees to demonstrate evolutionary relationships between extinct and modern species based on morphological comparisons, and using molecular analysis to help validate these relationships when possible. Mass extinctions that eliminate large percentages of species are also investigated to better understand causes and patterns of extinction and subsequent radiations as new species evolve to fill open niches.
1. The study examined the relative influence of local environmental conditions and regional spatial processes on the structure of aquatic plant communities in 98 lakes and ponds across Connecticut.
2. Using statistical analyses like partial canonical correspondence analysis and partial Mantel tests, the study found that aquatic plant community structure reflects both local environmental conditions like pH, water clarity, and depth, as well as regional spatial processes like dispersal.
3. While environmental conditions and spatial processes explained 27% of the variation in the plant community data, local environmental conditions accounted for 45% of the explained variation, and regional spatial processes like dispersal accounted for 40%, suggesting both play a role in structuring aquatic plant communities.
This study examines morphological changes in threespine stickleback over eight generations following their transplantation from a large lake habitat with diverse predators to a smaller pond habitat lacking fish predators. Measurements of 20 defence and trophic traits were taken on stickleback collected from the source lake and transplant pond population between 1994 and 2009. Significant reductions were observed in several plate and spine traits, gill raker number and length, and increases in jaw length and eye diameter within eight generations - representing about one-third of the differences typically seen between natural lake and pond populations. These changes occurred rapidly and in the directions predicted based on the different predator regimes and food sources between the habitats. Both natural selection and phenotypic plasticity likely contributed to the
This document summarizes a study investigating the relationship between genetic heterozygosity, fluctuating asymmetry (FA), length, and weight in two samples of Atlantic salmon with different timings of first active feeding. The study found that:
1) Fish in the early feeding sample (EA) exhibited smaller FA values, were longer, heavier, and more heterozygous at allozyme loci than fish in the late feeding sample (LA).
2) Within both samples, heterozygosity at allozyme loci was inversely related to FA and positively related to weight and length. However, no significant differences were found in microsatellite diversity between the EA and LA samples.
3) No association was observed between microsatellite variability and FA
Behavioral traits can change more quickly than morphology in response to environmental cues, allowing behaviors to lead evolutionary changes. A behavioral shift can occur faster than an adaptive change in physiology, taking the lead in evolution. While behaviors are flexible, they can also inhibit evolutionary changes by preventing natural selection from acting on other traits when environments change. A unified framework is needed to understand how behaviors can both drive and inhibit evolutionary processes through altering selection pressures.
Living organisms (even human) evolve to match with the climate or not and geo...MdAbdulAhad26
Living organisms (even human) evolve to match with the climate or not and geographical distribution (biogeography) opposite to Darwin’s theory or not
This document discusses whether living organisms evolve to match their climate and environment, as Darwin theorized, or if their geographical distributions oppose Darwin's theory of evolution. It provides several arguments against Darwin's view: 1) Plants and animals are beautifully adapted to changing environments but do not evolve in response. 2) When climate changes, organisms migrate to safer areas and later return, rather than evolving new adaptations. 3) If migration is impossible, species will go extinct rather than evolve new traits. 4) The presence of cosmopolitan species in different climates opposes climate influencing evolution.
Evolution occurs as traits are inherited from ancestors over thousands of years, resulting in changes to physical and behavioral traits that help organisms survive and reproduce in different environments. All living things share common ancestry according to Darwin's theory of evolution by natural selection, whereby organisms better suited to their environment tend to produce more offspring, passing on favorable traits that accumulate over generations and lead to the formation of new species. Key evidence for this includes the fossil record showing changes over time, similarities in DNA, and comparative anatomy revealing ancestral relationships between structures in different organisms.
This document summarizes key concepts in macroevolution, including how macroevolutionary changes occur through both gradualism and punctuated equilibrium. It discusses evidence from fossils and comparisons between living organisms. It also covers plate tectonics, morphological divergence and convergence, developmental patterns, comparative biochemistry, molecular clocks, and taxonomy.
O R I G I N A L A RT I C L Edoi10.1111evo.13631Two d.docxamit657720
This study examines evolutionary changes over two decades in the annual plant Brassica rapa in response to fluctuations in precipitation and drought in California. Seeds were collected from two populations at four time points spanning 18 generations. In resurrection experiments, plants from ancestral and descendant generations were grown under control and drought conditions to test for evolutionary changes in drought response traits. The study found evidence that B. rapa evolved earlier flowering and reduced stem diameter and water use efficiency in response to drought, but that occasional wet periods reversed these adaptations, potentially making the species less adapted to increasing drought severity. Evolutionary responses sometimes differed between the two populations.
O R I G I N A L A RT I C L Edoi10.1111evo.13631Two d.docxvannagoforth
O R I G I N A L A RT I C L E
doi:10.1111/evo.13631
Two decades of evolutionary changes in
Brassica rapa in response to fluctuations in
precipitation and severe drought
Elena Hamann,1,2 Arthur E. Weis,3 and Steven J. Franks1
1Department of Biological Sciences, Fordham University, Bronx, New York 10458
2E-mail: [email protected]
3Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, ON M5S 3B2, Canada
Received March 27, 2018
Accepted October 5, 2018
As climate changes at unprecedented rates, understanding population responses is a major challenge. Resurrection studies can
provide crucial insights into the contemporary evolution of species to climate change. We used a seed collection of two Californian
populations of the annual plant Brassica rapa made over two decades of dramatic precipitation fluctuations, including increasingly
severe droughts. We compared flowering phenology, other drought response traits, and seed production among four generations,
grown under drought and control conditions, to test for evolutionary change and to characterize the strength and direction of
selection. Postdrought generations flowered earlier, with a reduced stem diameter, and lower water-use efficiency (WUE), while
intervening wet seasons reversed these adaptations. There was selection for earlier flowering, which was adaptive, but delayed
flowering after wet years resulted in reduced total seed mass, indicating a maladaptive response caused by brief wet periods.
Furthermore, evolutionary changes and plastic responses often differed in magnitude between populations and drought periods,
suggesting independent adaptive pathways. While B. rapa rapidly evolved a drought escape strategy, plant fitness was reduced
in contemporary generations, suggesting that rapid shifts in flowering time may no longer keep up with the increasing severity
of drought periods, especially when drought adaptation is slowed by occasional wet seasons.
K E Y W O R D S : Drought escape, global change, phenotypic plasticity, phenology, rapid evolution, resurrection study.
There is now abundant evidence that climate change and altered
precipitation patterns (IPCC 2014) trigger large-scale species
losses, shifts in vegetation communities, and evolutionary plant
responses (Parmesan and Yohe 2003; Jump and Penuelas 2005;
Parmesan 2006; Franks et al. 2014). Particularly well documented
are worldwide shifts in flowering time following advanced spring-
time (Menzel et al. 2006; Miller-Rushing and Primack 2008;
Cleland et al. 2012). While much of the shift in this trait may be
due to the direct effects of temperature on developmental rate,
some could be due to an evolutionary response to selection im-
posed by a warmer environment (Nicotra et al. 2010; Hoffmann
and Sgro 2011; Merila and Hendry 2014; Gugger et al. 2015;
Stoks et al. 2016). Phenotypic plasticity, in which organisms re-
spond to changes in environmental conditions, is itself ge ...
Some of our most basic questions about the history of life concern w.pdfaloeplusint
Some of our most basic questions about the history of life concern when major events occurred.
Some evolutionary events can be dated from the fossil record But what options are available
when fossil data are missing? In at least some instances, it should be possible to address
questions about timing by analyzing molecular traits that change at a steady rate. This
hypothesis, called the molecular clock, @ originated with Emile Zuckerkandl and Linus Pauling
(1962). There are reasons to expect that some types of DNA sequences change in a clocklike
fashion. Many mutations change an individual's DNA but not its phenotype In most cases,
mutations like these are not exposed to natural selection. Instead, these neutral @ changes evolve
by a random process called genetic drift As we will discuss elsewhere, the neutral theory of
molecular evolution predicts that neutral changes in DNA should accumulate in populations at a
rate equal to the mutation rate If the mutation rate stays reasonably constant, and if generation
times remain similar, then the number of neutral molecular differences between two taxa should
be proportional to the age of their most recent common ancestor By counting distinct neutral
mutations observed in two species and multiplying by a calibration rate representing how
frequently neutral changes occur per million years, researchers can estimate when the species
diverged. Although the possibility of dating events from estimates of genetic divergence is
tantalizing, there are several important caveats. For example, it is critical to realize that the
mutation rate to neutral alleles will vary from gene to gene and lineage to lineage, and even from
base to base. For reasons explained in another chapter, silent site changes in the third positions of
codons are more likely to be neutral with respect to fitness, and thus to accumulate at a clocklike
rate, than replacement changes that occur at the first and second positions in codons. And if
allele frequencies change rapidly due to strong selection at a particular gene, it is unlikely that
the mutations involved are accumulating in a clocklike fashion. Finally rates of change calibrated
for a particular gene and lineage are unlikely to work for other groups, which may have different
generation times and selection histories (Martin et al. 1992; Martin and Palumbi 1993; Hillis et
al. 1996). Even if clocklike change occurs in a particular gene and lineage, how can the rate be
determined? Investigators have to rely on the fossil or geological records. The idea is to measure
the genetic distance between two taxa whose divergence date is known from fossil or geological
data and then to use this calibration to date the divergence times of groups that have no fossil
record. As an example of how researchers use molecular clocks to date events, consider work by
Ralf Kittler and colleagues (2003, 2004) on the origin of human body lice Body lice (Pediculus
humanus corporis) are similar to head lice (.
History and Usefulness of Evolution.pptxMaryam Riasat
Evolution refers to the process of change over generations in organisms through descent with modification. Darwin proposed the theory of evolution by natural selection, where organisms with variations better suited to the environment are more likely to survive and pass on their traits. Evidence for evolution includes fossils showing changes over time, homologous and vestigial structures, biogeography, and molecular biology. Neo-Darwinism integrated genetics and population genetics into an evolutionary synthesis. Evolution explains the diversity of life and relationships between organisms.
Biology - Chp 17 - History Of Life - PowerPointMr. Walajtys
This chapter discusses the history of life on Earth based on evidence from the fossil record. It covers how fossils are formed and dated, as well as how paleontologists interpret and classify fossils to understand past life forms and environments. The fossil record shows that life has changed over time, with more than 99% of all species becoming extinct. It also reveals patterns of evolution such as extinction events, adaptive radiation, convergent and coevolution. The chapter discusses theories of punctuated equilibrium and the role of developmental genes in transformations of body plans over long periods of evolutionary time.
What scientific mechanism for evolution did Charles Darwin and Alfred.pdfmeerobertsonheyde608
What scientific mechanism for evolution did Charles Darwin and Alfred Wallace propose in
1858? Describe how this concept works. What is the difference between microevolution and
macroevolution?
Solution
1. Charles Darwin during 1858 had proposed the Theory of Evolution by Natural Selection,
where he has said that organisms change over time as a result of changes in heritable physical or
behavioral traits. Changes that allow an organism to better adapt to its environment will help it
survive and have more offspring.
Alfred Wallace who is also called the Father of Biogeography, studied the geographical
distribution of animal species and said the evolution is caused due to environmental pressures on
varieties of species it forces different species to adapt themselves to the local conditions, leading
to different populations in different locations.
Both groups of scientists have said that natural selection is the single most and important factor
in evolutionary changes seen in species.
Scientifically, This concept works due to different phenomenon such as:
a)Mutation- It is the Sudden change in the genetic makeup of an organism.And it is the driving
force of evolution, which influences the population’s gene pool.The change in the genetic
makeup of organism which favors the life of species makes it the fittest to survive!
b)Genetic Drift-It can occur when a small group of individuals leaves a population and
establishes a new one in a geographically isolated region.It occurs in a large population and helps
the population to survive.
c)Natural selection-This happens when populations of organisms are subjected to the
environment. The fittest creatures are more likely to survive and pass their genes to their
offspring, producing a population that is better adapted to the environment.
2.MicroEvolution-Microevolution is defined as changes in gene frequency in a population from
one generation to the next.These small changes in species occur by recombining existing genetic
material within the group of same species.
Micro Evolution is caused due to Mutation, Migration or Gene Flow, Genetic Drift and Selection
(natural and artificial).
Macro Evolution: It refers to major evolutionary changes over time, the origin of new types of
organisms from previously existing, but different, ancestral types.It is an evolution on a larger
scale where we find the descent of many species from one common ancestor over billions of
years.It is very difficult to see the Macro Evolution happening because it takes a longer time to
happen. But Instead, we can reconstruct the history of life using all available evidence like
geology, fossils, and living organisms..
This study examined how local and regional processes structure dragonfly distributions across a habitat gradient with varying pond permanence and top predator types. Through experiments, the study found that habitat specialist dragonflies, which are restricted to permanent lakes, were more vulnerable to invertebrate predators and dispersed and colonized artificial ponds less frequently than generalist species. Additionally, habitat specialists traveled shorter distances than generalists. These results suggest that dispersal limitation plays a key role in restricting the distributions of habitat specialist species, and that predation may reinforce this pattern by excluding specialists from certain habitats. By examining both local predation and regional dispersal, the study provides evidence that species distributions can be shaped by processes operating at multiple spatial scales.
Why is it important to study reactions norms to understand phenotypi.pdfarrowmobile
Why is it important to study reactions norms to understand phenotypic plasticity?
Solution
Ans:
Phenotypic plasticity, the capacity of a single genotype to exhibit variable phenotypes in
different environments, is common in insects and is often highly adaptive. Phenotypic plasticity
is important because it expands the existing “genocentric” evolutionary theory, producing an
encompassing paradigm to explain life on earth. Plasticity was once considered “noise” but is
now widely recognized as potentially adaptive under a wide array of circumstances. As with any
major shift in scientific thinking, phenotypic plasticity engenders new ideas, causing us to ask
new questions and test hypotheses that would not otherwise be examined, leading us to
productive new scientific insights.
Phenotypic plasticity is counterbalance to mutation driven evolution: It is not surprising that
during the first half of the 20th Century, scientists, flushed with excitement about Mendelian
genetics, viewed evolution primarily as a mutational process. However, this bias largely ignored
an important reality of evolution – that natural selection selects not among genotypes, but among
phenotypes. Thus, the phenotype, and variation among phenotypes, plays a major role in
evolution. And, because the environment in which an individual develops determines its
phenotype, the environment also assumes a greater role in evolution, and may, in fact, produce
more viable phenotypic variation than do mutations. This is because mutations are not only rare,
but usually deleterious. In contrast, a single environmental factor may alter the phenotypes of an
entire population, providing natural selection with access to perhaps thousands of
environmentally altered individuals, as opposed to a single mutant individual. In addition,
mutations generally arise randomly with no correlation to specific environments, whereas new
environmentally induced phenotypes are both directional and highly correlated with the specific
new environment, allowing new environments to immediately produce and select among new
phenotypes.
Including phenotypic plasticity produces a better model: As suggested above, the inclusion of
phenotypic plasticity can result in a better model than mutation-allelic substitution alone in
explaining the production of organismal diversity. For example, the initial evolution of warning
color (aposematism), starting as a rare mutation is problematic because conspicuous prey should
be quickly found and removed by predators (Lindström et al. 2001). In contrast, evolution of
aposematism is easily explained by phenotypic plasticity (Sword 2002). Likewise, for
development, phenotypic plasticity explains the evolution of allometry and exaggerated
morphologies (Emlen and Nijhout 2000, Shingleton et al. 2007). For physiology, phenotypic
plasticity explains adaptive, beneficial plasticities such as acclimation and response to exercise
(Swallow et al. 2005), quite well. In ecology, it aids our un.
This document provides an introduction to ecology, including definitions of key terms and concepts. It explains that ecology is the study of the relationships between organisms and their environments. It describes the different levels of ecological organization from the biosphere down to the individual organism. It also defines biomes as large geographical areas with similar climates and ecosystems, and lists some major biomes such as tundra, taiga, and tropical rainforest. Finally, it discusses adaptations as traits that enhance an organism's survival and are maintained by natural selection.
[Ostrom, 2009] a general framework for analyzing sustainability of social-e...FiorellaIsabelCampos1
This document presents a framework for analyzing the sustainability of social-ecological systems (SESs). The framework identifies four core subsystems that interact within an SES: resource systems, resource units, governance systems, and users. It then identifies 10 variables within these subsystems that are frequently found to affect the likelihood of users self-organizing to manage resources and achieve sustainability. These variables influence the perceived costs and benefits of investing in governance systems, such as the size of the resource system, its productivity, clarity of system boundaries, and collective-choice arrangements. The framework is intended to facilitate multidisciplinary analysis of complex SESs by providing a common structure for organizing knowledge about factors influencing sustainability.
FOOD WEBS 31
INTRODUCTION
“The classic marine food chain –algae, zooplank-
ton, fish– can now be considered as a variable phe-
nomenon in a sea of microbes (Karl 1999).”
The food web is one of the earliest and most fun-
damental concepts in ecology. Darwin (1845) recog-
nized the existence of a pelagic food chain. Elton is
credited with first appreciating the importance of food
chain and food web concepts (Lawton 1989), but
major antecedents include Petersen’s (1918) quantita-
tive conceptual model of the food web that is support-
ed by eel grass, and Hardy’s (1924) conceptual model
of the herring food web. Elton, and later Hutchinson
and his students, developed both population and mate-
rials-flux approaches to food webs (Hagen 1992), but
the two approaches quickly diverged. Paine (1980)
showed that population interactions do not equate with
energy flux, and asserted that energy flux was unim-
portant and “has generated few insights into ecologi-
cal processes.” This put studies of population biology
and energy-flux on separate paths that only recently
have shown signs of beginning to merge into a unified
paradigm (e. g. McQueen, et al., 1986; Hunter and
Price 1992; Polis and Winemiller 1996). Ecologists
recognize that energy input matters, although they
agree less on when and how it matters. What clearly
matters is formulating tractable hypotheses about
ecosystem structure and function. Repeated attempts
to simplify the inherent complexity of food webs have
led to Sisyphus-like progress in which investigators
have made generalizations and then have been forced
to qualify them.
SCI. MAR., 65 (Suppl. 2): 31-40 SCIENTIA MARINA 2001
A MARINE SCIENCE ODYSSEY INTO THE 21st CENTURY. J.M. GILI, J.L. PRETUS and T.T. PACKARD (eds.)
Caught in the food web: complexity made simple?*
LAWRENCE R. POMEROY
Institute of Ecology, University of Georgia, Athens GA 30602-2202, USA. E-mail: [email protected]
SUMMARY: Several historically separate lines of food-web research are merging into a unified approach. Connections
between microbial and metazoan food webs are significant. Interactions of control by predators, defenses against predation,
and availability of organic and inorganic nutrition, not any one of these, shape food webs. The same principles of popula-
tion ecology apply to metazoans and microorganisms, but microorganisms dominate the flux of energy in both marine and
terrestrial systems. Microbial biomass often is a major fraction of total biomass, and very small organisms have a very large
ratio of production and respiration to biomass. Assimilation efficiency of bacteria in natural systems is often not as high as
in experimental systems, so more primary production is lost to microbial respiration than had been thought. Simulation has
been a highly useful adjunct to experiments in both population theory and in studies of biogeochemical mass balance, but it
does not fully encompass the complexity of real systems. A major challenge for the futu.
Milestones Navigating Late Childhood to AdolescenceFrom the m.docxjessiehampson
Milestones: Navigating Late Childhood to Adolescence
From the movie, Lila, Eight to Thirteen in this week's materials, identify 2–3 developmental milestones Lila reaches, and assess whether or not you think she successfully navigates her way through them as she prepares for adolescence. Support your assertions with evidence from your text and this week's materials.
.
Migration and RefugeesMany immigrants in the region flee persecu.docxjessiehampson
Migration and Refugees
Many immigrants in the region flee persecution and then return after they are liberated. For example, 700,000 Jews were allowed to leave the former Soviet Union and enter Israel in the 1990s. There has also been a migration of Palestinian people. Discuss the following:
Why do you think that Israel is such an important place for the Jews?
What is the importance of the area to the Palestinians?
What do you think the impact would be on you and your families if you participated in such long-distance migration?
No references needed, need response within 3 hours!
.
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O R I G I N A L A RT I C L E
doi:10.1111/evo.13631
Two decades of evolutionary changes in
Brassica rapa in response to fluctuations in
precipitation and severe drought
Elena Hamann,1,2 Arthur E. Weis,3 and Steven J. Franks1
1Department of Biological Sciences, Fordham University, Bronx, New York 10458
2E-mail: [email protected]
3Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, ON M5S 3B2, Canada
Received March 27, 2018
Accepted October 5, 2018
As climate changes at unprecedented rates, understanding population responses is a major challenge. Resurrection studies can
provide crucial insights into the contemporary evolution of species to climate change. We used a seed collection of two Californian
populations of the annual plant Brassica rapa made over two decades of dramatic precipitation fluctuations, including increasingly
severe droughts. We compared flowering phenology, other drought response traits, and seed production among four generations,
grown under drought and control conditions, to test for evolutionary change and to characterize the strength and direction of
selection. Postdrought generations flowered earlier, with a reduced stem diameter, and lower water-use efficiency (WUE), while
intervening wet seasons reversed these adaptations. There was selection for earlier flowering, which was adaptive, but delayed
flowering after wet years resulted in reduced total seed mass, indicating a maladaptive response caused by brief wet periods.
Furthermore, evolutionary changes and plastic responses often differed in magnitude between populations and drought periods,
suggesting independent adaptive pathways. While B. rapa rapidly evolved a drought escape strategy, plant fitness was reduced
in contemporary generations, suggesting that rapid shifts in flowering time may no longer keep up with the increasing severity
of drought periods, especially when drought adaptation is slowed by occasional wet seasons.
K E Y W O R D S : Drought escape, global change, phenotypic plasticity, phenology, rapid evolution, resurrection study.
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responses (Parmesan and Yohe 2003; Jump and Penuelas 2005;
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Cleland et al. 2012). While much of the shift in this trait may be
due to the direct effects of temperature on developmental rate,
some could be due to an evolutionary response to selection im-
posed by a warmer environment (Nicotra et al. 2010; Hoffmann
and Sgro 2011; Merila and Hendry 2014; Gugger et al. 2015;
Stoks et al. 2016). Phenotypic plasticity, in which organisms re-
spond to changes in environmental conditions, is itself ge ...
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Some evolutionary events can be dated from the fossil record But what options are available
when fossil data are missing? In at least some instances, it should be possible to address
questions about timing by analyzing molecular traits that change at a steady rate. This
hypothesis, called the molecular clock, @ originated with Emile Zuckerkandl and Linus Pauling
(1962). There are reasons to expect that some types of DNA sequences change in a clocklike
fashion. Many mutations change an individual's DNA but not its phenotype In most cases,
mutations like these are not exposed to natural selection. Instead, these neutral @ changes evolve
by a random process called genetic drift As we will discuss elsewhere, the neutral theory of
molecular evolution predicts that neutral changes in DNA should accumulate in populations at a
rate equal to the mutation rate If the mutation rate stays reasonably constant, and if generation
times remain similar, then the number of neutral molecular differences between two taxa should
be proportional to the age of their most recent common ancestor By counting distinct neutral
mutations observed in two species and multiplying by a calibration rate representing how
frequently neutral changes occur per million years, researchers can estimate when the species
diverged. Although the possibility of dating events from estimates of genetic divergence is
tantalizing, there are several important caveats. For example, it is critical to realize that the
mutation rate to neutral alleles will vary from gene to gene and lineage to lineage, and even from
base to base. For reasons explained in another chapter, silent site changes in the third positions of
codons are more likely to be neutral with respect to fitness, and thus to accumulate at a clocklike
rate, than replacement changes that occur at the first and second positions in codons. And if
allele frequencies change rapidly due to strong selection at a particular gene, it is unlikely that
the mutations involved are accumulating in a clocklike fashion. Finally rates of change calibrated
for a particular gene and lineage are unlikely to work for other groups, which may have different
generation times and selection histories (Martin et al. 1992; Martin and Palumbi 1993; Hillis et
al. 1996). Even if clocklike change occurs in a particular gene and lineage, how can the rate be
determined? Investigators have to rely on the fossil or geological records. The idea is to measure
the genetic distance between two taxa whose divergence date is known from fossil or geological
data and then to use this calibration to date the divergence times of groups that have no fossil
record. As an example of how researchers use molecular clocks to date events, consider work by
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Evolution refers to the process of change over generations in organisms through descent with modification. Darwin proposed the theory of evolution by natural selection, where organisms with variations better suited to the environment are more likely to survive and pass on their traits. Evidence for evolution includes fossils showing changes over time, homologous and vestigial structures, biogeography, and molecular biology. Neo-Darwinism integrated genetics and population genetics into an evolutionary synthesis. Evolution explains the diversity of life and relationships between organisms.
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What scientific mechanism for evolution did Charles Darwin and Alfred.pdfmeerobertsonheyde608
What scientific mechanism for evolution did Charles Darwin and Alfred Wallace propose in
1858? Describe how this concept works. What is the difference between microevolution and
macroevolution?
Solution
1. Charles Darwin during 1858 had proposed the Theory of Evolution by Natural Selection,
where he has said that organisms change over time as a result of changes in heritable physical or
behavioral traits. Changes that allow an organism to better adapt to its environment will help it
survive and have more offspring.
Alfred Wallace who is also called the Father of Biogeography, studied the geographical
distribution of animal species and said the evolution is caused due to environmental pressures on
varieties of species it forces different species to adapt themselves to the local conditions, leading
to different populations in different locations.
Both groups of scientists have said that natural selection is the single most and important factor
in evolutionary changes seen in species.
Scientifically, This concept works due to different phenomenon such as:
a)Mutation- It is the Sudden change in the genetic makeup of an organism.And it is the driving
force of evolution, which influences the population’s gene pool.The change in the genetic
makeup of organism which favors the life of species makes it the fittest to survive!
b)Genetic Drift-It can occur when a small group of individuals leaves a population and
establishes a new one in a geographically isolated region.It occurs in a large population and helps
the population to survive.
c)Natural selection-This happens when populations of organisms are subjected to the
environment. The fittest creatures are more likely to survive and pass their genes to their
offspring, producing a population that is better adapted to the environment.
2.MicroEvolution-Microevolution is defined as changes in gene frequency in a population from
one generation to the next.These small changes in species occur by recombining existing genetic
material within the group of same species.
Micro Evolution is caused due to Mutation, Migration or Gene Flow, Genetic Drift and Selection
(natural and artificial).
Macro Evolution: It refers to major evolutionary changes over time, the origin of new types of
organisms from previously existing, but different, ancestral types.It is an evolution on a larger
scale where we find the descent of many species from one common ancestor over billions of
years.It is very difficult to see the Macro Evolution happening because it takes a longer time to
happen. But Instead, we can reconstruct the history of life using all available evidence like
geology, fossils, and living organisms..
This study examined how local and regional processes structure dragonfly distributions across a habitat gradient with varying pond permanence and top predator types. Through experiments, the study found that habitat specialist dragonflies, which are restricted to permanent lakes, were more vulnerable to invertebrate predators and dispersed and colonized artificial ponds less frequently than generalist species. Additionally, habitat specialists traveled shorter distances than generalists. These results suggest that dispersal limitation plays a key role in restricting the distributions of habitat specialist species, and that predation may reinforce this pattern by excluding specialists from certain habitats. By examining both local predation and regional dispersal, the study provides evidence that species distributions can be shaped by processes operating at multiple spatial scales.
Why is it important to study reactions norms to understand phenotypi.pdfarrowmobile
Why is it important to study reactions norms to understand phenotypic plasticity?
Solution
Ans:
Phenotypic plasticity, the capacity of a single genotype to exhibit variable phenotypes in
different environments, is common in insects and is often highly adaptive. Phenotypic plasticity
is important because it expands the existing “genocentric” evolutionary theory, producing an
encompassing paradigm to explain life on earth. Plasticity was once considered “noise” but is
now widely recognized as potentially adaptive under a wide array of circumstances. As with any
major shift in scientific thinking, phenotypic plasticity engenders new ideas, causing us to ask
new questions and test hypotheses that would not otherwise be examined, leading us to
productive new scientific insights.
Phenotypic plasticity is counterbalance to mutation driven evolution: It is not surprising that
during the first half of the 20th Century, scientists, flushed with excitement about Mendelian
genetics, viewed evolution primarily as a mutational process. However, this bias largely ignored
an important reality of evolution – that natural selection selects not among genotypes, but among
phenotypes. Thus, the phenotype, and variation among phenotypes, plays a major role in
evolution. And, because the environment in which an individual develops determines its
phenotype, the environment also assumes a greater role in evolution, and may, in fact, produce
more viable phenotypic variation than do mutations. This is because mutations are not only rare,
but usually deleterious. In contrast, a single environmental factor may alter the phenotypes of an
entire population, providing natural selection with access to perhaps thousands of
environmentally altered individuals, as opposed to a single mutant individual. In addition,
mutations generally arise randomly with no correlation to specific environments, whereas new
environmentally induced phenotypes are both directional and highly correlated with the specific
new environment, allowing new environments to immediately produce and select among new
phenotypes.
Including phenotypic plasticity produces a better model: As suggested above, the inclusion of
phenotypic plasticity can result in a better model than mutation-allelic substitution alone in
explaining the production of organismal diversity. For example, the initial evolution of warning
color (aposematism), starting as a rare mutation is problematic because conspicuous prey should
be quickly found and removed by predators (Lindström et al. 2001). In contrast, evolution of
aposematism is easily explained by phenotypic plasticity (Sword 2002). Likewise, for
development, phenotypic plasticity explains the evolution of allometry and exaggerated
morphologies (Emlen and Nijhout 2000, Shingleton et al. 2007). For physiology, phenotypic
plasticity explains adaptive, beneficial plasticities such as acclimation and response to exercise
(Swallow et al. 2005), quite well. In ecology, it aids our un.
This document provides an introduction to ecology, including definitions of key terms and concepts. It explains that ecology is the study of the relationships between organisms and their environments. It describes the different levels of ecological organization from the biosphere down to the individual organism. It also defines biomes as large geographical areas with similar climates and ecosystems, and lists some major biomes such as tundra, taiga, and tropical rainforest. Finally, it discusses adaptations as traits that enhance an organism's survival and are maintained by natural selection.
[Ostrom, 2009] a general framework for analyzing sustainability of social-e...FiorellaIsabelCampos1
This document presents a framework for analyzing the sustainability of social-ecological systems (SESs). The framework identifies four core subsystems that interact within an SES: resource systems, resource units, governance systems, and users. It then identifies 10 variables within these subsystems that are frequently found to affect the likelihood of users self-organizing to manage resources and achieve sustainability. These variables influence the perceived costs and benefits of investing in governance systems, such as the size of the resource system, its productivity, clarity of system boundaries, and collective-choice arrangements. The framework is intended to facilitate multidisciplinary analysis of complex SESs by providing a common structure for organizing knowledge about factors influencing sustainability.
FOOD WEBS 31
INTRODUCTION
“The classic marine food chain –algae, zooplank-
ton, fish– can now be considered as a variable phe-
nomenon in a sea of microbes (Karl 1999).”
The food web is one of the earliest and most fun-
damental concepts in ecology. Darwin (1845) recog-
nized the existence of a pelagic food chain. Elton is
credited with first appreciating the importance of food
chain and food web concepts (Lawton 1989), but
major antecedents include Petersen’s (1918) quantita-
tive conceptual model of the food web that is support-
ed by eel grass, and Hardy’s (1924) conceptual model
of the herring food web. Elton, and later Hutchinson
and his students, developed both population and mate-
rials-flux approaches to food webs (Hagen 1992), but
the two approaches quickly diverged. Paine (1980)
showed that population interactions do not equate with
energy flux, and asserted that energy flux was unim-
portant and “has generated few insights into ecologi-
cal processes.” This put studies of population biology
and energy-flux on separate paths that only recently
have shown signs of beginning to merge into a unified
paradigm (e. g. McQueen, et al., 1986; Hunter and
Price 1992; Polis and Winemiller 1996). Ecologists
recognize that energy input matters, although they
agree less on when and how it matters. What clearly
matters is formulating tractable hypotheses about
ecosystem structure and function. Repeated attempts
to simplify the inherent complexity of food webs have
led to Sisyphus-like progress in which investigators
have made generalizations and then have been forced
to qualify them.
SCI. MAR., 65 (Suppl. 2): 31-40 SCIENTIA MARINA 2001
A MARINE SCIENCE ODYSSEY INTO THE 21st CENTURY. J.M. GILI, J.L. PRETUS and T.T. PACKARD (eds.)
Caught in the food web: complexity made simple?*
LAWRENCE R. POMEROY
Institute of Ecology, University of Georgia, Athens GA 30602-2202, USA. E-mail: [email protected]
SUMMARY: Several historically separate lines of food-web research are merging into a unified approach. Connections
between microbial and metazoan food webs are significant. Interactions of control by predators, defenses against predation,
and availability of organic and inorganic nutrition, not any one of these, shape food webs. The same principles of popula-
tion ecology apply to metazoans and microorganisms, but microorganisms dominate the flux of energy in both marine and
terrestrial systems. Microbial biomass often is a major fraction of total biomass, and very small organisms have a very large
ratio of production and respiration to biomass. Assimilation efficiency of bacteria in natural systems is often not as high as
in experimental systems, so more primary production is lost to microbial respiration than had been thought. Simulation has
been a highly useful adjunct to experiments in both population theory and in studies of biogeochemical mass balance, but it
does not fully encompass the complexity of real systems. A major challenge for the futu.
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From the movie, Lila, Eight to Thirteen in this week's materials, identify 2–3 developmental milestones Lila reaches, and assess whether or not you think she successfully navigates her way through them as she prepares for adolescence. Support your assertions with evidence from your text and this week's materials.
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Migration and RefugeesMany immigrants in the region flee persecu.docxjessiehampson
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Many immigrants in the region flee persecution and then return after they are liberated. For example, 700,000 Jews were allowed to leave the former Soviet Union and enter Israel in the 1990s. There has also been a migration of Palestinian people. Discuss the following:
Why do you think that Israel is such an important place for the Jews?
What is the importance of the area to the Palestinians?
What do you think the impact would be on you and your families if you participated in such long-distance migration?
No references needed, need response within 3 hours!
.
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International relation can be best understood through the various schools of thought or
rather theories. They are significant in giving a comprehensive detail of the constructs that make
international relations. Realism theory still remains one of the most influential tools in
understanding events related to international relations. This is because it provides a pragmatic
approach in examining current events in the sphere of international relations (Maghroori, pg. 17).
Realism is divided into three subdivisions, seeking to explain causes of state conflict. This
include classical realism that argues that the conflict comes from the nature of man, neorealist
which associates conflict the elements of the state, and neoclassical realism which associates it to
both human nature and elements of the state. This school of thought is grounded on some
fundamental principles that make the core of its arguments.
The first assumption in realism is the idea that a country, usually referred to as a state,
serves as the main actor in international relations. It acknowledges the fact that there are other
actors like individuals and organizations, which have limited influence (Maghroori 11).
Secondly, the state is considered a unitary player, which is expected to work harmoniously, with
regard to matters of national interest. In addition, realists believe that the people who make
decisions are rational players, since this rationality is required in pursuing the interest of the
nation. In essence, the leaders are believed to understand these assumptions regardless of their
Laci Hubbard-Mattix
90000004849605
But selfish
Laci Hubbard-Mattix
90000004849605
Laci Hubbard-Mattix
90000004849605
What do you mean by "work harmoniously"
Laci Hubbard-Mattix
90000004849605
It is not clear what this sentence means.
political position, so ensure their sustainability and continuity. Consequently, it is assumed that
states exist in an anarchy context, where there is no single international leader. In this
theorization, the role of nature in influencing human action is not ignored. It asserts that nature
influence people to continue acting in repetitive tendencies. In this assumption, it comes out that
people desire power because of the egoistic nature. The innate selfishness of human beings,
mistrust and their thirst for power explains the unpredicted consequences that can result from
their actions (Maghroori 20). Such human tendencies can explain the unending wars among
nations. Bearing the fact that nations are governed by human beings, their nature contributes
largely to their behavioral tendencies, which in turn influence its security.
Realist therefore assume that leaders have the responsibility to promote the security of
their country in all fronts. This can be realized through consta.
Miller, 1 Sarah Miller Professor Kristen Johnson C.docxjessiehampson
Miller, 1
Sarah Miller
Professor Kristen Johnson
CHID 230
2 April 2019
The Myth of Disability as Isolating in Tim Burton’s Edward Scissorhands
Jay Timothy Dolmage discusses the common disability myths that condition our
understanding of disability in his work Disability Rhetoric. He argues that these myths create the
perception that disabled people are “others”, through the portrayal of them as lesser, surplus, or
improper (Dolmage, 31). One of the myths that Dolmage examines is disability as isolating or
individualizing, which is perpetrated through narratives of disabled people living in isolation,
rarely having romantic relationships or friendships, and often being left alone at the end
(Dolmage, 43). This myth can be seen in the film Edward Scissorhands, directed by Tim Burton.
Edward is a human being created by an inventor, yet the inventor’s death before his completion
leaves him with scissor blades for hands. Edward lives in a gothic mansion atop a hill,
completely in isolation until local Avon saleswoman Peg Boggs visits. She is initially frightened
by his appearance, yet decides to take him home with her upon the realization that he is
harmless. Edward’s disability causes his transition into society to be largely unsuccessful, as he
is objectified and used by other people for their benefit, and at the end of the film he is forced to
return to living in isolation after their perception of him turns to one of fear and scorn.
Edward’s isolation from society is symbolically portrayed through many film design
techniques. The mansion in which he lives at the beginning and the end of the film starkly
contrasts the community in which the able-bodied society lives. The mansion is gothic, dark, and
partially in ruins, whereas the rest of the houses are brightly colored in pinks, yellows, and
Miller, 2
greens, all with perfectly manicured green lawns. His appearance also separates him from the
rest of society, as he has very pale skin, dark under-eyes, black untamed hair, and wears gothic
industrial clothes. The able-bodied individuals often wear colorful or light clothes and appear
quite “ordinary”. The contrast created between Edward and society through set, clothing,
makeup, and hair design work to portray Edward and his disability as unusual, creepy, and
“other”. Peg even attempts to “normalize” his appearance by giving him different clothes to wear
and attempting to cover his scars with makeup, in the hopes that it will ease his transition into the
community. This film phenomenon is discussed by Martin F. Norden in his book The Cinema of
Isolation: A History of Physical Disabilities in the Movies. He argues that filmmakers will
separate disabled characters from their able-bodied peers not only through the storyline, but also
through a number of design elements. He also states that this technique allows filmmakers to
reflect an able-bodied point of view and reduce d.
Migrating to the Cloud Please respond to the following1. .docxjessiehampson
"Migrating to the Cloud" Please respond to the following:
1. Imagine that you are a CIO and you have been tasked to examine the process of moving from one host server or storage location to another. Predict two foreseen challenges of migrating an application to the cloud in a live migration and high- availability setting. Propose a preventative measure or a solution for each of these challenges.
2. Imagine that you are the CIO for a midsized organization in this industry. Determine, in 10 or less steps, the timeline for a live migration to the cloud in your organization. Determine the three greatest risks in this deployment.
.
Mike, Ana, Tiffany, Josh and Annie are heading to the store to get.docxjessiehampson
Mike, Ana, Tiffany, Josh and Annie are heading to the store to get some snacks. Mike has $1, Ana has $2, Tiffany has $3, Josh has $4, and Annie has $5.
What's the average (mean) amount of cash the five kids have? What's the median? A few days later, Annie's family won the lottery, and the kids go together to the store to get some snacks again. This time Mike has $1, Ana has $2, Tiffany has $3, Josh has $4, and Annie has wad of cash totaling $5,000.
What's the average (mean) amount of cash the five kids have this time? What's the median?
From part a, how have the mean and the median changed?
Which one - the mean or the median - is a better reflection of how much money they have together? Take you time before answering.
.
Michelle Wrote; There are several different reasons why an inter.docxjessiehampson
Michelle Wrote;
There are several different reasons why an intervention fails, such as the wrong intervention being selected or trying to solve the wrong problem. It is important that when performing and intervention that every thing have been severely observed and taken into consideration. I worked with an organization that was a travel agency, and they operated off of the commission that was collected from the booking that are processed, but they also provided a discount to the members that was taken out of the commission total. The issue was that when they initially opened the department there was no budget plan done and no guidelines were given, the agents were told to use discretion, and all though the department was a huge success in booking reservations they were still failing, because they were not withholding enough commission for the organization to operate under. Where the intervention process failed is that they never had formal training, which would have been a focus group to define the exact percentage to give to customer and the amount the organization needed to cover their overhead. During the meeting process there should have been definite guidelines to lead employees and managers from the accounting department so that the employees did not need to play the guessing game. Although they had the meeting nothing changed, because the problem was not solved with the employees and managers and was not addressed by the accounting department. The business is now in danger of folding because of the poor communication practices.
William Wrote:
Although what I am going to talk about is not my workplace but the place that I volunteer my time to sit on the board of directors for a non profit agency. As a board member we oversee the agency as a whole but we also break down into small committee groups to address needs as they arise. One of the committees that I am on is the planning committee. A change that was implemented by administration, program staff, and the board was all departments would start entering all their own data. At the time the agency had two data entry personal that was entering all agency data. So the change we made was that instead of hiring another data entry person we would require all programs to enter their own data into the collection software. This ended up being a failure that could have been huge had we not pulled reports the first two quarters of the year. What we found was some programs were right on target with getting their information entered with the first quarter. The Executive Director addressed this with staff. When the second quarter reports were pulled the data did not get any better. As an agency this failed due to program staff just did not have the appropriate time to take on more data entry. The agency ended up where we should have to start off, hiring another data entry staff member. I will say with this failure it actually turned into a very positive experience over all.
.
Midterm Lad Report 7
Midterm Lab Report
Introduction
Cellular respiration refers to all the metabolic processes and chemical reactions that take place in living organisms, particularly at the cellular level. These processes focus on the extraction of energy from nutrients. It is also responsible for converting the biochemical energy into 'adenosine triphosphate' (ATP) by the breakdown of sugars in the cells (Bennet 58). Cellular respiration is also responsible for the process by which cells release chemical energy required for conducting cellular activities. The reactions and processes facilitate the release of waste products from the cells. This experiment seeks to conduct a study of the processes and reactions involved during cellular respiration. The experiment will include several activities, such as having a study on the amount of Carbon dioxide produced during the experiment.
The number of levels of the growth of a yeast medium as a dependent variable will also be monitored during the experiment. There are other several independent variables associated with the experiment. These independent variables include sugar and temperature, among others, and their role in the experiment were also monitored. The experiment design involved the use of airtight balloons capped over reaction chambers that were used to collect the Carbon dioxide produced during the experiment. The reaction chambers contained sugars and yeast medium, which facilitated the reactions. Thermometers and pH scale were used to monitor the changes in temperature and acidity levels during the experiment. The paper involves a lab design that institute steps such as arranging the bottles used on the experiment. Notably, a proper arrangement to make sure that all the carbon dioxide released during the respiration process is well tapped in the bottles for correct lab results
Methodology
The actual procedure for experimenting involved taking measurements and recording of all observations made during the experiment. For accurate results, measures were taken three times, and a mean measurement was calculated and recorded. Winzler asserts that the mean obtained from the measurements should be used to calculate the standard deviation, which in turn facilitated the calculation of uncertainty (276). Below are the steps for conducting the experiment. It is essential to read the instructions carefully safety and accuracy during the experiment. Notably, all the lab and experiment results were well observed and thus making sure that there are limited errors in the whole process.
Consequently, all the steps required in the lab report were also clearly followed to help in getting the correct data and even not to affect the whole experiment process. The experiment involved setting the apparatus as per the set standard and the requirement. As per this concept, all the apparatus were set in a proper way to avoid vague results. Notably, to get the correct measurement and results, it is import.
MicroEssay Identify a behavioral tendency that you believe.docxjessiehampson
MicroEssay
Identify a behavioral tendency that you believe you have inherited (one that is determined, at least in part, by your genetic make-up). Explain the ways you think this trait has been affected by your environment by applying the different types of gene x environment correlations to your example (passive, evocative, and active)? What does this suggest about the nature-nurture debate?
.
MILNETVisionMILNETs vision is to leverage the diverse mili.docxjessiehampson
MILNET
Vision
MILNETs vision is to leverage the diverse military experience of Crawford employees to create awareness opportunities that help forester an appreciation, understand, and respect for the military culture and members we serve
Benefits
· Know our Members
· Support recruiting and retention
· Facilitate transition from military to Crawford
· Centralized source to connect with peer veterans
· Provide Member Experience, Marketing, and other Crawford initiatives and expert knowledge base.
MILNET Leadership Team (Volunteer position)
· Event & Volunteer Lead- Plan and execute mandatory enterprise events
· Technology Lead- Maintain MILNET budget throughout the year and reports overview or expenses monthly
· MILNET Spouse Lead- Ensures connect of sites are up to date/accurate, to include Veteran/Military Spouse Registration
· Secretary-Manages relationships by identifying opportunism for partnership
· Communications/Marketing Lead- Communicates to the MILNET community regularly via multiple channels (Email, Internal Social) regarding upcoming events, announcement, and other communications.
Background
Grandfather Air force
Parents- Army
Myself- Army
Spouse Army
Skills
Knowledgeable
Passionate
Qualified
Education
-Associates Accounting
-Bachelor’s in business and HR
-MRA w/ HR concentration
1 – Paragraph for each question (Professional answers)
Question 1- What is your visions of MILNET?
Question 2-How would your selection impact the Leadership Team?
.
midtermAnswer all question with proper number atleast 1 and half.docxjessiehampson
midterm
Answer all question with proper number atleast 1 and half page
APA FORMAT SIZE 12
1. Why is culture important to political scientists?
2. How is political science an interdisciplinary major?
3. How can politics be treated as a science?
4. Describe how modern liberalism differs from classical liberalism and explain how modern conservatism related to classical liberalism?
5. Explain how nationalism can be dangerous to a nation. Use both theoretical ideas and concrete examples to support your claims
6.
Evaluate the "end of ideology" argument by considering the facts that fit and contradict this view on today's world
7. What are the means by which power is institutionalized? What makes for good institutions? Provide examples from the United States and one other country
8. Identify the purposes of constitutions and explain why they are necessary
9. Describe how the principle of separation of powers is manifested in the U.S. Constitution and explain how this principle has evolved over time in the United States.
10. Bonus Question: What are the 10 Bill of Rights
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Midterm QuestionIs the movement towards human security a true .docxjessiehampson
Midterm Question
Is the movement towards human security a true paradigm shift? In answering this question make sure to consider which of the authors whom you have read in Weeks one to four of the course support your view and which do not. *The sole use of attached readings is required for the midterm*
Midterm Assignment – Instructions (Read Carefully)
In university courses, assignments (or assessments) are meant to give students the opportunity to demonstrate what they have been learning in the course – and give instructors evidence that such learning is occurring within the classroom. Because of these objectives, it is imperative to incorporate the specifics of what you’ve been studying in the course into your writing assignments. You accomplish this by answering the Midterm question in the assessment via the course objectives and readings from the course. The midterm will cover the following objectives:
1. Describe the role of rapid globalization in changing perceptions of security
2. Identify key threats to human security (food security, personal security, environmental security)
3. Apply the concepts of human security
4. Compare and contrast traditional international relations approaches to security with the doctrine of human security.
Additional Instructions
To answer the Midterm question you will write an analytical essay. The analytical essay is a practical approach to solving a problem. So think of this essay question as you would an assignment from your boss: “I need you to take a look at this problem and solve it for me using things from your IR toolkit (what you have learned, or know). Present a well-written, concise answer to me in four pages. I need it by tomorrow morning.” This is how it happens in the real world, and this is what we want to prepare you to do. To achieve this structure of the essay please keep the following tips in mind:
1. Remember that the analytical essay is highly-structured. Each paragraph should look like the others in terms of style and substance. Writing to the limit of four pages is an art and something you need to learn to do. So, don’t write fewer than four pages and don’t write more. You may need to write over just a little and then edit away the extra parts of the essay to reach the concise four pages.
2. Review your submission and make sure that you have covered the requirements of the assignment using only material from the lessons and readings.
Format for the Essay:
1. Do not use a cover page. Instead, create a header with your name, assignment name, and date. To do this in Word, go to “insert” and then “header.” Do the same thing to insert a ‘footer’ and include page numbers. If you need help, use the ‘help’ function to learn more within Word.
2. Your submission should be four pages (no more, no less) and look like this:
a. Introduction: Introduce your topic & include a thesis. To help you set up your analytical essay include three reasons why you agree or disagree with the midterm quest.
MGT/526 v1
Wk 2 – Apply: Organizational Analysis
MGT/526 v1
Page 2 of 2
Wk 2 – Apply: Organizational AnalysisInstructions
Complete the worksheet based on your chosen organization. Use Business Source Complete and your selected company’s website, annual report, and other available sources. Part 1: Organization Information
Organization
Define your chosen company and its industry.
Mission and Vision
Identify the mission and vision of the organization.
Mission
Vision
Organizational Initiatives
Outline 1-2 major initiative for this organization. What are they currently doing to support these initiatives?
Organizational Plans
Describe the plans employed by the organization. Determine which types of managers create each type of plan.
Type of Plan
Description
Type of Manager
SWOT Analysis
There are various factors within the external environment of an organization that impacts its strategy.
Analyze the organization’s SWOT analysis. Identify the internal and external factors. Include a link to the SWOT analysis in the Reference section of this worksheet.
Internal Factors
External Factors
Part 2: Evaluation
Evaluate if the mission, vision, planning process, and SWOT analysis meets the current needs of the organization. Include the following in your evaluation:
· Describe the unmet need, (not limited to product or service, can be new demographic, new mode of delivery, etc.).
· Analyze your competitive advantages.
· Based upon the SWOT analysis, is there another business that is doing something similar that can be referred to? Provide examples.
· If there is not another business, describe how what you’re doing is a unique product or service offering.
· Propose a competitive business initiative to address the unmet need.
· Create a high-level timeline and operational steps necessary to implement your solution. References
Include a link to theSWOT analysis.
Copyright 2020 by University of Phoenix. All rights reserved.
Copyright 2020 by University of Phoenix. All rights reserved.
COUN 6785: Social Change in Action:
Prevention, Consultation, and Advocacy
Social Change Portfolio
M. Negrón
Contents
Introduction
Scope and Consequences
Social-ecological Model
Theories of Prevention
Diversity and Ethical Considerations
Advocacy
INTRODUCTIONAdressing Teen Pregnancy in Pittsburg, California
In more recent years, there has been an effort in my community to address teen pregnancy due to its growing rates. Over the years teen pregnancy rates have continued to rise in Contra Costa County as well as surrounding counties. Unfortanately, the town I come from is a small town within Contra Costa County so resources are limited. In order to address teen pregnancy there needs to be easier access to resources to prevent teen pregnancy from occurring. Teen pregnancy can lead to a number of different problems such as low socioeconomic status, greater chance of contracting a sexually transmitted infec.
Microsoft Word Editing Version 1.0Software Requirement Speci.docxjessiehampson
This document provides a software requirements specification for Microsoft Word 2016. It includes an introduction, purpose, scope, definitions, and overview. Use cases are defined for signing in, opening, creating new files, saving, saving as, exporting, printing, and changing fonts. Requirements cover performance, usability, supportability, configurability, and recoverability. The 8 use cases are then described in more detail with normal and alternate flows and screenshots.
Microsoft Windows implements access controls by allowing organiz.docxjessiehampson
Microsoft Windows implements access controls by allowing organizations to define users, groups, and object DACLs that support their environment. Organizations define the rules, and Windows enables those rules to be enforced.
Answer the following question(s):
Do you think access controls are implemented differently in a government agency versus a typical information technology company? Why or why not?
2. Do you think access controls differ among private industries, such as retail, banking, and manufacturing? Why or why not?
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MGT520
Critical Thinking Writing Rubric - Module 10
Exceeds
Expectation
Meets Expectation Below Expectation Limited Evidence
Content, Research, and Analysis
21-25 Points 16-20 Points 11-15 Points 6-10 Points
Requirements Exceeds
Expectation -
Includes all of the
required
components as
specified in the
assignment.
Meets Expectation-
Includes most of
the required
components as
specified in the
assignment.
Below Expectation-
Includes some of
the required
components as
specified in the
assignment.
Limited Evidence -
Includes few of the
required
components as
specified in the
assignment.
21-25 Points 16-20 Points 11-15 Points 6-10 Points
Content Exceeds
Expectation -
Demonstrates
substantial and
extensive
knowledge of the
materials, with no
errors or major
omissions.
Meets Expectation-
Demonstrates
adequate
knowledge of the
materials; may
include some
minor errors or
omissions.
Below Expectation-
Demonstrates fair
knowledge of the
materials and/or
includes some
major errors or
omissions.
Limited Evidence -
Fails to
demonstrate
knowledge of the
materials and/or
includes many
major errors or
omissions.
25-30 Points 19-24 Points 13-18 Points 7-12 Points
Analysis Exceeds
Expectation -
Provides strong
thought, insight,
and analysis of
performance
management
system, concepts
and applications.
Meets Expectation-
Provides adequate
thought, insight,
and analysis of
performance
management
system, concepts
and applications.
Below Expectation-
Provides poor
thought, insight,
and analysis of
performance
management
system, concepts
and applications.
Limited Evidence -
Provides little or no
thought, insight,
and analysis of
performance
management
system, concepts
and applications.
13-15 Points 10-12 Points 7-9 Points 4-6 Points
Sources Exceeds
Expectation -
Sources go above
and beyond
required criteria,
and are well
chosen to provide
effective
substance and
perspectives on
the issue under
examination.
Meets Expectation-
Sources meet
required criteria
and are adequately
chosen to provide
substance and
perspectives on the
issue under
examination.
Below Expectation-
Sources meet
required criteria,
but are poorly
chosen to provide
substance and
perspectives on the
issue under
examination.
Limited Evidence -
Source selection
and integration of
knowledge from
the course is
clearly deficient.
Mechanics and Writing
5 Points 4 Points 3 Points 1-2 Points
Demonstrates Exceeds Meets Expectation- Below Expectation- Limited Evidence -
MGT520
Critical Thinking Writing Rubric - Module 10
college-level
proficiency in
organization,
grammar and
style.
Expectation -
Project is clearly
organized, well
written, and in
proper format as
outlined in the
assignment. Strong
sentence and
paragraph
structure; contains
no errors in
grammar, spelling,
APA style, or APA
citations and
references..
Midterm PaperThe Midterm Paper is worth 100 points. It will .docxjessiehampson
Midterm Paper
The Midterm Paper is worth 100 points. It will consist of a 500 word written description and analysis of a work of art using terminology from Chapters 2-5.
For this assignment, you are to discuss the form, content, and subject matter of a work of art chosen from the list provided. This is an exercise in recognizing visual elements and principles of design in works of art and demonstrating an understanding of how they relate to each other to create meaning. This paper is about looking and seeing. This is not a research paper; you will not need to do additional research. Please follow the outline provided below.
First: Select a work of art
Select one of the following listed works of art:
Circle of Diego Quispe Tito.
The Virgin of Carmel Saving Souls in Purgatory
. Late 17th century. Fig. 1.22, pg. 17.
Henri Matisse.
Large Reclining Nude
. 1935. Fig. 4.24, pg. 85.
Faith Ringgold.
Tar Beach
. 1988. Fig. 13.18, pg. 219.
Henry Ossawa Tanner.
The Banjo Lesson
. 1893. Fig. 21.15, pg. 373
Andy Warhol.
Marilyn Diptych
. 1962. Fig. 24.23, pg. 447.
Format
Describe the use of each visual element and principle of design in the order they are listed in the outline. You can simply list each term and address how it is used in the painting. If you write in paragraph form be sure to identify each term clearly. Any term not addressed will receive 0 points. Provide specific examples. For example, don’t just say “there are lines,” give specific examples of how line is used in the piece you’ve selected.
Papers should be 500 words minimum (not including images), double-spaced, 10 or 12 point, with 1" margins. The preferred format is Microsoft Word (.doc or .docx). If these formats are not available, other acceptable formats are ASCII (.txt), rich text format (.rtf), Open Office (.odt), and PDF. Make sure you proofread your papers for incorrect grammar, spelling, punctuation, and other errors.
The Midterm Paper is due at 11:59 pm CT Sunday of Week 4.
Midterm Paper Outline
Introduction (First Paragraph)
In the first paragraph, called the introduction, you will include:
An identification of the work of art you selected: The name of the artist, title (which is underlined or italicized every time you use the title in your paper), date, and medium.
Your initial interpretation of the subject based on your initial observations.
Description
Describe how each of the following is used in the piece you selected.
Visual Elements
:
Line: what types of lines do you see in the piece? Provide examples.
Shape: what types of shapes do you see? Provide examples.
Mass: How is mass implied?
Space: How is the illusion of space created in the piece?
Time and Motion: Are time and motion evident in tis piece? How so?
Light: How is light used here?
Color: How does the artist use color?
Texture: How does the artist create the illusion of texture, or incorporate actual texture
Principles of Design
Unity and Variety: In what way is this pi.
Miami Florida is considered ground zero for climate change, in parti.docxjessiehampson
Miami Florida is considered ground zero for climate change, in particular rising seas will not only drown coastal sections of the city but will disrupt our local supply of drinking water.
Based on what you have learned so far from this class, discuss the following:
Explain where the drinking water from South Florida primarily comes from and why would rising sea levels disrupt this supply?
What efforts can be made and are being made to mitigate the effects of rising seas on our drinking water?
If you were a local politician, what advice would you give to state and federal officials on the best way to ensure residents in South Florida had a steady supply of drinking water for many years to come?
.
MGT230 v6Nordstrom Case Study AnalysisMGT230 v6Page 2 of 2.docxjessiehampson
MGT/230 v6
Nordstrom Case Study Analysis
MGT/230 v6
Page 2 of 2
Nordstrom Case Study Analysis
Nordstrom—“High Touch” with “High Tech”
How does Nordstrom stay profitable despite dips in consumer spending, changing fashion trends, and intense competition among retailers? One answer: Acute attention to detail and well-laid plans.
All in the Family
The fourth generation of family members that runs Nordstrom has brought the store’s time-honored and successful retail practices into a new era. “Nordstrom, it seems, is that rarity in American business: an enterprise run by a founding family that hasn’t wrecked it,” says one business writer. The company provides a quality customer experience via personalized service, a compelling merchandise offering, a pleasant shopping environment, and increasingly better management of its inventory.
Secret of Success
The secret of this company’s success lies in its strategic planning efforts and the ability of its management team to set broad, comprehensive, and longer-term action directions, all of which are focused on the customer experience. The current generation of Nordstrom family members was quick to spearhead an ultramodern multimillion-dollar, Web-based inventory management system. This upgrade helped the company meet two key goals: (1) correlate purchasing with demand to keep inventory as lean as possible, and (2) give customers and sales associates a comprehensive view of Nordstrom’s entire inventory, including every store and warehouse.
Demand Planning
Instead of relying on one-day sales, coupon blitzes, or marking down entire lines of product, Nordstrom discounts only certain items. “Markdown optimization” software assists in planning more profitable sale prices. According to retail analyst, Patricia Edwards, this helps Nordstrom calculate what will sell better at different discounts and forecast which single items should be marked down. If a style is no longer in demand, the company can ship it off to its Nordstrom Rack outlet stores. It’s all part of Nordstrom’s long-term investment in efficiency. “If we can identify what is not performing and move it out to bring in fresh merchandise,” says Pete Nordstrom, “that’s a decision we want to make.”
Inventory Planning
Although inventory naturally fluctuates, Nordstrom associates can easily locate any item in another store or verify when it will return to stock. Customers on their smart phones and associates behind sales counters see the same thing—the entire inventory of Nordstrom’s stores is presented as one selection, which the company refers to as perpetual inventory. “Customer service is not just a friendly, helpful, knowledgeable salesperson helping you buy something,” says Robert Spector, retail expert and author of The Nordstrom Way. “Part of customer service is having the right item at the right size at the right price at the right time. And that’s something perpetual inventory will help with.”
The upgraded inventory management system was an .
How to Build a Module in Odoo 17 Using the Scaffold MethodCeline George
Odoo provides an option for creating a module by using a single line command. By using this command the user can make a whole structure of a module. It is very easy for a beginner to make a module. There is no need to make each file manually. This slide will show how to create a module using the scaffold method.
Main Java[All of the Base Concepts}.docxadhitya5119
This is part 1 of my Java Learning Journey. This Contains Custom methods, classes, constructors, packages, multithreading , try- catch block, finally block and more.
LAND USE LAND COVER AND NDVI OF MIRZAPUR DISTRICT, UPRAHUL
This Dissertation explores the particular circumstances of Mirzapur, a region located in the
core of India. Mirzapur, with its varied terrains and abundant biodiversity, offers an optimal
environment for investigating the changes in vegetation cover dynamics. Our study utilizes
advanced technologies such as GIS (Geographic Information Systems) and Remote sensing to
analyze the transformations that have taken place over the course of a decade.
The complex relationship between human activities and the environment has been the focus
of extensive research and worry. As the global community grapples with swift urbanization,
population expansion, and economic progress, the effects on natural ecosystems are becoming
more evident. A crucial element of this impact is the alteration of vegetation cover, which plays a
significant role in maintaining the ecological equilibrium of our planet.Land serves as the foundation for all human activities and provides the necessary materials for
these activities. As the most crucial natural resource, its utilization by humans results in different
'Land uses,' which are determined by both human activities and the physical characteristics of the
land.
The utilization of land is impacted by human needs and environmental factors. In countries
like India, rapid population growth and the emphasis on extensive resource exploitation can lead
to significant land degradation, adversely affecting the region's land cover.
Therefore, human intervention has significantly influenced land use patterns over many
centuries, evolving its structure over time and space. In the present era, these changes have
accelerated due to factors such as agriculture and urbanization. Information regarding land use and
cover is essential for various planning and management tasks related to the Earth's surface,
providing crucial environmental data for scientific, resource management, policy purposes, and
diverse human activities.
Accurate understanding of land use and cover is imperative for the development planning
of any area. Consequently, a wide range of professionals, including earth system scientists, land
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vol. 191, no. 1 the american naturalist january 2018E-Articl.docx
1. vol. 191, no. 1 the american naturalist january 2018
E-Article
Thermoregulatory Behavior Simultaneously Promotes
and Forestalls Evolution in a Tropical Lizard
Martha M. Muñoz1,* and Jonathan B. Losos2
1. Department of Biological Sciences, Virginia Tech,
Blacksburg, Virginia 24060; 2. Department of Organismic and
Evolutionary Biology,
Harvard University, Cambridge, Massachusetts 02138
Submitted January 11, 2017; Accepted July 21, 2017;
Electronically published October 25, 2017
Online enhancements: appendix.
abstract: The role of behavior in evolution has long been
discussed,
with some arguing that behavior promotes evolution by
exposing
organisms to selection (behavioral drive) and others proposing
that
it inhibits evolution by shielding organisms from environmental
var-
iation (behavioral inertia). However, this discussion has
generally fo-
cused on the effects of behavior along a single axis without
considering
that behavior simultaneously influences selection in various
niche
dimensions. By examining evolutionary change along two
2. distinct
niche axes—structural and thermal—we propose that behavior
si-
multaneously drives and impedes evolution in a group of Anolis
lizards from the Caribbean island of Hispaniola. Specifically, a
behav-
ioral shift in microhabitat to boulders at high altitude enables
thermo-
regulation, thus forestalling physiological evolution in spite of
colder
environments. This same behavioral shift drives skull and limb
evolu-
tion to boulder use. Our results emphasize the multidimensional
ef-
fects of behavior in evolution. These findings reveal how, rather
than
being diametrically opposed, niche conservatism and niche
lability
can occur simultaneously. Furthermore, patterns of niche
evolution
may vary at different geographic scales: because of
thermoregulatory
behavior, lizards at high and low elevation share similar
microclimatic
niches (consistent with niche conservatism) while inhabiting
distinct
macroclimatic environments (consistent with niche divergence).
To-
gether, our results suggest that behavior can connect patterns of
niche
divergence and conservatism at different geographic scales and
among
traits.
Keywords: niche evolution, behavior, thermoregulation,
Caribbean,
3. lizard, Bogert effect.
Introduction
For nearly a century, behavior has been recognized as a
key pacemaker for evolution (discussed in Huey et al. 2003;
Duckworth 2009). On the one hand, behavior can facilitate
evolutionary change; as organisms explore novel environ-
ments, they experience new selective pressures that can lead
* Corresponding author; e-mail: [email protected]
Am. Nat. 2018. Vol. 191, pp. E000–E000. q 2017 by The
University of Chicago.
0003-0147/2018/19101-57488$15.00. All rights reserved.
DOI: 10.1086/694779
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to phenotypic adaptations and speciation (Miller 1956;
Lande 1981; Wyles et al. 1983; Bateson 1988; West-Eberhard
1989; Sol et al. 2005). Mayr (1963) proposed, for example,
that shifts in behavior were key initial changes precipitating
the prolific morphological diversification of tropical birds.
An alternative school of thought contends that behavior
often plays a constraining role in evolution (Bogert 1949;
Bartholomew 1964; Wake et al. 1983; Brandon 1988). Or-
ganisms can choose microhabitats to which they are already
well adapted, preventing exposure to directional selection,
even in the face of changing environmental conditions.
The Bogert effect (sensu Huey et al. 2003) is a prime exam-
ple of this process: rather than shifting their physiology,
ectotherms such as reptiles and insects use regulatory be-
haviors to buffer their physiology from selection, even as en-
vironments change across space or through time (Huey
1982; Angilletta 2009). In this context, behavior constrains
rather than facilitates evolutionary divergence.
Both perspectives have empirical support, but consider-
4. ing them a dichotomy overlooks the connectivity among
a species’ multiple niche dimensions (Hutchinson 1957;
Holt 2009). By simultaneously affecting diverse aspects of
the niche, a single behavior can mold the natural selection
pressures on multiple phenotypic traits (Lewontin 1983;
Levins and Lewontin 1985; Huey et al. 2003). For example,
a population may be adapted to eat a particular type of
food found in certain microhabitats with specific micro-
climates. By behaviorally shifting to eat a different food that
occurs in a separate microhabitat, the population may have
to adapt not just to the new diet item but also to new struc-
tural habitat and microclimatic niche conditions. In other
words, behavioral shifts in the use of any resource may have
multiple potentially contrasting effects on trait evolution
(Levins and Lewontin 1985; Huey et al. 2003), but these kinds
of effects remain empirically unexplored.
Anolis lizards provide a unique opportunity to test the
multidimensional effects of behavior on evolution because
both their physiology and morphology are tightly linked to
64.011.153 on October 28, 2017 07:17:22 AM
s and Conditions (http://www.journals.uchicago.edu/t-and-c).
E000 The American Naturalist
habitat choice (Losos 2009). The form-function relationship
between structural habitat (i.e., the matrix of structural sup-
ports these arboreal lizards utilize) and morphology is well
established (Losos 1990; Garland and Losos 1994; Irschick
and Losos 1999), supplying concrete hypotheses for how be-
havioral changes in microhabitat use will affect morphologi-
cal evolution. Similarly, the effects of thermal habitat use (i.e.,
the matrix of thermal patches in a habitat) on thermal phys-
iology in anoles are also well understood (Ruibal 1961; Rand
5. 1964; Hertz 1974, 1981, 1992a, 1992b; Huey et al. 2009), and
anoles are known to shift their habitat use at different ele-
vations (Rand 1964; Schoener and Schoener 1971; Hertz and
Huey 1981; Schwartz 1989). In other words, habitat choice by
anolesmayimpact boththeirphysiologyandtheirmorphology.
We focus our study on the Anolis cybotes species complex
(the cybotoids) from the Caribbean island of Hispaniola.
The thermal biology of these lizards makes them an excel-
lent subject for the study of the role of behavior in evolu-
tion. Despite being distributed across a broad altitudinal
range (from sea level to more than 2,500 m), lizards in this
complex have quite similar body temperatures (Hertz and
Huey 1981; Muñoz et al. 2014). Similarly, heat tolerance
(CTmax) remains nearly constant (∼407C) across this eleva-
tional range (Muñoz et al. 2014). In many lizards, including
many species of anoles, both body temperature and thermal
tolerance are labile, often shifting substantially over shal-
lower thermal clines (Hertz 1979; van Berkum 1988; Labra
et al. 2009; Rodríguez-Serrano et al. 2009; Bonino et al. 2011;
Muñoz et al. 2016). In contrast, the minimal change in body
temperature in the cybotoids over such large elevational tran-
sects suggests that behavioral thermoregulation, potentially
involving shifts in microhabitat use, may be occurring.
We address four specific hypotheses in this study. We
first predicted that thermoregulatory behavior changes with
altitude, such that montane lizards behaviorally compensate
for their colder environments. Second, we predicted that ther-
moregulation at high elevation is accomplished by a shift in
structural microhabitat away from these lizards’ ancestral ar-
boreal niche to capitalize on more exposed and thus warmer
substrates, namely boulders (Rand 1964; Schoener and Schoe-
ner 1971; Hertz and Huey 1981; Schwartz 1989). Third, we
tested the hypothesis that behavioral shifts in microhabitat
use involve adaptive shifts in skull, limb, and toe pad morphol-
6. ogy. Specifically, we predicted that boulder-dwelling lizards
evolved a flattened body form and fewer toe pad scales as well
as differences in hind limb length. Lizards utilizing boulders
and cliff faces are typically characterized by dorsoventral flat-
tening, which eases their movements into narrow crevices and
tight spaces under boulders (Vitt et al. 1997; Arnold 1998a;
Revell et al. 2007; Goodman and Isaac 2008). Anoles perching
close to the ground tend to have fewer adhesive toe scales (la-
mellae) than arboreal anoles because lamellae assist with cling-
ing to vertical surfaces (Collette 1961; Glossip and Losos 1997;
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Stuart et al. 2014). Shifts in relative limb length are also often
associated with boulder use in lizards, although the direction
of change varies, with longer hind limbs evolving in some lin-
eages and shorter hind limbs evolving in others (Leal et al.
2002;
Goodman 2007; Revell et al. 2007; Goodman et al. 2008; Collar
et al. 2011; Muñoz et al. 2015; Blom et al. 2016). Why limb
length evolves in different directions is not fully understood
(Cartmill 1985; Pounds 1988; Losos and Sinervo 1989; Losos
et al. 1993; Losos and Irschick 1996; Arnold 1998b). As a re-
sult, we predict that the shift to using boulders will lead to a
change in limb length, but we do not specify the predicted
direction of this change. Finally, using a common garden ex-
perimental design, we tested whether such morphological
shifts are evolved or due to phenotypic plasticity.
Material and Methods
Species and Study Sites
The Anolis cybotes species complex is found throughout the
island of Hispaniola (Schwartz 1989; Glor et al. 2003), across
habitats ranging from tropical lowland forests and lowland
xeric scrub forest to cold montane pine forests. Importantly
for this study, A. cybotes and its close relatives are distributed
7. across wide elevational ranges in the island’s two principal
mountain chains, the Cordillera Central and the Sierra de
Baoruco, or the eastern and western mountains, respectively
(fig. A1; figs. A1, A2 are available online). Lizards in this
group found at elevations higher than 1500 m have been
designated as separate species—Anolis shrevei in the eastern
mountains and Anolis armouri in the western mountains
(Schwartz 1989)—but these populations are phylogenetically
nested within populations described as A. cybotes (Glor et al.
2003; Wollenberg et al. 2013). Anolis armouri and A. shrevei
are each more closely related to lowland populations of
A. cybotes in the same mountain range than they are to each
other (Glor et al. 2003; Wollenberg et al. 2013), suggesting
that the invasion of high-elevation environments occurred in-
dependently. We studied A. cybotes at four sites in this study,
one low- and one high-elevation site in each mountain chain.
Study sites in the Sierra de Baoruco were located in Los Patos,
Barahona Province (45 m), and in Loma del Toro, Sierra de
Baoruco National Park, Independencia Province (2,258 m),
and sampling days were June 11–16, 2012. Study sites in
the Cordillera Central were located in the Francisco Alberto
Caamaño Deñó National Park, Azua Province (13 m), and
in Valle Nuevo National Park, La Vega Province (2,450 m),
and sampling days were June 3–8, 2012 (table 1).
In contrast to the mesic, broadleaf forests found near sea
level, the montane habitats of Hispaniola are covered in
pine forest stands interspersed with savannas and rocky
outcrops (Martin et al. 2011). Previous research has found
that both A. armouri and A. shrevei are notably absent from
64.011.153 on October 28, 2017 07:17:22 AM
s and Conditions (http://www.journals.uchicago.edu/t-and-c).
Behavioral Inertia and Behavioral Drive E000
8. the deep forest and instead primarily occur in savannas and
other clearings (Hertz and Huey 1981; Williams 1983). To
gather body and operative temperature data, we centered our
study sites on the open clearings adjacent to the deep forest.
Nonetheless, during our experiments (described below) we
periodically searched for lizards in deep forest. None of these
searches yielded any lizards, confirming that they were pri-
marily restricted to clearings and forest edges.
Measurement of Thermoregulation Efficiency
Behavioral thermoregulation occurs when an organism uses
behavior to actively maintain a particular mean and variance
of body temperature (Cowles and Bogert 1944; Huey 1982).
To determine whether lizards were actively regulating their
body temperature, we measured the efficiency (E) of thermo-
regulation, which estimates how well organisms are able to
stay within their preferred temperature range, given their lo-
cal thermal conditions (Hertz et al. 1993). To estimate E (de-
scribed in detail below), we needed to know field-measured
body temperatures (Tb), the preferred thermal conditions
for the lizards (Tsel), and the environmental (operative) tem-
perature (Te) at each of the sites, which we measured as
follows.
Operative Temperature. The operative environmental tem-
perature (Te) is the steady-state temperature of an organism
in the absence of physiological thermoregulation (Bakken
1992). To estimate Te at each site, we deployed 44 lizard
models containing an iButton temperature sensor (DS1921G,
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Maxim). Models were built from electroformed copper match-
ing the shape and color of an adult male A. cybotes (Bakken
and Angilletta 2014; Muñoz et al. 2014). We examined lizard
color by measuring reflectivity of the head, body, and tail using
an Ocean Optics USB 2000 spectrometer. Reflectivity values
9. were recorded as percent reflectance relative to a barium sul-
phate white standard using an Ocean Optics R400 ultraviolet-
visible (UV-VIS) reflectance probe, which was attached to a
DT-1000 tungsten halogen/deuterium UV-VIS light source
(Ocean Optics, Dunedin, FL). Note that we are assuming that
the reflectivity in the visible range is also representative of the
near infrared. The matte brown paint color for lizard models
was selected by comparing sampled lizard reflectance to var-
ious paint samples. More details for building model are pro-
vided by Muñoz et al. (2014).
To establish transects for placing copper lizard models
on perches, we walked a certain number of meters into
the study site (with number of meters determined using a
random number table). From that point we then picked a
direction in which to walk (07–3607 in 457 increments), with
direction also chosen using a random number table. On the
basis of previous work (Schwartz 1989; Glor et al. 2003;
Wollenberg et al. 2013), these lizards are most commonly
observed on boulders and arboreal vegetation within 1.5 m
of the ground. Following Hertz (1992a), we placed copper
models on tree trunks and boulders in roughly equal num-
bers at each site. Substrate choice (trunk vs. boulder), device
height (0–150 cm in 15-cm increments), and orientation for
perches (07–3607 in 457 increments) in each transect were all
chosen using a random number table. Models recorded Te
Table 1: Summary environmental information for localities
visited and for lizards’ preferred temperature ranges
Locality
Coordinates
Altitude
(m)
Mean Te 5 1
10. SD (7C)
64.011.153 o
s and Condit
Preferred thermal range
Tsel (7C; N)
n October 28, 2017 07:17:22 A
ions (http://www.journals.uchic
Thermal habitat quality
de 5 1 SD (7C)
Cordillera Central (east):
Low: Caamaño
18726.1190N,
70735.5460W
13
30.4 5 3.9
29.1–31.3 (17)
3.27 5 2.17
High: Valle Nuevo
18743.8000N,
70736.0000W
2,450
23.6 5 10.1
26.8–31.4 (19)
7.68 5 5.76
11. Sierra de Baoruco (west):
Low: Los Patos
17757.3000N,
71711.2880W
45
31.3 5 4.7
28.3–30.4 (21)
3.65 5 3.54
High: Loma de Toro
18717.2440N,
71742.7490W
2,320
18.2 5 6.3
28.8–32.7 (23)
12.99 5 5.21
Note: This study was conducted at low- and high-elevation sites
in the two main mountain ranges—the Cordillera Central (east)
and the Sierra de Baoruco
(west)—on the Caribbean island of Hispaniola. The mean
operative temperature, Te (i.e., the expected temperature of
nonthermoregulating lizards), is substan-
tially lower at high elevation than near sea level. Despite this
12. environmental difference, lizards at high elevation have
preferred thermal ranges (Tsel) nearly iden-
tical to their low-elevation counterparts, indicating a greater
mismatch between their preferred temperature and the available
thermal habitat (higher de). To
compensate for their colder habitats, high-elevation lizards are
predicted to thermoregulate more effectively than their low-
elevation counterparts (table 2). Mea-
surement of Te and Tsel are described in “Material and
Methods.” Thermal habitat quality, de, describes the deviation
between the operative temperature (Te) and
the preferred temperature range (Tsel), such that a value of 0
indicates that Te falls within Tsel (high thermal quality), and
progressively greater numbers indicate
increasing mismatch between Te and Tsel (low thermal quality).
M
ago.edu/t-and-c).
E000 The American Naturalist
every 10 min from 0600 to 1900 hours for 6 continuous days,
providing an average of 1,714 copper model hours of sam-
pling per site.
As discussed above, the high-elevation lizards (A. armouri
and A. shrevei) are found in open clearings adjacent to deep
forest (in which they are notably absent; Hertz and Huey
1981; Williams 1983). We centered our study sites in the sa-
vannas and rocky clearings where these lizards occur. We did
not explicitly quantify habitat availability (i.e., the relative
abundance of boulders and trees) in our study sites, because
it would have been arbitrary to decide which trees were avail-
able to lizards. Hence, we measured the operative tempera-
tures of boulders and trees found in the savannas and in the
surrounding forest edges. Thus, our study design does not al-
13. low us to explicitly quantify lizardpreference for forest or clear-
ings or for the perches within them. However, our approach
does allow us to compare the distribution of operative tem-
peratures on the two most commonly utilized substrates in the
habitats where lizards occur and to test whether lizard perch
use in those habitats confers a thermoregulatory advantage.
Body Temperature. On the same days that we recorded Te,
we also measured Tb from wild adult male lizards. Following
established methods (Hertz 1992a; Muñoz et al. 2014), two
researchers walked slowly through each habitat and cap-
tured adult lizards using a noose tied to the end of a 3–
3.7-m telescopic panfish pole (Cabela’s, Sidney, NE). We
measured Tb (to the nearest 0.17C) through the cloaca using
a thermocouple (type T, 5SC series, 36 gauge [0.13-mm di-
ameter], OMEGA Engineering) connected to a temperature
logger (HH603A; Omega). We marked lizards using non-
toxic paint to prevent resampling during our experimental
period.
Preferred Temperature Range. The preferred temperature
range or the set point range (Tsel) refers to the central 50%
of body temperatures measured in lizards that have been
put in a thermal gradient and allowed to choose where to
sit (i.e., in the absence of environmental constraints; Huey
1982; Hertz et al. 1993). We measured Tsel from wild-caught
adult male lizards from all localities (N p 17–23 per popu-
lation; table 1). Before experiments, lizards were maintained
in environmentally controlled reptile facilities at Harvard
University. Lizards were kept at 29.47C and under a 12L∶ 12D
cycle for less than a week after capture. Lizards were not fed in
the 24 h before experiments. The thermal arena consisted of a
pine box with eight identical lanes (91 cm#15 cm#14 cm)
covered with chicken wire. We laid a bed of gravel over heat
cables (150 W; Zoo Med) in an air-conditioned room, creat-
ing a stable temperature range of 187–407C in each lane. Be-
14. fore experiments, we inserted a thermocouple (type T, 5SC
series, 40 gauge [0.076-mm diameter], OMEGA Engineer-
ing) ∼1 cm into the cloaca of each lizard and secured it to
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the base of the tail using medical tape. The probe was con-
nected to a digital temperature logger (HH147U, Omega).
Lizards acclimated to the experimental chamber for 30 min
before the beginning of the experiment. Temperature loggers
recorded Tb every 10 min during each 4-h experimental
trial. Trials started at 0830 and 1330 hours, and we put equal
numbers of lizards from each population into each time
trial.
Thermoregulatory Efficiency. We calculated thermoregula-
tory efficiency (E) for adult males observed in the wild as
follows: E p 1 2 db=de, where db and de refer to the mean
deviation of Tb and Te from Tsel, respectively (Hertz et al.
1993). A highly efficient thermoregulator would have a low
db (meaning they are effective at maintaining temperatures
within their preferred range), even in habitats in which op-
erative temperatures diverge from the preferred range (i.e.,
high de), leading to values of E close to 1. As organisms be-
come more behaviorally passive with respect to tempera-
ture, the ratio between db and de increases, and E approaches
0. Following Hertz et al. (1993), we set confidence intervals
on E through bootstrap resampling of our empirical distri-
butions of Te and Tb. Specifically, we computed de and db
by randomly drawing samples (with replacement) of n ob-
servations of Te and Tb, where n is the total number of
observations. We estimated the confidence interval from
1,000 replicates of random resampling.
Structural Microhabitat Use
For each lizard captured for Tb measurement (described
above), we also recorded two key aspects of microhabitat
15. use at first observation: substrate type and perch height.
Perch types included rocky substrates (boulders, ledges),
woody substrates (branches, logs, roots, trunks, and twigs),
leafy substrates (vines and leaves), man-made substrates
(fence posts and walls), and the bare ground (soil). We com-
pared boulder use with arboreal perch use (woody and leafy
substrates) using the G-test of goodness-of-fit. We excluded
a few observations of lizards on unusual perches, such as a
coconut (n p 1) and a conch shell (n p 1). We also excluded
man-made substrates (posts and gates) that were only avail-
able at low elevation. All other perches (boulders, branches,
leaves, logs/stumps, tree roots, tree trunks, and twigs) were
found in abundance at all sites. Following previous studies
(Hertz 1981; Muñoz et al. 2014), we also recorded how ex-
posed to sunlight the lizards were; lizards were considered
to be exposed if 50% or more of their bodies was in direct
sunlight. We compared observations of exposed (full sun),
partially exposed (filtered), and shaded lizards among pop-
ulations using a G-test for goodness-of-fit. To avoid resam-
pling, all captured lizards were marked using nontoxic ink.
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Behavioral Inertia and Behavioral Drive E000
Morphological Analysis
To obtain morphological measurements, we captured adult
male lizards (n p 20–31) from each locality. We took digital
images (1200 dpi) of toe pads using a flatbed scanner, from
which we measured lamella number (number of adhesive
scales under the fourth toe). We then generated full skeletal
images using a portable X-ray machine (Kodex). From each
image, we measured body size (represented by snout-vent
length [SVL]; the distance from the tip of the snout to the
16. cloaca), femur length, tibia length, fourth metatarsal length,
and fourth hind toe length. All skeletal measurements were
gathered using the Object-J plugin in Image-J (ver. 1.47; Abra-
moff et al. 2004). We measured head height (the distance from
under the base of the jaw to the top of the head) using digital
calipers (Mitutoyo) to the nearest 0.01 mm. All morphological
traits were log transformed before analysis. We tested for dif-
ferences in morphological traits between high- and low-
elevation
lizards within transects using ANCOVA, with morphologi-
cal traits as separate dependent variables, SVL as the covar-
iate, and elevation (high vs. low) as the fixed effect.
Common Garden Experiment
In July 2012, we collected 20 gravid females each from the
high- (Loma de Toro) and low- (Los Patos) elevation sites
in the Sierra de Baoruco. We kept these females in environ-
mentally controlled facilities (room temperature p 29:47C)
at Harvard University. Females were housed with one male
from their native locality. Each enclosure consisted of a 30-cm3
polyester/nylon mesh cage (BioQuip Products) and included
a felt cage carpet, two bamboo dowels for perching, and a
small pot with moist soil for laying eggs. Cages were illumi-
nated with full spectrum lighting, which was maintained at
a 12L∶ 12D cycle. Lizards were misted twice daily and fed
1.25-cm vitamin-dusted crickets three times per week.
Cages were checked three to four times per week for eggs
from August to December 2012. Despite several months of
effort (through May 2013), only two eggs were laid by fe-
males from the high-elevation population (Loma de Toro)
under our laboratory conditions, and no changes in hus-
bandry induced the females to become gravid again. Thus,
we chose to rear young from eggs excised directly from
17. females. In June 2013, we collected 10 heavily gravid fe-
males from Loma de Toro and surgically removed the shelled
eggs from the oviduct. Eggs were immediately transferred to
cell culture dishes containing moistened vermiculite, where
they remained until hatching. Hatchlings were immediately
placed in individual cages with temperature, lighting, feed-
ing, and misting conditions as described above. We fed juve-
nile lizards vitamin-dusted Drosophila twice weekly until
they reached approximately 20 mm in body size (SVL), at
which point they were switched to a diet of 1.25-cm vitamin-
dusted crickets.
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We reared male lizards to adulthood (8–10 months) and
then measured skull and limb dimensions, as described
above. As with adults, we tested for differences in morpho-
logical traits between high- and low-elevation laboratory-
reared lizards using ANCOVAs, with SVL as the covariate
and elevation (high vs. low) as the fixed effect.
Results
Thermoregulatory Efficiency Increased with Altitude
The set-point range (Tsel) varied little between high- and
low-elevation populations of lizards (table 1). On average,
operative temperatures (Te) in montane environments are
about 77C below lizards’ preferred temperature range (table 1;
fig. 1), indicating lower thermal quality (higher de) at high el-
evation. Lizards in these cool montane habitats are effective
thermoregulators (E p 0:93; table 2), meaning that they
maintain their body temperatures within their preferred
thermal range, despite the available environment tending
away from the preferred range (table 2). This near-perfect
behavioral efficiency appears to stem from montane lizards
effectively utilizing the relatively narrow sliver of their hab-
itat matching their ancestral preferred thermal conditions
18. (fig. 1C, 1D). At low elevation, in contrast, lizards are more
behaviorally passive (lower E; table 2), with the distribution
of body temperatures closely mirroring the distribution of
operative temperatures (fig. 1A, 1B). This correspondence
likely results because their warmer habitats more closely
match their thermal preferences: at low elevation, operative
temperatures were, on average, 17C warmer than the pre-
ferred range (i.e., lower de) than at high elevation (fig. 1; ta-
ble 1). In other words, lowland lizards do not need to ther-
moregulate as precisely to remain within their preferred
range, whereas montane lizards must thermoregulate to ac-
complish the same feat and appear to do so effectively.
Microhabitat Use Differed between
High- and Low-Elevation Sites
At high elevation, most operative temperatures (Te) were
below lizards’ preferred range, but there were clear differ-
ences in thermal habitat quality between boulders and trees.
Nearly all (93%) of the Te’s measured on trees were below
lizards’ preferred range, in contrast to 70% of the Te’s mea-
sured on boulders (fig. 1). Whereas lowland lizards predom-
inantly (68% of observations) perched on arboreal vegeta-
tion (tree trunks, roots, leaves, branches), most lizards at
high elevation were observed on rocky substrates (83% of
observations; tables 2, A1; tables A1, A2 are available online;
G-test for differences in boulder use between high- and low-
elevation lizards; P ! :001 for both mountain chains). Con-
comitant with the substrate switch, nearly all lizards at high
elevation occupied habitats exposed to the sun (table A2;
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E000 The American Naturalist
19. fig. A2; G-test for differences in perch exposure between
high- and low-elevation lizards: P ! :001 for both mountain
chains). In short, perching on boulders provides montane
lizards with a thermoregulatory advantage, enabling them
to be highly efficient thermoregulators in relatively cold
habitats.
Montane Lizards Evolved Flatter Skulls
and Shorter Hind Limbs
Morphological analyses of wild-caught adults revealed that
montane cybotoid anoles have more dorsoventrally flat-
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tened skulls relative to lowland lizards (table 3; fig. 2). Mon-
tane lizards also had relatively shorter hind limbs than low-
land lizards, a result that stems from reduction in the length
of the femur, tibia, and metatarsus but not the fourth toe
(table 3; fig. 2). High- and low-elevation lizards did not dif-
fer in lamella number.
The common garden experiment revealed that the mor-
phological differences observed among wild-caught lizards
were evident in the laboratory-reared generation as well (ta-
ble 3; fig. 2). As with wild-caught adults, laboratory-reared
lizards from high-elevation habitats had dorsoventrally flat-
tened heads, shorter femora, and shorter tibiae than low-
A B
C D
0.
04
0.
08
22. 0 10 20 30 40 50 60 0 10
20 30 40 50 60
F
re
qu
en
cy
F
re
qu
en
cy
F
re
qu
en
cy
Tsel Tsel
Tsel
Tb lizards
23. Te trees
Te boulders
Tb lizards
Te trees
Te boulders
Tb lizards
Te trees
Te boulders
Tb lizards
Te trees
Te boulders
2450 m.a.s.l.
43 m.a.s.l.
Figure 1: Montane lizards thermoregulate efficiently, despite
living in thermally challenging habitats. A, B, Low-elevation
sites. C, D, High-
elevation sites. In each panel, the distribution of field-measured
lizard body temperatures is shown in black, with the preferred
temperature
range measured in the laboratory indicated with red dashed
lines. The distribution of Te measured on boulders (gray) and
trees (green) are
overlaid. At low elevation (A, B), the temperature range of
boulders, trees, and body temperatures overlap with Tsel,
indicating that lizards do
not need to thermoregulate precisely to maintain temperatures
within their preferred range. In contrast, at high elevation
lizards maintain body
temperatures warmer than their local ambient conditions. The
distribution of substrate temperatures shows that boulders fall
24. within the pre-
ferred range more often than trees (insets). CC, Cordillera
Central; SB, Sierra de Baoruco.
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Behavioral Inertia and Behavioral Drive E000
elevation lizards, although metatarsus length was not found
to differ (table 3).
Discussion
The Bogert Effect Occurs through Shifts in Habitat Use
The extent to which ectotherms, such as lizards, can use be-
havior to buffer the effects of thermal variation has received
renewed attention in light of global climate warming (Huey
et al. 2009; Kearney et al. 2009; Sunday et al. 2014; Buckley
et al. 2015). This topic, first discussed in the middle of the
last century (Bogert 1949; Huey et al. 2003), has become
particularly relevant in studies of tropical forest lizards be-
cause these organisms are currently functioning near their
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upper limits (Huey et al. 2009). Our results suggest that be-
havior can be remarkably effective at buffering lizards from
thermal variation. Specifically, the Bogert effect—physio-
logical buffering through thermoregulation—is occurring
in the high-elevation cybotoids from Hispaniola because
these lizards appear to have adjusted their behavior, rather
than their physiology, to their colder habitats.
The proximate mechanism underlying the Bogert effect
in Anolis armouri and Anolis shrevei involves behavioral
shifts in habitat use. Specifically, we found that lizards at
25. high elevation are found in relatively more open habitats,
where they perched primarily on boulders. Whereas nearly
all temperatures measured on trees were below the set point
range, rocky perches were nearly 57C warmer, providing a
distinct thermoregulatory advantage to montane lizards.
Table 2: Summary data for behavioral variables used to
determine thermoregulatory efficiency
Locality
N
Boulder and arboreal
use (%)
Mean perch height
5 1 SD (cm)
64.011.153 on Octob
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Mean Tb
51 SD (7C)
er 28, 2017 07:17:22
p://www.journals.uc
Thermoregulatory efficiency
E (95% CI)
Cordillera Central (east):
Low
104
11.3, 66.2
26. 42.5 5 41.8
30.3 5 3.6
.368 (.364–.371)
High
58
77.6, 5.2
30.2 5 29.7
29.7 5 2.6
.930 (.929–.931)
Sierra de Baoruco (west):
Low
83
2.3, 69.8
102.3 5 41.9
29.8 5 2.3
.670 (.693–.697)
High
27. 68
88.2, 2.9
26.1 5 29.7
28.9 5 2.4
.929 (.928–.930)
Note: Lizards at high elevation are effective at maintaining
body temperatures (Tb) as high as their lowland counterparts,
despite living in considerably colder
habitats (see also table 1; fig. 1). Consequently, their
thermoregulatory efficiency E (the ability of lizards to maintain
body temperatures within the preferred
range, given its thermal habitat) is nearly perfect (i.e., E is
close to 1). Lizards at high elevation utilize boulders more often
than arboreal perches, although perch
height remains roughly the same across elevation. Full substrate
use given in table A1, available online. Measurement of body
temperature, perch use, and ther-
moregulatory efficiency are described in “Material and
Methods.” CI, confidence interval.
Table 3: Regression results for comparisons of morphological
variables across elevation
West: wild caught
East: wild caught
West: common garden
Coefficient
F2, 54
P
31. 13.05
.336
Lamella no. 2
.001 5 .007
2.84b
.894
.021 5 .019
.579c
.279
. . .
. . .
. . .
Note: ANCOVA for individual morphological traits showing
differences in skull and limb dimensions between lowland and
highland lizards. Montane
lizards have more dorsoventrally flattened skulls and shorter
femora, tibiae, and metatarsi relative to lowland lizards. All
differences persisted in the common
garden experiment, except for metatarsus length. For each
regression, the model coefficient (51 SEM), F value with
degrees of freedom, and significance value
(P) are given. Separate analyses were performed for wild-caught
32. lizards from each mountain chain (east and west) and for
laboratory-reared lizards. For wild-
caught lizards, the dependent variables examined were head
height, head width, femur, tibia, metatarsus, fourth toe, and
lamella number. For laboratory-reared
lizards, the same variables except for lamella number were
included (because no differences were detected in the wild-
caught lizards).
a F2, 53.
b F2, 40.
c F2, 39.
).
E000 The American Naturalist
Behavioral Inertia Meets Behavioral Drive
While rendering evolutionary shifts in physiology unneces-
sary (Muñoz et al. 2014), behavioral thermoregulation oc-
curred through a shift in structural supports, setting the
stage for morphological adaptation (Huey et al. 2003). In
other words, the same behavioral shift that precludes diver-
gence in thermal physiology (behavioral inertia) may have
concurrently driven divergence in morphology (behavioral
drive).
Although Anolis cybotes is commonly known as the
large-headed anole, we found that montane populations
in this group have compressed skulls compared with their
low-elevation counterparts. The adaptive value of this mor-
phology is clear:rock-dwelling lizards—including thesemon-
tane anoles (Hertz and Huey 1981)—often quickly scramble
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2017 07:17:22 AM
33. All use subject to University of Chicago Press Terms and
Conditions (http://www.journals.uchicago.edu/t-and-c).
-
d
s
f
h
e
into crevices and under boulders when threatened (Cooper
et al. 2000; Stapley and Keogh 2004), a behavior facilitated
by a flattened body shape. In contrast, arboreal lizards tend
to flee up and around vegetation (Williams 1983; Cooper
1997; Cooper and Whiting 2007), a strategy unaffected by
head size. Both A. armouri and A. shrevei are also known to
retreat under rocks and logs under inclement weather condi-
tions (Schwartz 1989).
In addition to skull compression, we also found that
montane lizards evolved relatively shorter hind limbs, a re-
sult that echoes findings in Petrosaurus, another rock-
dwelling lizard group (Revell et al. 2007). Relatively short
hind limbs are thought to assist with climbing by main-
taining the lizard’s center of balance close to the surface
(Sinervo and Losos 1991; Losos et al. 1993; Van Damme
et al. 1997; Zaaf and Van Damme 2001). However, further
A B C
D E F
Figure 2: Skull and limb dimensions diverged between high-
and low-elevation populations of lizards, with population means
for size
corrected residuals (51 SEM) shown. Dark blue indicates wild-
caught lizards from high elevation, and light blue indicates
laboratory-reare
lizards. Dark red indicates wild-caught lizards from low
34. elevation, and pink indicates laboratory-reared lizards. Circles
correspond to lizard
from the western mountains (Sierra de Baoruco) and squares to
lizards from the eastern mountains (Cordillera Central). The
utilization o
boulders at higher elevation corresponds to a significant
decrease in head height (A), femur length (B), tibia length (C),
and metatarsus lengt
(D), but no shifts were detected in the fourth toe (E) or in
lamella number (F). All shifts except metatarsus length were
also found in th
common garden experiment. Expanded ANCOVA results are
given in table 3.
Behavioral Inertia and Behavioral Drive E000
study on how montane lizards move on boulders (running,
jumping, climbing, and clinging) is needed to illuminate the
mechanistic relationship between boulder use, locomotor
patterns, and limb length evolution.
Whereas changes in skull and limb morphology accom-
pany the shift to boulders at high elevation, we detected no
differences in toe scalation among populations. This may be
due, in part, to the fact that high- and low-elevation lizards
perched equally high off the ground and within the ex-
pected range for trunk-ground lizards (i.e., !1.5 m above
the ground; table 2). Our findings of morphological diver-
gence in limb and skull dimensions further suggest that
physiological similarity is not due to gene flow precluding
differentiation (e.g., Peterson 1995).
The adaptive radiation of Anolis lizards in the Caribbean
is characterized by two distinct phases: morphological ad-
aptation to structural habitat early in the radiation (i.e.,
35. the evolution of ecomorphs), followed by physiological ad-
aptation within ecomorph clades to different thermal and
hydric environments (reviewed in Williams 1972; Losos
2009). As a result, closely related anoles often exhibit mor-
phological similarity and physiological differentiation (Hertz
et al. 2013). Our results show, however, that this pattern does
not hold in all cases. Rather than diverging physiologically
to occupy different habitats, the cybotoids took an alterna-
tive tack, diverging in thermoregulatory behavior rather than
physiology. The different path of the cybotoids compared
with other anoles, however, is a result of geographic context.
Whereas the cybotoid taxa are allopatric and adapt to broad-
scale environmental differences across elevation (Schwartz
1989; Glor et al. 2003; Wollenberg et al. 2013), the diver-
gence in thermal microhabitat among closely related species
usually occurs among sympatric species that are thought to
partition thermal habitat to minimize interspecific interac-
tions (Ruibal 1961; Rand 1964).
Niche Conservatism, Behavior, and the
Multidimensional Niche
Our central finding is that regulatory behavior—in this case,
thermoregulation—and the evolutionary stasis it promotes
can concomitantly lead to trait divergence, emphasizing the
dual role of behavior in evolution. In light of these con-
clusions, we argue that investigating how behavior shapes
the evolutionary process should involve an “exploration of
the vectors and tensions that link these multidimensional
roles” (Huey et al. 2003, p. 363). Behavioral shifts to preferred
thermal microclimates are a widespread phenomenon in ver-
tebrates (Rand 1964; Hertz 1981; Adolph 1990; Hertz et al.
1993; Harvey and Weatherhead 2010) and invertebrates
(Watt 1968; Kingsolver 1983; Sømme 1989; Merrill et al.
2008; Ashton et al. 2009), setting the stage for both behavioral
inertia and behavioral drive to be at play. For example, mir-
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36. All use subject to University of Chicago Press Term
roring our finding in Hispaniolan anoles, spiny lizards (Sce-
loporus) in the western United States are arboreal at low ele-
vation and ground-dwelling at high elevation; this shift in
habitat use is thought to relate to behavioral thermoreg-
ulation (Adolph 1990). Although the Bogert effect has most
often been examined in the context of thermal physiology, be-
havioral buffering can also affect other aspects of physiology.
For example, Appalachian salamanders distributed across al-
titudinal gradients behaviorally shift habitat use to seek their
preferred hydric microclimates, which sets up the possibility
that buffering behaviors for hydric physiology may also influ-
ence evolution in nonphysiological parameters (Farallo and
Miles 2016).
Especially since Hutchinson’s seminal work (1957), the
multidimensionality of the niche has been recognized (Holt
2009). Less appreciated, however, is that niche dimensions,
although distinct, are not independent of each other. As a
result, shifts in resource use can affect multiple niche axes,
simultaneously impinging on multiple selective pressures
(Levins and Lewontin 1985; Huey et al. 2003). Our findings
thus provide an important perspective on the ongoing de-
bate about niche conservatism. The extent to which species’
niches are labile or conserved is a topic of keen interest in
evolutionary biology (reviewed in Wiens et al. 2010). On
the one hand, the tendency for ecological niches to remain
stable across evolutionary timescales is a common pattern
in nature and has been implicated in a variety of evolution-
ary phenomena, including allopatric speciation and latitu-
dinal diversity patterns (Wiens and Graham 2005; Kozak
and Wiens 2006; Buckley et al. 2010). On the other hand,
rapid shifts in species’ niches have also been detected, leading
to communities of closely related species with little ecologi-
cal overlap (discussed in Losos 2008; Pearman et al. 2008).
Patterns of niche divergence or conservatism are often
37. inferred from macroclimatic data. We argue that behavioral
inertia can make such inferences misleading. For example,
on the basis of macroclimatic data (e.g., climate layers re-
solved to 1 km2; Hijmans et al. 2005), thermal niche overlap
between A. cybotes and the two highland taxa, A. armouri
and A. shrevei, is low (Schoener’s D p 0:396 and 0.135, re-
spectively; see appendix), indicating that the thermal con-
ditions between high and low elevation differ substantially.
However, the thermal niches for these species are consider-
ably less differentiated at the microclimatic scale because of
behavioral thermoregulation (fig. 1). In other words, exam-
ination of macroclimatic data alone would lead to interpre-
tation of niche divergence among taxa, whereas consider-
ation of microclimatic and behavioral data suggests niche
conservatism.
Even when species’ behavior and physiology are taken into
consideration, we emphasize the need for caution about ex-
trapolating evolutionary patterns from a single aspect of a
species’ niche to overall patterns of niche evolution. Given
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E000 The American Naturalist
that different aspects of a species’ niche can evolve in differ-
ent ways, we suggest that the tendency to study niche evolu-
tion along a single axis may often be too narrowly focused.
Indeed, several studies have found that different niche axes
can evolve in different ways, suggesting that niche conserva-
tism and divergence can occur simultaneously (Böhning-
Gaese and Oberrath 1999; Vitt et al. 1999; Lindeman 2000;
Freckleton et al. 2002). This finding may stem, in part, from
the multiple interacting roles of behavior in evolution.
38. More broadly, the connection between behavioral drive
and behavioral inertia offers an opportunity to unify studies
of climate, behavioral ecology, and phenotypic evolution
into a common framework. For example, behavioral shifts
are one of the key ways in which organisms can respond
to climate warming (Williams et al. 2008; Kearney et al.
2009; Huey et al. 2012). While serving to buffer organisms,
a less explored corollary is that such behavioral shifts may
simultaneously expose organisms to distinct selective pres-
sures along different niche axes, in turn impelling evolu-
tion. Multiple potentially contrasting selective pressures
can be operating simultaneously, leading to patterns of both
stasis and divergence, and behavior may be the bridge that
links these outcomes.
Acknowledgments
We thank K. Boronow, E. Cook, M. Landestoy, and I.
Shields for field assistance and the Ministerio de Medio
Ambiente y Recursos Naturales (Dominican Republic) for
permits. We thank J. L. Bronstein, M. Kearney, and two re-
viewers for their thoughtful feedback, which greatly im-
proved the manuscript. A.Algar and Y. Stuart provided feed-
back and support with methods on the manuscript. Funding
was provided by the National Science Foundation (DEB
award 0918975 and a Doctoral Dissertation Improvement
Grant) and a Sigma Xi Grant-In-Aid.
Literature Cited
Abramoff, M. D., P. J. Magalhaes, and S. J. Ram. 2004. Image
pro-
cessing with ImageJ. Biophotonics International 11:36–42.
Adolph, S. C. 1990. Influence of behavioral thermoregulation
on micro-
habitat use by two Sceloporus lizards. Ecology 71:315–327.
39. Angilletta, M. J. 2009. Thermal adaptation: a theoretical and
empir-
ical synthesis. Oxford University Press, Oxford.
Arnold, E. N. 1998a. Cranial kinesis in lizards: variations, uses,
and
origins. Evolutionary Biology 30:323–357.
———. 1998b. Structural niche, limb morphology and
locomotion
in lacertid lizards (Squamata, Lacertidae): a preliminary survey.
Bulletin of the Natural History Museum 6:63–89.
Ashton, S., D. Gutiérrez, and R. J. Wilson. 2009. Effects of
temperature
and elevation on habitat use by a rare mountain butterfly:
implica-
tions for species responses to climate change. Ecological
Entomol-
ogy 34:437–446.
This content downloaded from 130.0
All use subject to University of Chicago Press Term
Bakken, G. S. 1992. Measurement and application of operative
and
standard operative temperatures in ecology. American Zoology
25:933–943.
Bakken, G. S., and M. J. Angilletta. 2014. How to avoid errors
when
quantifying thermal environments. Functional Ecology 28:96–
107.
Bartholomew, G. A. 1964. The roles of physiology and
behaviour in
the maintenance of homeostasis in the desert environment. Sym-
40. posia of the Society for Experimental Biology 18:7–29.
Bateson, P. 1988. The active role of behaviour in evolution.
Pages
191–207 in M.-W. Ho and S. W. Fox, eds. Evolutionary
processes
and metaphors. Wiley, Chichester.
Blom, M. P. K., P. Horner, and C. Moritz. 2016. Convergence
across
a continent: adaptive diversification in a recent radiation of
Aus-
tralian lizards. Proceedings of the Royal Society B
283:20160181.
Bogert, C. M. 1949. Thermoregulation in reptiles, a factor in
evolu-
tion. Evolution 3:195–211.
Böhning-Gaese, K., and R. Oberrath. 1999. Phylogenetic effects
on
morphological, life-history, behavioural and ecological traits of
birds. Evolutionary Ecology Research 1:347–364.
Bonino, M. F., D. L. Moreno Azócar, M. J. Tulli, C. S. Abdala,
M. G.
Perotti, and F. B. Cruz. 2011. Running in cold weather:
morphol-
ogy, thermal biology, and performance in the southernmost
lizard
clade in the world (Liolaemus lineomaculatus section:
Liolaemini:
Iguania). Journal of Experimental Zoology A 315:495–503.
Brandon, R. N. 1988. The levels of selection: a hierarchy of
interactors.
41. Pages 51–71 in H. C. Plotkin, ed. The role of behavior in
evolution.
MIT Press, Cambridge, MA.
Buckley, L. B., T. J. Davies, D. D. Ackerly, N. J. B. Kraft, S. P.
Harrison,
B. L. Anacker, H. V. Cornell, et al. 2010. Phylogeny, niche
conser-
vatism and the latitudinal diversity gradient in mammals. Pro-
ceedings of the Royal Society B 277:2131–2138.
Buckley, L. B., J. C. Ehrenberger, and M. J. Angilletta Jr. 2015.
Thermo-
regulatory behaviour limits local adaptation of thermal niches
and
confers sensitivity to climate change. Functional Ecology
29:1038–1047.
Cartmill, M. 1985. Climbing. Pages 73–99 in M. Hildebrand, D.
M.
Bramble, K. F. Liem, and D. B. Wake, eds. Functional
vertebrate
morphology. Belknap, Cambridge, MA.
Collar, D. C., J. A. Schulte, and J. B. Losos. 2011. Evolution of
ex-
treme body size disparity in monitor lizards (Varanus).
Evolution
65:2664–2680.
Collette, B. B. 1961. Correlations between ecology and
morphology
in anoline lizards from Havana, Cuba and southern Florida. Bul-
letin of the Museum of Comparative Zoology 5:137–162.
Cooper, W. E., Jr. 1997. Escape by a refuging prey, the broad-
42. headed
skink (Eumeces laticeps). Canadian Journal of Zoology 75:943–
947.
Cooper, W. E., Jr., J. H. Van Wyk, P. L. F. N. Mouton, A. M.
Al-
Johany, J. A. Lemos-Espinal, M. A. Paulissen, and M. Flowers.
2000. Lizard antipredator behaviors preventing extraction from
crevices. Herpetologica 2000:394–401.
Cooper, W. E., and M. J. Whiting. 2007. Universal optimization
of
flight initiation distance and habitat-driven variation in escape
tac-
tics in a Namibian lizard assemblage. Ethology 113:661–672.
Cowles, R. B., and C. M. Bogert. 1944. A preliminary study of
the
thermal requirements of desert reptiles. Bulletin of the
American
Museum of Natural History 83:267–296.
Duckworth, R. A. 2009. The role of behavior in evolution: a
search
for mechanism. Evolutionary Ecology 23:513–531.
Farallo, V. R., and D. B. Miles. 2016. The importance of
microhabitat:
a comparison of two microendemic species of Plethodon to the
widespread P. cinereus. Copeia 104:67–77.
64.011.153 on October 28, 2017 07:17:22 AM
s and Conditions (http://www.journals.uchicago.edu/t-and-c).
Behavioral Inertia and Behavioral Drive E000
43. Freckleton, R. P., P. H. Harvey, and M. Pagel. 2002.
Phylogenetic anal-
ysis and comparative data: a test and review of evidence.
American
Naturalist 160:712–726.
Garland, T., Jr., and J. B. Losos. 1994. Ecological morphology
of lo-
comotor performance in squamate reptiles. Pages 240–302 in P.
C.
Wainwright and S. M. Reilly, eds. Ecological morphology:
integra-
tive organismal biology. University of Chicago Press, Chicago.
Glor, R. E., J. J. Kolbe, R. Powell, A. Larson, and J. B. Losos.
2003.
Phylogenetic analysis of ecological and morphological
diversifica-
tion in Hispaniolan trunk-ground anoles (Anolis cybotes group).
Evolution 57:2383–2397.
Glossip, D., and J. B. Losos. 1997. Ecological correlates of
number of
subdigital lamellae in anoles. Herpetologica 53:192–199.
Goodman, B. A. 2007. Divergent morphologies, performance,
and
escape behaviour in two tropical rock-using lizards (Reptilia:
Scincidae). Biological Journal of the Linnean Society 91:85–98.
Goodman, B. A., and J. L. Isaac. 2008. Convergent body
flattening in
a clade of tropical rock-using lizards (Scincidae: Lysominae).
Bio-
logical Journal of the Linnean Society 94:399–411.
44. Goodman, B. A., D. B. Miles, and L. Schwarzkopf. 2008. Life
on the
rocks: habitat use drives morphological and performance evolu-
tion in lizards. Ecology 89:3462–3471.
Harvey, D. S., and P. J. Weatherhead. 2010. Habitat selection as
the
mechanism for thermoregulation in a northern population of
mas-
sasauga rattlesnakes (Sistrurus catenatus). Ecoscience 17:411–
419.
Hertz, P. E. 1974. Thermal passivity of a tropical forest lizard,
Anolis
polylepis. Journal of Herpetology 8:323–327.
———. 1979. Sensitivity to high temperatures in threeWest
Indian grass
anoles (Sauria, Iguanidae), with a review of heat sensitivity in
the ge-
nus Anolis. Comparative Biochemistry and Physiology A
63:217–222.
———. 1981. Adaptation to altitude in two West Indian anoles
(Reptilia: Iguanidae): field thermal biology and physiological
ecol-
ogy. Journal of Zoology 195:25–37.
———. 1992a. Temperature regulation in Puerto Rican Anolis
lizards: a field test using null hypotheses. Ecology 73:1405–
1417.
———. 1992b. Thermal resource partitioning by sympatric
lizards
Anolis cooki and A. cristatellus: a field test using null
hypotheses.
45. Oecologia 90:127–136.
Hertz, P. E., Y. Arima, A. Harrison, R. B. Huey, J. B. Losos,
and R. E.
Glor. 2013. Asynchronous evolution of physiology and
morphol-
ogy in Anolis lizards. Evolution 67:2101–2113.
Hertz, P. E., and R. B. Huey. 1981. Compensation for
altitudinal
changes in the thermal environment by some Anolis lizards on
Hispaniola. Ecology 62:515–521.
Hertz, P. E., R. B. Huey, and R. D. Stevenson. 1993. Evaluating
tem-
perature regulation by field-active ectotherms: the fallacy of the
inappropriate question. American Naturalist 142:796–818.
Hijmans, R. J., S. E. Cameron, J. L. Parra, P. G. Jones, and A.
Jarvis.
2005. Very high resolution interpolated climate surfaces for
global
land areas. International Journal of Climatology 25:1965–1978.
Holt, R. D. 2009. Bringing the Hutchinsonian niche into the
21st cen-
tury: ecological and evolutionary perspectives. Proceedings of
the
National Academy of Sciences of the USA 106:19659–19665.
Huey, R. B. 1982. Temperature, physiology, and the ecology of
rep-
tiles. Pages 25–91 in C. Gans and F. H. Pough, eds. Biology of
the
Reptilia. Academic Press, New York.
46. Huey, R. B., C. A. Deutsch, J. J. Tewksbury, L. J. Vitt, P. E.
Hertz,
H. J. Álvarez Perez, and T. Garland. 2009. Why tropical forest
lizards are vulnerable to climate warming. Proceedings of the
Royal
Society B 276:1939–1948.
This content downloaded from 130.0
All use subject to University of Chicago Press Term
Huey, R. B., P. E. Hertz, and B. Sinervo. 2003. Behavioral drive
versus
behavioral inertia in evolution: a null model approach.
American
Naturalist 161:357–366.
Huey, R. B., M. R. Kearney, A. Krockenberger, J. A. M.
Holtum, M.
Jess, and S. E. Williams. 2012. Predicting organismal
vulnerability
to climate warming: roles of behaviour, physiology, and
adaptation.
Philosophical Transactions of the Royal Society B 367:1665–
1679.
Hutchinson, G. E. 1957. Concluding remarks. Cold Spring
Harbor
Symposium 22:415–427.
Irschick, D. J., and J. B. Losos. 1999. Do lizards avoid habitats
in which
performance is submaximal? the relationship between sprinting
ca-
pabilities and structural habitat use in Caribbean anoles.
American
Naturalist 154:293–305.
Kearney, M., R. Shine, and W. P. Porter. 2009. The potential
47. for be-
havioral thermoregulation to buffer “cold-blooded” animals
against
climate warming. Proceedings of the National Academy of
Sciences
of the USA 106:3835–3840.
Kingsolver, J. G. 1983. Thermoregulation and flight in Colias
but-
terflies: elevational patterns and mechanistic limitations.
Ecology
64:534–545.
Kozak, K. H., and J. J. Wiens. 2006. Does niche conservatism
promote
speciation? a case study in North American salamanders.
Evolution
60:2604–2621.
Labra, A., J. Pienaar, and T. F. Hansen. 2009. Evolution of
thermal
physiology in Liolaemus lizards: adaptation, phylogenetic
inertia,
and niche tracking. American Naturalist 174:204–220.
Lande, R. 1981. Models of speciation by sexual selection on
poly-
genic traits. Proceedings of the National Academy of Sciences
of
the USA 78:3721–3725.
Leal, M., A. K. Knox, and J. B. Losos. 2002. Lack of
convergence in
aquatic Anolis lizards. Evolution 56:785–791.
Levins, R., and R. Lewontin. 1985. The dialectical biologist.
48. Harvard
University Press, Cambridge, MA.
Lewontin, R. C. 1983. The organism as the subject and object of
evo-
lution. Scientia 118:63–82.
Lindeman, P. V. 2000. Resource use of five sympatric turtle
species: ef-
fects of competition, phylogeny, and morphology. Canadian
Journal
of Zoology 78:992–1008.
Losos, J. B. 1990. The evolution of form and function:
morphology
and locomotor performance in West Indian Anolis lizards.
Evolu-
tion 44:1189–1203.
———. 2008. Phylogenetic niche conservatism, phylogenetic
signal
and the relationship between phylogenetic relatedness and
ecolog-
ical similarity among species. Ecology Letters 11:995–1007.
———. 2009. Lizards in an evolutionary tree: ecology and
adaptive
radiation of anoles. University of California Press, Berkeley.
Losos, J. B., and D. J. Irschick. 1996. The effects of perch
diameter on
the escape behavior of Anolis lizards: laboratory based
predictions
and field tests. Animal Behavior 51:593–560.
Losos, J. B., and B. Sinervo. 1989. The effects of morphology
49. and
perch diameter on sprint performance of Anolis lizards. Journal
of Experimental Biology 245:23–30.
Losos, J. B., B. M. Walton, and A. F. Bennett. 1993. Trade-offs
be-
tween sprinting and clinging ability in Kenyan chameleons.
Func-
tional Ecology 7:281–286.
Martin, P. H., T. J. Fahey, and R. E. Sherman. 2011. Vegetation
zo-
nation in a Neotropical montane forest: environment,
disturbance
and ecotones. Biotropica 43:533–543.
Mayr, E. 1963. Animal species and evolution. Belknap,
Cambridge,
MA.
64.011.153 on October 28, 2017 07:17:22 AM
s and Conditions (http://www.journals.uchicago.edu/t-and-c).
E000 The American Naturalist
Merrill, R. M., D. Gutiérrez, O. T. Lewis, J. Gutiérrez, S. B.
Díez, and
R. J. Wilson. 2008. Combined effects of climate and biotic
interac-
tions on the elevational range of a phytophagous insect. Journal
of
Animal Ecology 77:145–155.
Miller, A. H. 1956. Ecological factors that accelerate formation
of
races and species of terrestrial vertebrates. Evolution 10:262–
50. 277.
Muñoz, M. M., K. E. Crandell, S. Campbell-Staton, K.
Fenstermacher,
H. Kim-Frank, P. Van Middlesworth, M. Sasa, J. B. Losos, and
A.
Herrel. 2015. Multiple paths to aquatic specialization in four
species
of Central American Anolis lizards. Journal of Natural History
49:1717—1730.
Muñoz, M. M., G. M. Langham, M. C. Brandley, D. Rosauer, S.
E.
Williams, and C. Moritz. 2016. Basking behavior predicts the
evo-
lution of heat tolerance in Australian rainforest lizards.
Evolution
70:2537–2549.
Muñoz, M. M., M. A. Stimola, A. C. Algar, A. Conover, A.
Rodriguez,
M. A. Landestoy, G. S. Bakken, and J. B. Losos. 2014.
Evolutionary
stasis and lability in thermal physiology in a group of tropical
lizard.
Proceedings of the Royal Society B 281:20132433.
Pearman, P. B., A. Guisan, O. Broennimann, and C. F. Randin.
2008.
Niche dynamics in space and time. Trends in Ecology and
Evolu-
tion 23:149–158.
Peterson, M. A. 1995. Phenological isolation, gene flow, and
develop-
mental differences among low- and high-elevation populations
51. of
Euphilotes enoptes (Leptidoptera: Lycaenidae). Evolution
49:446–455.
Pounds, J. A. 1988. Ecomorphology, locomotion, and
microhabitat
structure: patterns in a tropical mainland Anolis community.
Eco-
logical Monographs 58:299–320.
Rand, A. S. 1964. Ecological distribution in anoline lizards of
Puerto
Rico. Ecology 45:745–752.
Revell, L. J., M. A. Johnson, J. A. Schulte III, J. J. Kolbe, and
J. B. Losos.
2007. A phylogenetic test for adaptive convergence in rock-
dwelling lizards. Evolution 61:2898–2912.
Rodríguez-Serrano, E., C. A. Navas, and F. Bozinovic. 2009.
The com-
parative field body temperature among Liolaemus lizards:
testing
the static and the labile hypotheses. Journal of Thermal Biology
34:306–309.
Ruibal, R. 1961. Thermal relations of five species of tropical
lizards.
Evolution 15:98–111.
Schoener, T. W., and A. Schoener. 1971. Structural habitats of
West
Indian Anolis lizards. II. Puerto Rican uplands. Breviora 35:1–
39.
Schwartz, A. 1989. A review of the cybotoid anoles (Reptilia:
52. Sauria:
Iguanidae) from Hispaniola. Milwaukee Public Museum Contri-
butions in Biology and Geology 78:1–32.
Sinervo, B., and J. B. Losos. 1991. Walking the tight rope: a
comparison
of arboreal sprint performance among populations of Sceloporus
occidentalis. Ecology 72:1225–1237.
Sol, D., D. G. Stirling, and L. Lefebvre. 2005. Behavioral drive
or be-
havioral inhibition in evolution: subspecific diversification in
Hol-
arctic passerines. Evolution 59:2669–2677.
Sømme, L. 1989. Adaptation of terrestrial arthropods to the
alpine
environment. Biological Reviews 64:367–407.
Stapley, J., and J. S. Keogh. 2004. Exploratory antipredator
behaviours
differ between territorial and nonterritorial male lizards. Animal
Behavior 68:841–846.
Stuart, Y. E., T. S. Campbell, P. A. Hoehenlohe, R. G.
Reynolds, L. J.
Revell, and J. B. Losos. 2014. Rapid evolution of a native
species
following invasion by a congener. Science 346:463–465.
This content downloaded from 130.0
All use subject to University of Chicago Press Term
Sunday, J. M., A. E. Bates, A. R. Kearney, R. K. Colwell, N. K.
Dulvy,
J. T. Longino, and R. B. Huey. 2014. Thermal-safety margins
and
the necessity of thermoregulatory behavior across latitude and
53. el-
evation. Proceedings of the National Academy of Sciences of
the
USA 111:5610–5615.
van Berkum, F. H. 1988. Latitudinal patterns of the thermal
sensitiv-
ity of sprint speed in lizards. American Naturalist 132:327–343.
Van Damme, R., P. Aerts, and B. Vanhooydonck. 1997. No
trade-off
between sprinting and climbing in two populations of the lizard
Podarcis hispanica (Reptilia: Lacertidae). Biological Journal of
the
Linnean Society 60:493–503.
Vitt, L. J., J. P. Caldwell, and P. A. Zani. 1997. The role of
habitat
shift in the evolution of lizard morphology: evidence from
tropical
Tropidurus. Proceedings of the National Academy of Sciences
of
the USA 94:3828–3832.
Vitt, L. J., P. A. Zani, and M. C. Espósito. 1999. Historical
ecology of
Amazonian lizards: implications for community ecology. Oikos
87:286–294.
Wake, D. B., G. Roth, and M. H. Wake. 1983. On the problem
of stasis
in organismal evolution. Journal of Theoretical Biology
101:211–
224.
Watt, W. B. 1968. Adaptive significance of pigment
54. polymorphisms
in Colias butterflies. I. Variation in melanin pigments in
relation
to thermoregulation. Evolution 22:437–458.
West-Eberhard, M. J. 1989. Phenotypic plasticity and the
origins of
diversity. Annual Review of Ecology and Systematics 20:249–
279.
Wiens, J. J., D. D. Ackerly, A. P. Allen, B. L. Anacker, L. B.
Buckley,
H. V. Cornell, E. I. Damschen, et al. 2010. Niche conservatism
as
an emerging principle in ecology and conservation biology.
Ecol-
ogy Letters 13:1310–1324.
Wiens, J. J., and C. H. Graham. 2005. Niche conservatism:
integrat-
ing evolution, ecology, and conservation biology. Annual
Review
of Ecology, Evolution, and Systematics 36:519–539.
Williams, E. E. 1972. The origin of faunas. Evolution of lizard
congeners
in a complex island fauna: a trial analysis. Evolutionary
Biology 6:47–
89.
———. 1983. Ecomorphs, faunas, island size, and diverse end
points
in island radiation of Anolis. Pages 326–370 in R. B. Huey, E.
R.
Pianka, and T. W. Schoener, eds. Lizard ecology: studies of a
model
55. organism. Harvard University Press, Cambridge, MA.
Williams, S. E., L. P. Shoo, J. L. Isaac, A. A. Hoffmann, and G.
Langham.
2008. Towards an integrated framework for assessing the
vulnerabil-
ity of species to climate change. PLoS Biology 6:2621–2626.
Wollenberg, K. C., I. J. Wang, R. E. Glor, and J. B. Losos.
2013. Deter-
minism in the diversification of Hispaniolan trunk-ground
anoles
(Anolis cybotes species complex). Evolution 67:3175–3190.
Wyles, J. S., J. G. Kunkel, and A. C. Wilson. 1983. Birds,
behavior,
and anatomical evolution. Proceedings of the National Academy
of Sciences of the USA 80:4394–4397.
Zaaf, A., and R. Van Damme. 2001. Limb proportions in
climbing
and ground-dwelling geckos (Lepidosauria, Gekkonidae): a
phylo-
genetically informed analysis. Zoomorphology 121:45–53.
Associate Editor: Michael Kearney
Editor: Judith L. Bronstein
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s and Conditions (http://www.journals.uchicago.edu/t-and-c).
Discussion question
As an advanced practice nurse assisting physicians in the
diagnosis and treatment of disorders, it is important to not only
understand the impact of disorders on the body, but also the
56. impact of drug treatments on the body. The relationships
between drugs and the body can be described by
pharmacokinetics and pharmacodynamics.
Pharmacokinetics describes what the body does to the drug
through absorption, distribution, metabolism, and excretion,
whereas pharmacodynamics describes what the drug does to the
body.
When selecting drugs and determining dosages for patients, it is
essential to consider individual patient factors that might impact
the patient’s pharmacokinetic and pharmacodynamic processes.
These patient factors include genetics, gender, ethnicity, age,
behavior (i.e., diet, nutrition, smoking, alcohol, illicit drug
abuse), and/or pathophysiological changes due to disease.
For this Discussion, you reflect on a case from your past
clinical experiences and consider how a patient’s
pharmacokinetic and pharmacodynamic processes may alter his
or her response to a drug.
To Prepare
· Review the Resources for this module and consider the
principles of pharmacokinetics and pharmacodynamics.
· Reflect on your experiences, observations, and/or clinical
practices from the last 5 years and think about how
pharmacokinetic and pharmacodynamic factors altered his or
her anticipated response to a drug.
· Consider factors that might have influenced the patient’s
pharmacokinetic and pharmacodynamic processes, such as
genetics (including pharmacogenetics), gender, ethnicity, age,
behavior, and/or possible pathophysiological changes due to
disease.
· Think about a personalized plan of care based on these
influencing factors and patient history in your case study.
QUESTION
Post a description of the patient case from your experiences,
observations, and/or clinical practice from the last 5 years.
Then, describe factors that might have influenced
57. pharmacokinetic and pharmacodynamic processes of the patient
you identified. Finally, explain details of the personalized plan
of care that you would develop based on influencing factors and
patient history in your case. Be specific and provide examples.
MODULE 1
REQUIRED READING:
Rosenthal, L. D., & Burchum, J. R. (2018). Lehne’s
pharmacotherapeutics for advanced practice providers. St.
Louis, MO: Elsevier.
· Chapter 1, “Prescriptive Authority” (pp. 1–3)
· Chapter 2, “Rational Drug Selection and Prescription Writing”
(pp. 5–9)
· Chapter 3, “Promoting Positive Outcomes of Drug Therapy”
(pp. 11–16)
· Chapter 4, “Pharmacokinetics, Pharmacodynamics, and Drug
Interactions” (pp. 17–40)
· Chapter 5, “Adverse Drug Reactions and Medical Errors” (pp.
41–49)
· Chapter 6, “Individual Variation in Drug Response” (pp. 51–
56)
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https://www.deadiversion.usdoj.gov/drugreg/index.html
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