MOOC Innovative methods to assess the distribution of marine ecosystems

MODULE 2
Marine ecosystems characterization

2.1 Introduction to Marine ecosystems characterization

In previous lessons, we have noticed the “surprises” hidden by those ecosystems that
make the transition between terrestrial and marine areas. At the same time, we realize
the potential variability of species, habitats, ranges of distribution and ecological
functions they perform.
Certainly, we are talking about a set of ecosystems that contribute in a very important
way to the so-called “Global Biodiversity”.
Cliffs, dunes, subtidal and intertidal reefs, seagrass meadows, soft bottoms, and salt
marshes make up the biological mosaic of many coastal areas, where the singularities
of each type of ecosystem give way to numerous interactions, whose characterization is
based on common concepts and techniques that are the main object of this course.
Diversity, in its broadest sense, constitutes a good starting point to discover different
ways of characterizing these ecosystems and analyzing their spatial and temporal
distribution patterns.
This is the main goal of the second module.

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2.2 Approaches to biodiversity characterisation

Life,Variability, Biological Diversity, Species Diversity, Ecological Diversity,

Biodiversity…. Are we talking about the same concept?
This is the first important question that we must clarify.
In this lesson, I will try to show, first, how man's perception of the organization of natural
world has evolved over time. This overview will allow us to understand the relationship
between the different terms that we currently use to explain the approaches to what we
now call "biodiversity".
Since Aristoteles’ work Scala Naturae, different cultures, philosophical schools and
disciplines have shown their interest in the distinction and classification of natural entities
and levels of organization recognized in nature. The study of morphology and the
improvement in knowledge about the distribution of animals and plants served to
advance in the recognition of the “Variability of living entities” and the establishment of
systems to group them.
At the same time, different theories arose to justify the relationships of the identified
organisms with their close environment in the process of discovering: 1) the existing
variability of organisms, 2) the functions they play in nature and 3) the applications they
have for humans.
A good example of those preliminary services of plant species is their use as therapeutic
remedies started by Dioscorides, a Greek physician, botanist, and author of De materia
medica, which can be considered the pillar of current pharmacology.
However, the organization of the natural world, as we know it today, emerged in the 18th
century, because of the study of the diversification of living forms and the relationships
of living beings through time. We must refer to the rigorous work of Karl von Linnaeus
(1707-1788), one of the naturalists who laid the foundations for the modern classification
system, establishing for the first time the relationships between living beings. This led to
the proposal of Nature organized as a hierarchical system, initially based on seven
levels: empire, kingdom, class, order, genus, species and variety.
This was the starting point of two disciplines: the Taxonomy, whose main objective was
the classification and naming of organisms, and The Systematics, involved in the
evolutionary relationships of organisms. Both disciplines deal with the distinction,
grouping and classification of life and are the base of what we currently know as
Biological Diversity, or simply Species’ Richness.
Accordingly, Taxonomy and Anatomy were the two branches of Botany and Zoology that
received the most attention in the early development of these sciences. The publication
of the first floras and faunas, including information on the places where the different
species lived, their changing morphologies or their reproduction cycles, allowed a better
understanding of the “Biology of the species”, according to its definition by Lamarck.
Later on, other well-known naturalists formulated interesting hypotheses and discussed
the concepts and factors responsible for the variability, diversification, evolution, or
selection of species. Unfortunately, such an interesting debate is outside the scope of
the introductory lesson on the biodiversity, but may link to the next step in my
presentation.

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At the end of the 18th century and during the first half of the 19th century, the
fundamentals of the Ecology were established.
Scientific expeditions, such as that of Darwin (1831-1836) or those of Humboldt and
Wallace, made possible to expand the knowledge of the distribution of species and
establish the regulation of populations and their evolution, giving a new approach to
Natural History.
In 1859, Darwin's work The Origin of Species appeared and a few years later, Haeckel
(1866) defined the “Ecology”, in his work General Morphology of Organisms with the
environment.
Furthermore, in the second half of the 19th century, E. Forbes (1815-54) was one of the
first naturalist to recognize the existence of particular associations of species because
of the interrelation between biotic and abiotic factors.
The analysis of the taxonomic diversity at the biocenosis and community levels revealed
an important aspect, the inequalities observed between the abundances of the different
species, establishing the basis for the definition of what we nowadays identify as
Species’ Diversity.
This new concept integrates two variables:
1) the species’ richness, a concept equivalent to biological diversity or the number of
different species registered,
2) the evenness, or relative distribution of organisms, normally expressed in terms of
their abundance (e.g. number of individuals, biomass) of the different species observed.
The most important novelty of this type of diversity is the use of expressions based on
Information Theory, whose extensive use through different indexes allows its application
at different levels of organization, such as organisms, populations, biocenosis or the
ecosystem.
At the same time, these indices have also been the subject of numerous debates
regarding the ecological interpretation of the relative abundance of species instead of
the simple richness.
A major concern is that do not incorporate any consideration of species' identity, their
functions within the system, or their exclusivity, scoring similar values for very distinct
samples.
A more recent approach to solve this limitation has led to the development of a new
concept, the Functional Diversity.
New techniques, as the Taxonomic distinctness, that describe community structure and
composition were also developed, using information from the hierarchical taxonomic tree
upon which species ‘identities are based.
As we can see, the term Biodiversity has been absent in the entire scientific debate
around this topic until the celebration of the Convention on Biological Diversity in Rio de
Janeiro (1992). In this meeting, the growing concern about the loss of nature values at a
global level was highlighted and its use was accepted as a synthetic concept that
includes approaches to taxonomy, ecology or biogeography, among other disciplines.

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At that international meeting, Biodiversity became a key issue both from the
environmental point of view and from the perspective of the sustainability of the Planet.
As we may figure out, the term Biodiversity results from a combination of the words
“biological” and “diversity”. It essentially refers to the variety of life on Earth. The CBD
define this term as “the variability among living organisms from all sources including,
inter alia, terrestrial, marine and other aquatic ecosystems and the ecological complexes
of which they are part”. This includes diversity within species, between species and of
ecosystems.
Biodiversity can be understood as a fundamental property of all living systems of being
different from each other, because of unique ecological and evolutionary processes. But,
at the same time, it is a complex concept that go beyond levels of organization, including
ecological, organismal and genetic components, as well as different scales of space and
time.
According to this complexity, 3 main levels of analysis of biodiversity are recognized: 1)
genetic diversity (within species), 2) diversity of species or number of species and 3)
ecological diversity, recorded within communities in which the species live, the
ecosystems in which the communities are integrated and the interactions between these
levels.
This last level represents an equally complex area in which complementary approaches
can be found, such as those related to the functionality of the different species, with the
ecosystem services provided by the different levels of organization or with their
contribution to the natural landscape level.
In the following lessons, some of these approaches are explained in more detail as
essential techniques for the characterization of marine ecosystems.

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2.3 Genetic diversity

Genetic diversity represents the lowest but most important level of biodiversity. It is the
variability in the DNA sequence within and between populations and/or species. There
are four evolutionary forces that drive the genetic diversity among species, populations
and communities: Mutation, gene flow, natural selection and genetic drift.
Mutation is an alteration in the nucleotide sequence in the DNA of an organism. Gene
flow is the introduction of genetic material from one population of a species to another
population. Natural Selection is the process through which populations of living
organisms adapt and change. Genetic drift is the change in the frequency of an existing
gene variant (allele) in a population due to random sampling of organisms.
The effect of genetic drift is amplified and more dangerous in small populations. To easily
explain this important concept I will use as an example the Mediterranean limpet Patella
caerulea. Suppose there is a limpet population made up of 10 individuals. Suppose there
is another smaller limpet population with only 5 individuals. In each population, each
limpet is genetically different (just like each human is genetically distinct). If we walk to
the shore and we randomly collect three limpets from both populations, the loss of the
population’s genes in the genetic pool will be proportionally higher in the smaller
population instead compared to the bigger one. Therefore, large populations are not
greatly affected by the impact of genetic drift. This is a very important concept in terms
of species conservation.
You may now wonder why it is important maintain and preserve genetic diversity.
Because variation within a species or population increases the chance of survival when
conditions change. Species and populations are nowadays affected by several direct
and indirect anthropogenic pressures. The main effects of these pressures are the
reduction of the size of the populations and the increase of their isolation. This can lead
to an increased inbreeding effect. What is inbreeding effect? ... It is when relatives mate
between each other. A progressive increase in inbreeding lead to a decreasing of genetic
variation that consequently reduce the survival and therefore the population effective
size. Population effective size is represented as the n of individuals that can reproduce
in a population. If population effective size decrease, also reproduction effort decreases
and further also population size decreases. Summarizing populations affected by any
kind of pressure can be trapped within this reinforcing feedback loop: the extinction
vortex. The fate of these populations is to go extinct soon.
Man-made structures can cause a loss of genetic variability. That is because, these
structures are often built in areas which are otherwise soft sediment habitats. These
artificial substrata provide an alternative to soft-bottom substrates and promote the
establishment of hard-bottom species. These small and new populations have a low
genetic diversity compared to older and well-establish populations. We call this low
diversity the Founder Effect.
From 2002 to 2004, we sampled Mediterranean limpets - Patella caerulea - all along the
Adriatic Sea. The Adriatic is the northernmost arm of the Mediterranean Sea both in
terms of natural intertidal environment as well as artificial structures for coastal defense
namely breakwaters. We observed that genetic diversity was significantly higher in
natural habitat than in artificial habitat. The expansion of urban structures along the
coasts is altering the genetic diversity of the populations leading to loss of diversity at
regional scales. These changes in the genetic makeup of the populations at the regional

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scale can then have implication for the functioning of coastal marine systems at all spatial
scales.
Severe and sudden reductions in population size caused by mass mortality events linked
to local sea temperature increases during heatwaves, could also lead to a loss of genetic
diversity. We call this effect the Genetic Bottleneck. In a study from 2018 we showed that
the mass mortality of red gorgonian populations at the beginning of 2000 along the
Ligurian coasts of the Mediterranean Sea led to a strong genetic drift effect. This was
observed as a drastic reduction of the red gorgonian effective population size in most of
the populations studied.
Now that we know how important genetic diversity is, how do we can conserve, protect
and increase it? Management and conservation of the resource such as spatial
management of marine protected areas can help with this. Effective management also
requires good understanding of changes in genetic diversity through space and time,
which is achieved through species, population, and community monitoring. Finally,
everyone has a role to play in reducing the impact of their resource use on genetic
diversity by switching or promoting the switch to more sustainable extractive practices.

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2.4 Species diversity

In this lesson we continue talking about diversity, but in this case, we will dive deeper
into the key concept of Species Diversity.
So, what is species diversity? Species diversity is defined as the number of species and
abundance of each of them living in a certain location.
In ecology, the number of species is named as species richness and abundance is the
number of individuals of each species.
Species diversity is all the differences within and between Communities, as well as
between species. It is the most commonly used representation of ecological diversity,
but it is not the only measure as we have seen in the previous lessons and we will see
in the next ones.
How many different species make up life on Earth?
The two central paradigms of marine diversity are that there is a latitudinal gradient of
increasing richness from poles to tropics and that species increases with depth to a
maximum around 2,000 meters and thereafter decreases. There are approximately 1.8
million species classified on Earth and only 16 percent of all named species are marine.
Given the vast size of the ocean, it is impossible to know the exact number of species
that inhabit it and new species are discovered each day.
Scientists estimate that 91 percent of ocean species have yet to be classified, and that
more than 80 percent of the ocean is unmapped, unobserved and unexplored. For
instance, in 2014 approximately fifteen hundred marine species were added to this
growing list of known species.
But, Why species diversity is important? Biodiversity increases productivity and each
species, no matter how small, has an important role to play in the ecosystem related to
the essential connections and dependencies that are established between species.
For example, marine mammals play important ecological roles as predators at most
trophic levels and, as prey for sharks and other larger marine mammals. Imagine, what
would happen if marine mammals became extinct? The structure and function of certain
communities would change completely and the ocean would no longer be able to perform
many of its essential functions.
So, How to measure species diversity? Traditionally, there have been two approaches
to measure it.
1. The first approach constructs mathematical indices broadly known as diversity indices.
Here, we can see examples of 3 diversity indices, commonly used in ecology:
The Shannon-Wiener index (H) symbolizes species diversity in a community, through
the proportion of species, relative to the total number of species. The Simpson Index (D)
measures the degree of concentration when individuals are classified into types. And the
Pielou Index (J´), measures the proportion of the diversity observed in relation to the
maximum diversity expected.
In the year 1960, Whittaker proposed a hierarchical system to partition species diversity
into spatial scales:

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point diversity to define the diversity within a microhabitat.

Alfa (α) diversity within a habitat.
Gamma (γ) diversity within a landscape level, and
Epsilon (ε) diversity at the scale of biogeographical regions.
2. The second approach to measure species diversity involves comparing observed
patterns with theoretical models of species abundance, as shown in the example in the
figure.
There are numerous approaches to measuring species diversity, and every now and
then, it is possible to hear about new approaches, but the ones presented in this lesson
represent some of the most commonly used.

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2.5 Functional diversity

In this activity, we will introduce you to the concept of Functional Diversity.

You have already learned about Biodiversity, which concerns to the variation within each
species, quantified in the communities or ecosystems where they are found.
From that, it is easy to understand that, due to intrinsic characteristics, each species may
perform different activities in the ecosystem, and that different species are able to
perform also similar activities to other species. Each species participates in more than a
process in the ecosystem and each process may have contributions from more than a
single species. Kelps may provide shelter but also to store CO2 and be used as food. If
we go for a terrestrial example, pollination may be performed by birds or insects. And
each of those, may perform other functions in the ecosystem.
With this in mind, we may say that Functional Diversity is a component of biodiversity
that generally concerns to a range of things, different types of processes that organisms
do in ecosystems. An example may come from phytoplankton, which is the source of
energy and organic matter to zooplankton and other organisms in the water column, at
the same time it provides the removal of dissolved or suspended chemical elements,
contributing to its cleaning. In a simple way, Functional Diversity concerns to the different
type of processes, performed by biological diversity, that influence how ecosystems
function.
Functional Diversity, which is a subset of biological diversity, is measured from the traits
having relative importance to one or more aspects of the ecosystem functioning. Traits
are the appropriate functional information attributed to each species that allow them to
perform specific processes in the ecosystem or to show specific populational
characteristics.
For fish fauna is common to consider traits related to the habitat they use, mentioned as
ecological traits, and expressed as estuarine resident, diadromous, or marine seasonal
species, to indicate a few. Or traits characterizing their usual position in the water
column, or the trophic position. Some common traits relate to the way an organism
obtains energy (e.g., autotrophy), or to body size, life span, reproduction, position on
food web, etc. A range is defined for each trait and, depending on established criteria
and on the position on the scale of a specific process, a value is attributed to each
species.
Functional Diversity is usually measured based in two components:
1) functional richness – the number of functional traits found in a considered area;
2) functional evenness – the evenness of abundance distribution in traits found in the
study area.
Under a pressure gradient, functional trait richness decreases with the increase of
pressure. It means that less functions will be present in degraded ecosystems.
Concerning the distribution of taxa through different trait category, it is usual to have
more taxa in each trait when environmental degradation is lower, and less taxa per trait
when degradation is high.
Functional diversity is of ecological importance because, by definition, it is the
component of diversity that influences ecosystem structure and dynamics, stability,
productivity, nutrient balance, and other aspects of ecosystem functioning.
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It is important for the ecosystem to have a robust number of processes supported by

traits of species that are found on it. Also, a process is more susceptible to be preserved
when a higher number of species (and individuals) is able to perform it.
Functional diversity may be seen as a principal component of ecosystem functioning. A
decrease in functional richness and evenness forces ecosystem productivity and stability
also to decrease.
The identification of functional traits prevailing in the ecosystem allows the estimation of
its richness and evenness, which may reveal the functional ability of the ecosystem to
react to pressures.
The impact of human-induced pressures is huge on ecosystems and on organisms
inhabiting there. The habitat loss is a direct consequence of land use, and contributes to
biodiversity decrease. Those effects are mainly visible through the reduction of
taxonomic diversity, which can be measured from species richness.
An ecosystem with high biodiversity, where a specific process is fulfilled by several
species, presents more capacity to deal with a reduction in the number of species.
Redundancy of functional processes, when different species ensure the performance of
existing processes, allows any eliminated species to be replaced on its activity by a new
one without significantly affecting the functioning of the ecosystem.

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2.6 Landscape diversity

In this lesson, I will try explaining the ecological meaning of the landscape and the
concept of landscape diversity.
There are four major components of biological diversity, ranging from genes, species,
ecosystems and landscapes in a gradient of spatial scales. Landscape has been defined
in various ways partly because the word ‘landscape’ means quite different things to
people with different scientific and cultural backgrounds. Nevertheless, few disagree that
landscapes are compositionally diverse, spatially heterogeneous and functionally
multivariate.
Landscape can be defined as: “Spatial pattern that repeats across space in a similar
way, characterized by structure, dynamics and function, resulting from the interaction
between natural and human forces”
Since an ecological point of view, landscapes are defined as ‘spatial mosaics of
interacting biophysical and socioeconomic components on a range of scales’. This
ecological meaning focuses on two issues:
1) First, in mapping the structure, dynamics and functional properties of landscape
mosaics across terrestrial and sea domains (i.e. patches and corridors distributed across
a matrix of the physical environment, and the species that inhabit them, including
humans);
2)·Second, in assessing the relationship between the spatial pattern of these landscape
mosaics and the ecological processes and disturbances driving them on a multitude of
scales and organizational levels.
In this ecological context of diverse, heterogeneous and multifunctional mosaics,
landscape diversity is decomposed in patch, pattern and landscape type diversity.
Patch diversity identifies the number, type, size, shape, perimeter-area ratio,
fragmentation and edge of landscape patches.
Pattern diversity is a measure of the relative arrangement of patch types across
landscape mosaics and identifies their spatial pattern and linkage, connectivity and
neighbourhood effects. Pattern diversity considers not only physical distance between
patches but also the functional or cost connectivity among them. It is closely related to
the concept of landscape heterogeneity, that connects patch and pattern diversity (i.e.
number and proportion of different patch types and their complex spatial arrangement)
and has received increasing attention across all landscape types because of its
implications in management and conservation.
Finally, landscape type diversity relates to the distribution, richness and proportion of
patch types, allowing defining e.g. forestry, agricultural or marine landscapes.
Landscape diversity is driven by two different mechanism: first, the complexity and
diversity of living organisms, abiotic elements and ecological processes and, second,
their interaction with human populations since historical times.
Across all landscapes, nature and human societies are specifically linked and
interdependent, creating dynamic socioecosystems that require understanding
landscape diversity at two levels:

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- First, assessing the diversity of living organisms, abiotic elements and dominant
ecological processes:
- Second, assessing their interaction with human populations that will determine biotic
and abiotic fluxes such as species distributions and vegetation productivity, or water
runoff, erosion control, nutrient cycling and energy flows.
In marine areas, landscape diversity have been only measured in intertidal zones, more
accessible than the many still unexplored sea bottoms across the Earth.
Environmental management and planning need to urgently tackle the sustainable use of
natural resources not only by preserving an array of landscape patches, but also their
ecological function and connectivity. The role of landscape diversity is therefore
paramount for environmental management, landscape planning and biodiversity
conservation.
Resource managers and policy makers need to urgently tackle the sustainable use of
landscapes and their natural resources under a growing pressure of Earth population.
For doing so, it is not only necessary to manage and preserve an array of landscape
patches, but also to preserve landscape diversity, including function and connectivity
patterns, across an array of future scenarios of global change.

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2.7 Biogeographical patterns.

Following the review of possible approaches to assess what, in broad terms, we have
called the “diversity of coastal ecosystems”, in this lesson we will take a step further in
the knowledge of the distribution patterns of the organisms that colonize our coasts and,
consequently, of the distribution of coastal populations, communities and ecosystems.
In the first module of this course, we highlighted the importance of some environmental
factors as responsible for the distribution gradients of the organisms that inhabit coastal
systems.
Thus, we learned that salinity, light, tide or the type of substrate play a determining role
in specific environments, such as estuaries, subtidal zones, intertidal areas or sand
dunes.
We also learned that the geographic distribution of these ecosystems conformed to other
types of patterns dependent, in many cases, on variations in other variables, such as
temperature, or simply on gradients of these same variables at a larger scale.
The most important fact is that, usually, these gradients are beyond our ability to observe
on a local or regional scale. Hence, we must apply other strategies to gather the correct
information to perceive these type of patterns.
At this point, we face a simple misleading question: why do organisms live nowadays
where they live?
Answers to this question must be sought in Biogeography, a discipline that explores the
distribution of species on large scales, how assemblages of such species form different
ecosystems and their geographical limits. This branch of the Geography can provide
different reasons for answering the question we have previously posed.
On the one hand, Analytical biogeography would explain these gradients in terms of
characteristics associated with the biology of the species. For instance, with their life
cycle, or with their reproductive capacity and dispersal means.
On the other hand, Ecological biogeography or macro-ecology would seek answers in
the relations between life forms and their abiotic (physical) and biotic (biological)
environments.
Finally, Historical biogeography would focus on two basic ideas: 1) the dispersal capacity
of species from its theoretical “center of origin” and 2) the importance of geological and
climate processes in the splitting of a population into isolated groups.
For sure that most species’ distributions result from a combination of all these factors.
However, ecological biogeographers will maintain their interest 1) in the effects of
environmental changes in constraining species ranges, 2) in the role of past
environmental changes in shaping species ranges, and 3) in predicting species
distributions under future scenarios of change.
Meanwhile, historical biogeographers will keep their attention in finding centres of origin
and dispersal routes of various groups of organisms, or interpreting biogeographical
history through splitting events or vicariance events.
At this module, we may be interested in both approaches.

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Species and higher taxa tend to form characteristic groupings by regions around the
globe. Biogeography has been of interest to both terrestrial and marine biologists for
more than a century, since early botanists began to match vegetation structures to
regional climate types. A series of life zones were established according to
environmental characteristics, such as temperature, precipitation or radiation.
On land, characteristic animal and plant communities are associated with nine basic
climatic types, known as life zones, biomes or eco-regions, including from the Polar and
Subpolar zones, associated with tundra vegetation, to the Humid Tropical ones, that
support the evergreen tropical rain forest.
Similar advances were not easy in the marine environment where mismatches in
different aspects of biogeographic knowledge occurred, even in the most accessible
systems. Field observations on variations in the distribution of the different populations
and communities in the marine environment face a greater complexity. This is mainly
due to either the relative tiny extent of some of the coastal ecosystem, their poor
accessibility by land or sea or their location in deep underwater zones. This is the case
of the vast majority of coastal and marine systems.
From a practical point of view, the analysis of large-scale distribution patterns in marine
systems depends on experimental designs (techniques, sampling units, replicates,
proxis), introducing a significant degree of uncertainty. In the last decades, important
advances in sampling techniques, remote characterization and modelling of marine
ecosystems have result in a better understanding of biogeographic distributions of
organisms.
Anyway, large-scale patterns in coastal and marine systems may resemble those
recorded on land, mainly due to the importance of large-scale solar radiation and (air
and water) temperature gradients in global distributions of biological populations.
Similarly to terrestrial classifications, several attempts to identify biogeographical regions
in world’s seas have led to the proposal of different climatic zones or marine ecozones.
The three main marine ecozones include:
1) Polar zones, characterized by greenish, cold and low salinity waters covered by ice in
winter.
2) Temperate zones, including a great variability in hydrological conditions.
3) Tropical zones, generally characterized by blue, warm and high salinity waters.
Some authors distinguish between cold and warm temperate zones, and proposed
further subdivisions of these ecozones according to hemisphere, identifying regions
within each ecozone.
In total, up to 23 regions were delimited. The boundaries of each group of
biogeographical regions are primarily recognized by drastic changes in the composition
of the coastal flora and fauna. Furthermore, those boundaries use to follow certain
surface waters isotherms.
Currently, detailed information on average and extreme values of different physic and
chemical variables is available at a global level that allows classifications to be made
based on objective criteria. Concrete examples of the application of these criteria will be
presented throughout the course.

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Each species has a characteristic life history, reproduction rate, dispersal capacity and
respond to a range of environmental conditions, including both its physic and chemical
environment (abiotic conditions) and its living surroundings (biotic interactions).
Accordingly, all species have a geographical range or distribution. But, under
experimental conditions many species can endure environmental conditions far in
excess of the conditions in which their populations naturally are found. There may be
different reasons to explain why species do not distribute over the full range of their
actual physiological tolerance limit. Among these are different processes associated with
historical biogeography, such as processes of speciation, diversification (e.g. adaptive
radiation) or extinction.
In summary, the current distribution of species in coastal ecosystems represents the
synthesis of a set of biological processes associated with the biology of each species,
ecological processes derived from the interactions of organisms with their physical and
biological environment, and biogeographical processes associated with the evolutionary
history of the species. This generates very different distribution patterns, both spatially
and temporally, which is reflected in the existence of cosmopolitan species, distributed
throughout the seas, or endemic species, whose presence is associated with a specific
coastal area (regional sea, island, country). In addition, these patterns may present more
or less continuous or fragmented colonization patterns, which are also the result of the
response of the species to the same type of processes.

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2.8 Vertical zonation of marine species at local scales

In this lesson we will dive deeper into the concept of zonation. So what is zonation?
Zonation is the distribution of plant and animal communities in bands or groupings in a
habitat. We will discuss some of the factors that determine zonation.
The intertidal zone is a particularly interesting place to observe zonation because of the
sharp gradient that exists between the terrestrial land and the sea. Clear zonation
patterns can be seen in all intertidal habitats like saltmarshes, mangroves and rocky
shores.
In the previous lesson about rocky shores, you would’ve learnt that the rocky shore can
be divided into the splash zone, the upper intertidal, the middle intertidal, the lower
intertidal and the subtidal zone. You might recall that the intertidal is a challenging place
for organisms to live in because of the strong gradients in the physical factors that exist
here. Physical factors are also known as abiotic or environmental factors. Can you
remember what these abiotic factors are?
Firstly, the ebb and flow of the tide causes a desiccation gradient, with the upper zones
experiencing higher desiccation. Secondly, the exposed and higher parts of the shore
can experience higher temperature as the rocks are heated by the sun during the day.
Thirdly, the upper zones can experience higher salinity levels during low tide when
seawater evaporates from heat or wind.
In summary, the salinity, temperature and desiccation stress exist on a gradient with
increasing stress at higher elevations. These abiotic factors determine the upper edge
of an intertidal organism’s habitat range. For example, unlike barnacles, mussels cannot
live in the upper intertidal because this zone is too dry and too hot.
To cope with these challenges, different organisms that live here have developed unique
coping mechanisms that allow them to live in specific zones within this habitat. Some of
these strategies are covered in the rocky shores lesson.
If abiotic factors determine the intertidal organism’s upper range limit, what factors
determine its lower limit? To answer this question, we will introduce you to Dr. Joseph
Connell.
In the 1960s, Connell observed that the barnacle community, on the shores of the Isle of
Cumbrae in Scotland, were distributed in distinct bands. The species Chthamalus
stellatus grows at a higher elevation than the larger Semibalanus balanoides. The
scientific understanding at that time, was that species distributions were driven solely by
abiotic factors, which means that Chthamalus doesn’t grow lower because it is sensitive
to being submerged, while the Semibalanus doesn’t grow higher because it is sensitive
to desiccation. But Connell had other ideas, and he designed a series of experiments to
test his hypothesis.
He took rocks covered with Chthamalus from the Chthamalus zone and transplanted
them to the lower Semibalanus zone, and took rocks covered with Semibalanus and
transplanted them to the higher Chthamalus zone. He continued observing the rocks for
about a year, and this is what he found.

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The Semibalanus that were moved to a higher elevation died out over time, indicating
that their upper limit is indeed governed by desiccation. However, the Chthamalus
transplants gave different results. He found that the Chthamalus were slowly completely
replaced by Semibalanus. This happens when a Semibalanus larva settles adjacent to
a Chthamalus. As it grows, it wedges its shell under the Chthamalus shell, eventually
dislodging the Chthamalus. However, on half of the rocks, Connell actively removed any
invading Semibalanus with a needle before they could take hold. And on these rocks,
the Chthamalus continued to survive with no problems.
This experiment showed that while physical factors like desiccation and heat determine
the upper edge of an intertidal organism’s range, its lower limit is usually determined by
interactions with other organisms (biotic factors), in this case, competition. The less
physically stressed an environment is the more crowded it gets, and the organism
experiences more competition for space and resources as well predation pressure.
You can click on this link to read about another famous experiment that shows another
important biotic factor at play, that of predation.

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2.9 Temporal scale in species distribution:Phenology

After analyzing the main sources of spatial variation of coastal ecosystems at the
biogeographic and local scales, we will introduce a new factor of variation in the
distribution of organisms inhabiting coastal ecosystems: the temporal variations.
In this case, we can also recognize different scales, from the seasonal changes that
occur every year to those variations, also cyclical, but related to other global phenomena
of interannual character. Among the latter, we can mention, due to their regularity of
occurrence and interest for many coastal regions, the El Niño or La Niña phenomena.
Although the final result of both types of variation is the adaptation of populations and,
consequently, modifications in presences and abundances of different species of fauna
and flora that make up coastal ecosystems, in this lesson we will focus on the short-term,
seasonal changes.
These represent an important aspect of population’s dynamics regulated by their life
cycles, determining the variation of the ecological, functional and landscape diversity of
coastal communities.
Coastal and marine organisms show a great variety of life cycles, combining, for
instance, haploid and diploid generations. This implies the development of gametes in
independent multicellular forms, which are also haploid, before the formation of a diploid
zygote. This is the case of many macroalgae, in which we find generations of haploid
individuals, called gametophytes, which can be morphologically identical to the diploid
sporophytes, in the case of Gelidium corneum, or completely different, in the case of
kelps species. This set of individuals, generations and phases that make up the biological
cycle of a species can combine microscopic and macroscopic, floating (pelagic) and
sessile (benthic) forms.
In the case of ecosystems structured by "flowering plants", such as dunes, mangroves,
saltmarshes or seagrass meadows, the variability of their cycles translates into the
development of deciduous reproductive bodies in the form of inflorescences. These
structures transform the external appearance, or physiognomy, of their populations and
condition the annual productive cycle and the presence and abundance of other species
of fauna and flora.
The analysis of this type of variability in coastal environments could be the subject of a
complete and exciting course, but beyond the scope of this lesson. At this point, we must
introduce a new perspective of coastal ecosystems. I am referring to their phenology.
Phenology is the discipline related to the study of periodic events in biological life cycles
and how these are influenced by seasonal and interannual variations in climate.
Many phenological responses in coastal environments are triggered by temperature,
while others are more responsive to day length or specific weather and other related
factors that occur in the different climatic regions or ecozones, depending on latitude and
oceanographic conditions of regional seas.
One important singularity of some coastal ecosystems, as dunes, saltmarshes or
intertidal reefs, whose distribution spans the interface between terrestrial and aquatic
spaces, is their dependence on both aerial and aquatic environmental conditions (e.g.
air temperature/water temperature). This is a very important issue for distribution of these

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ecosystems in the actual situation of more frequent extreme events because of climate
change.
Such a continuous change in the ecological structure of the different communities implies
evident changes in their physiognomy. Consequently, we can observe those structural
changes mainly in the landscape of intertidal ecosystems, although seasonal variations
occur similarly in subtidal communities. Coastal ecosystems in temperate regions of both
hemispheres show the greatest temporal variability. An additional aspect is that these
changes affect the functional aspects of the marine ecosystem, as a whole, taking part
of the biogeochemical cycling.
Generally, so-called ‘spring advancement’ is seen in hundreds of plant and animal
species in many world regions, including marine species. Changes in phenology affect
the growing season and, thus, ecosystem functioning and productivity.
Climate warming affects the life cycles of all species. Populations at the northern range
margins of a species’ distribution may benefit from this change, whereas populations at
the southern margins may encounter increasing pressure on their life cycles.
Mild winters and the earlier onset of spring allow for an earlier onset of reproduction and,
in some species, the development of extra generations during the year. However, under
unfavorable autumn conditions, the attempted additional generation can result in high
mortality.
Changes in phenology are having an impact on the ecology of many coastal ecosystems,
with implications for the timing and intensity of different ecological processes.
In the case of a phenological decoupling of species interactions in an ecosystem (e.g.
reduced pressure from predators), certain populations may reach very high abundances
that attain or exceed damage thresholds in managed ecosystems. There is robust
evidence that generalist species with a high adaptive capacity are favored, whereas
specialist species will be affected mostly negatively.
In summary, it must be recognized that the comprehensive characterization of temporal
variability of these ecosystems must take into account the biological cycles of the
structuring species and those of the entire cohort of perennial, seasonal and ephemeral
species that make up them. Somehow, the balance between their biological cycles
determines the physiognomy of each biological community along the year.

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