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Ecomorphology or ecological morphology is the study of the relationship between the ecological role of an individual and its morphological adaptations. The term "morphological" here is in the
anatomical Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
context. Both the morphology and
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps w ...
exhibited by an organism are directly or indirectly influenced by their environment, and ecomorphology aims to identify the differences. Current research places emphasis on linking morphology and
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
by measuring the performance of traits (i.e. sprint speed, bite force, etc.) associated behaviours, and fitness outcomes of the relationships. Current ecomorphological research focuses on a functional approach and application to the science. A broadening of this field welcomes further research in the debate regarding differences between both the ecological and morphological makeup of an
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fung ...
.


Development of ecomorphology

The roots of ecomorphology date back to the late 19th century.Bock, W. J. 1994. Concepts and methods in ecomorphology. Journal of Biosciences 19:403–413.

/ref> Then, description and comparison of morphological form, primarily for use in Bird, avian
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
, was focal point of morphological research. However, during the 1930s and 40s morphology as a field shrank. This was likely due to the emergence of new areas of biological inquiry enabled by new techniques. The 1950s brought about not only a change in the approach of morphological studies, resulting in the development of evolutionary morphology in the form of theoretical questions, and a resurgence of interest in the field.Beer, G. 1954. Archaeopteryx and evolution. The Advancement of Science 11: 160–170

/ref> High-speed photography, High-speed cinematography and x-ray cinematography began to allow for observations of movements of parts while
electromyography Electromyography (EMG) is a technique for evaluating and recording the electrical activity produced by skeletal muscles. EMG is performed using an instrument called an electromyograph to produce a record called an electromyogram. An electromyog ...
allowed for observation of the integration of muscle activities. Together, these methodologies allowed morphologists to better delve into the intricacies of their study. It was then, in the 1950s and 60s, that ecologists began to use morphological measures to study evolutionary and ecological questions. This culminated in Karr and James coining the term "ecomorphology" in 1975.Karr, J.R. and James, F.C. 1975. Eco-morphological configurations and convergent evolution of species and communities; in Ecology and Evolution of Communities (eds). M.L. Cody and J.M. Diamond. Cambridge, Massachusetts: Harvard University Press. 258–291 The following year the links between
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
morphology and
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps w ...
were finally established creating the foundations of modern ecomorphology.Bock, W. 1977. Toward an ecological morphology. Vogelwarte 29: 127–135
/ref>Leisler, B. 1977. Morphological Aspects of Ecological Specializations in Bird Genera. American Zoologist 19(3): 1014–101

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Ecomorphology


Ecomorphology and functional morphology

Functional morphology differs from ecomorphology in that it deals with the features arising from form at varying levels of Biological organisation, organisation.Bock, W. Van Walhert, J. 1965. The role of adaptive mechanisms in the origin of higher levels of Biological organisation, organisation. Systematic Biology 14(4): 272–287

/ref> Ecomorphology, on the other hand, refers to those features which can be shown to derive from the ecology surrounding the species. In other words, functional morphology focuses heavily on the relationship between form and function whereas ecomorphology is interested in the form and the influences from which it arises. Functional morphology studies often investigate relationships between the form of
Skeletal muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
and physical properties such as force generation and joint mobility.Bock, W., G. Lanzavecchia, and R. Valvassori. 1991. Levels of complexity and organismal organization. Selected Symposia and Monographs UZI. Vol 5 181–212

/ref> This means that functional morphology experiments may be done under laboratory conditions whereas ecomorphological experiments may not. Moreover, studies of functional morphology themselves provide insufficient data upon which to make conclusions regarding environmental adaptations of a species. The data provided from these studies can, however, support and enrich the understanding of a species' ecomorphological adaptations. For instance, the relationship between the organization of the jaw lever-arm system, mouth size, and jaw muscle force generation and the feeding behaviour of sunfish has been investigated. Work of this variety lends scientific support to seemingly intuitive concepts. For instance, increases in mouth size correspond to an increase in prey size. However, less obvious trends also exist. The prey-size of fish does not seem to correlate so much to body size as to the characteristics of the feeding apparatus.


Behavioural studies

The work above is just one example of an ecomorphology based behavioural study. Studies of this variety are becoming increasingly important in the field. Behavioural studies interrelate functional and eco-morphology. Features such as Animal locomotion, locomotory ability in foraging birds have been shown to affect dietary preferences by studies of this type.Moermond, T., and J. Denslow. 1983. Fruit choice in neotropical birds: effects of fruit type and accessibility on selectivity. The Journal of Animal Ecology 52(2): 407–420

/ref> Behavioural studies are particularly common in
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, bot ...
and in studying birds.Sibbing, F., L. Nagelkerke, and J. Osse. 1994. Ecomorphology as a tool in fisheries-Identification and ecotyping of Lake Tana Barbs (Barbus-Intermedius COmplex), Ethiopia. Netherlands Journal of Agricultural Science 42:77–85. Royal Netherlands Soc Agr Sc

/ref> Other studies attempt to relate ecomorphological findings with the dietary habits of species. Griffen and Mosblack (2011) investigated differences in diet and consumption rate as a function of Gastrointestinal tract, gut ecomorphology.Griffen, B. D., and H. Mosblack. 2011. Predicting diet and consumption rate differences between and within species using gut ecomorphology. The Journal of animal ecology 80:854–6

/ref> Indeed, gut volume was found to correlate positively to increasing metabolic rate. Ecomorphological studies can often be used to determine to presence of parasites in a given temporospatial context as Parasitism, parasite presence can alter host
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
use.Goodman, B. A., and P. T. J. Johnson. 2011. Ecomorphology and disease: cryptic effects of parasitism on host habitat use, thermoregulation, and predator avoidance. Ecology 92:542–54

/ref> Other current work within ecomorphology focuses on broadening the knowledge base to allow for ecomorphological studies to incorporate a wider range of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s, Taxon, taxa, and systems. Much current work also focuses on the integration of ecomorphology with other comparative fields such as
phylogenetics In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
and ontogenetics to better understand evolutionary morphology.Norton, S. F., J. J. Luczkovich, and P. J. Motta. 1995. The role of ecomorphological studies in the comparative biology of fishes. Environmental Biology of Fishes 44:287–30

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Applications of ecomorphology

An understanding of ecomorphology is necessary when investigating both the origins of and reasons for
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
within a
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. Ecomorphology is fundamental for understanding changes in the
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of a
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
in which subsets occupy different
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s, demonstrate different reproductive techniques, and have various sensory modalities. Studies conducted on
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
with high
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
frequently investigate the extent to which species morphology is influenced by their ecology.
Bony fishes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilage ...
are often used to study ecomorphology due to their long evolutionary history, high
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
, and multi-stage life cycle. Studies on the morphological diversity of African
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted this ...
s conducted by Fryer and Iles were some of the first to demonstrate ecomorphology, . This is largely due to cichlids having great
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
, wide distribution, the ability to occupy various ecological niches, and obvious morphological differences.Fryer, G. T. and Iles, T. D. The cichlid fishes of the great lakes of Africa: their biology and evolution. 1972. Oliver and Boyd, Cornell Universit

/ref> Ecomorphology is also often used to study the paleohabitat of a species and/or its evolutionary morphology.


Paleohabitat determination from ecomorphology

The history of how a species has undergone morphological adaptations to better suit its ecological role can be used to draw conclusions about its paleohabitat. The morphologies of paleo-species found at a location help to make inferences about the previous appearance and properties of that habitat. Research using this approach has been widely conducted using
bovid The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, th ...
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
s due to their large skeletons and extensive species radiation.Scott, R. S., and W. A. Barr. 2014. Ecomorphology and phylogenetic risk: Implications for habitat reconstruction using fossil bovids. Journal of human evolution 73:47–5

/ref> Plummer and Bishop conducted a study using extant African bovids to investigate the animal’s paleoenvironment based on their habitat preference.Plummer, T. W., Bishop, L. C., Hertel, F. 2008. Habitat preference of extant African bovids based on astragalus morphology: operationalizing ecomorphology for palaeoenvironmental reconstruction. Journal of Archaeological Science 35(11): 3016–302

/ref> The strong correlation found between bovid
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
and habitat preference suggests that linking morphology and habitat is
taxon In biology, a taxon ( back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular na ...
dependent. Evidence also suggests that further study of the ecomorphology of previously existing habitats may be useful in determining the phylogenetic risk associated with species living in a specific habitat.


Evolutionary morphology

The study of evolutionary morphology concerns changes in species morphology over time in order to become better suited to their environment. These studies are conducted by comparing the features of species groups to provide a historical narrative of the changes in morphology observed with changes in habitat. A background history of a species features and homology must first be known before a history of evolutionary morphology can be observed. This area of biology serves only to provide a nominal explanation of evolutionary biology, as a more in depth explanation of species history is required to provide a thorough explanation of evolution within a species.


Ecomorphology versus habitat preference

Suggestions have been made that the correlations between species biodiversity and particular environments may not necessarily be due to ecomorphology, but rather a conscious decision made by species to relocate to an ecosystem to which their morphologies are better suited. However, there are currently no studies that provide concrete evidence to support this theory. Studies have been conducted to predict fish habitat preference based on body morphology, but no definitive distinction could be made between correlation and causation of fish habitat preference.Chan, M. D. 2001. Fish ecomorphology: predicting habitat preferences of stream fishes from their body shape. Virginia Polytechnic Institute and State Universit

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References

{{reflistBetz, O. (2006), Ecomorphology: Integration of form, function, and ecology in the analysis of morphological structures, Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie 15, 409-416. Comparative anatomy