Environmental Biology of Fishes 44 : 2 8 7-304,1995 .

©1995 Kluwer Academic Publishers . Printed in the Netherlands .

The role of ecomorphological studies in the comparative biology of fishes

Stephen F Norton', Joseph J . Luczkovich"2 & Philip J . Motta3

Department of Biology & 2 Institute for Coastal and Marine Resources, East Carolina University, Greenville,
NC 27834, U.S.A.
3
Department of Biology, University of South Florida, Tampa, FL, 33620, USA .

Received 8 .4.1995 Accepted 25 .4.1995

Key words: Performance, Functional morphology, Ecophysiology, Behavioral ecology, Evolutionary ecology,
Fitness, Ontogeny, Realized niche, Fundamental niche, Phenotypic plasticity, Hypothesis testing

Synopsis

The goal of an ecomorphological study is to understand the interactions between the morphology of orga-
nisms and their ecology. Both the morphology and the ecology presented by an organism are directly or
indirectly under the influence of the environmental conditions that the organism experiences and its heritable
composition . The development and interpretation of the central element of ecomorphological studies, the
comparison between patterns of variation of morphological and ecological characters, depends heavily on the
mechanistic framework provided by functional morphological and biomechanical studies. The cause-and-
effect hypotheses derived from this comparison can be tested with performance trials . Ecomorphology forms
an integral part of comparative biology, along with ecophysiology, behavioral ecology, and evolutionary ecol-
ogy. Current issues in ecomorphological research that are addressed in this volume include application of a
more functional approach to the choice of characters, integration of morphological, behavioral, and physio-
logical information to address adaptation, and the expansion of spatial and temporal (ontogenetic and evolu-
tionary) scales of ecomorphological questions . Future directions for ecomorphology include broadening the
knowledge base, further integration of information from other disciplines, examination of the role of envi-
ronmental and genetic factors in producing and maintaining ecological and morphological diversity, and ap-
plication of ecomorphological insights to questions of community structure.

Introduction ecomorphological study consists of a comparison

between patterns of variation in ecological charac-
Ecomorphology is a comparative, often strictly ob- teristics and patterns of variation in morphological
servational, approach to biology. While its antece- characteristics measured among the taxonomic
dents have a long history, its modern integrated units (e .g . genera, life history intervals, allopatric
form is relatively young (Motta et al . 1995b) . Eco- populations) . These comparisons may be strictly
morphological comparisons are pitched at two lev- qualitative (e .g . `Species with morphology A tend
els: (1) among taxonomic units or life history inter- to be found in habitat 1 . . .') or highly quantitative
vals and (2) between their phenotypic characteris- (e .g . `The results of a canonical correlation analysis
tics (e.g . their morphology) and their use of ecolog- showed a correlation between morphological fea-
ical resources (i .e. their realized niche) in a tures X, Y, and Z and ecological features 2 and 4').
particular environment. The core element of an After developing these comparisons, ecomorpho-
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logical studies may involve experimental tests of taxonomic or ecological characteristics may not be
the inferred causal relationships or tracking chang- the best choices . The decision concerning charac-
es in morphological and ecological characters ters in the analysis should be based on the results of
through evolutionary history . functional morphological and biomechanical stud-
There have been a variety of attempts to concep- ies, which may also provide a causal framework for
tualize the interactions between morphology and interpreting the ecomorphological correlations
ecology (e .g . Bock & von Wahlert 1965, Bock 1980, (e .g . Norton & Brainerd 1993) . Development of an
1990, Barel 1983, Balon 1985, Liem 1991,1993, Mot- explicit cause-and-effect hypothesis between the
ta & Kotrschal 1992, Reilly & Wainwright 1994 ; see morphological and ecological characters is critically
also Motta et al. 1995b, Smirnov et al . 1995) . In our important in order to separate spurious correla-
view, the phenotype of an organism and its realized tions or autocorrelations from those with a poten-
niche are both products of complex interactions be- tially causal basis.
tween the environment and factors intrinsic to an Performance tests, conducted either in the lab-
organism (Fig . 1) . These intrinsic factors, which in- oratory or in the field, can be used to test this cause-
clude its unique genotype, epigenetic interactions, and-effect hypothesis. Usually, one presents indi-
and maternal effects, would lead to a defined set of viduals of different morphotypes with the same ec-
phenotypic characteristics if independent of eco- ological challenges and then monitors their per-
logical influences . For many characters, the influen- formance under these conditions (e.g. Moody et al .
ce of the environment may modify the phenotypic 1983, Wainwright 1988, Norton 1991,1995, Hernan-
expression of the intrinsic factors directly (i .e. trig- dez & Motta personal communication) . An alterna-
gering genetically-based phenotypic plasticity - Co- tive strategy is to manipulate morphotypes experi-
nover & Kynard 1981) or indirectly (through use- mentally and then to present them with the ecolog-
induced changes in phenotype under specific envi- ical challenge, but this has been used rarely because
ronmental conditions - Meyer 1987, Wainwright et of the technical difficulties (but see Sinervo & Licht
al . 1991, Wimberger 1991) . The realized niche of a 1991, Sinervo et al . 1992) . One of the assumptions of
species depends on the interactions of the pheno- performance tests is that the ecological challenges
typic characteristics of an individual (defining its being presented mimic conditions that would pro-
potential niche) and the physical and biological duce differences in fitness among the morphotypes
conditions present in the environment . The realized under natural conditions (Arnold 1983, Lauder et
niche can influence environmental conditions al . 1993) . Ideally, performance experiments should
through biogenic changes in the environment (e .g . be designed to manipulate direct measures of fit-
changes in the prey community, alterations of local ness : fecundity and survivorship (e .g. Basolo 1990,
hydrological conditions) . The realized niche can in- Bronmark & Miner 1992, Moller 1992) . Where this
fluence phenotype through use-induced changes . is not possible Arnold (1983) has proposed a two-
The realized niche can also influence intrinsic fac- step process in which the correlation between
tors through its differential effects on fitness morphological variation and performance is deter-
(growth and survival) among genetically-varying mined first and then the relationship between varia-
individuals over many generations (e .g . changes in tion in performance and fitness is measured . The
gill raker numbers due to a competitor - Crowder appropriate performance standards will depend on
1986, see also Robinson & Wilson 1994). the functional relationship under review and have
In ecomorphological studies as in other multiva- included growth rate (e.g . Schluter 1994, 1995), lo-
riate studies, the choices of ecological characteris- comotory rates (e .g . Webb 1986, Jayne & Bennett
tics (i.e. aspects of the realized niche) and morph- 1989, Losos 1990a,b, Nicoletto 1991, Garland & Lo-
ological characteristics (i.e . aspects of the pheno- sos 1994), and capture efficiency (e .g . Moody et al .
type) are critical and should reflect those characters 1983, Wainwright 1988, Norton 1991, 1995) among
that the researcher regards as most important to the others .
functional relationship being examined . Traditional Implicit in the invocation of fitness in perform-


289

Intrinsic factors Envir nment

ance studies is the concept of adaptation, that the
l
correlation between morphological and ecological
characteristics is the product of the natural selec- 11 Morphology.
.-a . Realized niche 1
11
tion among the taxa under examination . This eco- Natural ---- " Consumption
selection
morphological correlation may result either from
direct selection on the phenotypic characteristics of Fig. 1.
Aspects of an organism's morphology (i .e. its phenotype)
taxa to fit the unique physical and biotic character- may be the result of the interaction between the environment, its
istics of a particular site or from selective invasion use of resources, and intrinsic factors (e .g . genotype) . Its mor-
phology and the environment interact to produce its realized
from a broader species pool of those members
niche. The consumption of resources by an individual may alter
whose phenotypic characteristics (evolved else- its environment . Its relative success at turning resources into
where) are appropriate to the new site . In either components of fitness (survival and reproduction) will affect the
case, several authors (e.g . Gould & Lewontin 1979, genotypes of subsequent generations via natural selection .
Brooks & McLennan 1991, Lauder 1992, Lauder et
al . 1993) have argued that to describe a specific from these three interdisciplinary fields, which fo-
morphological feature as having evolved to access cus on the present, is evolutionary ecology which
an ecological resource requires demonstration that attempts to trace the historical patterns in the evo-
these historical transitions in morphology are coin- lution of these phenotypic characteristics and to
cident with ecological transitions . Discrepancies link them to environmental conditions where pos-
may be examples of exaptations (Gould & Vrba sible (e .g . Brooks & McLennan 1991, Nitecki 1990) .
1982), but this is a topic of debate (Reeve & Sher- Clearly a complete understanding of the links be-
man 1993) . Techniques to address the patterns of tween form (or any other phenotypic characteris-
historical transitions in ecology and morphology tic), function, and realized niche requires a syner-
are proliferating quickly (e.g . Pagel & Harvey 1988, gistic attack incorporating ecology, behavior, physi-
Brooks & McLennan 1991, Losos & Miles 1994, ology, morphology, genetics, and evolution . Very
Westneat 1995) . few systems have been studied in such a holistic
way, but several of the most complete examples can
be found in fishes . Block and her coworkers have
Ecomorphology of fishes and comparative biology examined the biological implications of regional
endothermy among scombroid fishes . They have
Ecomorphology forms part of the broader develop- identified the morphological and physiological spe-
ing field of comparative biology . The goal of this cializations that make regional endothermy possi-
growing field is to explore the interactions between ble ; they have demonstrated the behavioral and ec-
the intrinsic characteristics of individuals or taxo- ological consequences of regional endothermy in
nomic units and their environment . Other interdis- the field and have traced its historical development
ciplinary fields that embrace this goal include eco- among scombroid fishes (e.g. Block 1991, Block et
physiology and behavioral ecology. Of these three al . 1993) . Endler, Reznick, and their coworkers
fields, ecomorphology has tended to be more cor- have manipulated the environmental conditions in
relative and observational (see Wainwright & Reil- streams to understand the reproductive biology and
ly 1994), behavioral ecology has been driven more life-history styles of guppies (e.g . Endler 1983,
by optimality models (e.g . Krebs & Davis 1978) and Houde & Endler 1990, Reznick et al . 1990, Rodd &
ecophysiology by laboratory experimentation (e .g . Reznick 1991) . This work has been especially in-
Feder et al . 19 :37) . Regardless of these differences in sightful in documenting the dynamic tension that
emphasis, each field examines the implications of exists between sexual selection and natural selec-
individual variation in a phenotypic character on tion in field populations . Freshwater sticklebacks
the ecological differences between species or other have been the focus of a variety of interdisciplinary
taxonomic units or the implications of ecological studies (see Bell & Foster 1994) . A large portion of
differences on phenotypic characters . Emerging these studies highlight stickleback foraging biology
290

(e .g. Lavin & McPhail 1985, 1986, Schluter & that we have emphasized . Clearly ecomorphology,
McPhail 1992, Schluter 1994,1995) and their morph- especially of fishes, is a dynamic field .
ological defenses to predators (e .g . Reist 1980a, b,
Bell et al . 1985, Reimchen et al . 1985, Baumgartner
1992, Reimchen 1992, Baker et al . 1995) . The forag- (1) How good is the basic fit between an organism's
ing biology of the North American centrarchids has structure and its use of ecological resources when tra-
also received extensive attention (e .g . Werner & ditional taxonomic and ecological variables are
Hall 1979, Mittelbach et al . 1992, Wainwright & used?
Lauder 1992, Norton & Brainerd 1993, Wainwright
& Richard 1995) . The use of traditional morphological and ecological
variables may obscure ecomorphological relation-
ships because of the confounding influence of evo-
Contributions from the ecomorphology of fishes lutionary history or because these variables are not
volume relevant to important functional relationships . In
many ecomorphological studies the initial correla-
It is our view that further progress in ecomorpho- tional analysis is based on morphological characters
logical research will come from the broader context derived from systematic studies and ecological
of comparative biology . This perspective was a ma- characters derived from phylogeny (e .g . prey types
jor consideration as we organized a symposium for by taxa) or from broad habitat categories (e .g . ben-
the 1992 Annual Meeting of the American Society thic versus midwater) . Often clear functional con-
of Ichthyologists and Herpetologists and is reflect- nections between the ecological and morphological
ed in the contributions to this volume . The diversity characters are lacking . Such a `shotgun' approach
of opinions and approaches included here clearly (e.g . Gatz 1979a, b) may produce correlations be-
indicates the vigor of comparative biology, especial- tween morphology and ecology, but separating
ly that subset that focuses primarily on the relation- meaningful correlations from spurious correlations
ships between an organism's morphology, its real- or autocorrelations is difficult . Potential ecomorph-
ized niche, and the environment . The contributors ological relationships may be swamped by phyloge-
include individuals whose primary research focus is netic influences (Norton 1995) . Even after narrow-
the relationship between ecology and morphology ing the choice of morphological characters to those
and others whose primary focus may be described found important by other studies, Motta et al .
as more physiological, biomechanical, or evolution- (1995a) detected little overall ecomorphological
ary. While the individual papers contribute a wealth correlation in a seagrass fish assemblage that spans
of new information and new ideas concerning spe- a large part of the phylogenetic breadth of bony
cific ecomorphological relationships, particular fishes . Some morphological associations did
taxa and communities, and biological systems, we emerge when subsets of the ecological and morph-
have synthesized their more general contributions ological characters were analyzed for a part of the
and conclusions on the ecomorphology of fishes . assemblage (Felley 1985) . Motta et al . (1995a) also
We center our discussion around five critical issues highlighted the morphological characters that most
in current ecomorphological research . Ten years often appear to be influencing ecomorphological
ago these questions would have been quite differ- associations in their study and others . In contrast to
ent, emphasizing the search for broad correlations the findings of Motta et al . in a taxonomically di-
between large numbers of ecological and morpho- verse system, Winemiller et al . (1995) have been
logical variables or for patterns of morphology and successful in using a broad character set to docu-
ecology consistent with the idea of a Hutchinsonian ment clear ecomorphological patterns for a single
niche. Quite likely in another ten years the most family of fishes, the Cichlidae, living in 3 riverine
pressing questions will be different from the five drainages : in central Africa, in Central America,
and in South America . They also document pat-
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terns of diversification and convergence in morph- biomechanical perspective ; specifically he examin-

ological structure and ecological niche among these ed the role of variation in the notochord and its bio-
cichlid faunas. Thus, the `shotgun approach' ap- mechanical properties on the swimming perform-
pears to be more useful in ecomorphological stud- ance of white sturgeon . One result of his study was a
ies when the suite of characters is narrowed to those lack of correlation between swimming performance
that demonstrate some functional relevance and and those aspects of notochord morphology that he
when they are applied with a narrow taxonomic fo- measured (e.g . curvature, angular stiffness, and dia-
cus (see also Douglas & Matthews 1992) . meter), yet the biomechanical properties of the no-
tochord regions were correlated with performance .
His conclusion was that this apparent contradiction
(2) Can we improve our conceptual or statistical fit may be resolved by modeling the notochord of the
between ecology and morphology by choosing non- sturgeon as a pressurized cylinder, rather than a
traditional characters that reflect a clearer under- simple beam, thus requiring the measurement of
standing of the functional potentials of specific struc- different morphological characteristics . Alterna-
tures and the ecological complexities facing fishes? tively, the performance of the swimming apparatus
was due to the emergent properties of the locomo-
The characters chosen in an ecomorphological tor morphology taken as a whole, rather than as
study should be restricted to those for which we separate elements . His research highlights the crit-
have some knowledge of the potential functional ical need to identify relevant morphological fea-
relationship.. The choice of appropriate morpholog- tures as part of both performance tests and broader
ical characters requires a clear understanding of the ecomorphological studies .
basic function of a particular structure, of the func- One important difference in recent ecomorpho-
tional significance of morphological variation, and logical studies is a trend to rely less on purely corre-
of the relationship between function and biological lational approaches and to move towards the gener-
role (Bock & van Wahlert 1965, Bock 1980, Long ation of predictions that can be tested in perform-
1995, Westneat 1995) . Beginning in the 1970's, bio- ance studies . Both Wainwright & Richard (1995)
mechanical and experimental functional morpho- and Norton (1995) use insights derived from func-
logical studies have increased greatly our under- tional morphological studies of fish feeding to high-
standing of the potential ecological role of partic- light a restricted set of characteristics of the oral
ular morphological structures (e .g . Levine & Mac- jaws of the predators that appears to influence the
Nichol 1979., Sibbing 1982, 1988, Weihs & Webb prey composition in fish diets . Both of these studies
1983, Webb 1984, Sanderson et al . 1991, Liem 1993, make explicit, testable predictions of the relation-
and see Motta et al . 1995b) . Kotrschal's (1995) re- ship between morphological variation and ecolog-
view of the current knowledge of the functional ical variation . The biomechanical model of Wainw-
properties of solitary chemosensory cells in gadids right & Richard (1995) assesses the importance of
and other fishes highlights the importance of func- variation in jaw morphology for the kinematics of
tional morphology at the cellular level in interpret- mouth opening versus mouth closing ; this model
ing ecomorphological diversity. The types of stimuli has clear ecological implications that are consistent
to which these diverse cells respond in the laborato- with field diet data from a broad spectrum of fishes .
ry provide critical insights into their potential bio- In Norton (1995), predictions regarding the rela-
logical role in the field and into the design of appro- tionship between mouth size and feeding ecology in
priate performance tests . Similarly, van der Meer et cottid fishes were supported by the results of per-
al . (1995) show that in haplochromine cichlids in- formance studies in which elusive and non-elusive
terspecific differences at the cellular level can be re- prey were presented to various species of cottids
lated to differences in visual sensitivity and resolu- and their capture success was recorded . In another
tion, although not to ecological differences . Long experimental study, Mensinger (1995) addresses the
(1995) approaches ecomorphology from a primarily burglar alarm hypothesis for the evolution of biolu-
292

minescence in marine algae . In these innovative (3) When a structure plays a role in several biological
laboratory experiments he demonstrated the value contexts, does its form represent a compromise
of the concordance between the luminescence pro- among optimal sulutions to these contexts or are
duced by dinoflagellates and the visual system of some ecological roles more important than others in
the midshipman to the foraging ecology of this fish . the evolution of form?
These fishes are capable of using the light flashes
that are produced by dinoflagellates in response to It is unlikely that observed ecomorphological rela-
disturbance by swimming mysids to increase feed- tionships will be perfectly matched to some theoret-
ing efficiency on these mysids . These studies repre- ical optimum . Structures operate not in isolation,
sent a shift in ecomorphological studies away from a but in coordination with other structures and often
purely correlative, post hoc approach to a more pre- in multiple contexts (Barel 1983) . Deviation from
dictive, a priori approach . some hypothetical ecomorphological optimum may
Just as a clear understanding of the functional reflect the constraints that other features impose on
roles of morphological features is critical to a prop- our targeted structure-function-ecology relation-
er ecomorphological analysis, a clear understand- ship . Alternatively, deviation from the hypothetical
ing of the demands placed on an organism by a par- optimum may be due to interactions among the
ticular environment is also critical . A poor eco- multiple functions that a single structure must per-
morphological correlation may be due to improper form; here the observed morphology represents a
categorization or superficial understanding of the compromise among several functions and not the
environmental conditions . In addressing the diffi- optimum solution to any single function .
culties in ecological categorization, Luczkovich et The morphological variation present in allopatric
al . (1995), Norton (1995) and Wainwright & Ri- populations of sticklebacks has proven to be a fer-
chard (1995) have all argued that prey types in the tile area to examine these relationships . Baker et al .
diet of a predator should be assorted by the func- (1995) investigated the potential constraining role
tional challenges that the prey pose to the predator, of a defensive structure (the pelvic girdle complex)
not by their taxonomic affinities (see also Motta on the reproductive ecology of three-spined stickle-
1988) . As an example of underlying ecological com- backs. They had proposed that reductions in the de-
plexities, Martin (1995) demonstrates that the vol- velopment of the pelvic girdle could lead to in-
untary emergence of fishes from the water may be creased clutch volumes of female sticklebacks .
better considered as several distinct ecological However, individuals with a fully developed pelvic
styles in response to very different environmental girdle actually had larger clutch volumes than indi-
conditions . The activities of amphibious marine viduals lacking this defensive structure. They argue
fishes range from very active lifestyles by species for a more holistic examination of the factors driv-
that may forage or court when emerged to those of ing variation in clutch volume . Foster et al . (1995)
other species that show only minimal activity when had predicted that populations of the three-spined
emerged . These different styles have very divergent stickleback with larger mouth sizes (a potential ad-
implications for the kinds of morphological struc- aptation for foraging on benthic prey) would be
tures that would be required to support these activ- more predisposed toward courtship cannibalism of
ities (e.g . locomotion, respiration) . Thus, an alter- eggs, a life-history style, than would populations
native to the `shotgun' approach is to focus on only characterized by smaller mouth sizes . While com-
a few morphological and ecological characters parisons among populations that differ in mouth
which are relevant to important functional chal- size demonstrated no correlation between gape and
lenges in the environment . courtship cannibalism, body size differences within
populations did appear to be linked with courtship
cannibalism . They present several alternative hy-
potheses for this pattern.
Similarly, Cech & Massingill (1995) explore the
 293

potentially conflicting constraints operating on the For many fish species ontogenetic transitions in
pharyngeal apparatus of Sacramento blackfish in their feeding ecology follow similar general trajec-
which the gill bars function not only in support of tories that appear to be correlated with changes in
respiration, but also as the primary prey-capture body size and correlated structures (Norton 1991,
surface for these suspension-feeding fishes . Black- Wainwright & Richard 1995) . Not all fishes follow
fish altered a number of gill ventilation behaviors in this general pattern however ; Luczkovich et al .
response to both hypoxia and feeding. Regional (1995) document contrasting ontogenetic changes
separation of respiratory surfaces and prey capture in the trophic ecology and morphology of two im-
surfaces within the gill apparatus allow this species portant marine fishes. For snook, changes in body
to forage efficiently under both normoxic and hy- size lead to little change in the functional groups in-
poxic conditions . Understanding the potential con- cluded in their diet, but growth by pinfish leads to
straints provided by other structures or other eco- dramatic shifts in functional groups in the diet : from
logical roles should be one of the major focuses of prey types that can be captured by ram-feeding or
ecomorphological research . suction to prey that can only be captured by biting .
The importance of a clear understanding the impli-
cations of relative versus absolute size relationships
(4) How responsive are ecomorphological patterns between predators and their prey have also been
to changes in scale, either temporal (e.g. during the discussed by Wainwright (1988), Galis (1990), and
ontogeny of any individual or over the evolutionary Wainwright & Richard (1995) .
history of a Glade) or spatial (e.g. between different Intraspecific ecomorphological differences are
communities of fishes or different habitats)? not confined to ontogenetic changes alone ; many
studies have documented morphological and eco-
Ecomorphological relationships do change as we logical differences among the individuals of a single
expand either temporal or spatial scales . As a result nominal species (e.g . Turner & Grosse 1980, Crowd-
of their often complex life-histories and pattern of er 1986, Lavin & McPhail 1986, Ehlinger 1990, Mit-
indeterminate growth, fishes provide abundant op- telbach et al . 1992) . The focus may be on allopatric
portunities to investigate ontogenetic changes in populations of a putative species complex (e .g. Fos-
ecomorphological relationships . Individuals of ter & Baker 1995, Baker et al . 1995), on allotopic
many freshwater and marine fishes begin their lives populations of a species (e.g . Baumgartner 1992,
as planktonic embryos and larvae, settle out of the Chapman & Liem 1995), or on sympatric morphs of
water column into specific juvenile or nursery hab- a single species (e .g . Kornfield et al . 1982, Sundland
itats and then gradually move into their adult hab- et al . 1992, Hori 1993). For example, the study by
itats. These different habitats present dramatically Chapman & Liem (1995) clearly documents differ-
different abiotic and biotic challenges, including ences in gill morphology between populations of a
changes in the hydrodynamic regime and habitat small cyprinid that inhabit a hypoxic swamp and
structure and a vastly different array of potential populations that inhabit a well-oxygenated river .
prey, competitors, and predators (e .g . Balon 1986, These intraspecific studies put the spotlight on the
Crawford & Balon 1994, Wainwright & Richard degree of morphological and ecological plasticity
1995) . Separate from these ecological transitions present in a species and may shed insights on the
during growth, individual structures or groups of interaction between genetic, developmental, and
structures may demonstrate non-isometric growth, environmental contributions to the morphological
modifying their mechanical relationships to other phenotype and niche breadth of a species .
structures and altering the functional abilities of Through much of its development, one of the
fishes . Taken together, the often-abrupt ecological central focuses in morphological research has been
changes and smoother phenotypic changes may al- the desire to understand the patterns of historical
ter dramatically the ecomorphological patterns ob- change in morphological features and the evolu-
served in fishes. tionary forces driving these changes (Liem 1991) .
294

For the most part, ecology has been far less con- (5) In a broader context, how does an organism's
cerned with evolutionary history, and yet ecological morphology interact with other phenotypic charac-
interactions among species are subject to evolution- teristics (e.g . behavioral repertoire, physiology) to
ary pressures and are the product of their unique define the fundamental niche and with environmen-
evolutionary histories . One potential outcome of tal conditions to produce the realized niche?
ecomorphological studies is an understanding of
the degree of co-evolution of morphology and ecol- With rare exception, morphological structures re-
ogy. However, morphological and ecological char- quire appropriate behaviors and physiological sup-
acteristics may have completely independent evo- port to function properly and therefore to meet the
lutionary histories . This creates both conceptual challenges presented by the environment . We have
and practical problems for ecomorphological ana- argued earlier that morphology, behavior, and
lyses . On a conceptual level, separating co-evolu- physiology must be considered together to fully un-
tion of morphological and ecological characteristics derstand the abilities of an individual to survive in a
from chance historical co-occurence of these char- specified environment . Any one of these aspects
acteristics is a difficult task (Gould & Vrba 1982, may limit its ability to respond to changes in envi-
Huey & Bennett 1986, Douglas & Matthews 1992) . ronmental conditions . In many cases there are tight
On a practical level, the statistical tests used to es- correlations among morphology, behavior, and
tablish the initial ecomorphological correlation as- physiology to meet a specific extrinsic challenge
sume that each data point (e.g . species) in the mor- (e.g . Block 1991, Martin 1995) . For example, the
phology-ecology correlation matrix represents an structure of the midshipman eye, the physiology of
independent historical event, an unlikely occur- its photoreceptors and photosystems, and its behav-
rence when analyzing closely-related species (see ioral repertoire are coordinated to use biolumine-
Felsenstein 1985, Losos 1990a) . The incorporation scence to improve prey capture success (Mensinger
of explicit phylogenetic information is becoming 1995) . Sacramento blackfish may alter several re-
the new standard in the diverse fields that comprise spiratory behaviors (e.g . ventilation rate) that are
comparative biology, including ecomorphology integrated with structural features (e.g . gape) to en-
(e .g . Wainwright & Reilly 1994) . Methods of ad- able them to survive and even to feed under hypoxic
dressing these problems are developing rapidly conditions (Cech & Massingill 1995) . These types of
(e .g. Brooks & McLennan 1991) . Westneat (1995) interdisciplinary studies highlight the importance
presents several alternative solutions to these evo- and the advantages of integrating ecophysiology,
lutionary problems in ecomorphological studies . behavioral ecology, and ecomorphology.
He illustrates their application by examining the
evolution of the feeding ecology and feeding mor-
phology among cheline wrasses . In his analysis The future of ecomorphological research in fishes
Westneat traces the multiple evolution of suites of
jaw characteristics and dietary patterns in this line- Ecomorphology is a relatively young discipline
age. The value of this evolutionary approach is also (Motta et al . 1995b) and we foresee tremendous
clearly demonstrated by Winemiller et al . (1995) . In gains in a number of different areas. First, we need
this study information from morphology and feed- to add to the knowledge base. We need to apply eco-
ing ecology was incorporated into the existing phy- morphology analyses to additional functional sys-
logeny to identify ecomorphological convergence tems, to broaden the taxonomic coverage, to in-
and to compare the rate of evolutionary change clude a wider range of habitats, and to incorporate
among three independent cichlid faunas . The scope all life history intervals . This comprehensive ap-
of ecomorphological research should be expanded proach will allow us to estimate which ecomorph-
to encompass broader spatial scales, longer tempo- ological interactions are general and which apply
ral scales, and all stages of development . only narrowly to particular taxa, environments, life
intervals, or systems. Second, we need to integrate
 295

the results of ecomorphological studies with those 1992, Luczkovich et al . 1995) . Often linked with spe-
from the other comparative fields : ecophysiology, cific dental features, digestive adaptations, includ-
behavioral ecology and evolutionary ecology. ing the structure and length of the intestinal tract,
Clearly an understanding of phylogenetic relation- have clear correlations with the digestive demands
ships of the taxa under investigation is fast becom- that particular prey present to the predator (e .g . De
ing a new sine qua non in ecomorphological re- Silva et al . 1980, Zihler 1982, Verigina 1991, Sturm-
search . Third, we need to apply the insights gained bauer et al . 1992) . Among visual systems, intra and
from ecomorphology to address the formation and interspecific differences in ocular structures, retinal
maintenance of morphological and ecological di- pigments, and retinal organization are strongly cor-
versity and the development of new species . Finally, related with the spectral environment and habitat
ecomorphological studies can be critical to an un- characteristics (e .g . Levine & MacNicol 1979, Hob-
derstanding of community organization . In partic- son et al . 1981, Herring 1982, Partridge 1989, Hueter
ular, ecomorphological studies can contribute to as- 1990, Mas-Riera 1991, McFarland 1991) .
sessing the relative contributions to community The incompletely known or poorly known sys-
structure by stochastic processes (e .g . chance inva- tems await the development of a better understand-
sion of species from another community) versus de- ing of the functional role of morphological diversity
terministic processes (e.g . interspecific differences and its ecological implications . Often basic descrip-
in ability to invade or persist in a local habitat) . This tions of the environmental conditions (e .g. odor
information can also be used to predict changes in plumes, hydrodynamic regime) have proven to be
community structure after disturbance . difficult. Compared to our understanding of the vi-
sual system, the ecomorphological relationships for
other sensory systems, such as mechanoreception
Additions to the knowledge base and olfaction, are poorly known . For example, there
is tremendous diversity among fishes in the place-
Progress in our understanding of ecomorphological ments and numbers of lateral line and cephalic ca-
diversity among the various functional systems of nals, in degree to which these canals are open or
fishes has been uneven . The relationships between closed, and in the distribution of canal versus super-
patterns of morphological diversity and patterns of ficial neuromasts (e .g . Blaxter 1987, Janssen et al.
ecological diversity are well known in some systems 1987, Coombs et al . 1988, Kalmijn 1989, Webb 1989) .
(e .g . vision, digestion, and foraging), incompletely The functional significance and ecological corre-
known for some systems (e.g . locomotion and elec- lates of this structural diversity is an area of re-
troreception) and poorly known in others (e .g., search only now undergoing explosive growth (e .g .
sound detectiion, chemosensory abilities, and life- Jones & Janssen 1992, Denton & Gray 1993) . The
history characters) . For example, the role of mouth importance of chemoreception of environmental
size in determining diet has been explored by many cues by olfactory organs or through solitary chemo-
studies (e .g . Schmitt & Holbrook 1984, Motta 1988, sensory cells is another fast-developing areas of
Schluter & McPhail 1992, Luczkovich et al . 1995, sensory biology (e .g. Kotrschal et al. 1993, Whitear
Norton 1995, Wainwright & Richard 1995) . Inter- & Moate 1994, Kotrschal 1995) . Progress in under-
specific differences in dentition have clear associ- standing the interactions of morphological diversity
ations with interspecific differences in diet (e .g . and ecological role in these systems will follow the
Lauder 1983, Kotrschal & Goldschmid 1983, Stoner development of a firm understanding of the biome-
& Livingston 1984, Norton 1988, Motta 1989) . Other chanics and functional morphology of these sys-
studies have documented similar results for intra- tems .
specific differences in dentition and diet (e.g . Sage A majority of ecomorphological studies have
& Selander 1975, Turner & Grosse 1980, Kornfield been conducted in temperate freshwater lakes and
et al . 1982, Turner et al . 1983, Liem & Kaufman 1984, streams from Europe and North America and in the
Galis 1990, Wainwright et al . 1991, Mittelbach et al . tropical African Rift lakes (especially the cichlid
296

fauna). Surprisingly, there have been general eco- tions may lead to unexpected threshold effects (Ga-
morphological surveys of only a few tropical reef lis 1990, Luczkovich et al . 1995) . Tremendous poten-
systems (e.g . Davis & Birdsong 1973, Emery 1973, tial exists for additional studies . The transition from
Motta 1988,1989, Bellwood & Choat 1990, Purcell planktonic larva to benthic juvenile is likely to be a
& Bellwood 1993, Wainwright & Richard 1995, fruitful area for in-depth investigation as the pace of
Westneat 1995) . This effort pales in comparison to morphological change is often increased during this
the challenge of understanding the interrelation- abrupt habitat transition.
ships among the diverse, complex faunas present in The majority of ecomorphological studies have
modern reef systems . Many other habitats have re- been conducted on actinopterygian fishes, especial-
ceived only cursory treatments, including pelagic ly those groups either in or related to the Perci-
habitats (Ebeling & Cailliet 1974), meso- and bathy- formes (e .g . Centrarchidae - Keast & Webb 1966,
pelagic environments (Casinos 1978, McLellan Werner 1977, Werner & Hall 1979, Wainwright et al .
1977), the non-cichlid faunas of tropical lakes (Na- 1991, Mittelbach et al . 1992 ; Cichlidae - Hooger-
gelkerke et al . 1994), and the tremendously season- houd et al . 1983, Witte et al . 1990, Winemiller 1991,
al river systems of South America (Winemiller 1991, Hori 1993, Winemiller et al . 1995 ; and the Labridae
Winemiller et al . 1995), Asia (Wikramanayake 1980, - Wainwright 1988, Sanderson 1990, Westneat 1990,
Moyle & Senanayake 1984, Watson & Balon 1984), 1995) . Non-percomorph taxa that have been exam-
and Australia . ined include the salmonids (e .g . Malmquist et al .
Most ecomorphological studies have focused on 1992) and the cyprinids faunas of North America
adult fishes . More studies are needed that examine (e .g . Felley 1984, Douglas & Matthews 1992) and
communities composed of earlier life-history inter- Europe (Kryzhanovsky 1949, Lammens & Hoogen-
vals. Largely unexplored are the ecomorphological boezem 1991), but large open horizons remain. For
relationships that may exist among the planktonic example, the morphologically and ecologically di-
larvae of many fishes . There is clearly tremendous verse catfishes and characins of Central and South
morphological diversity among fish embryos, lar- America should be a very fertile area for ecomorph-
vae and juveniles (Lasker 1981, Balon 1985), but the ological research . Similarly, the diverse clinid and
ecological correlates have not been well-establish- gobiid fishes of temperate and tropical marine wa-
ed. Functional studies and performance tests in- ters are ripe candidates for this approach . While the
volving fish embryos and larvae are just beginning feeding mechanics of bony fishes is well under-
(Drost 1987, Drost et al . 1988, Osse 1990, Coughlin stood, experimental functional morphological stud-
1991) . ies of the feeding of elasmobranchs (Wu 1995, Mot-
Only a few ecomorphological studies have been ta et al . personal communication) is now in progress
conducted that have examined changes during the and will add to our understanding of ecomorpho-
entire ontogeny of a species ; a quick overview of the logical relationships among these fishes.
results of these studies reveals a diversity of re-
sponses in the interactions between morphological
and ecological characteristic during ontogeny, not a Integration with other comparative fields
consensus. Some features (e .g . visual pigments)
demonstrate quick discrete morphological changes, As we have already pointed out, there are tremen-
apparently in response to the changing environ- dous advantages to viewing ecomorphological re-
ment during habitat shifts (Boehlert 1978,1979) . As search within the synergistic framework of compar-
Luczkovich et al . (1995) and Wainwright & Richard ative biology. Morphological structures rarely oper-
(1995) have demonstrated some species may show ate in isolation ; the appropriate behavioral and
gradual changes in their ecomorphological rela- physiological supports are necessary for their prop-
tionships during ontogeny, while other species do er function . For example, interspecific morpholog-
not. Further, the interaction of morphological ical specializations among centrarchids have been
changes during ontogeny and environmental condi- accompanied by modifications of motor patterns
 297

used during the strike (Wainwright & Lauder 1986) gree of concordance between morphological and
that lead to differences in patterns of subambient ecological characteristics can be examined during
pressure generation (Norton & Brainerd 1993) and the ontogeny of a species either throughout its
produce in terspecific differences in diet (Werner range or locally (see above) . In the second, eco-
1977, Werner & Hall 1979, Wainwright & Richard morphological comparisons can be made among al-
1995) . Among some embiotocids, the development lopatric populations of a species especially if these
of winnowing, a specialized prey handling activity, allopatric sites appear to show great differences in
is the result of coordination of behavioral and their abiotic or biotic environments (Endler 1983,
morphological specializations that produce a Lavin & McPhail 1985, Wainwright et al. 1991, Mit-
unique diet in these species (Laur & Ebeling 1983, telbach et al. 1992, Swain 1992a, b, Baker et al . 1995,
Drucker & Jensen 1991) . Ontogenetic changes in Chapman & Liem 1995) . Finally, the presence in
the diets of pinfish are accompanied by morpholog- sympatry of multiple morphs of a single fish species
ical, behavioral, and physiological changes (Stoner has garnered considerable scientific attention (re-
& Livingston 1984, Luczkovich & Stellwag 1993, viewed by Robinson & Wilson 1994) . Intraspecific
Luczkovich et al . 1995) . trophic polymorphisms have been identified in a
These ecological, morphological, behavioral, and variety of taxa including salmonids (Malmquist et
physiological changes need not appear simultane- al . 1992, Sandlund et al . 1992), sticklebacks (Lavin
ously. Tracing the co-evolution of these changes re- & McPhail 1985, 1986, Schluter & McPhail 1992,
quires an understanding of the historical relation- Schluter 1994), cichlids (Sage & Selander 1975,
ships among taxa . This information is critical to Kornfield et al . 1982, Witte et al . 1990, Hori 1993),
identifying morphological and ecological conver- centrarchids (Ehlinger & Wilson 1988), characoids
gence and divergence within a taxon (e.g . Wainw- (Roberts 1974), and goodeids (Turner & Grosse
right & Lauder 1992, Winemiller et al . 1995, West- 1980, Turner et al . 1983) . The predominance of
neat 1995) . In the future, ecomorphological studies freshwater, lacustrine examples, and the lack of ma-
will need to address the phylogenetic history of its rine or riverine examples poses an interesting ques-
taxa as an integral part of the analysis. tion : is there something unique to the lakes that
leads to the development of polymorphisms or does
the abundance of examples from lakes simply re-
The generation of ecological and morphological flect the preponderance of ecomorphological stud-
diversity ies in these lacustrine habitats .
The relative contributions of genetic versus envi-
It is clear from examining the contributions of this ronmental factors in producing the various forms of
volume and of other investigations that there is a phenotypic plasticity remains unclear for many taxa
split in ecomorphological studies between those (see review in Robinson & Wilson 1994) . At one
that focus on intraspecific ecomorphological pat- end of the spectrum, Hori (1993) has demonstrated
terns and those that compare ecomorphological re- that the handedness polymorphism in a scale-eating
lationships between species, either because they are cichlid is clearly under tight genetic control, but the
closely related or because they are members of a frequency of different morphs depends on frequen-
local assemblage. The intrinsic advantage of intra- cy-dependent selection. At the other end, in pump-
specific studies is a closer adherence to the ceteris kinseed sunfishes the development of molariform
paribus assumption central to ecomorphological in- morphs depends on the availability of durophagous
vestigations and freedom from some of the statisti- prey in the environment and the incorporation of
cal difficulties in interspecific comparisons (see these prey into the diet (Wainwright et al. 1991, Mit-
Westneat 1995) . telbach et al . 1992) . The role of intrinsic, environ-
Intraspecific variation in ecomorphological pat- mental, and use-driven factors (see Fig . 1) in pro-
terns can be studied at three scales, ontogenetically, ducing polymorphisms will be an active area for fu-
allopatrically, and sympatrically . In the first, the de- ture research .
298

A major unanswered question concerns the role and needed by fish are to some extent predictable .
of polymorphisms, especially in panmictic sympa- Winemiller (1991) in a survey of five independently-
tric populations, in inhibiting or promoting morph- derived fish assemblages distributed from boreal
ological and ecological diversity . The development, regions to the tropics has identified consistent, con-
fixation and ultimate reproductive isolation of dis- vergent ecomorphological patterns among these as-
tinct morphs due to disruptive selection may pro- semblages . Winemiller et al . (1995) reports similar
mote the development of morphological and eco- results from a comparison of riverine cichlid assem-
logical diversity in a community . However, the pres- blages in Africa, Central America, and South
ence of a polymorphic species may inhibit invasion America . Functional morphological, ecological,
by other taxa because the polymorphic species is oc- and morphological studies have demonstrated con-
cupying a wider ecological niche and thus usurping vergence in the foraging biology of cichlids and cen-
ecological resources . For example, are the four dis- trarchids (e .g . Norton & Brainerd 1993) . Schluter &
tinct morphs of Arctic charr described by Malm- McPhail (1992) present serveral examples of mul-
quist et al . (1992) and Sandland et al . (1992) on their tiple invasions by sticklebacks from the marine en-
way to forming distinct species or do they form a vironment into freshwater lakes that have been ac-
stable metaspecies that excludes other species from companied by consistent, predictable changes in
inhabiting these lakes? The potential role of poly- morphology and ecology . Robinson & Sloan (1994)
morphisms in generating species diversity among in their review of character release and displace-
fishes was discussed extensively in Echelle & Korn- ment in fishes have argued that niches in part are
field (1984) and remains a controversial topic today . properties of the environment into which new fish
species invade or evolve . These studies would argue
that deterministic elements do play an important
Composition of ecological communities role in the structure of these fish communities.
However, the importance of deterministic elements
For over seventy-five years two contrasting views may decline in communities dominated by frequent
have dominated the debate over the organization of disturbance, broad dispersal potential, and high di-
communities (Jackson et al . 1992) . Clements (1916) versity.
argued that communities are discrete assemblages Finally, insights from ecomorphological studies
of species with tight biotic connections; in contrast, may be used to predict the impacts on existing com-
Gleason (1926) argued that communities are munities by biotic or abiotic disturbance, including
chance associations of species which overlap in the invasion of new species . Several studies have
some abiotic requirements, but lack tight intercon- documented dramatic shifts in community compo-
nections . The contentious nature of this debate can sition and the morphology of community members
be seen in arguments over stochastic versus deter- after invasions . Witte et al . (1990) recorded dramat-
ministic patterns of community structure (e.g . ic changes in the abundance of haplochromine ci-
Grossman et al . 1982 versus Rahel et al. 1984 and chlids after the introduction of nile perch as well as
Yant et al . 1984 or Gladfelter & Gladfelter 1978 ver- shifts in several morphological characters . Similar-
sus Sale 1978) and random versus nonrandom as- ly, Crowder (1986) documented ecological and
semblages (e.g . Gilpin & Diamond 1984 versus morphological shifts in the trophic biology of bloa-
Connor & Simberloff 1984) . ter, Coregonus hoyi, that followed dramatic in-
Ecomorphological studies can play a central role creases in the populations of the introduced alewife,
in these debates . Ecomorphological studies not Alosa pseudoharengus, and simultaneous changes
only contribute insights into the role of biotic and in zooplankton resources . All of these changes can
abiotic factors in determining the distributional be interpreted after the fact as a result of ecomorph-
limits of individual species but also can address the ological interactions between the invaders and the
integration of species within communities, especial- existing community. Ecomorphological studies can
ly among competitors. The resources available to also be applied in a more pro-active way to predict
 299

changes in community structure that would result Bell, M .A. & S .A . Foster. (ed) 1994. Evolutionary biology of the
threespine stickleback. Oxford University Press, Oxford . 571
from species introductions or other anthropogenic
PP.
manipulations (e.g . Balon et al . 1986, Sibbing et al . Bellwood, D.R . & J .H. Choat . 1990. A functional analysis of
1994, Smirnov et al . 1995) . For example, Balon et al . grazing in parrotfishes (family Scaridae) : the ecological impli-
(1986) used the results from a broad ecomorpholog- cations . Env. Biol . Fish. 28:189-214.
ical study to predict several changes in European Blaxter, J.H .S. 1987 . Structure and development of the lateral
fish communities that would be produced from the line. Biol . Rev. 62 :471-514.
Block, B.A . 1991 . Evolutionary novelties : how fish have built a
mixture of faunas after the construction of a ship heater out of muscle . Amer . Zool. 31 : 726-742.
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the Danube.. Main, and Rhein rivers . Ecomorpho- lution of endothermy in fish : mapping physiological traits on a
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aptation. Amer. Zool. 20: 217-227 .
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communities and at the forces driving the evolution function complex. Evolution 19 : 269-299 .
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single and double cones in the teleost retina. Science 202 : 309-
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area of research will continue to grow and to con- juvenile Sebastes diploproa: adaptations to a changing photic
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Bronmark, C. & J .G . Miner. 1992 . Predator-induced phenotyp-
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