Body Size Diversity and Frequency Distributions of Neotropical Cichlid Fishes (Cichliformes: Cichlidae: Cichlinae)

Body size is an important correlate of life history, ecology and distribution of species. Despite this, very little is known about body size evolution in fishes, particularly freshwater fishes of the Neotropics where species and body size diversity are relatively high. Phylogenetic history and body size data were used to explore body size frequency distributions in Neotropical cichlids, a broadly distributed and ecologically diverse group of fishes that is highly representative of body size diversity in Neotropical freshwater fishes. We test for divergence, phylogenetic autocorrelation and among-clade partitioning of body size space. Neotropical cichlids show low phylogenetic autocorrelation and divergence within and among taxonomic levels. Three distinct regions of body size space were identified from body size frequency distributions at various taxonomic levels corresponding to subclades of the most diverse tribe, Geophagini. These regions suggest that lineages may be evolving towards particular size optima that may be tied to specific ecological roles. The diversification of Geophagini appears to constrain the evolution of body size among other Neotropical cichlid lineages; non-Geophagini clades show lower species-richness in body size regions shared with Geophagini. Neotropical cichlid genera show less divergence and extreme body size than expected within and among tribes. Body size divergence among species may instead be present or linked to ecology at the community assembly scale.     

Body Size and Termite Evolution

Termites are a monophyletic lineage within the paraphyletic Blattaria, with xylophagous cockroaches in the genus Cryptocercus as sister group. Given this ancestry, termite divergence involved a substantial leap in body plans, as they are pale, fragile, and miniaturized relative to most cockroaches. Here I suggest that the evolutionary transition to an altricial morphotype in termites is grounded in the economics of utilizing a wood diet, and occurred via a series of sequential steps associated with modifications in social behavior. The chief benefit of an altricial morphotype is suggested to be nitrogen made available by decreasing individual body size and by dispensing with a heavy, melanized cuticle. The primary costs lie in increased vulnerability to environment hazards, including predators, pathogens, and desiccation. Termites tipped the evolutionary scale via cooperative behaviors that mitigate the cost or amplify the benefit of a small, fragile morphotype, and were present in rudimentary form in their cockroach relatives. These include building behavior, cooperative feeding, allogrooming, and most notably, trophallaxis. It was the directed circulation of nitrogenous reserves via trophallactic feeding among units of the superorganism rather than their progressive storage in a large, heavily armored body that was the foundation of termite evolutionary success.     

Evolution, Ecology, and Multimodal Distributions of Body Size

The sizes of organisms are determined by their interactions with their environment and related ecological and evolutionary processes. Recent studies of body size distributions across communities show evidence for multimodality. The multiple modes were originally explained as a consequence of textural discontinuities in habitat structure. Because communities consist of species that are drawn from lineages, body size patterns within lineages will affect those that are expressed in communities. We used a cellular automation model to argue that multimodality in body sizes within lineages can arise from a few fundamental evolutionary mechanisms alone. We tested the hypothesis using body size data for 138 fish genera and found strong support for the idea that evolution structures body size distributions. The results suggest, first, that we should expect the distribution of body sizes within lineages to be multimodal and second, that a coherent theory of community body size distributions will need to combine both evolutionary and ecological perspectives. Received 28 January 2002; accepted 21 March 2002     

A Morphospace for Reef Fishes: Elongation Is the Dominant Axis of Body Shape Evolution

Tropical reef fishes are widely regarded as being perhaps the most morphologically diverse vertebrate assemblage on earth, yet much remains to be discovered about the scope and patterns of this diversity. We created a morphospace of 2,939 species spanning 56 families of tropical Indo-Pacific reef fishes and established the primary axes of body shape variation, the phylogenetic consistency of these patterns, and whether dominant patterns of shape change can be accomplished by diverse underlying changes. Principal component analysis showed a major axis of shape variation that contrasts deep-bodied species with slender, elongate forms. Furthermore, using custom methods to compare the elongation vector (axis that maximizes elongation deformation) and the main vector of shape variation (first principal component) for each family in the morphospace, we showed that two thirds of the families diversify along an axis of body elongation. Finally, a comparative analysis using a principal coordinate analysis based on the angles among first principal component vectors of each family shape showed that families accomplish changes in elongation with a wide range of underlying modifications. Some groups such as Pomacentridae and Lethrinidae undergo decreases in body depth with proportional increases in all body regions, while other families show disproportionate changes in the length of the head (e.g., Labridae), the trunk or caudal region in all combinations (e.g., Pempheridae and Pinguipedidae). In conclusion, we found that evolutionary changes in body shape along an axis of elongation dominates diversification in reef fishes. Changes in shape on this axis are thought to have immediate implications for swimming performance, defense from gape limited predators, suction feeding performance and access to some highly specialized habitats. The morphological modifications that underlie changes in elongation are highly diverse, suggesting a role for a range of developmental processes and functional consequences.     

Latitudinal Variations in Body Size and Species Diversity in Marine Decapod Crustaceans of the Continental Shelf

The number of species of caridean, brachyuran and anomuran decapods increases from high to low latitudes in both hemispheres. This is shown to be due mainly to changes in the abundance of small species. Possible explanations for these variations in abundance are discussed. The other infraorders of decapods are confined to mid and low latitudes.     

The Evolution of Thyroidal Function in Fishes

Although the thyroid gland evolved from the gut, there is noevidence that thyroxine functions as part of the gastro-intestinalendocrine system nor does it have any major function analogousto the control of glucose by the pancreatic islets. The controlof the thyroid evolved from the pituitary control of the gonadsuggesting that an early role of thyroxine was in reproduction.This idea is supported by the presence of cycles of thyroidactivity associated with reproduction in both elasmobranchsand teleosts. In teleosts thyroxine is necessary for gonadalmaturation. The numerous other effects of thyroxine in teleostsmay have evolved from this maturational effect or have beenadded to it during the course of teleost evolution.     

Evolution and Diversification of Antarctic Notothenioid Fishes

Antarctica supported fossil ichthyofaunas during the Devonian,Jurassic, Cretaceous and Eocene/Oligocene. These faunas arenot ancestral to each other, nor are they related to any componentof the modern fauna. About one hundred species of notothenioidsdominate a modern fauna of over 200 species of bottom fishes.This highly endemic perciform suborder is not representedinthe fossil record of Antarctica. Notothenioids may have evolvedin situ on the margins of the Antarctic continent while graduallyadapting to cooling conditions during the Tertiary. Cladisticstudies indicate that notothenioids are a monophyletic group,but a sister group has not been identified among perciform fishes.With relatively few non-notothenioid fishes in Antarctic waters,notothenioids fill ecological roles normally occupied by taxonomicallydiverse fishes in temperate waters. There are six notothenioidfamilies: Bovichtidae, Nototheniidae, Harpagiferidae, Artedidraconidae,Bathydraconidae and Channichthyidae. Aspects of theirbiologyare briefly considered with emphasis on the Nototheniidae, themost speciose family. Evolutionary diversification within thisfamily allows recognition of species which are pelagic, cryopelagic,benthopelagic and benthic.     

Body Size Evolution in Insular Speckled Rattlesnakes (Viperidae: Crotalus mitchellii)

Background

Speckled rattlesnakes (Crotalus mitchellii) inhabit multiple islands off the coast of Baja California, Mexico. Two of the 14 known insular populations have been recognized as subspecies based primarily on body size divergence from putative mainland ancestral populations; however, a survey of body size variation from other islands occupied by these snakes has not been previously reported. We examined body size variation between island and mainland speckled rattlesnakes, and the relationship between body size and various island physical variables among 12 island populations. We also examined relative head size among giant, dwarfed, and mainland speckled rattlesnakes to determine whether allometric differences conformed to predictions of gape size (and indirectly body size) evolving in response to shifts in prey size.

Methodology/Principal Findings

Insular speckled rattlesnakes show considerable variation in body size when compared to mainland source subspecies. In addition to previously known instances of gigantism on Ángel de la Guarda and dwarfism on El Muerto, various degrees of body size decrease have occurred frequently in this taxon, with dwarfed rattlesnakes occurring mostly on small, recently isolated, land-bridge islands. Regression models using the Akaike information criterion (AIC) showed that mean SVL of insular populations was most strongly correlated with island area, suggesting the influence of selection for different body size optima for islands of different size. Allometric differences in head size of giant and dwarf rattlesnakes revealed patterns consistent with shifts to larger and smaller prey, respectively.

Conclusions/Significance

Our data provide the first example of a clear relationship between body size and island area in a squamate reptile species; among vertebrates this pattern has been previously documented in few insular mammals. This finding suggests that selection for body size is influenced by changes in community dynamics that are related to graded differences in area over what are otherwise similar bioclimatic conditions. We hypothesize that in this system shifts to larger prey, episodic saturation and depression of primary prey density, and predator release may have led to insular gigantism, and that shifts to smaller prey and increased reproductive efficiency in the presence of intense intraspecific competition may have led to insular dwarfism.     

The Physiology and Evolution of Urea Transport in Fishes

This review summarizes what is currently known about urea transporters in fishes in the context of their physiology and evolution within the vertebrates. The existence of urea transporters has been investigated in red blood cells and hepatocytes of fish as well as in renal and branchial cells. Little is known about urea transport in red blood cells and hepatocytes, in fact, urea transporters are not believed to be present in the erythrocytes of elasmobranchs nor in teleost fish. What little physiological evidence there is for urea transport across fish hepatocytes is not supported by molecular evidence and could be explained by other transporters. In contrast, early findings on elasmobranch renal urea transporters were the impetus for research in other organisms. Urea transport in both the elasmobranch kidney and gill functions to retain urea within the animal against a massive concentration gradient with the environment. Information on branchial and renal urea transporters in teleost fish is recent in comparison but in teleosts urea transporters appear to function for excretion and not retention as in elasmobranchs. The presence of urea transporters in fish that produce a copious amount of urea, such as elasmobranchs and ureotelic teleosts, is reasonable. However, the existence of urea transporters in ammoniotelic fish is curious and could likely be due to their ability to manufacture urea early in life as a means to avoid ammonia toxicity. It is believed that the facilitated diffusion urea transporter (UT) gene family has undergone major evolutionary changes, likely in association with the role of urea transport in the evolution of terrestriality in the vertebrates.     

Locomotor Patterns in the Evolution of Actinopterygian Fishes

SYNOPSIS. Locomotor adaptations in actinopterygian fishes aredescribed for (a) caudal propulsion, used in cruising and sprintswimming, acceleration, and fast turns and (b) median and pairedfin propulsion used for slow swimming and in precise maneuver.Caudal swimming is subdivided into steady (time independent)and unsteady (time dependent acceleration and turning) locomotion. High power caudal propulsion is the major theme in actinopterygianswimming morphology because of its role in predator evasionand food capture. Non-caudal slow swimming appears to be secondaryand is not exploited before the Acanthopterygii. Optimal morphological requirements for unsteady swimming are(a) large caudal fin and general body area, (b) deep caudalpeduncle, often enhanced by posterior dorsal and anal fins,(c) an anterior stabilizing body mass and\or added mass, (d)flexible body and (e) large ratio of muscle mass to body mass.Optimal morphological requirements for steady swimming are (a)high aspect ratio caudal fin, (b) narrow caudal peduncle, (c)small total caudal area, (d) anterior stabilizing body massand added mass, and (e) a stiff body. Small changes in morphologycan have large effects on performance. Exclusive morphological requirements for steady versus unsteadyswimming are partially overcome using collapsible fins, butcompromises remain necessary. Morphologies favoring unsteadyperformance are a recurring theme in actinopterygian evolution.Successive radiations at chondrostean, halecostome and teleosteanlevels are associated with modifications in the axial and caudalskeleton. Strength of ossified structures probably limited maximum propulsionforces early in actinopterygian evolution, so that specializationsfor fast cruising (carangiform and thunmform modes) followedstructural advances especially in the caudal skeleton. No suchlimits apply to eel-like forms which consequently recur in successiveactinopterygian radiations. Slow swimming using mainly non-caudal propulsion probably firstoccurred among neopterygians in association with reduced andneutral buoyancy. Slow swimming adaptations can add to and extendthe scope of caudal swimming, but specialization is associatedwith reduced caudal swimming performance. Marked exploitationof slow swimming opportunities does not occur prior to the anterodorsallocation of pectoral and pelvic girdles and the vertical rotationof the base of the pectoral fin, as found in the Acanthopterygii.     

Large-Scale Diversity of Slope Fishes: Pattern Inconsistency between Multiple Diversity Indices

Large-scale studies focused on the diversity of continental slope ecosystems are still rare, usually restricted to a limited number of diversity indices and mainly based on the empirical comparison of heterogeneous local data sets. In contrast, we investigate large-scale fish diversity on the basis of multiple diversity indices and using 1454 standardized trawl hauls collected throughout the upper and middle slope of the whole northern Mediterranean Sea (36°3′- 45°7′ N; 5°3′W - 28°E). We have analyzed (1) the empirical relationships between a set of 11 diversity indices in order to assess their degree of complementarity/redundancy and (2) the consistency of spatial patterns exhibited by each of the complementary groups of indices. Regarding species richness, our results contrasted both the traditional view based on the hump-shaped theory for bathymetric pattern and the commonly-admitted hypothesis of a large-scale decreasing trend correlated with a similar gradient of primary production in the Mediterranean Sea. More generally, we found that the components of slope fish diversity we analyzed did not always show a consistent pattern of distribution according either to depth or to spatial areas, suggesting that they are not driven by the same factors. These results, which stress the need to extend the number of indices traditionally considered in diversity monitoring networks, could provide a basis for rethinking not only the methodological approach used in monitoring systems, but also the definition of priority zones for protection. Finally, our results call into question the feasibility of properly investigating large-scale diversity patterns using a widespread approach in ecology, which is based on the compilation of pre-existing heterogeneous and disparate data sets, in particular when focusing on indices that are very sensitive to sampling design standardization, such as species richness.     

Size, Age and Demography of Metamorphosis and Sexual Maturation in Fishes

Understanding what determines the timing of transitions betweenlife-history stages in fishes is crucial to an understandingof their demography. Most theoretical treatments assume thatthese transitions are age dependent, but evidence accumulatingfrom a variety of organisms indicates that size is an importantvariable also. In the starry flounder, Platichthys slellatus(Pallas),the onset of metamorphosis is more closely related to size thanage. In the platyfish, Xiphophorus maculatus (Günther),the onset of sexual maturation depends on both size and age.I compare these results with those reported for other organisms,and outline the expected ecological correlates of age- and sizedependenttransitions between life history stages in general.     

Diet and Diversification in the Evolution of Coral Reef Fishes

The disparity in species richness among evolutionary lineages is one of the oldest and most intriguing issues in evolutionary biology. Although geographical factors have been traditionally thought to promote speciation, recent studies have underscored the importance of ecological interactions as one of the main drivers of diversification. Here, we test if differences in species richness of closely related lineages match predictions based on the concept of density-dependent diversification. As radiation progresses, ecological niche-space would become increasingly saturated, resulting in fewer opportunities for speciation. To assess this hypothesis, we tested whether reef fish niche shifts toward usage of low-quality food resources (i.e. relatively low energy/protein per unit mass), such as algae, detritus, sponges and corals are accompanied by rapid net diversification. Using available molecular information, we reconstructed phylogenies of four major reef fish clades (Acanthuroidei, Chaetodontidae, Labridae and Pomacentridae) to estimate the timing of radiations of their subclades. We found that the evolution of species-rich clades was associated with a switch to low quality food in three of the four clades analyzed, which is consistent with a density-dependent model of diversification. We suggest that ecological opportunity may play an important role in understanding the diversification of reef-fish lineages.     

The Morphology and Evolution of the Ear in Actinopterygian Fishes

SYNOPSIS. In this paper we consider various aspects of the anatomyand ultrastructure of the actinopterygian ear and make a numberof suggestions on the possible adaptive significance of thestructural specializations. The focus of the arguments is basedupon the substantial inter-specific variation in teleost auditorysystems as measured anatomically, behaviorally, and physiologically.It is potentially of considerable significance that the majorpoints of inter-specific variation in the teleost ear are associatedwith the gross morphology and ultrastructure of the otolithicorgan most often implicated in sound detection, the sacculus.Analysis of patterns of sacculus ultrastructure has led to theconclusion that there are, in effect, only about five differentsaccular ultrastructural patterns but that these patterns arebroadly found throughout the teleost fishes. Based upon patternsof inter-specific variation in the sacculus and in other aspectsof the ear and more peripheral auditory structures (e.g., swimbladder),it is argued that adaptations encountered in the teleost auditorysystem cannot be used as reliable taxonomic indicators amongfishes. Rather, it is proposed that the teleost auditory systemis quite maleable in the evolutionary sense, and that interspecificsimilarities in many features of the auditory system reflectconvergent evoluuon, rather than phylogenetic affinities. Theactual selective pressures operating in the evoluuon of thefish auditory system are still essentially unknown. In addition,we cannot be certain that similar ear patterns in differentspecies reflect convergent evolution (or common ancestry), orthat conversely, different ear patterns among species reflectdifferences in auditory function.     

The Evolution of Male and Female Parental Care in Fishes

In this paper we propose an explanation for (a) the predominanceof male care in fishes, and (b) the phylogenies and transitionsthat occur among care states. We also provide a general evolutionarymodel for studying the conditions under which parental careevolves. Our conclusions are as follows: (i) Parental care hasonly one benefit, the increased survivorship of young. It may,however, have three costs: a "mating cost," an "adult survivorshipcost," and a "future fertility cost." (ii) On average, malesand females will derive the same benefit from care. They probablyalso pay the same adult survivorship cost. However, their matingcost and future fertility costs may differ, (iii) A mating costusually applies only to males. However, this cost may be reducedby male territoriality and, in some situations, be entirelyremoved. Under this condition, natural selection on presentreproductive success is equivalent for males and females, (iv)When fecundity accelerates with body size in females, whilemale mating success follows a linear relationship with bodysize, future fertility costs of parental care are greater forfemales than males. Although further tests are needed, a preliminaryanalysis suggests this often may be the case in fishes. Thus,the predominance of male parental care in fishes is not explainedby males deriving greater benefits from care, but by males payingsmaller future costs. Males thus accrue a greater net fitnessadvantage from parental care (see expressions [6] and [12]).(v) The evolution of biparental care from uniparental male caremay occur because male care selects for larger egg sizes andincreased embryo investment by females. This increases the benefitto the female of parental care, (vi) By contrast, uniparentalfemale care may originate from biparental care when males areselected to desert. This occurs when female care creates a matingcost to males. In some cases male desertion may "lock" femalesinto uniparental care. However, in many other cases femalesmay be selected to desert, giving rise to "no care." (vii) Theorigin of uniparental female care from no care is rare in externallyfertilizing fishes. This is because the benefits of care rarelyoutweigh a female's future fertility costs (expression [9]).For internally fertilizing species, however, the benefit ofcare is high whereas the cost is probably low. Most of thesespecies have evolved embryo retention, (viii) When parentalcare begins with male care and moves to biparental care, ouranalysis suggests that care evolution will include cyclicaldynamics. Parental care in some fishes may thus be seen as transitionaland changing through evolutionary time rather than as an evolutionarilystable state. In theory, "no care" may be a phylogeneticallyadvanced state.     

Tradeoffs in the Evolution of Caste and Body Size in the Hyperdiverse Ant Genus Pheidole

The efficient investment of resources is often the route to ecological success, and the adaptability of resource investment may play a critical role in promoting biodiversity. The ants of the “hyperdiverse” genus Pheidole produce two discrete sterile castes, soldiers and minor workers. Within Pheidole, there is tremendous interspecific variation in proportion of soldiers. The causes and correlates of caste ratio variation among species of Pheidole remain enigmatic. Here we test whether a body size threshold model accounts for interspecific variation in caste ratio in Pheidole, such that species with larger body sizes produce relatively fewer soldiers within their colonies. We evaluated the caste ratio of 26 species of Pheidole and found that the body size of workers accounts for interspecific variation in the production of soldiers as we predicted. Twelve species sampled from one forest in Costa Rica yielded the same relationship as found in previously published data from many localities. We conclude that production of soldiers in the most species-rich group of ants is regulated by a body size threshold mechanism, and that the great variation in body size and caste ratio in Pheidole plays a role in niche divergence in this rapidly evolving taxon.     

Genome Size Evaluation in Tetraodontiform Fishes from the Neotropical Region

Smooth pufferfish of the family Tetraodontidae had become pure genomic models because of the remarkable compaction of their genome. This trait seems to be the result of DNA loss following its divergence from the sister family Diodontidae, which possess larger genomes. In this study, flow cytometry was used for estimate the genome size of four pufferfish species from the Neotropical region. Cytogenetic data and confocal microscopy were also used attempting to confirm relationships between DNA content and cytological parameters. The haploid genome size was 0.71?±?0.03 pg for Sphoeroides greeleyi, 0.34?±?0.01 pg for Sphoeroides spengleri, 0.82?±?0.03 pg for Sphoeroides testudineus (all Tetraodontidae), and 1.00?±?0.03 pg for Chilomycterus spinosus (Diodontidae). These differences are not related with ploidy level, because 46 chromosomes are considered basal for both families. The value for S. spengleri represents the smallest vertebrate genome reported to date. Since erythrocyte cell and nuclear sizes are strongly correlated with genome size, the variation in this last is considered under both adaptive and evolutionary perspectives.     

Introgressive Hybridization and the Evolution of Lake-Adapted Catostomid Fishes

Hybridization has been identified as a significant factor in the evolution of plants as groups of interbreeding species retain their phenotypic integrity despite gene exchange among forms. Recent studies have identified similar interactions in animals; however, the role of hybridization in the evolution of animals has been contested. Here we examine patterns of gene flow among four species of catostomid fishes from the Klamath and Rogue rivers using molecular and morphological traits. Catostomus rimiculus from the Rogue and Klamath basins represent a monophyletic group for nuclear and morphological traits; however, the Klamath form shares mtDNA lineages with other Klamath Basin species (C. snyderi, Chasmistes brevirostris, Deltistes luxatus). Within other Klamath Basin taxa, D. luxatus was largely fixed for alternate nuclear alleles relative to C. rimiculus, while Ch. brevirostris and C. snyderi exhibited a mixture of these alleles. Deltistes luxatus was the only Klamath Basin species that exhibited consistent covariation of nuclear and mitochondrial traits and was the primary source of mismatched mtDNA in Ch. brevirostris and C. snyderi, suggesting asymmetrical introgression into the latter species. In Upper Klamath Lake, D. luxatus spawning was more likely to overlap spatially and temporally with C. snyderi and Ch. brevirostris than either of those two with each other. The latter two species could not be distinguished with any molecular markers but were morphologically diagnosable in Upper Klamath Lake, where they were largely spatially and temporally segregated during spawning. We examine parallel evolution and syngameon hypotheses and conclude that observed patterns are most easily explained by introgressive hybridization among Klamath Basin catostomids.