Trophic specialization influences the rate of environmental niche evolution in damselfishes (Pomacentridae)

The rate of environmental niche evolution describes the capability of species to explore the available environmental space and is known to vary among species owing to lineage-specific factors. Trophic specialization is a main force driving species evolution and is responsible for classical examples of adaptive radiations in fishes. We investigate the effect of trophic specialization on the rate of environmental niche evolution in the damselfish, Pomacentridae, which is an important family of tropical reef fishes. First, phylogenetic niche conservatism is not detected in the family using a standard test of phylogenetic signal, and we demonstrate that the environmental niches of damselfishes that differ in trophic specialization are not equivalent while they still overlap at their mean values. Second, we estimate the relative rates of niche evolution on the phylogenetic tree and show the heterogeneity among rates of environmental niche evolution of the three trophic groups. We suggest that behavioural characteristics related to trophic specialization can constrain the evolution of the environmental niche and lead to conserved niches in specialist lineages. Our results show the extent of influence of several traits on the evolution of the environmental niche and shed new light on the evolution of damselfishes, which is a key lineage in current efforts to conserve biodiversity in coral reefs.     

Modular diversification of the locomotor system in damselfishes (Pomacentridae)

As fish move and interact with their aquatic environment by swimming, small morphological variations of the locomotor system can have profound implications on fitness. Damselfishes (Pomacentridae) have inhabited coral reef ecosystems for more than 50 million years. As such, habitat preferences and behavior could significantly constrain the morphology and evolvability of the locomotor system. To test this hypothesis, we used phylogenetic comparative methods on morphometric, ecological and behavioral data. While body elongation represented the primary source of variation in the locomotor system of damselfishes, results also showed a diverse suite of morphological combinations between extreme morphologies. Results show clear associations between behavior, habitat preferences, and morphology, suggesting ecological constraints on shape diversification of the locomotor system. In addition, results indicate that the three modules of the locomotor system are weakly correlated, resulting in versatile and independent characters. These results suggest that Pomacentridae is shape may result from the interaction between (1) integrated parts of morphological variation that maintain overall swimming ability and (2) relatively independent parts of the morphology that facilitate adaptation and diversification. J. Morphol. 277:603–614, 2016. © 2016 Wiley Periodicals, Inc.     

Diversification of the pectoral fin shape in damselfishes (Perciformes, Pomacentridae) of the Eastern Pacific

Fin shape strongly influences performance of locomotion across all swimming styles. In this study, we focused on the diversity of the pectoral fin morphology in damselfishes of the Eastern Pacific. Underwater observations and a review of literature allowed the characterization of ten behavioral groups. Territorial and non-territorial species were discriminated easily with traditional morphometrics. Five ecomorphological groups were recognized by geometric morphometric analyses. Geometric data segregated the outgroup from the damselfishes and allowed the distinction of mean morphologies from extreme ones within territorial and non-territorial species. Additionally, geometric morphometric data split Abudefduf into two groups: (1) A. troschelii is similar to C. atrilobata and (2) A. concolor and A. declivifrons are close to Stegastes. Solitary territorial species (e.g., Stegastes) show rounded and high fins whereas non-territorial species living in groups (e.g., Chromis) present long and curved pectoral fins. In the range of morphological variation, the morphologies of Microspathodon (elongate with highly curved hydrodynamic trailing edge) and Azurina (long, slender and angular) represent the extreme morphologies within territorial and non-territorial species, respectively. Our study revealed a strong relationship between the pectoral fin shape and the behavioral diversification in damselfishes.     

Feeding patterns in eastern tropical Pacific blennioid fishes (Teleostei: Tripterygiidae,Labrisomidae, Chaenopsidae,Blenniidae)

Summary In terms of species number (47) and numerical abundance, blennioids are the most important primary resident rocky reef fishes in the Gulf of California, Mexico. We present the feeding patterns of the 34 most abundant species of blennioid fishes, 8 of which are Gulf endemics. A total of 2,144 specimens were sampled at 51 anaesthetic stations in 9 areas throughout the Gulf. Four feeding guilds were distinguished: 1) The majority (29 of 34 species) are microcarnivores exhibiting a number of different feeding strategies (ambush and stalking predators, active foragers, pickers, etc.). The more important prey categories were mobile invertebrates, and to some extent also sedentary fauna. Algae were of no importance for most of the latter species. 2) Hypsoblennius brevipinnis and H. gentilis are two omnivorous species, browsing mainly on sessile items including 52% and 13% (Vol.) algae in their diets. 3) Entomacrodus chiostictus and Ophioblennius steindachneri are herbivores, grazing on fine algae. 4) Plagiotremus azaleus specializes in cropping mucus and scales from the body surface of other fishes.Crustaceans account for 58.6% of the total volume of prey items in the 34 species investigated. Benthic amphipods were most important and made up 26% of the total volume of all prey items.Cluster analysis of percentage volumetric data using Squared Euclidian Distance and Horn's Index of Overlap produced distinct subgroups which coarsely reflected taxonomic grouping.The species are separated either by their geographic ranges, habitat and microhabitat preferences, feeding, or a combination thereof. Only rarely do sympatric species significantly overlap in diet.Trophic diversity as measured by the Shannon-index provides a tool for distinguishing: 1) specialists (6 species) from 2) low diversity feeders (18 species) and 3) high diversity generalists (10 species). Two different types of specialists can be distinguished: those which feed on the same items as the generalists but utilize only a very restricted prey spectrum (Stathmonotus sinuscalifornici and the chaenopsids Chaenopsis alepidota and Emblemaria hypacanthus). A second group of specialists (Entomacrodus chiostictus and Ophioblennius steindachneri as well as Plagiotremus azaleus) feed on items not utilized by any of the generalists.There is some evidence that high diversity generalists are numerically more abundant than the other trophic groups.In the labrisomids and blenniids a phylogenetic trend from microcarnivory towards feeding on sessile items appears to be expressed.     

Habitat use and behavioural ecology of the juveniles of two sympatric damselfishes (Actinopterygii: Pomacentridae) in the south-western Atlantic Ocean

Spatial distribution, microhabitat use and territorial and feeding behaviours were compared between the juveniles of two sympatric territorial damselfishes Stegastes variabilis and Stegastes fuscus on a small tropical reef in the south-western Atlantic Ocean. Juvenile S. variabilis were most abundant at sites subject to stronger hydrodynamics and with mixed benthic cover, whereas juvenile S. fuscus were most abundant at sheltered sites with dense turf algae cover. No differences regarding feeding habits were detected, with both species preferentially feeding on turf algae. Also, despite similarities in territory area and agonistic encounter rates, the identity and proportion of intruders involved in agonistic interactions differed significantly between species. These interdependent traits suggest strong asymmetric competition, with juvenile S. fuscus dominating high-quality sites and evicting juvenile S. variabilis to low quality, marginal areas of the reef.     

Evolution of ontogenetic allometry shaping giant species: a case study from the damselfish genus Dascyllus (Pomacentridae)

The evolution of body size, the paired phenomena of giantism and dwarfism, has long been studied by biologists and paleontologists. However, detailed investigations devoted to the study of the evolution of ontogenetic patterns shaping giant species are scarce. The damselfishes of the genus Dascyllus appear as an excellent model for such a study. Their well understood phylogeny reveals that large‐bodied species have evolved in two different clades. Geometric morphometric methods were used to compare the ontogenetic trajectories of the neurocranium and the mandible in both small‐bodied (Dascyllus aruanus and Dascyllus carneus; maximum size: 50–65 mm standard length) and giant (Dascyllus trimaculatus and Dascyllus flavicaudus; maximum size: 90–110 mm standard length) Dascyllus species. At their respective maximum body size, the neurocranium of the giant species is significantly shorter and have a higher supraoccipital crest relative to the small‐bodied species, whereas mandible shape variation is more limited and is not related to the ‘giant’ trait. The hypothesis of ontogenetic scaling whereby the giant species evolved by extending the allometric trajectory of the small‐bodied ones (i.e. hypermorphosis) is rejected. Instead, the allometric trajectories vary among species by lateral transpositions. The rate of shape changes and the type of lateral transposition also differ according to the skeletal unit among Dascyllus species. Differences seen between the two giant species in the present study demonstrate that giant species may appear by varied alterations of the ancestor allometric pattern. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 99–117.     

Patterns of morphological evolution in the mandible of the house mouse Mus musculus (Rodentia: Muridae)

The worldwide distributed house mouse, Mus musculus, is subdivided into at least three lineages, Mus musculus musculus, Mus musculus domesticus, and Mus musculus castaneus. The subspecies occur parapatrically in a region considered to be the cradle of the species in Southern Asia (‘central region’), as well as in the rest of the world (‘peripheral region’). The morphological evolution of this species in a phylogeographical context is studied using a landmark‐based approach on mandible morphology of different populations of the three lineages. The morphological variation increases from central to peripheral regions at the population and subspecific levels, confirming a centrifugal sub‐speciation within this species. Furthermore, the outgroup comparison with sister species suggests that M. musculus musculus and populations of all subspecies inhabiting the Iranian plateau have retained a more ancestral mandible morphology, suggesting that this region may represent one of the relevant places of the origin of the species. Mus musculus castaneus, both from central and peripheral regions, is morphologically the most variable and divergent subspecies. Finally, the results obtained in the present study suggest that the independent evolution to commensalism in the three lineages is not accompanied by a convergence detectable on jaw morphology. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 635–647.     

A catalogue of skulls and jaws of eastern tropical Pacific blennioid fishes (Blennioidei: Teleostei): A proposed evolutionary sequence of morphological change

Skulls and jaws are compared in 35 species of tropical eastern Pacific (mainly Gulf of California) blennioid fishes (5 species of Tripterygiidae, 13 Labrisomidae, 12 Chaenopsidae, 5 Blenniidae). Morphotypes are arranged in a series from relatively large-mouthed fishes (tripterygiids and a few labrisomids) with protrusible jaws and conical teeth to species with small, non-protrusible jaws and a single row of incisiform teeth. This polarity of arrangement probably reflects the direction of phylogenetic development, as suggested by outgroup comparison with other perciform fishes. Distribution patterns of morphotypes in a simple morphospace, obtained by combining dentition, shape of the jaw arch and jaw protrusibility, are discussed within an adaptive context.     

Trophic morphology of the coral-associated genus Gobiodon (Teleostei: Gobiidae) from the Red Sea

Trophic morphology plays a major role in niche diversification among the highly diverse fish fauna of coral reefs. It gives insight into feeding types and is useful in taxonomy, as it can be used to differentiate between species and to estimate phylogenetic relationships. The gobiid genus Gobiodon comprises obligate coral dwelling species that are widespread on Red Sea and Indo-Pacific reefs. Yet very little is known about their ecology and behaviour of feeding. In this study, viscerocranial and oral jaw morphology was investigated in seven species of Gobiodon that occur in the Red Sea. Osteological analyses were carried out on enzyme cleared and alizarin red stained specimens. Shape variation of the viscerocranium was analyzed using a landmark-based geometric morphometric method on digital scans of hand drawings and visualized in thin-plate splines. Shape was determined by placing landmarks on skeletal elements in their articulated state. Morphology of the dentary and premaxilla was examined in further detail after disarticulation. A combination of landmarks and semilandmarks was applied to digital photographs of the oral jaws. It was shown that orbit size influences viscerocranial morphology by delimiting the lower jaw articulation angle. The premaxilla differs both in its extent and direction of curvature and in the structure of the articular process. Form changes in the dentary are mainly caused by variation of curvature and depth. A significant relationship between oral jaw shape and the number of anterior rows of teeth was found. Morphological diversity of selected elements is used to generate hypotheses concerning the trophic biology and niche diversification of Gobiodon . Brief dietary and in situ behavioural observations were carried out to further develop and test hypotheses.     

Ontogenetic shape changes in Pomacentridae (Teleostei,Perciformes) and their relationships with feeding strategies: a geometric morphometric approach

The present study explores the shape changes of cranial structures directly involved in food capturing during growth after reef settlement in two species of Pomacentridae (Dascyllus aruanus and Pomacentrus pavo). Landmark‐based geometric morphometrics were used to study allometric patterns and related shape changes in four skeletal units: neurocranium, suspensorium and opercle, mandible and premaxilla. At settlement, the larvae of both species have a relatively similar morphology, especially with respect to the mandible. Their shapes suggest a feeding mode defined as ram/suction‐feeding. Ontogenetic shape changes show a shift to a suction feeding mode of prey capture. The main transformations involved are an increase in height of the suspensorium and the opercle, an elevation of the supraoccipital crest, a relative shortening of the mandible, and a lengthening of the ascending process of the premaxilla. Shape changes of the mandible in the two studied species also reflect an increase of biting capacities. The high disparity between adult shape results from differences in the rate and in the length of ontogenetic trajectories, from divergence of the ontogenetic trajectories (neurocranium, mandible, and premaxilla) and parallel shifts of the trajectories in the size‐shape space (suspensorium and opercle). In an evolutionary context, allometric heterochronies during ontogeny of different skeletal unit of the head may be considered as a basis for the explanation of the diversity of damselfishes. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 92–105.     

Modularity in attachment organs of African Cichlidogyrus (Platyhelminthes: Monogenea: Ancyrocephalidae) reflects phylogeny rather than host specificity or geographic distribution

Cichlidogyrus spp. (Monogenea, Ancyrocephalidae) are common parasites of cichlid fishes from Africa and the Levant. They display important morphological variation in their attachment apparatus and infect a broad host spectrum throughout a wide geographic range. Thus, they offer an interesting model to investigate to what extent the phenotypic variability of the attachment organ among congeners is related to host specificity, geographic/environmental components, or phylogeny. A geometric morphometric approach was carried out to analyse the shape variation of sclerotized structures of the attachment organ within 66 African species of the genus Cichlidogyrus. The interspecific shape comparison supports the presence of three main morphological configurations, each consisting of a given combination of particular sclerite shapes. Moreover, data emphasize strong coordination and integration (shape co‐variation) among the different sclerites jointly forming the attachment organ. Although attachment apparatuses are usually considered to be the result of adaptive processes and must be adapted to the hosts and local environmental conditions, we found no relationship between these clusters and host specificity or geographical distribution. Nevertheless, groups are partially congruent with those obtained with the molecular phylogeny of a subset of species, suggesting a phylogenetic constraint rather than an adaptation to either hosts or environment. Because of the necessity to form a functional entity, modularity within attachment organ imposes important evolutionary constraint. This provides new insights into the evolvability of attachment organs, as well as into the morphological basis of host specificity and host–parasite co‐evolutionary interaction in helminth parasites. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 694–706.     

Correlates of Environmental Variables with Patterns in the Distribution and Abundance of Two Anemonefishes (Pomacentridae: Amphiprion) on an Eastern Australian Sub-tropical Reef System

The relationship between characteristics of the reef environment and variations in the distribution and abundance of the anemonefishes Amphiprion akindynos and A. latezonatus was investigated at North Solitary Island, a sub-tropical rocky reef system on the east coast of Australia. During the summers of 1994 and 1995, fish densities and host sea-anemone cover were assessed on replicate 25m transects at sites where host sea-anemones form semi-contiguous mats throughout the 6–21m depth range. Multiple regression analyses indicated that environmental variables accounted for 65–71% and 61–80% of the variations in the number of A. akindynos and A. latezonatus among-sites, respectively. Among-habitat comparisons indicated that A. latezonatus densities were positively correlated with depth (r=0.45–0.90), whereas A. akindynos showed no consistent depth-related abundance patterns. Poor correlations (p>0.05) between the densities of each species on transect lines suggested that present-day competition was unlikely to determine the preference of A. latezonatus for deeper depths. Correlations between host sea-anemone cover and fish densities at the within-habitat (depth) scale were comparatively stronger than correlations at among-habitats in both species. These results suggest that among-habitat comparisons can confound finer scale fish-habitat associations within habitat (depth) zones. Evidence suggests that while sea-anemone cover does, to an extent, regulate the local ecology of anemonefishes, other factors are also likely to interact to limit their densities.     

Tooth morphology of Echimyidae (Rodentia,Caviomorpha): homology assessments,fossils, and evolution

Echimyidae constitute the most important radiation of caviomorph rodents in the Neotropical region, represented by 20 extant genera and several extinct species. Both in extant and fossil forms, this diversity is reflected by a significant morphological variation found in crown structures of the cheek teeth. Different hypotheses of primary homology have been proposed for these structures, which, in turn, support diverse dental evolutionary hypotheses. In this contribution we inspect the main structures (cusps and lophids) of the lower deciduous teeth and molars in extinct and extant Echimyidae, and establish their topological correspondences. Comparisons with cusps and lophids of Erethizontidae are emphasized. We explore the testing of alternative primary hypotheses of lophid correspondences in a cladistic context. Following a ‘dynamic’ approach, we select the hypothesis of primary homology, which produced the more parsimonious results, and evaluate the evolutionary transformations of the dental characters analysed. In this context, the phylogenetic relationships of living Myocastor coypus (Molina, 1782) with the extinct Tramyocastor and Paramyocastor are tested. Our results indicate that pentalophodonty is the derived condition for the lower molars in Echimyidae, that trilophodonty evolved independently at least three times during the evolutionary history of these rodents, and that tetralophodonty represents the plesiomorphic condition. This study shows that dental evolution in echimyids can be better understood when occlusal structures are expressed as reliably comparable characters, and when fossils are taken into account. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 451–480.     

Trophic level and overlap of sea lions (Zalophus californianus) in the Gulf of California, Mexico

Stable isotope and scat analyses were used in concert to determine trophic level and dietary overlap among California sea lions from different rookeries in the Gulf of California. Isotopic analysis of the fur of sea lion pups revealed differences in δ¹⁵N and δ¹³C values among rookeries during the breeding season. Mean δ¹⁵N and δ¹³C values varied from 20.2‰ to 22.4‰ and from −15.4‰ to −14.0‰, respectively. The pattern of differences among rookeries was similar between years in most cases. Isotopic variations among rookeries were associated with differences in prey consumption. There was a significant correlation between δ¹⁵N value and trophic level, as determined by scat analysis. Joint application of isotopic and scat analyses allowed us to identify how the feeding habits of sea lions vary with location. Our results suggest the presence of spatial structure in available prey as well as the localized use of prey by sea lions across the Gulf of California.     

Host choice and fitness of anemonefish Amphiprion ocellaris (Perciformes: Pomacentridae) living with host anemones (Anthozoa: Actiniaria) in captive conditions

In this study, we investigated the host choice of naïve Amphiprion ocellaris, a specialist, at two different stages of development (newly settling juveniles and post-settlement juveniles). The fish were exposed to their natural and unnatural host species in the laboratory and their fitness was assessed in terms of activity and growth rate. Newly settling juveniles exhibited little host preference, while post-settlement juveniles immediately associated with their most common host in the wild. The analysis of fish activity confirmed that A. ocellaris is diurnal; they are most active in the morning, less at midday and barely move at night. The average travelling distance of juveniles was shorter in the groups living with their natural host, increasing in the groups living with an unnatural host and was highest in groups that did not become associated with any other unnatural host species. Post-settlement juveniles living with the natural host species grew better than those living with unnatural hosts or without anemone contact. These results suggest that the welfare of A. ocellaris in captivity will be optimized by keeping them with their natural anemone host species, although more generalist Amphiprion species may survive in association with other hosts.     

Head shape evolution in monitor lizards (Varanus): interactions between extreme size disparity,phylogeny and ecology

Characterizing patterns of observed current variation, and testing hypotheses concerning the potential drivers of this variation, is fundamental to understanding how morphology evolves. Phylogenetic history, size and ecology are all central components driving the evolution of morphological variation, but only recently have methods become available to tease these aspects apart for particular body structures. Extant monitor lizards (Varanus) have radiated into an incredible range of habitats and display the largest body size range of any terrestrial vertebrate genus. Although their body morphology remains remarkably conservative, they have obvious head shape variation. We use two‐dimensional geometric morphometric techniques to characterize the patterns of dorsal head shape variation in 36 species (375 specimens) of varanid, and test how this variation relates to size, phylogenetic history and ecology as represented by habitat. Interspecific head shape disparity is strongly allometric. Once size effects are removed, principal component analysis shows that most shape variation relates to changes in the snout and head width. Size‐corrected head shape variation has strong phylogenetic signal at a broad level, but habitat use is predictive of shape disparity within phylogenetic lineages. Size often explains shape disparity among organisms; however, the ability to separate size and shape variation using geometric morphometrics has enabled the identification of phylogenetic history and habitat as additional key factors contributing to the evolution of head shape disparity among varanid lizards.     

Arboreallty and morphological evolution in ground beetles (Carabidae: Harpalinae): testing the taxon pulse model

Abstract.— One-third to two-thirds of all tropical carabids, or ground beetles, are arboreal, and evolution of arboreality has been proposed to be a dead end in this group. Many arboreal carabids have unusual morphological features that have been proposed to be adaptations for life on vegetation, including large, hemispheric eyes; an elongated prothorax; long elytra; long legs; bilobed fourth tarsomeres; adhesive setae on tarsi; and pectinate claws. However, correlations between these features and arboreality have not been rigorously tested previously. I examined the evolution of arboreality and morphological features often associated with this habitat in a phylogenetic context. The number and rates of origins and losses of arboreality in carabids in the subfamily Harpalinae were inferred with parsimony and maximum-likelihood on a variety of phylogenetic hypotheses. Correlated evolution in arboreality and morphological characters was tested with concentrated changes tests, maximum-likelihood, and independent contrasts on optimal phylogenies. There is strong evidence that both arboreality and the morphological features examined originated multiple times and can be reversed, and in no case could the hypothesis of equal rates of gains and losses be rejected. Several features are associated with arboreality: adhesive setae on the tarsi, bilobed tarsomeres, and possibly pectinate claws and an elongated prothorax. Bulgy eyes, long legs, and long elytra were not correlated with arboreality and are probably not arboreal adaptations. The evolution of arboreal carabids has not been unidirectional. These beetles have experienced multiple gains and losses of arboreality and the morphological characters commonly associated with the arboreal habitat. The evolutionary process of unidirectional character change may not be as widespread as previously thought and reversal from specialized lifestyles or habitats may be common.     

Patterns of diversification in two species of short‐tailed bats (Carollia Gray, 1838): the effects of historical fragmentation of Brazilian rainforests

The small‐sized frugivorous bat Carollia perspicillata is an understory specialist and occurs in a wide range of lowland habitats, tending to be more common in tropical dry or moist forests of South and Central America. Its sister species, Carollia brevicauda, occurs almost exclusively in the Amazon rainforest. A recent phylogeographic study proposed a hypothesis of origin and subsequent diversification for C. perspicillata along the Atlantic coastal forest of Brazil. Additionally, it also found two allopatric clades for C. brevicauda separated by the Amazon Basin. We used cytochrome b gene sequences and a more extensive sampling to test hypotheses related to the origin and diversification of C. perspicillata plus C. brevicauda clade in South America. The results obtained indicate that there are two sympatric evolutionary lineages within each species. In C. perspicillata, one lineage is limited to the Southern Atlantic Forest, whereas the other is widely distributed. Coalescent analysis points to a simultaneous origin for C. perspicillata and C. brevicauda, although no place for the diversification of each species can be firmly suggested. The phylogeographic pattern shown by C. perspicillata is also congruent with the Pleistocene refugia hypothesis as a likely vicariant phenomenon shaping the present distribution of its intraspecific lineages. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 527–539.     

Evolution of molar shape in didelphid marsupials (Marsupialia: Didelphidae): analysis of the influence of ecological factors and phylogenetic legacy

The diversity of items consumed by modern didelphids, varying from mostly fruits in Caluromys Allen to mostly small vertebrates in Lutreolina O. Thomas, may cause changes in molar size and shape. We evaluated the morphometric variation of the first and third upper and lower molars of 16 genera of didelphid marsupials, with the aim of assessing the relationship between molar shape change, diet and phylogeny. We used a geometric morphometric approach to analyse how shape changes with diet. We mapped shape onto the phylogeny of the group to reconstruct ancestral states and analyse the evolution of molar shape. Finally, we statistically estimated the effect of size, diet and phylogeny on molar shape. All the analyses indicated little correlation between diet and molar shape and a strong correlation between the position of each genus on the phylogeny and molar shape. We believe that the wide ecological niche used by most of the groups (at least regarding diet) makes the evolutionary changes not strong enough to override pre‐existing differences that occur among clades, and the absence of highly diet‐specialized species (e.g. hypercarnivory or obligate folivory) causes the need for retaining a molar shape that can be useful to process different kinds of food items. © 2014 The Linnean Society of London     

Tidal influence on the evolution of egg size in the grunions (Leuresthes,Atherinidae)

Synopsis The duration of natural in-beach-sand incubation of Gulf of California and California grunion embryos is dictated by the tidally-driven wave activity in their respective habitats. Analyses of these tidal regimes reveal that the incubation period can be three times longer on beaches of the California coast than on those of the northern Gulf.In response to the more irregular tides of California, Leuresthes tenuis, the less primitive of the two species, has evolved an egg volume 310% larger than the Gulf grunion, L. sardina. The. larger egg not only supplies greater yolk reserves for differentiation, growth and maintenance, but also produces a larger embryo. The more abundant yolk stores and greater embryo size helps California grunion embryos to remain viable in the sand longer than those of the Gulf grunion.