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.     

Post-natal ontogeny of the mandible and ventral cranium in <Emphasis Type="Italic">Marmota</Emphasis> species (Rodentia,Sciuridae): allometry and phylogeny

Post-natal ontogenetic variation of the marmot mandible and ventral cranium is investigated in two species of the subgenus Petromarmota (M. caligata, M. flaviventris) and four species of the subgenus Marmota (M. caudata, M. himalayana, M. marmota, M. monax). Relationships between size and shape are analysed using geometric morphometric techniques. Sexual dimorphism is negligible, allometry explains the main changes in shape during growth, and males and females manifest similar allometric trajectories. Anatomical regions affected by size-related shape variation are similar in different species, but allometric trajectories are divergent. The largest modifications of the mandible and ventral cranium occur in regions directly involved in the mechanics of mastication. Relative to other anatomical regions, the size of areas of muscle insertion increases, while the size of sense organs, nerves and teeth generally decreases. Epigenetic factors, developmental constraints and size variation were found to be the major contributors in producing the observed allometric patterns. A phylogenetic signal was not evident in the comparison of allometric trajectories, but traits that allow discrimination of the Palaearctic marmots from the Nearctic species of Petromarmota are present early in development and are conserved during post-natal ontogeny.     

Ontogeny and phyletic size change in living and fossil lemurs

Lemurs are notable for encompassing the range of body‐size variation for all primates past and present—close to four orders of magnitude. Benefiting from the phylogenetic proximity of subfossil lemurs to smaller‐bodied living forms, we employ allometric data from the skull to probe the ontogenetic bases of size differentiation and morphological diversity across these clades. Building upon prior pairwise comparisons between sister taxa, we performed the first clade‐wide analyses of craniomandibular growth allometries in 359 specimens from 10 lemuroids and 176 specimens from 8 indrioids. Ontogenetic trajectories for extant forms were used as a criterion of subtraction to evaluate morphological variation, and putative adaptations among sister taxa. In other words, do species‐level differences in skull form result from the differential extension of common patterns of relative growth? In lemuroids, a pervasive pattern of ontogenetic scaling is observed for facial dimensions in all genera, with three genera also sharing relative growth trajectories for jaw proportions (Lemur, Eulemur, Varecia). Differences in masticatory growth and form characterizing Hapalemur and fossil Pachylemur likely reflect dietary factors. Pervasive ontogenetic scaling characterizes the facial skull in extant Indri, Avahi, and Propithecus, as well as their larger, extinct sister taxa Mesopropithecus and Babakotia. Significant interspecific differences are observed in the allometry of indrioid masticatory proportions, with variation in the mechanical advantage of the jaw adductors and stress‐resisting elements correlated with diet. As the growth series and adult data are largely coincidental in each clade, interspecific variation in facial form may result from selection for body‐size differentiation among sister taxa. Those cases where trajectories are discordant identify potential dietary adaptations linked to variation in masticatory forces during chewing and biting. Although such dissociations highlight selection to uncouple shared ancestral growth patterns, they occur largely via transpositions and retention of primitive size‐shape covariation patterns or relative growth coefficients. Am. J. Primatol. 72:161–172, 2010. © 2009 Wiley‐Liss, Inc.     

Patterns of cranial ontogeny in lacertid lizards: morphological and allometric disparity

We explored the ontogenetic dynamics of the morphological and allometric disparity in the cranium shapes of twelve lacertid lizard species. The analysed species (Darevskia praticola, Dinarolacerta mosorensis, Iberolacerta horvathi, Lacerta agilis, L. trilineata, L. viridis, Podarcis erhardii, P. melisellensis, P. muralis, P. sicula, P. taurica and Zootoca vivipara) can be classified into different ecomorphs: terrestrial lizards that inhabit vegetated habitats (habitats with lush or sparse vegetation), saxicolous and shrub‐climbing lizards. We observed that there was an overall increase in the morphological disparity (MD) during the ontogeny of the lacertid lizards. The ventral cranium, which is involved in the mechanics of jaw movement and feeding, showed higher levels of MD, an ontogenetic shift in the morphospace planes and more variable allometric patterns than more conserved dorsal crania. With respect to ecology, the allometric trajectories of the shrub‐climbing species tended to cluster together, whereas the allometric trajectories of the saxicolous species were highly dispersed. Our results indicate that the ontogenetic patterns of morphological and allometric disparity in the lacertid lizards are modified by ecology and functional constraints and that the identical mechanisms that lead to intraspecific morphological variation also produce morphological divergence at higher taxonomic levels.     

Dietary partitioning promotes the coexistence of planktivorous species on coral reefs

Theories involving niche diversification to explain high levels of tropical diversity propose that species are more likely to co‐occur if they partition at least one dimension of their ecological niche space. Yet, numerous species appear to have widely overlapping niches based upon broad categorizations of resource use or functional traits. In particular, the extent to which food partitioning contributes to species coexistence in hyperdiverse tropical ecosystems remains unresolved. Here, we use a molecular approach to investigate inter‐ and intraspecific dietary partitioning between two species of damselfish (Dascyllus flavicaudus, Chromis viridis) that commonly co‐occur in branching corals. Species‐level identification of their diverse zooplankton prey revealed significant differences in diet composition between species despite their seemingly similar feeding strategies. Dascyllus exhibited a more diverse diet than Chromis, whereas Chromis tended to select larger prey items. A large calanoid copepod, Labidocera sp., found in low density and higher in the water column during the day, explained more than 19% of the variation in dietary composition between Dascyllus and Chromis. Dascyllus did not significantly shift its diet in the presence of Chromis, which suggests intrinsic differences in feeding behaviour. Finally, prey composition significantly shifted during the ontogeny of both fish species. Our findings show that levels of dietary specialization among coral reef associated species have likely been underestimated, and they underscore the importance of characterizing trophic webs in tropical ecosystems at higher levels of taxonomic resolution. They also suggest that niche redundancy may not be as common as previously thought.     

Ladder webs in orb‐web spiders: ontogenetic and evolutionary patterns in Nephilidae

Spider web research bridges ethology, ecology, functional morphology, material science, development, genetics, and evolution. Recent work proposes the aerial orb web as a one‐time key evolutionary innovation that has freed spider‐web architecture from substrate constraints. However, the orb has repeatedly been modified or lost within araneoid spiders. Modifications include not only sheet‐ and cobwebs, but also ladder webs, which secondarily utilize the substrate. A recent nephilid species level phylogeny suggests that the ancestral nephilid web architecture was an arboricolous ladder and that round aerial webs were derived. Because the web biology of the basalmost Clitaetra and the derived Nephila are well understood, the present study focuses on the webs of the two phylogenetically intervening genera, Herennia and Nephilengys, to establish ontogenetic and macroevolutionary patterns across the nephilid tree. We compared juvenile and adult webs of 95 Herennia multipuncta and 143 Nephilengys malabarensis for two measures of ontogenetic allometric web changes: web asymmetry quantified by the ladder index, and hub asymmetry quantified by the hub displacement index. We define a ‘ladder web’ as a vertically elongated orb exceeding twice the length over width (ladder index ≥ 2) and possessing (sub)parallel rather than round side frames. Webs in both genera allometrically grew from orbs to ladders, more so in Herennia. Such allometric web growth enables the spider to maintain its arboricolous web site. Unexpectedly, hub asymmetry only increased significantly in heavy‐bodied Nephilengys females, and not in Herennia, challenging the commonly invoked gravity hypothesis. The findings obtained in the present study support the intrageneric uniformness of nephilid webs, with Herennia etruscilla webs being identical to H. multipuncta. The nephilid web evolution suggests that the ancestor of Nephila reinvented the aerial orb web because the orb arises at a much more inclusive phylogenetic level, and all intervening nephilids retained the secondarily acquired substrate‐dependent ladder web. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 849–866.     

Evolution of ontogeny in the hippopotamus skull: using allometry to dissect developmental change

Allometry describes the effect of size change on aspects of an organism's form and can be used to summarize the developmental history of growing parts of an animal. By comparing how allometric growth differs between species, it is possible to reveal differences in their pathways of development. The ability to compare and categorize developmental change between species is demonstrated here using morphometric methods. This involves the interspecific statistical comparison of a large number of bivariate relationships that summarize ontogenetic trajectories. These linear ontogenetic trajectories can be modified as they evolve in any of three ways: ontogenetic scaling indicative of change in the duration of growth, lateral shifts indicative of changes in prenatal development, and directional change indicative of novel modes of postnatal growth. I apply this analysis to skulls of the common hippopotamus ( Hippopotamus amphibius ) and the pygmy hippopotamus ( Hexaprotodon liberiensis ). The number of allometric changes falling into each category was statistically determined and Jolicoeur's multivariate generalization of simple allometry was used to provide an overview of cranial variation. For these skulls, directional change was not found to be statistically significant, but ontogenetic scaling and lateral shifts were both common. This indicates that conserved patterns of growth covariance (ontogenetic scaling) can be separated from novel or derived patterns (directional change and/or lateral shifts). This study demonstrates that He. liberiensis is not simply an ontogenetically scaled version of its larger relative. The evolutionary implications of allometric growth variation are discussed in the light of these findings and those of other studies.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 625–638.     

A test of Rensch's rule in varanid lizards

In a model group of giant reptiles, we explored the allometric relationships between male and female body size and compared the effects of sexual and fecundity selection, as well as some proximate causes, on macroevolutionary patterns of sexual size dimorphism (SSD). Monitor lizards are a morphologically homogeneous group that has been affected by extreme changes in body size during their evolutionary history, resulting in 14‐fold differences among the body sizes of recent species. Here, we analysed data concerning the maximum and/or mean male and female snout–vent lengths in 42 species of monitor lizard from literary sources and supplemented these data with measurements made in zoos. There was a wide scale of SSD from nearly monomorphic species belonging mostly to the subgenus Odatria and Prasinus group of the Euprepriosaurus to apparently male‐larger taxa. The variable best explaining SSD was the body size itself; the larger the species, the higher the SSD. This pattern agrees with the currently discussed Rensch's rule, claiming that the relationship between male and female body size is hyperallometric, i.e. the allometric exponent of this relationship exceeds unity and thus SSD increases with body size in the case of male‐larger taxa. All our estimates of the reduced major axis regression slopes of this relationship ranged from 1.132 to 1.155. These estimates are significantly higher than unity, and thus unequivocally corroborate the validity of Rensch's rule in this reptilian group. In spite of our expectation that the variation in SSD can be alternatively explained by variables reflecting the strength of sexual selection (presence of male combat), fecundity selection (e.g. clutch size and mass) and/or proximate ecological factors (habitat type), none of these variables had consistent effects on SSD, especially when the data were adjusted to phylogenetic dependence and/or body size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 293–306.     

Dealing with allometry in linear and geometric morphometrics: a taxonomic case study in the Leporinus cylindriformis group (Characiformes: Anostomidae) with description of a new species from Suriname

To achieve maximum efficacy, taxonomic studies that seek to distinguish amongst species must first account for allometric shape variation within species. Two recently developed software packages (SMATR and MorphoJ) offer regression‐based allometric approaches that are notable for their statistical power and ease of use and that may prove highly useful to taxonomists working with linear or geometric morphometric data. We investigate species delimitation of the slender‐bodied fishes in the Leporinus cylindriformis group using these programs and demonstrate the utility of the allometric corrections that they provide. Without allometric correction, many pairs of species are difficult to distinguish on the basis of morphometrics, but once regressions are used to account for marked allometric variation within species, most of the recognized species in this group can be readily distinguished with linear or geometric morphometrics, particularly using variation in the depth of the body. Both approaches returned congruent patterns of separation amongst putative species, but the geometric approach in MorphoJ distinguished amongst four more pairs of species than did the linear approach in SMATR and appears to provide slightly more statistical power. Based on distinctive morphometrics, meristics, and coloration, a highly elongate species of Leporinus from the Suriname, Corantijn, and Coppename rivers of Suriname is described herein as a new species, Leporinus apollo sp. nov. The unique L. cylindriformis holotype from Porto de Moz, Brazil differs in morphology, meristics, and pigmentation from specimens commonly referred to that species from the main basin of the Amazon; the latter specimens may represent an additional undescribed species. The L. cylindriformis holotype itself may represent a rare species or a specimen collected at the edge of its native range. Measurements of the holotype and paratype of Leporinus niceforoi, which were collected in the Amazonian slope of Colombia, differ substantially from similarly pigmented and putatively conspecific specimens from Amazonian portions of Ecuador and Peru. Recently collected specimens from Colombia are needed to determine whether the observed morphometric variation encompassed by the current concept of L. niceforoi indicates a morphocline within a single species, suggests the presence of multiple cryptic species, or results from shrinkage of the types. In all these cases, linear or geometric morphometric data can reliably differentiate amongst species, but only after one accounts for allometric shape variation. The new SMATR and MorphoJ software packages both offer easy and effective approaches to such allometrically informed taxonomy, and may prove useful to any systematist working on taxa that change shape as they grow. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 103–130.     

Daily age determination and growth rates of freshwater fish throughout a regulated lotic system of the Murray‐Darling Basin Australia

To facilitate future research in freshwater fish recruitment response to environmental flow delivery, size‐at‐age and growth models are presented for eight fish species occurring in south‐eastern Australia; three small‐bodied species and the juvenile 0+ age classes of five large‐bodied species. Otolith increments were used to estimate the daily age of golden perch Macquaria ambigua, bony bream Nematalosa erebi, common carp Cyprinus carpio; Murray cod Maccullochella peelii, freshwater eel‐tailed catfish Tandanus tandanus, Australian smelt Retropinna semoni, un‐specked hardyhead Craterocephalus stercusmuscarum fulvus and Murray–Darling rainbowfish Melanotaenia fluviatilis. Linear growth models provided the best fit for length‐at‐age data of juvenile 0+ age large‐bodied species; whereas von Bertalanffy growth functions provided the best fit to length‐at‐age data of small‐bodied species. The results provide novel baseline data for future research in this area.     

Allometric comparison of skulls from two closely related weasels, Mustela itatsi and M. sibirica

We conducted an interspecific comparison of skulls from two closely related but differently sized mustelid species, Mustela itatsi and M. sibirica (Mammalia, Carnivora, Mustelidae); a sexual comparison within the latter species showed remarkable size dimorphism. We clarified several differences in skull proportion related to size using allometric analyses and qualitative comparisons. Allometric analysis revealed that the skulls of male M. itatsi (the smaller species) have a relatively long palate; a slender viscerocranium and postorbital constriction; a broad, short, and low neurocranium; small carnassials; and a short mandible with a thin body and small ramus compared to the skulls of male M. sibirica (the larger species). Similar results were obtained when male M. itatsi were compared to female M. sibirica, although the male M. itatsi had a broader viscerocranium than female M. sibirica. A sexual comparison in M. sibirica revealed a larger skull size among the males with a relatively wide viscerocranium; wide postorbital constriction; a slender, long, and high neurocranium; short and wide auditory bullae; short carnassials; and a long and high mandible compared to females. Qualitative comparisons revealed changes in a few characters depending on skull size or with respect to some cranial components in each species. The interspecific differences observed were clearly larger than the intraspecific differences for three qualitative characters. The allometric and qualitative differences detected between these species suggest that each species is not simply the dwarf and/or giant morph of the other, and complicated differences were clarified.     

Sex combs,allometry, and asymmetry in Drosophila

There has been recent debate about the expected allometry of sexually‐selected traits. Although sexually‐selected traits exhibit a diversity of allometric patterns, signalling characters are frequently positively allometric. By contrast, insect genitalia tend to be negatively allometric, although the allometry of nongenital sexually‐selected characters in insects is largely unknown (with some notable exceptions). It has also been suggested that there should be a negative association between the asymmetry and size of bilaterally‐paired, sexually‐selected traits, although this claim is controversial. We assessed the allometry and asymmetry (fluctuating asymmetry, FA) of a nongenital contact–courtship structure, the sex comb, in replicate populations of three species of Drosophila (we also measured wing FA). Sex combs are sexually‐selected characters used to grasp the female's abdomen and genitalia and to spread her wings prior to and during copulation. Although species differed in the size of the sex combs, all combs were positively allometric, and comb allometry did not generally differ significantly between species or populations. Comb and wing asymmetry did vary across species, although not across populations of the same species. However, FA was trait specific and was never negatively associated with trait size. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 923–934.     

Plasticity of ontogenetic trajectories in cyprinids: a source of evolutionary novelties

Research in evolutionary developmental (evo‐devo) biology is making an increasingly important contribution to our understanding of the molecular mechanisms underlying the establishment of complex morphological traits. Deciphering the ontogenetic trajectories leading to the differentiation of sister species (and the existence of hybrids) is a new challenge in our understanding of speciation processes. In the present study, we characterized the ontogenetic trajectory of lower lip morphology in two cyprinid species and their hybrids. Chondrostoma toxostoma has an arched lower lip and a generalist diet. Chondrostoma nasus has a straight lower lip and a specialist diet. An analysis of 99 C. toxostoma, 99 C. nasus and 25 first‐generation (F₁) hybrid individuals demonstrated that the difference between arched and straight lip morphology was found to depend strongly on the height/width ratio of the lower lip. A comparison of the ontogenetic trajectories of these morphometric traits showed that the height of the lower lip was isometric to body length in both species, whereas developmental changes involving an acceleration and a hypermorphosis of the widening of the lower lip led to a straight lip morphology in C. nasus. F₁ hybrids were characterized by an extreme phenotype resulting from a rate of lower lip widening slower than that in the two parent species. Therefore, we rejected a codominance hypothesis and concluded that the first stage of hybridization provides the foundations of evolutionary novelty. These results have important evolutionary implications because lower lip shape is linked to dietary behaviour in many fish species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 342–355.     

Identification of cryptic species of Miniopterus bats (Chiroptera: Miniopteridae) from Madagascar and the Comoros using bioacoustics overlaid on molecular genetic and morphological characters

The number of Miniopterus bat species on Madagascar and the nearby Comoros islands (Malagasy region) has risen from four to 11. These recently described cryptic taxa have been differentiated primarily based on molecular markers and associated a posteriori morphological characters that corroborate the different clades. Members of this Old World genus are notably conservative in morphology across their range. Several sites on Madagascar hold up to four small‐bodied taxa of this genus that are morphologically similar to one another, although they can be distinguished based on the tragus, an ear structure associated with echolocation. Miniopterus often emit species‐specific calls. In the present study, we analyze the bioacoustics of the 11 species of Miniopterus currently recognized from the Malagasy region, with an initial identification of the 87 recorded and collected individuals based on molecular markers and certain morphological characters. In most cases, bioacoustic parameters differentiate species and have taxonomic utility. Miniopterus griveaudi populations, which occur on three islands (Madagascar, Anjouan, and Grande Comore), showed no significant differences in peak echolocation frequencies. After running a discriminant function analysis based on five bioacoustic parameters, some mismatched assignments of Malagasy species were found, which include allopatric sister‐taxa and sympatric, phylogenetically not closely‐related species of similar body size. Because the peak echolocation frequencies of two species (Miniopterus sororculus and Miniopterus aelleni) were independent of body size, they were acoustically distinguishable from cryptic sympatric congeners. The small variation around the allometric relationship between body size and peak echolocation frequency of Malagasy Miniopterus species suggests that intraspecific communication rather than competition or prey detection may be the driver for the acoustic divergence of these two species. Our well‐defined echolocation data allow detailed ecological work to commence aiming to test predictions about the relative roles of competition, prey availability, and social communication on the evolution of echolocation in Malagasy Miniopterus species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 284–302.     

Ontogeny of skull size and shape changes within a framework of biphasic lifestyle: a case study in six <Emphasis Type="Italic">Triturus</Emphasis> species (Amphibia,Salamandridae)

As with many other amphibians, Triturus species are characterized by a biphasic life cycle with abrupt changes in the cranial skeleton during metamorphosis. The post-metamorphic shape changes of the cranial skeleton were investigated using geometric morphometric techniques in six species: Triturus alpestris, T. vulgaris, T. dobrogicus, T. cristatus, T. carnifex, and T. karelinii. The comparative analysis of ontogenetic trajectories revealed that these species have a conserved developmental rate with divergent ontogenetic trajectories of the ventral skull shape that mainly reflect phylogenetic relatedness. A striking exception in the ontogenetic pattern was possibly found in T. dobrogicus, characterized by a marked increase in the developmental rate compared to the other newt species. The size-related shape changes explained a large proportion of shape change during post-metamorphic growth within each species, with marked positive allometric growth of skull elements related to foraging.     

Paedomorphosis in two small species of toothed whales (Odontoceti): how and why?

To evaluate and assess the ontogenetic background for paedomorphosis in phocoenids, samples of 144 harbour porpoises, 81 white‐beaked dolphins, and 130 Commerson's dolphins were compared in terms of the development of epiphyseal fusion, cranial suture fusion, and ontogeny of cranial shape. Harbour porpoises and Commerson's dolphins terminated growth and development of all investigated traits sooner than white‐beaked dolphins, leading to lesser degrees of fusion of skeletal elements and less postnatal allometric development. The latter occurred even though shape in the two paedomorphic species developed at twice the rate relative to the size of white‐beaked dolphins. These observations imply that progenetic evolution has occurred convergently in phocoenid and Cephalorhynchus ancestors. The truncated ontogenies allow sexual maturity to be attained earlier and provide a greater reproductive potential. Both species inhabit similar temperate productive habitats and, hence, ecological factors are proposed to have supplied the selection pressures leading to progenesis. Constant prey availability must be a prerequisite for the observed phenomena because frequent food‐intake is necessitated by the limited capacity for energy storage and high heat‐loss entailed by the resulting small body sizes. Progenesis has rarely been proposed in mammal species. This may reflect rarity or that mammalian expressions of progenesis are less obvious. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 278–295.     

Three-dimensional geometric morphometric analysis of the humerus: Comparative postweaning ontogeny between fossorial and semiaquatic water voles (Arvicola)

Different types of locomotion in phylogenetically close rodent species can lead to significantly different growth patterns of certain skeletal structures. In the present study, we compared the allometric and phenotypic trajectories of the humerus in semiaquatic (Arvicola sapidus) and fossorial (Arvicola scherman) water vole taxa, using three-dimensional geometric morphometrics, to investigate the relationships between functional and ontogenetic differences. Results revealed shared humerus traits between A. sapidus and A. scherman, specifically an expansion of the epicondylar and deltopectoral crests along postnatal ontogeny. In both species, the humerus of young specimens is more robust than in adults, possibly as a compensatory response for lower bone stiffness. However, significant interspecific differences were detected in all components of allometric and phenotypic trajectories. Noticeably divergent allometric trajectories were observed, probably as a result of different functional pressures exerted on this bone. Important differences in the form of the adult humerus between taxa were also found, particularly in features located in muscle insertion zones. Furthermore, the allometric regression revealed certain shape variation not associated with size in A. scherman, suggesting mechanical stress produced by the persistent digging activity during adulthood. A. scherman is a chisel-tooth digger that shares several traits in the humerus morphology with scratch-digger rodent species. Nevertheless, these shared characteristics are less pronounced in fossorial water voles, which is congruent with the different implications of the forelimb in the digging activity in these two types of diggers.     

Variation in the ontogenetic allometry of horn length in bovids along a body mass continuum

Allometric relationships describe the proportional covariation between morphological, physiological, or life‐history traits and the size of the organisms. Evolutionary allometries estimated among species are expected to result from species differences in ontogenetic allometry, but it remains uncertain whether ontogenetic allometric parameters and particularly the ontogenetic slope can evolve. In bovids, the nonlinear evolutionary allometry between horn length and body mass in males suggests systematic changes in ontogenetic allometry with increasing species body mass. To test this hypothesis, we estimated ontogenetic allometry between horn length and body mass in males and females of 19 bovid species ranging from ca. 5 to 700 kg. Ontogenetic allometry changed systematically with species body mass from steep ontogenetic allometries over a short period of horn growth in small species to shallow allometry with the growth period of horns matching the period of body mass increase in the largest species. Intermediate species displayed steep allometry over long period of horn growth. Females tended to display shallower ontogenetic allometry with longer horn growth compared to males, but these differences were weak and highly variable. These findings show that ontogenetic allometric slope evolved across species possibly as a response to size‐related changes in the selection pressures acting on horn length and body mass.     

Plant stages with biotic,indirect defences are more palatable and suffer less herbivory than their undefended counterparts

Plants have evolved several anti‐herbivory strategies, including direct defences, such as mechanical and chemical defences, and indirect or biotic defences, such as the recruitment of defending animals. We examined whether the investment plants make in direct defences differs between those which do and do not invest in biotic defences, by comparing standing herbivory and palatability of congeneric species with and without indirect defences at two ontogenetic stages: before and after the onset of indirect defences. We used Cordia alliodora and Croton suberosus as the species with indirect defences and Cordia elaeagnoides and Croton pseudoniveus as the species without indirect defences. We predicted that herbivores would prefer to eat species and stages with indirect defences to those without them. As predicted, we found that herbivores preferred species and ontogenetic stages with indirect defences in all cases. Overall, however, natural levels of herbivory were lower in species with indirect defences. We conclude that indirect defences offer effective protection against herbivores and posit that their recruitment allows plants to reduce investment in other defence mechanisms. Our results support the notion that plants trade‐off between direct and indirect defensive strategies. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 536–543.