Differential sexual dimorphism: size and shape in the cranium and pelvis of grey foxes (Urocyon)

Patterns of sexual size dimorphism (SSD) and cranial dimorphism are well documented. However, limited examinations exist of the contrasts in the patterns and nature of dimorphism across body regions (e.g. cranium, pelvis), particularly when these regions have different sex-specific functions (e.g. display in mating, locomotion, and reproduction). Using landmark-based morphometric techniques, we investigated size and shape dimorphism variation in the crania and pelves of two closely-related fox species within the genus Urocyon . Although we found no significant size and shape dimorphism in the crania of either species, we did find significant dimorphism in the pelvis: its size was dimorphic in Urocyon littoralis (but not in Urocyon cinereoargenteus ) and its shape was dimorphic in both species (though more pronounced in U. littoralis ). The observation of greater dimorphism in the pelvis than in the cranium suggests that factors such as offspring size and locomotor mode play a greater role in sexual dimorphism than simple 'whole body' allometric affects associated with dimorphism in body size.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 339–353.     

Female-biased sexual size dimorphism in tinamous: a comparative test fails to support Rensch's rule

Rensch's rule states that degree of sexual dimorphism increases with body size in species with larger males, and decreases with body size in those with larger females. To test this rule, we assessed the pattern of sexual size dimorphism in tinamous using a comparative analysis of independent contrasts. Tinamous are a monophyletic group of primitive birds comprising at least 47 ground dwelling species with prominent or exclusive paternal care of eggs and offspring. Although the size of females exceeded that of males in most considered species, we found an isometric relationship between males and females, instead of the negative allometric one predicted by Rensch's rule. Previous studies in Strigiformes and Falconiformes found positive allometric and isometric relationships respectively, and, considering these findings with our results, we conclude that Rensch's rule is not supported by birds with exclusively female-biased sexual dimorphism in size.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 519–527     

Size, shape and structural versatility of the skull of the subterranean rodent Ctenomys (Rodentia, Caviomorpha): functional and morphological analysis

Morphological analysis of the skull of the subterranean rodent Ctenomys , a highly speciose genus which uses both claws and teeth when digging, shows that for a broad range of species size, scaling was associated with both variation and maintenance of shape. Our results show that the angle of incisor procumbency (AIP), a character largely viewed as an adaptation to digging with teeth, is highly variable. We found a non-significant relationship between AIP and basicranial axis (basioccipital + basisphenoid) length, a measure of overall skull size. Accordingly, both small and large Ctenomys species possess either high or low AIP. A significant relationship between AIP and diastema length, given the rostral allometry seen in Ctenomys , suggests that hypermorphosis to a certain extent influences AIP. However, the roots of the incisor are lateral to those of the cheek teeth and their position may thus shift freely. This observation supports the notion that skull structural design, and to a certain extent rostral allometry, underlies variation in AIP. On the other hand, the positive allometry of incisor width and thickness indicates that, in larger species, proportionately powerful incisors are able to resist greater bending forces. We found that the out-lever arm of the jaw adductor muscles scales with positive allometry against basicranial axis length. However, we found an isometric relationship between in- and out-lever arms. In this case, conservation of skull proportions, regardless of variation in size, is a feature possibly related to the maintenance of an effective tooth digging capability. Functional and ecological data are discussed when assessing the implications of size and shape variation in the skull of Ctenomys .  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 78 , 85−96.     

Ecological and phylogenetic dimensions of cranial shape diversification in South American caviomorph rodents (Rodentia: Hystricomorpha)

Caviomorph rodents represent an excellent model to explore morphological diversification on a macroevolutionary scale, as they are ecologically and morphologically diverse. We analysed cranial shape variation using geometric morphometrics and phylogenetic comparative methods. Most variation involved the shape of the rostrum, basicranium, and cranial vault, and clearly matched the phylogenetic structure. At the same time, a strong allometric pattern was associated with the length of the rostrum and cranial vault, size of the auditory bulla, and depth of the zygomatic arch. After accounting for size influence, and taking phylogenetic structure into account, shape variation was significantly associated with habitat. Our results highlight the presence of complex relationships between morphological, phylogenetic, and ecological dimensions in the diversification of the caviomorph cranium. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 898–913.     

Quantifying shape and ecology in avian pedal claws: The relationship between the bony core and keratinous sheath

Terrestrial tetrapods use their claws to interact with their environments in a plethora of ways. Birds in particular have developed a diversity of claw shapes since they are often not bound to terrestrial locomotion and have heterogeneous body masses ranging several orders of magnitude. Numerous previous studies have hypothesized a connection between pedal claw shape and ecological mode in birds, yet have generated conflicting results, spanning from clear ecological groupings based on claw shape to a complete overlap of ecological modes. The majority of these studies have relied on traditional morphometric arc measurements of keratinous sheaths and have variably accounted for likely confounding factors such as body mass and phylogenetic relatedness. To better address the hypothesized relationship between ecology and claw shape in birds, we collected 580 radiographs allowing visualization of the bony core and keratinous sheath shape in 21 avian orders. Geometric morphometrics was used to quantify bony core and keratinous sheath shape and was compared to results using traditional arc measurements. Neither approach significantly separates bird claws into coarse ecological categories after integrating body size and phylogenetic relatedness; however, some separation between ecological groups is evident and we find a gradual shift from the claw shape of ground‐dwelling birds to those of predatory birds. Further, the bony claw core and keratinous sheath are significantly correlated, and the degree of functional integration does not differ across ecological groups. Therefore, it is likely possible to compare fossil bony cores with extant keratinous sheaths after applying corrections. Finally, traditional metrics and geometric morphometric shape are significantly, yet loosely correlated. Based on these results, future workers are encouraged to use geometric morphometric approaches to study claw geometry and account for confounding factors such as body size, phylogeny, and individual variation prior to predicting ecology in fossil taxa.     

Allometry of the baculum and sexual size dimorphism in American martens and fishers (Mammalia: Mustelidae)

Genitalia are among the most variable of morphological traits, and recent research suggests that this variability may be the result of sexual selection. For example, large bacula may undergo post‐copulatory selection by females as a signal of male size and age. This should lead to positive allometry in baculum size. In addition to hyperallometry, sexually selected traits that undergo strong directional selection should exhibit high phenotypic variation. Nonetheless, in species in which pre‐copulatory selection predominates over post‐copulatory selection (such as those with male‐biased sexual size dimorphism), baculum allometry may be isometric or exhibit negative allometry. We tested this hypothesis using data collected from two highly dimorphic species of the Mustelidae, the American marten (Martes americana) and the fisher (Martes pennanti). Allometric relationships were weak, with only 4.5–10.1% of the variation in baculum length explained by body length. Because of this weak relationship, there was a large discrepancy in slope estimates derived from ordinary least squares and reduced major axis regression models. We conclude that stabilizing selection rather than sexual selection is the evolutionary force shaping variation in baculum length because allometric slopes were less than one (using the ordinary least squares regression model), a very low proportion of variance in baculum length was explained by body length, and there was low phenotypic variability in baculum length relative to other traits. We hypothesize that this pattern occurs because post‐copulatory selection plays a smaller role than pre‐copulatory selection (manifested as male‐biased sexual size dimorphism). We suggest a broader analysis of baculum allometry and sexual size dimorphism in the Mustelidae, and other taxonomic groups, coupled with a comparative analysis and with phylogenetic contrasts to test our hypothesis. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 955–963.     

The evolution of waving displays in fiddler crabs (Uca spp., Crustacea: Ocypodidae)

Male fiddler crabs are commonly recognized by the presence of a single massive claw used in a variety of contexts, including territorial defence, agonistic interactions, and courtship behaviour. The most common behavioural context involving these enlarged chelipeds is their use in waving displays, which are remarkably diverse among species. Although the waving display is one of the most obvious behavioural features of male fiddler crabs, little is known about their main evolutionary trends during the diversification of the genus. The present study employed phylogenetic comparative methods to investigate the evolution of waving behaviour in a sample of 19 species of Uca from Central and North America. Digital recordings were used to quantify the temporal dynamics of waving behaviour in each species. Multivariate ordination methods were used to assess whether different elements of the display showed distinct evolutionary dynamics, particularly with respect to body size and the environment where species are most commonly found. Most of the interspecific variation in displays involves differences in the overall waving velocity, with no correspondence to their local environments, nor their body size. Interestingly, despite the strong concentration of variance in the first two ordination axes, there was no statistically significant evidence for phylogenetic signals in their respective scores. These results suggest that the overall structure of waving displays is evolutionarily labile, at the same time as being concentrated in a few particular axes of variation, possibly indicating evolution along lines of least resistance. The approach employed in the present study highlights the utility of phylogenetic comparative methods for elucidating the evolution of complex behavioural characteristics, such as the waving display in male fiddler crabs. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 307–315.     

Bilateral linkage of monomorphic and dimorphic limb sizes in fiddler crabs

In this study, the subject of whether investment in one bilateral structure was linked to investment in the homologous bilateral opposite structure was investigated. Male fiddler crabs (genus Uca, family Ocypodidae) displayed strong bilateral claw differentiation of function and size, which are used for feeding (minor claw) or display/combat (major claw). Females had similar‐sized feeding claws. Linkage between claw size was investigated by estimating the deviations from an overall fitted regression of claw length to body size. The positive correlations of the deviations of claw size for major and minor claws of males and between right and left claws of females, relative to body size, suggested a linkage in investment between one claw and the corresponding claw on the other side of the body, for both monomorphic females and dimorphic males. A signal to send resources may be effectively gated to the claw complex, suggesting that positively correlated resources are allocated to both claws. Positive correlations were also found at the interspecific level. The fiddler crab model, described here, gives access to study the linkage in symmetric and asymmetric bilateral structures in the same species with a connection to the macroevolutionary level.     

Phylogenetic relationships within Plantago (Plantaginaceae): evidence from nuclear ribosomal ITS and plastid trnL-F sequence data

A molecular phylogenetic study of Plantago L. (Plantaginaceae) analysed nucleotide variation in the internal transcribed spacers (ITS) of nuclear ribosomal and plastid trnL-F regions. Included are 57 Plantago species, with two Aragoa species as the ingroup and three Veronica species as the outgroup. Phylogenetic analysis using maximum parsimony identified five major clades, corresponding to the taxonomic groups Plantago subgenera Plantago, Coronopus, Psyllium, Littorella and Bougueria . Aragoa is sister to genus Plantago . Plantago subgenus Littorella is sister to the other subgenera of Plantago . The results are in general correlated with a morphological phylogenetic study and iridoid glucoside patterns, but Plantago subgenus Albicans is paraphyletic and should be included in Plantago subgenus Psyllium sensu lato to obtain a monophyletic clade with six sections. Plantago section Hymenopsyllium is more closely related to section Gnaphaloides than to section Albicans . Plantago subgenus Bougueria is sister to subgenus Psyllium s.l. section Coronopus in Plantago subgenus Coronopus is subdivided in two series. Only some of the sections can be resolved into series. DNA variation within genus Plantago is high, a result that would not have been predicted on the basis of morphology, which is relatively stereotyped. If we calibrate a molecular clock based on the divergence of P. stauntoni , endemic to New Amsterdam in the southern Indian Ocean, we calculate the time of the split between Plantago and Aragoa to be 7.1 million years ago, which is congruent with the fossil record.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 323–338.     

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.     

Falling apart and merging: diversification of slender salamanders (Plethodontidae: Batrachoseps) in the American West

The plethodontid genus Batrachoseps , the slender salamanders, is the most diverse clade of salamanders in western North America, but it has posed taxonomic difficulties because it contains many morphologically cryptic species. A segment of the mitochondrial DNA gene cytochrome b was studied for 278 individuals densely sampled from throughout the range of all 18 described species and several undescribed species. Phylogenetic analyses of the mtDNA data identify six major clades, one corresponding to the subgenus Plethopsis and five within a monophyletic subgenus Batrachoseps. All major clades and most species within these clades display strong phylogeographic structuring. Comparisons of mtDNA and allozyme data show that several allozymically cohesive groups are not monophyletic with respect to mtDNA. We suggest that this phenomenon results from fragmentation of populations, divergence in allopatry, and then recontact and gradual merging of units caused predominantly by male-mediated gene flow. The mtDNA offers evidence that populations were once more isolated than they are now, while the patterns of allozyme variation reflect recent and current interactions among populations. The complex patterns of morphological, allozymic and mtDNA variation associated with the constantly changing geological landscape give insight into the nature of processes responsible for species formation in Batrachoseps .  © 2002 The Linnean Society of London. Biological Journal of the Linnean Society , 2002, 76 , 361–391.     

Evolution and determinants of host specificity in the genus Lamellodiscus (Monogenea)

The evolution and determinants of host specificity in Lamellodiscus species (Monogenea, Diplectanidae) were investigated. The 20 known Mediterranean species were studied, all parasites of fishes from the family Sparidae (Teleostei). An index of specificity, which takes into account the phylogenetic relationships of their fish host species, was defined. The link between specificity and its potential determinants was investigated in a phylogenetic context using the method of independent contrasts. Host specificity in Lamellodiscus species appeared to be highly constrained by phylogeny, but also linked to host size. Mapping specificity onto the parasite phylogenetic tree suggests that specialist species do not represent an evolutionary dead end, and that specialization is not a derived condition. It is hypothesized that the ability to be generalist or specialist in Lamellodiscus is controlled by intrinsic, phylogenetically-related characteristics, and that specialist species tend to use large hosts, which may be more predictable.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 431−443.     

Morphological Variation of the Forelimb and Claw in Neotropical Sigmodontine Rodents (Rodentia: Cricetidae)

The limbs of mammals exhibit a variety of morphologies that reflect the diversity of their habitats and their functional needs, including subtle structural differences in their distal limb integumentary appendages (hooks, claws, adhesive pads). Little is known about structure and function of claws of sigmodontine rodents. Here, we analyze claw shape and forelimb skeleton morphology of 25 species of sigmodontine rodents with different locomotory types (ambulatory, fossorial, natatorial, quadrupedal saltatorial, and scansorial), taking into account their phylogenetic affinities. Qualitative differences in claw shape were examined using digital photographs, and quantitative measurements were made for length, height, and curvature of the claws of all digits, and dimensions of other forelimb skeletal elements. Our results show that both phylogeny and ecological categories explain substantial components of the morphological variation in sigmodontine rodents. Qualitative analysis reveals that non-specialized forms (ambulatory, quadrupedal saltatorial, and scansorial) tend to have high and strongly curved claws, whereas highly specialized forms (fossorial and natatorial) tend to have elongate and smoothly curved claws. However, the quantitative analysis differentiated the fossorial and scansorial by variables related to claw, and natatorial by variables related to bones of the forelimb. No variables that could differentiate ambulatory or quadrupedal saltatorial forms were found, demonstrating that these forms show a generalized morphological pattern. This study indicates that both historical and ecological factors contribute to the evolution of claw length in these groups.     

Influence of sociality on allometric growth and morphological differentiation in sponge-dwelling alpheid shrimp

Eusocial societies are defined by a reproductive division of labour between breeders and nonbreeders that is often accompanied by morphological differentiation. Some eusocial taxa are further characterized by a subdivision of tasks among nonbreeders, often resulting in morphological differentiation among different groups (subcastes) that specialize on different sets of tasks. We investigated the possibility of morphological castes in eusocial shrimp colonies ( Zuzalpheus , formerly part of Synalpheus ) by comparing growth allometry and body proportions of three eusocial shrimp species with three pair-forming species (species where reproductive females and males occur in equal sex ratios). Allometry of eusocial species differed in several respects from that of pair-forming species in both lineages. First, allometry of fighting claw size among individuals other than female breeders was steeper in eusocial than in pair-forming species. Second, breeding females in eusocial colonies had proportionally smaller weapons (fighting claws) than females in pair-forming species. Finally, claw allometry changed with increasing colony size in eusocial species; large colonies showed a diphasic allometry of fighting claw and finger size, indicating a distinctive group of large individuals possessing relatively larger weapons than other colony members. Shrimp are thus similar to other eusocial animals in the morphological differentiation between breeders and nonbreeders, and in the indication that some larger nonbreeders might contribute more to defence than others.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 527–540.     

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.     

PATTERNS OF QUANTITATIVE VARIATION IN LEPIDOPTERAN WING MORPHOLOGY: THE CONVERGENT GROUPS HELICONIINAE AND ITHOMIINAE (PAPILIONOIDEA: NYMPHALIDAE)

Wing morphology has historically been a major focus in taxonomic and evolutionary studies of lepidopterans. However, general patterns of quantitative variation and diversification in wing sizes and shapes and the factors underlying them have been unexplored. A morphometric study of wing variation in the convergent heliconine and ithomine butterflies reveals remarkable similarities, both in their morphologies at a given size and in their patterns of allometry and variability. The groups differ primarily in the relative lengths of inner and outer forewing margins, with larger species being more similar across groups than smaller ones. Allometric size-scaling variation accounts for more than 90% of the total morphological variation in the two groups and thus seems to be the major determinant of wing shape. Forewings and hind wings are isometric in size (area) with respect to one another; however, wing shape within and among groups is significantly allometric, resulting in considerable shape differences between small and large species. A strong trend of increasing variability from anterior to posterior along the wings is consistent with hypotheses of aerodynamic constraint. Wings and bodies represent classical morphological “character suites” in that size and shape variation are more tightly correlated within suites than among them. Such complexes argue against the overriding importance of aerodynamic factors, such as wing load and muscle development, in constraining gross morphology.     

The evolution of armament strength: evidence for a constraint on the biting performance of claws of durophagous decapods

Performance data for the claws of six sympatric species of Cancer crabs confirmed a puzzling pattern reported previously for two other decapod crustaceans (stone crabs, Menippe mercenaria, and lobsters, Homarus americanus): Although biting forces increased, maximum muscle stresses (force per unit area) declined with increasing claw size. The negative allometry of muscle stress and the stress at a given claw size were fairly consistent within and among Cancer species despite significant differences in adult body size and relative claw size, but were not consistent among decapod genera. Therefore, claw height can be used as a reliable predictor of maximum biting force for the genus Cancer, but must be used with caution as a predictor of maximum biting force in wider evolutionary and biogeographical comparisons of decapods. The decline in maximum muscle stress with increasing claw size in Cancer crabs contrasts with the pattern in several other claw traits. Significantly, three traits that affect maximal biting force increased intraspecifically with increasing claw size: relative claw size, mechanical advantage, and sarcomere length of the closer muscle. Closer apodeme area and angle of pinnation of the closer muscle fibers varied isometrically with claw size. The concordant behavior of these traits suggests selection for higher biting forces in larger crabs. The contrast between the size dependence of muscle stress (negative allometry) and the remaining claw traits (isometry or positive allometry) strongly suggests that an as yet unidentified constraint impairs muscle performance in larger claws. The negative allometry of muscle stress in two distantly related taxa (stone crabs and lobsters) further suggests this constraint may be widespread in decapod crustaceans. The implications of this performance constraint for the evolution of claw size and the "arms-race" between decapod predators and their hard-shelled prey is discussed.     

Relationships, classification and evolution of Scleranthus (Caryophyllaceae) as inferred from analysis of morphological characters

Study of floral anatomy, micromorphology, palynology and onotogeny has revealed new characters for phylogenetic analysis in the genus Scleranthus . Cladistic analysis of these characters, along with those previously available, suggests that the genus consists of Eurasian/Mediterranean and Australasian sister clades. Gynoecial morphology and development are closely similar in all species, suggesting the genus is monophyletic despite its disjunct northern and southern hemisphere distribution. Variation in pollen:ovule ratios and their implications for the evolution of Scleranthus species are also discussed and it is concluded that a range of breeding strategies intermediate between autogamy and xenogamy exists in the genus. Single-stamened species of Scleranthus are likely to be obligate autogams, despite their comparatively high pollen:ovule ratios in relation to autogamic species of other genera.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 15–29.     

Cranial shape variation and molecular phylogenetic structure of crested newts (Triturus cristatus superspecies: Caudata,Salamandridae) in the Balkans

In the present study, we investigated the degree of congruence between phylogeny, as inferred from mitochondrial (mt)DNA sequences, and cranium shape variation of crested newts (Triturus cristatus superspecies) in the Balkans. These newts belong to four phylogenetic clades defined by mtDNA analysis, and significantly differed in cranial shape. Allometry explained a high percentage of shape variation in crested newts. The clade‐specific allometric slopes significantly diverged for both the ventral cranium and dorsal cranium, indicating that differences in shape between clades could not be a simple consequence of their difference in size. The analysis of hierarchical and spatial variation showed similarity in the patterns of global and spatially localized hierarchical variation of cranial shape. We also found significant congruence between the pattern of cranial shape variation and molecular phylogeny. The differences in morphology of Triturus dobrogicus in comparison to other crested newt clades, including marked differences in cranium shape, is discussed in the context of the evolution and ecology of crested newts. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 348–360.