Size constraints and the evolution of display complexity: why do large lizards have simple displays?

Social, environmental, and perceptual factors have been suggested to account for the evolution of visual signal diversity in lizards. Previous investigations have inferred that signal complexity may also be related to body size. In this study, we use three complementary comparative analyses to investigate whether body size has influenced macro-evolutionary trends in display modifier repertoire size for 110 species of iguanian lizards. We found evidence that signal complexity, as measured by repertoire size, is negatively associated with body size. However, this relationship was not strictly linear. Rather, body size seems to impose a threshold on signal evolution. Specifically, the evolution of large repertoire size appears to be less likely above a particular size threshold, which results in large-bodied lizards having a significantly lower probability of evolving elaborate displays. This relationship may reflect the influence of body size on resource use and the emergent social dynamics it promotes. Large lizards tend to be herbivorous and typically do not defend foraging patches. Consistent with this hypothesis is the previously reported finding of a similar size threshold dividing herbivorous from insectivorous lizards. We suggest to fully understand the evolutionary processes acting on communicative systems, it is important that we identify both the selective forces involved and the nature of their influence. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 145–161.     

Relations between microhabitat use and limb shape in phrynosomatid lizards

With the exception of the well-documented case for anoline lizards, recent studies have found few evolutionary relationships between morphology and habitat use in lizards despite clear-cut biomechanical predictions. One of the factors typically hampering these analyses is the clustering of habitat use within evolutionary lineages. In the present study, body shape was quantified for male and female lizards of 30 species of phrynosomatid lizards. This group was selected as little clustering of ecological variables seemed to be present. The results of traditional analyses indicate that evolutionary correlates of habitat use were prominent in the hindlimbs of both sexes. Species living in open habitats are characterized by longer femurs, and longer hindlimbs relative to the forelimb. Moreover, males from ground-dwelling species utilizing open habitats have longer toes on the hind foot than males from climbing species. Phylogenetic analyses indicated strong evolutionary associations between habitat use and the relative length of front and hindlimbs, with species from open terrestrial habitats having significantly shorter frontlimbs relative to their hindlimb than rock or tree climbing species. Evolutionary associations between morphology and habitat use were generally stronger for male lizards, indicating a potentially important contribution of sexual selection to the evolution of differences in limb proportions.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 149–163.     

Patagial complex evolution in hummingbirds and swifts (Apodiformes): a molecular phylogenetic perspective

Studies of the role of flight in vertebrate evolution often have focused on the propatagial muscle complex because this structure forms the wing's leading edge. However, historical narratives about the evolution of flight anatomy are compromised because traditional higher-level taxonomies typically are based in part on the propatagium itself. To avoid this circularity, I used a consensus molecular phylogeny to examine propatagial evolution in the highly aerial sister groups, hummingbirds and swifts (Apodiformes). Mapping of anatomy on molecular-based phylogeny indicates that structural variation in M .  tensor propatagialis pars brevis (TPB) is congruent with the major subclades of both hummingbirds and swifts. However, the humeral tendon and broad attachment of the fleshy belly of TPB with M .  extensor metacarpi radialis (EMR) most likely underwent parallel change in hummingbirds and swifts, while the distal tendon present only in hummingbirds changed from a thin sheet to a strong tendon. The combination of divergent (within hummingbirds or swifts) and parallel (between hummingbirds and swifts) evolutionary patterns implies that the taxonomic value of the propatagial complex in apodiformes depends on anatomical component and level of divergence. The congruence of anatomy with molecular phylogeny provides independent criteria for designating relatively ancestral versus derived clades of apodiformes. Based on these polarities, living hummingbirds and swifts express additional parallel trends from arboreal to more aerial foraging styles, and from depauperate to species-rich clades. Within apodiformes, the link of flight anatomy with taxonomic and ecologic diversity suggests that elaboration of locomotor modes was important for apodiform diversification, echoing a similar pattern for birds relative to their reptilian ancestors.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 211–219.     

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.     

Revising the definition of the crustacean seta and setal classification systems based on examinations of the mouthpart setae of seven species of decapods

The crustacean cuticle has numerous projections and some of these projections, the setae, have important mechanical as well as sensory functions. The setae display a wide diversity in their external morphology, which has led to great problems separating setae from other projections in the cuticle and problems in making a consistent classification system. Here, the cuticular projections on the mouthparts of seven species of decapods are examined by scanning and transmission electron microscopy. A new definition is given: a seta is an elongate projection with a more or less circular base and a continuous lumen; the lumen has a semicircular arrangement of sheath cells basally. From the details of the external morphology the mouthpart setae are divided into seven types: pappose, plumose, serrulate, serrate, papposerrate, simple and cuspidate setae, which are suggested to reflect mechanical functions and not evolutionary history. This classification system is compared with earlier systems.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 233–252.     

Ecological diversity and adaptive tendencies in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to climbing and epiphytic habits

Among the basal fern families, the Hymenophyllaceae, with more than 600 species, display a high diversity in terms of their morphology and the habitats that they occupy. We have chosen to focus on Trichomanes L., a clearly defined genus for which a phylogeny is presently being developed, to investigate the appearance of the climbing and epiphytic habits, as well as the related supposed adaptive characters. In this study we present the first review of the different ecological types within the genus: terrestrial, climbing (divided into hemi-epiphytic forms and true lianas), and epiphytic types. The study of several features concerning stem morphology and leaf size allows a proposal on relationships between ecology and plant morphology. Terrestrial species display a thick monocaulous rhizome with robust roots and short internodes. Climbing species are characterized by a branched, thick, creeping rhizome with long internodes. Epiphytic species also exhibit long, creeping and branching stems with long internodes but the rhizome is fine to filiform. Under these circumstances, there is a reduction of root system and frond size leading to dwarfism in numerous instances. This may be related to an extreme hygrophilous epiphytic strategy. Finally, hypotheses on the evolution of these habits and hence on the evolutionary relationships between ecology and characters are presented and discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 41–63.     

Mechanisms of heterochronic change and stasis for clutch size in swifts (Apodiformes)

Since the mid‐1800s, explanations in evolution have focused on adaptivity and natural selection, largely at the expense of factors internal to the organism. Today, the importance of internal factors is no longer in dispute. Progress continues to lag, however, on the design of a framework that would allow the integration of their action in evolution. Herein, as part of a wider effort aimed at testing the functionality of the general principle based upon the embryological concepts of induction, competence, and determination (dubbed developmental determination), I present a model for the evolution of clutch size in swifts. It is structured around the physiological mechanism controlling the trait. In indeterminate laying species, it involves an exogenous signal to the female, or induction, and the acquired ability to respond to this signal, or competence. The model explains, with the action of natural selection, how a 2‐day shift in the onset of competence induces change in clutch size among species. Through the origin of induction, it predicts the evolutionary transition from indeterminate to determinate laying. Finally, in some species that are indifferent to the general increase in clutch size with latitude, it reveals how both types of developmental transformation may operate to create stasis. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 1067–1079.     

Song diversification and complexity in canaries and seedeaters (Serinus spp.)

Bird song is a composite trait that shows great variation in syntax and phonology between taxa, which hampers the precise quantification of complexity. This should be more feasible for closely-related species that share aspects of song organization. We conducted a comparative study of song and syllable characteristics in the largest fringillid genus, Serinus (canaries and seedeaters), with two objectives: (1) to describe and quantify song complexity in the genus and (2) to assess evolutionary dynamics of song, in particular whether interspecific song diversification evolved along intraspecific lines of covariation between syllable characteristics. We document a pattern of intense, very labile evolution of song in the genus. Well-known vocal constraints such as the relationship between frequency and body size did not appear to limit this evolution, and intraspecific trade-offs between different aspects of syllable complexity did not constrain interspecific diversification. Factor analysis returned an axis of variation that comprehensively expresses song complexity in the genus, and is defined by song speed, repertoire size, and also various aspects of syllable phonology. The species body size is not related to this axis of complexity, but to a minor axis that may indicate better respiratory capacity of larger birds. We discuss the pattern of song evolution in Serinus spp. in the context of the evolutionary dynamics of sexually selected traits.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 183–194.     

Character divergences and convergences in canopy-dwelling Loranthaceae

The Loranthaceae is the largest plant family with aerial branch parasites termed mistletoes. Three genera of Loranthaceae are terrestrial root parasites and the remaining 72 genera are aerial parasites. Several characters, including habit, haustorial type, germination pattern, pollen morphology, chromosome number, inflorescence morphology and flower merosity, fusion, symmetry and size, are considered to reflect evolutionary relationships within the family. Convergence is a common evolutionary pattern and can confound interpretations of evolution. We investigated character evolution by mapping character states onto a phylogenetic tree based on the nuclear ITS and chloroplast trnL–trnF regions. Convergences in form were found in several characters, including habit, haustorial type, flower symmetry and merosity. These convergences typically correspond to ecological parameters such as pollination syndrome or stresses associated with the canopy habit. Other characters such as chromosome number and germination pattern illustrate divergent evolution among clades.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 101–113.     

Ontogenetic transition from unicuspid to multicuspid oral dentition in a teleost fish: Astyanax mexicanus, the Mexican tetra (Ostariophysi: Characidae)

Teleost fishes display a remarkable diversity of adult dentitions; this diversity is all the more remarkable in light of the uniformity of first-generation dentitions. Few studies have quantitatively documented the transition between generalized first-generation dentitions and specialized adult dentitions in teleosts. We investigated this transition in the Mexican tetra, Astyanax mexicanus (Characidae), by measuring aspects of the dentition in an ontogenetic series of individuals from embryos to 160 days old, in addition to adults of unknown age. The first-generation dentition and its immediate successors consist of small, unicuspid teeth that develop extraosseously. Multicuspid teeth first appear during the second tooth replacement event, and are derived from single tooth germs, rather than from the fusion of multiple conical tooth germs. We document that the transition from unicuspid to multicuspid teeth corresponds to a change in the location of developing tooth germs (from extraosseous to intraosseous) and in patterns of tooth replacement (from haphazard to simultaneous within a jaw quadrant). In addition, while the size of the largest teeth scales with positive allometry to fish size, the transition to multicuspid teeth is accompanied by an exceptionally large increase in tooth size.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 523–538.     

Modelling intraspecific resting egg shape variation in a freshwater fairy shrimp Tanymastix stagnalis (L., 1758) (Crustacea, Branchiopoda)

The shape of the resting eggs of a large branchiopod crustacean, the Anostraca Tanymastix stagnalis , is represented very accurately by analytical expressions. The occurrence of atypical shape of some T. stagnalis eggs may be viewed as a simple change of the analytical expression describing the usual egg shape. Their unusual shape may be explained by a higher embryo volume within an envelope of a given size. Biological implications are briefly discussed and hypothesized in an evolutionary point of view.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 55–60.     

Disentangling composite colour patterns in a poison frog species

A phylogenetic approach was performed to infer whether variation in conspicuous colour-patterns of a poison frog (Dendrobatidae: Dendrobates tinctorius ) has evolved neutrally or under selection. Colour and pattern were split into components that were separately analysed and subsequently re-grouped via principal component analysis. This revealed four different 'displayed' factors on the dorsal and lateral views versus one 'concealed' factor on the ventral view. Based on the assumption that current patterns of trait variation contain information about the evolutionary history of the phenotype, we correlated these trait components to a neutrally evolving gene fragment (cytochrome b ). The concealed factor was significantly correlated with the marker fragment, which identified it as having evolved under genetic drift. Noncorrelation of all displayed factors with the marker may indicate the evolution of colour patterns on dorsum and flanks under selection. In our example, colour pattern should therefore be regarded as a multicomponent signal system.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 433–444.     

Evolution of skull and body shape in Triturus newts reconstructed from three‐dimensional morphometric data and phylogeny

To explore the relationship between morphological change and species diversification, we reconstructed the evolutionary changes in skull size, skull shape, and body elongation in a monophyletic group of eight species that make up salamander genus Triturus. Their well‐studied phylogenetic relationships and the marked difference in ecological preferences among five species groups makes this genus an excellent model system for the study of morphological evolution. The study involved three‐dimensional imagery of the skull and the number of trunk vertebrae, in material that represents the morphological, spatial, and molecular diversity of the genus. Morphological change largely followed the pattern of descent. The reconstruction of ancestral skull shape indicated that morphological change was mostly confined to two episodes, corresponding to the ancestral lineage that all crested newts have in common and the Triturus dobrogicus lineage. When corrected for common descent, evolution of skull shape was correlated to change in skull size. Also, skull size and shape, as well as body shape, as inferred from the number of trunk vertebrae, were correlated, indicating a marked impact of species' ecological preferences on morphological evolution, accompanied by a series of niche shifts, with the most pronounced one in the T. dobrogicus lineage. The presence of phylogenetic signal and correlated evolutionary changes in skull and body shape suggested complex interplay of niche shifts, natural selection, and constraints by a common developmental system. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 243–255.     

The evolution of release and advertisement calls in green toads (Bufo viridis complex)

In the present paper we tested the hypothesis that differences in the acoustic communication system of diploid and tetraploid green toads ( Bufo viridis complex) might be due to selection for reproductive character displacement. We recorded two acoustic signals of the toad repertoire − the advertisement call (a long range mate-attracting signal) and the release call (a short-range signal mediating male–male interactions) − from six sympatric Central Asian populations (three diploid and three tetraploid populations) as well as from three allopatric diploid populations from Italy, and compared their patterns of variation with the pattern of among-population genetic distances. Although release and advertisement calls share the same morpho-physiological constraints, they show significantly different patterns of variation. Release calls vary congruently with the pattern of genetic distances, suggesting that mutation and genetic drift have been the major forces responsible for their change both in time and space. By contrast, the pattern of advertisement-call variation is not consistent with the phylogeny of the group, because the advertisement calls of Asian diploid and tetraploid populations differ from each other more than their genetic distances would predict. These results strongly support the hypothesis that selection acted on the advertisement calls of either or both Asian taxa , possibly, to favour reproductive isolation.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 379–391.     

Phylogenetic relationships between parthenogens and their sexual relatives: the possible routes to parthenogenesis in animals

In theory, parthenogenetic lineages have low evolutionary potential because they inexorably accumulate deleterious mutations and do not generate much genotypic diversity. As a result, most parthenogenetic taxa occupy the terminal nodes of phylogenetic trees. The rate and mode of development of parthenogenesis are important factors to consider when assessing its costs and benefits since they determine both the level of genetic diversity and the ecological adaptability of the resulting lineages. The origin of parthenogenesis is polyphyletic in many taxa, suggesting that genetic systems maintaining sexuality are often labile. In addition, the loss of sex may be achieved in several ways, leading to parthenogenetic lineages with distinct genetic profiles. This could then influence not only the fate of such lineages in the long term, but also the outcome of competition with their sexual counterparts in the short term. In this paper, we review the possible evolutionary routes to parthenogenesis based on a survey of the phylogenetic relationships between sexual and parthenogenetic lineages in a broad range of animals. We also examine the different mechanisms by which parthenogenetic lineages could arise, and discuss the influence of these mechanisms on both the genetic properties and the ecological life styles of the resulting lineages.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 79 , 151–163.     

Parallel evolution in molar outline of murine rodents: the case of the extinct Malpaisomys insularis (Eastern Canary Islands)

The lava mouse Malpaisomys was part of Pleistocene and Holocene faunas of the eastern Canary Islands; it became extinct during historical times. In order to evaluate the evolutionary processes of this endemic species, we set out to identify its relationships with possible mainland relatives. Its dental morphology was compared to a set of fossil and modern murine rodents from various phylogenetic groups, characterized by different diets and dental patterns, using a quantitative method based on a Fourier analysis of the outline of the first upper and lower molars. This morphometric analysis identified different evolutionary grades that are independent of the phylogenetic group. The first cluster is associated with primitive, asymmetric dental outlines, the second with intermediate forms linked to a more herbivorous diet. Highly differentiated forms diverge not only from the second cluster but also from each other. Our investigations reveal Malpaisomys to have had an intermediate dental pattern, a result which confirms previous palaeoecological interpretations of this taxon. However, conclusions about its closest mainland relative remain tentative. Based on a comparison of dental size and shape, as well as geographical considerations, a possible mainland ancestor could be either Paraethomys (North-African Pliocene) or Occitanomys (South-western European Pliocene). Such results support the hypothesis of a Pliocene colonization event, in which case Malpaisomys would display a normal evolutionary rate in dental size and shape, in spite of the insular context. In contrast, a late colonization by a modern representative of the African fauna would imply exceptionally high evolutionary rates compared to the background morphological evolution and a decrease in size that is unlikely under insular conditions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 555–572.     

Exploring the genomic mysteries of polyploidy in cotton

For several years allopolyploid cottons have been the subject of evolutionary investigations into the genomic mysteries of polyploidy. An array of genomic interactions have been documented, including interlocus concerted evolution, differential rates of genomic evolution and intergenomic sequence transfer. Substantial alterations in gene expression have occurred in response to allopolyploidization, including gene silencing and expression changes that vary by organ. Some of the molecular phenomena occurring in polyploids appear to be non-Mendelian. Many of the genomic and expression alterations have occurred on an evolutionary timescale, whereas others reflect more immediate consequences of genomic merger.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 573–581.     

Evolutionary and ecological implications of genome size in the North American endemic sagebrushes and allies (Artemisia, Asteraceae)

The genome size of 51 populations of 20 species of the North American endemic sagebrushes (subgenus Tridentatae ), related species, and some hybrid taxa were assessed by flow cytometry, and were analysed in a phylogenetic framework. Results were similar for most Tridentatae species, with the exception of three taxonomically conflictive species:  Artemisia bigelovii Gray,  Artemisia pygmaea Gray, and  Artemisia rigida Gray. Genome size homogeneity (together with the high morphological, chemical, and karyological affinities, as well as low DNA sequence divergence) could support a recent diversification process in this geographically restricted group, thought to be built upon a reticulate evolutionary framework. The Tridentatae and the other North American endemic Artemisia show a significantly higher genome size compared with the other subgenera. Our comparative analyses including genome size results, together with different kinds of ecological and morphological traits, suggest an evolutionary change in lifestyle strategy linked to genome expansion, in which junk or selfish DNA accumulation might be involved. Conversely, weed or invasive behaviour in Artemisia is coupled with lower genome sizes. Data for both homoploid and polyploid hybrids were also assessed. Genome sizes are close to the expected mean of parental species for homoploid hybrids, but are lower than expected in the allopolyploids, a phenomenon previously documented to be related with polyploidy.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 631–649.     

Post-Pleistocene evolution of Bornean shrews Crocidura foetida (Mammalia, Soricidae)

Fossil mandibles of the Bornean shrew Crocidura foetida recovered from excavations at the west mouth of Niah cave, Sarawak, Malaysia, show that the late Pleistocene population at this lowland location was comparable in size with the large subspecies Crocidura foetida doriae , presently occurring at inland, upland locations. Two Holocene specimens fall in the size range of the smaller lowland subspecies C. f. foetida . Comparable post-Pleistocene size-reduction is known among other mammals of Borneo, but this is the first instance of dated examples. The evolutionary trend conforms with Bergmann's 'rule' but, other than climate change, no selective agent is apparent.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 413–419.     

Genetic diversity patterns in Curcuma reflect differences in genome size

The relationships between genome size and the systematic and evolutionary patterns in vascular plants are equivocal, although a close relationship between genome size and evolutionary patterns has been previously reported. However, several studies have also revealed the dynamic nature of genome size evolution and its considerable ‘ups’ and ‘downs’. Thus, in this study, the phylogenetic relationships among three previously revealed genome size groups and among species of the highly polyploid genus Curcuma were evaluated using AFLP. Our results suggest two main lineages within Indian Curcuma reflecting evolution of genome size. The first one includes hexaploids and higher polyploids of the previously recognized genome size group I, and the second one includes mainly hexaploids of genome size groups II and III. Within genome size group I, relationships among species seem to be influenced by reticulate evolution and higher polyploids are likely to be of allopolyploid origin. Reproductive systems in Indian Curcuma vary considerably among ploidy levels and these differences considerably affect morphological and genetic variation. In general, clonally reproducing species are expected to exhibit low genotypic diversity, but, at the same time, species of allopolyploid origin are expected to maintain higher levels of heterozygosity compared with their progenitors. We investigated intra‐populational genetic variability in Curcuma spp. to evaluate whether mode of reproduction or ploidy represent the main factor influencing the degree of genetic diversity. We found that hexaploid species exhibited significantly higher genetic diversity than higher polyploids (9x, 15x). Our results suggest that this genetic diversity pattern is largely influenced by the mode of reproduction, as higher polyploids reproduce exclusively vegetatively, whereas hexaploids reproduce mainly sexually. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 388–401.