How common are cranial sesamoids among squamates?

Sesamoids are elements that originate as intratendinous structures due to genetic and epigenetic factors. These elements have been reported frequently in vertebrates, although cranial sesamoids have been recorded almost exclusively in non‐tetrapod Osteichthyes. The only tetrapod cranial sesamoids reported until now have been the transiliens cartilage (of crocodiles and turtles), and another one located in the quadrate‐mandibular joint of birds. Here, we examined seven squamate species using histological sections, dissections of preserved specimens, dry skeletons, cleared and stained specimens, computed tomographies (CT), and report the presence of other cranial sesamoids. One is attached to the cephalic condyle of the quadrate, embedded in the bodenaponeurosis and jaw adductor muscles of Ophiodes intermedius (Anguidae). The other sesamoid is found at the base of the basicranium of several squamates, capping the sphenoccipital tubercle, on the lateral side of the basioccipital–basisphenoid suture. This bone has previously been reported as “element X.” We reinterpret it as a basicranial sesamoid, as it is associated with tendons of the cranio‐cervical muscles. This bone seems to have the function of resisting tension‐compression forces generated by the muscle during flexion the head. This element was previously known in several squamates, and we confirmed its presence in three additional squamate families: Gymnophthalmidae, Gekkonidae, and Pygopodidae. The evidence suggests that cranial sesamoids are a widespread character in squamates, and it is possible that this feature has been present since the origin of the group.     

Sesamoid bones in tuatara (Sphenodon punctatus) investigated with X‐ray microtomography,and implications for sesamoid evolution in Lepidosauria

Sesamoids bones are small intra‐tendinous (or ligamentous) ossifications found near joints and are often variable between individuals. Related bones, lunulae, are found within the menisci of certain joints. Several studies have described sesamoids and lunulae in lizards and their close relatives (Squamata) as potentially useful characters in phylogenetic analysis, but their status in the extant outgroup to Squamata, tuatara (Sphenodon ), remains unclear. Sphenodon is the only living rhynchocephalian, but museum specimens are valuable and difficult to replace. Here, we use non‐destructive X‐ray microtomography to investigate the distribution of sesamoids and lunulae in 19 Sphenodon specimens and trace the evolution of these bones in Lepidosauria (Rhynchocephalia + Squamata). We find adult Sphenodon to possess a sesamoid and lunula complement different from any known squamate, but also some variation within Sphenodon specimens. The penultimate phalangeal sesamoids and tibial lunula appear to mineralize prior to skeletal maturity, followed by mineralization of a sesamoid between metatarsal I and the astragalocalcaneum (MTI‐AC), the palmar sesamoids, and tibiofemoral lunulae around attainment of skeletal maturity. The tibial patella, ulnar, and plantar sesamoids mineralize late in maturity or variably. Ancestral state reconstruction indicates that the ulnar patella and tibiofemoral lunulae are synapomophies of Squamata, and the palmar sesamoid, tibial patella, tibial lunula, and MTI‐AC may be synapomorphies of Lepidosauria. J. Morphol. 278:62–72, 2017. ©© 2016 Wiley Periodicals,Inc.     

Sesamoids in tetrapods: the origin of new skeletal morphologies

Along with supernumerary bones, sesamoids, defined as any organized intratendinous/intraligamentous structure, including those composed of fibrocartilage, adjacent to an articulation or joint, have been frequently considered as enigmatic structures associated with the joints of the skeletal system of vertebrates. This review allows us to propose a dynamic model to account for part of skeletal phenotypic diversity: during evolution, sesamoids can become displaced, attaching to and detaching from the long bone epiphyses and diaphysis. Epiphyses, apophyses and detached sesamoids are able to transform into each other, contributing to the phenotypic variability of the tetrapod skeleton. This dynamic model is a new paradigm to delineate the contribution of sesamoids to skeletal diversity. Herein, we first present a historical approach to the study of sesamoids, discussing the genetic versus epigenetic theories of their genesis and growth. Second, we construct a dynamic model. Third, we present a summary of literature on sesamoids of the main groups of tetrapods, including veterinary and human clinical contributions, which are the best‐studied aspects of sesamoids in recent decades. Finally, we discuss the identity of certain structures that have been labelled as sesamoids despite insufficient formal testing of homology. We also propose a new definition to help the identification of sesamoids in general. This review is particularly timely, given the recent increasing interest and research activity into the developmental biology and mechanics of sesamoids. With this updated and integrative discussion, we hope to pave the way to improve the understanding of sesamoid biology and evolution.     

An Automated Technique for the Radiographic Determination of Bone Age

We have developed an automated procedure for determining bone-age in the monkey, Macaca fascicularis. The status of epiphysial junctions and sesamoids were determined from radiographs. Epiphyses were scored as either open or closed, and sesamoids were scored as either present or absent. Bone-age was calculated by a computer program that stored and used a data base of probabilities for epiphysial closure and sesamoid appearance. The simplicity of scoring, combined with the speed and accuracy of the calculations, make this technique potentially adaptable for use in a variety of primate species.     

Postnatal skeletal ontogeny in Callisaurus draconoides and Uta stansburiana (Iguania: Phrynosomatidae).

The mineralization of the skeleton from hatching to near maximum size in two phrynosomatid lizards, Callisaurus draconoides and Uta stansburiana, is described in detail. Observed patterns in the appearance of epiphyseal secondary centers, ossification centers, apophyseal ossifications, and calcifications, the distribution of sesamoids, and the timing of fusions, are compared and contrasted with observations of other squamates available in the literature. Overall, Callisaurus and Uta exhibit an advanced state of ossification in the hatchling relative to other squamate neonates and share a similar sequence of braincase fusions and appearance of secondary centers. Preliminary observations suggest that patterns of postnatal skeletal development are highly conserved and independent of patterns of prenatal morphogenesis, and thus a potentially rich source of character data for systematic investigations.     

The potential utility of postnatalskeletal developmental patterns in squamate phylogenetics

Postnatal patterns of skeletal development, includingthe sequence of appearance of ossification centres and the distributionof sesamoids, appear to be highly conserved species-level phenomenain squamates. As such, they are a potential source of charactersfor phylogenetic inquiry. These patterns, from 21 species representing14 crown squamate clades, form the basis for two analyses. In thefirst, the sequence of postnatal skeletal events is coded as charactersusing the sequence unit approach. This analysis reveals that thesequence of postnatal skeletal events might be useful for determiningrelationships at or above the level of crown clades, but not amongthem. The second analysis utilizes discrete data from postnatalskeletal development, such as the presence/absence of sesamoidsand the number of secondary centres in epiphyseal cartilages. Thesediscrete data appear capable of recovering the deeper divergenceswithin Squamata, but evolve too slowly to be informative at thelevel of crown clades. Thus, patterns of postnatal skeletal developmenthave the potential to help illuminate relationships throughout thesquamate tree. Further progress in this area will require the examinationof additional squamate species, the exploration of alternative codingschemes for developmental sequences, and comparable postnatal datafor Sphenodon .  © 2002 The Linnean Societyof London, Zoological Journal of the Linnean Society, 2002, 136 ,277−313.     

The distribution of adult blow-flies (Diptera: Calliphoridae) along an altitudinal gradient in Central Spain

The distribution of Calliphoridae along an altitudinal gradient was investigated in Central Spain using carrion-baited traps. Significant differences were found between elevation and mean abundances of almost all species of blow-flies. Several species of flies could be grouped according to their altitudinal preferences so that samples at high elevations are defined by Calliphora vomitoria and Calliphora vicina while samples at low elevations are defined by two thermophilous species: Lucilia sericata and Chrysomya albiceps. The remaining species show preferences for mid-elevations where wooded areas are more characteristic along the altitudinal gradient. Calliphora vomitoria and Chrysomya albiceps are the most abundant species representing the 87.74 % of all captures. Both species are spatially segregated along the altitudinal gradient. The changing patterns of abundance are discussed in relation to differences in climate conditions along the altitudinal gradient concluding that the environmental variables that influence the seasonality of many species also play an important role to explain the spatial distribution.     

Development of the cypriniform protrusible jaw complex in Danio rerio: Constructional insights for evolution

Studies on the evolution of complex biological systems are difficult because the construction of these traits cannot be observed during the course of evolution. Complex traits are defined as consisting of multiple elements, often of differing embryological origins, with multiple linkages integrated to form a single functional unit. An example of a complex system is the cypriniform oral jaw apparatus. Cypriniform fishes possess an upper jaw characterized by premaxillary protrusion during feeding. Cypriniforms effect protrusion via the kinethmoid, a synapomorphy for the order. The kinethmoid is a sesamoid ossification suspended by ligaments attaching to the premaxillae, maxillae, palatines, and neurocranium. Upon mouth opening, the kinethmoid rotates as the premaxillae move anteriorly. Along with bony and ligamentous elements, there are three divisions of the adductor mandibulae that render this system functional. It is unclear how cypriniform jaws evolved because although the evolution of sesamoid elements is common, the incorporation of the kinethmoid into the protrusible jaw results in a function that is atypical for sesamoids. Developmental studies can show how biological systems are assembled within individuals and offer clues about how traits might have been constructed during evolution. We investigated the development of the protrusible upper jaw in zebrafish to generate hypotheses regarding the evolution of this character. Early in development, the adductor mandibulae arises as a single unit. The muscle divides after ossification of the maxillae, on which the A1 division will ultimately insert. A cartilaginous kinethmoid first develops within the intermaxillary ligament; it later ossifies at points of ligamentous attachment. We combine our structural developmental data with published kinematic data at key developmental stages and discuss potential functional advantages in possessing even the earliest stages of a system for protrusion. J. Morphol. 2010. © 2010 Wiley‐Liss, Inc.     

Dispersal and Predation Behavior in Larvae of <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> and <Emphasis Type="Italic">Chrysomya megacephala</Emphasis> (Diptera: Calliphoridae)

Chrysomya albiceps and Chrysomya megacephala are exotic blowfly species known by producing myiasis in humans and other animals and by transmitting pathogens mechanically. C. albiceps stand out by being a facultative predator of other dipteran larvae. In this paper we investigated the influence of larval predation on the dispersal of larvae of C. albiceps and C. megacephala single and double species for three photophases. An experimental acrylic channel graduated and covered with wood shavings was used to observe the larval dispersal. The results showed that C. albiceps attacks C. megacephala larvae during dispersal and keeps an aggregated pattern close to the release point, in single and double species, independently of the different photophases. Chrysomya megacephala single species exhibited the same pattern, but in double species this was changed to a random distribution.     

Dispersal and Burial Behavior in Larvae of <Emphasis Type="Italic">Chrysomya megacephala</Emphasis> and <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> (Diptera,Calliphoridae)

Blowflies use discrete and ephemeral substrates to feed their larva. After they run out of food, the larvae begin to disperse in order to find adequate places for pupation or additional food sources, a process named post-feeding larval dispersal. Briefly state the aspects and why they are important were studied in a circular arena of 25 cm in diameter and covered with wood shavings to a height of 40 cm allowing post-feeding dispersal from the center of the arena. Larvae of both Chrysomya albiceps and C. megacephala were used in five experiments for each species. For each pupa location, determined as distance from the center, depth, and weight were evaluated. Statistical tests were done to verify the relation between weight, depth and distance for pupation and for larvae of two species shows that the media distance is significantly different for two species and for C. megacephala this distance is greater than the distance for C. albiceps. The depth too is different for each species, as the larvae of C. megacephala buries deeper than C. albiceps. With relation of weight, there is no statistic evidence that have any difference between weights for pupation for each species.     

Foraging behaviour of bird pollinators on Embothrium coccineum (Proteaceae) trees in forest fragments and pastures in southern Chile

Abstract We investigated the effects of forest patch size on the behaviour of birds feeding on the flower nectar of the proteaceous tree Embothrium coccineum J. R. et G. Forster, which is typically restricted to forest edges in agricultural landscapes in southern Chile. We quantified reproductive parameters of trees (no. inflorescences per branch, total and open flowers per inflorescence) in forest fragments varying from 1 ha (small), to 20 ha (medium) and to >150 ha (large), and in remnant trees in pastures. Visits to flowers by nectar‐feeding birds were recorded during 30‐min observation periods, spread throughout the day during two flowering seasons, November 1992 and 1993 (n = 242 periods overall). Aggressive encounters among flower visitors were recorded in 1992. We expected less visits to trees in pastures and small forest patches because abundances of Embothrium's main pollinators, the flycatcher Elaenia albiceps and the hummingbird Sephanoides sephaniodes, decreased in smaller patches. We found, however, that pollinator visiting rates were negatively correlated with forest patch area and were highest for pasture trees. This trend was largely due a decline in the number of visits by E. albiceps, the main flower visitor, in larger patches. Hummingbird visits did not change with forest patch size. Lower visitation rates to flowering trees in larger forest fragments seemed to be a consequence of territorial defence by E. albiceps and were unrelated to differences in floral display.     

Group living in squamate reptiles: a review of evidence for stable aggregations

How sociality evolves and is maintained remains a key question in evolutionary biology. Most studies to date have focused on insects, birds, and mammals but data from a wider range of taxonomic groups are essential to identify general patterns and processes. The extent of social behaviour among squamate reptiles is under‐appreciated, yet they are a promising group for further studies. Living in aggregations is posited as an important step in the evolution of more complex sociality. We review data on aggregations among squamates and find evidence for some form of aggregations in 94 species across 22 families. Of these, 18 species across 7 families exhibited ‘stable’ aggregations that entail overlapping home ranges and stable membership in long‐term (years) or seasonal aggregations. Phylogenetic analysis suggests that stable aggregations have evolved multiple times in squamates. We: (i) identify significant gaps in our understanding; (ii) outline key traits which should be the focus of future research; and (iii) outline the potential for utilising reproductive skew theory to provide insights into squamate sociality.     

Foraging behaviour by an intraguild predator blowfly, <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> (Diptera: Calliphoridae)

Optimal foraging theory assumes that predators use different prey types to maximize their rate of energetic gain. Studies focusing on prey preference are important sources of information to understand the foraging dynamics of Chrysomya albiceps. The purpose of this investigation is to determine the influence of larval starvation in C. albiceps on the predation rate of different prey blowfly species and instars under laboratory conditions. Our results suggest that C. albiceps prefers Cochliomyia macellaria larvae to Chrysomya megacephala under non-starvation and starvation conditions. Nevertheless, predators gained more weight consuming C. macellaria. This result suggests that C. albiceps profit more in consuming C. macellaria rather than C. megacephala. The foraging behaviour displayed by C. abiceps on their prey and the consequences for the blowfly community are also discussed.     

An exploration of differences in the scaling of life history traits with body mass within reptiles and between amniotes

Allometric relationships linking species characteristics to body size or mass (scaling) are important in biology. However, studies on the scaling of life history traits in the reptiles (the nonavian Reptilia) are rather scarce, especially for the clades Crocodilia, Testudines, and Rhynchocephalia (single extant species, the tuatara). Previous studies on the scaling of reptilian life history traits indicated that they differ from those seen in the other amniotes (mammals and birds), but so far most comparative studies used small species samples and also not phylogenetically informed analyses. Here, we analyzed the scaling of nine life history traits with adult body mass for crocodiles (n = 22), squamates (n = 294), turtles (n = 52), and reptiles (n = 369). We used for the first time a phylogenetically informed approach for crocodiles, turtles, and the whole group of reptiles. We explored differences in scaling relationships between the reptilian clades Crocodilia, Squamata, and Testudines as well as differences between reptiles, mammals, and birds. Finally, we applied our scaling relationships, in order to gain new insights into the degree of the exceptionality of the tuatara's life history within reptiles. We observed for none of the life history traits studied any difference in their scaling with body mass between squamates, crocodiles, and turtles, except for clutch size and egg weight showing small differences between these groups. Compared to birds and mammals, scaling relationships of reptiles were similar for time‐related traits, but they differed for reproductive traits. The tuatara's life history is more similar to that of a similar‐sized turtle or crocodile than to a squamate.     

Sarcosaprophagous Diptera assemblages in natural habitats in central Spain: spatial and seasonal changes in composition

The composition and spatial distribution of sarcosaprophagous Diptera assemblages were studied using carrion‐baited traps along a bioclimatic gradient of natural habitats in central Spain throughout the different seasons during 1 year. Calliphoridae and Muscidae were the most abundant families, accounting for, respectively, 41.9% and 35.1% of all Diptera specimens collected. Other abundant families were Heleomyzidae (8.4%), Sarcophagidae (6.9%) and Piophilidae (5.1%). Fly assemblage compositions differed among bioclimatic levels, with Chrysomya albiceps (Wiedemann) (Diptera: Calliphoridae) being the dominant species in mesomediterranean habitats, Muscina levida (Harris) (Diptera: Muscidae) the dominant species in supramediterranean habitats, and Prochyliza nigrimana (Meigen) (Diptera: Piophilidae) the dominant species in oromediterranean habitats. Differences in assemblage composition were also found among seasons. Thermophobic species such as Calliphora vicina Robineau‐Desvoidy (Diptera: Calliphoridae) and some species of Heleomyzidae were well represented during autumn, winter and spring in the three bioclimatic levels sampled. By contrast, thermophilic species such as Ch. albiceps and Lucilia sericata (Meigen) (Diptera: Calliphoridae) and most Muscidae and Sarcophagidae species were more abundant during summer and in mesomediterranean habitats located at lower elevations. Knowledge of the preferences of some species for certain habitats may be of ecological and forensic value and may establish a starting point for further research.     

Reproductive mode evolution in lizards revisited: updated analyses examining geographic,climatic and phylogenetic effects support the cold‐climate hypothesis

Viviparity, the bearing of live young, has evolved well over 100 times among squamate reptiles. This reproductive strategy is hypothesized to allow maternal control of the foetus' thermal environment and thereby to increase the fitness of the parents and offspring. Two hypotheses have been posited to explain this phenomenon: (i) the cold‐climate hypothesis (CCH), which advocates low temperatures as the primary selective force; and (ii) the maternal manipulation hypothesis (MMH), which advocates temperature variability as the primary selective force. Here, we investigate whether climatic and geographic variables associated with the CCH vs. the MMH best explain the current geographical distributions of viviparity in lizards while incorporating recent advances in comparative methods, squamate phylogenetics and geospatial analysis. To do this, we compared nonphylogenetic and phylogenetic models predicting viviparity based on point‐of‐capture data from 20 994 museum specimens representing 215 lizard species in conjunction with spatially explicit bioclimatic and geographic (elevation and latitude) data layers. The database we analysed emphasized Nearctic lizards from three species‐rich genera (Phrynosoma, Plestiodon and Sceloporus); however, we additionally analysed a less substantial, but worldwide sample of species to verify the universality of our Nearctic results. We found that maximum temperature of the warmest month (and, less commonly, elevation and maximum temperature of the driest quarter) was frequently the best predictor of viviparity and showed an association consistent with the CCH. Our results strongly favour the CCH over the MMH in explaining lizard reproductive mode evolution.     

Cannibalistic Behavior and Functional Response in Chrysomya albiceps (Diptera: Calliphoridae)

Chrysomya albiceps is a facultative predator and cannibal species during the larval stage. Very little is known about cannibalism and prey size preference, especially in blowflies. The purpose of this investigation was to determine the influence of prey size and larval density on cannibalism by third-instar larvae of C. albiceps under laboratory conditions. Our results indicate that no cannibalism occurs by third-instar larvae on first- and second-instar larvae, but third-instar larvae do eat second-instar larvae. The functional response on second-instar larvae is consistent with Holling type II. The consequences of consuming second-, compared to first- or third-, instar larvae as well as the implications of cannibalism for the population dynamics of C. albiceps are discussed.     

Ovipositional Behavior in Predator and Prey Blowflies

In this study we analyzed the ovipositional behavior of C. albiceps, C. megacephala and L. eximia in response to previous presence of larvae of different species, both predator and prey. The preference for substrates that previously had had no larvae was predominant for all species. However, the experiments showed that C. megacephala and L. eximia avoid laying eggs principally in patches with previous presence of C. albiceps larvae. The implications of these results for the necrophagous Diptera community dynamics are discussed.