Life in Burrows Channelled the Morphological Evolution of the Skull in Rodents: the Case of African Mole-Rats (Bathyergidae,Rodentia)

African mole-rats are fossorial rodents that consist of five chisel-tooth digging genera (Heterocephalus, Heliophobius, Georychus, Fukomys, and Cryptomys) and one scratch digger (Bathyergus). They are characterized by striking physiological, morphological, and behavioral adaptations intimately related to their subterranean life. The influence of their mode of life in shaping the cranial morphology has yet to be evaluated in comparison to other Ctenohystrica, especially fossorial genera, which include the subterranean genera Spalacopus and Ctenomys. In our study, we seek to determine to what extent subterranean life affects the morpho-functional properties of the skull among fossorial ctenohystricans. 3D geometric morphometric analyses were performed on 277 skulls, encompassing 63 genera of Ctenohystrica, and complemented by biomechanical studies. African mole-rats and other subterranean Ctenohystrica, especially chisel-tooth diggers, have a short snout, a wide cranium with enlarged zygomatic arches, and a strongly hystricognathous mandible. Even if convergences are also manifest between most fossorial Ctenohystrica, subterranean rodents departed from the main ctenohystrican allometric trends in having a skull shape less size-dependent, but under stronger directional selection with intense digging activity as a major constraint. African mole-rats, notably chisel-tooth diggers, show important mechanical advantage for the temporalis muscles favoring higher forces at the bite point, while mechanical advantage of the superficial masseter muscles is lower compared to other Ctenohystrica. If subterranean species can be clearly discriminated based on their skull morphology, the intrinsic mosaic of anatomical characters of each genus (e.g., skull, teeth, and muscles) can be understood only in the light of their ecology and evolutionary history.     

Geometric morphometrics of the skull of various murid rodents (Mammalia, Rodentia): relation of the skull shape to the trophic specialization

Presented is a brief overview of basic principles and notions of geometric morphometrics, a new approach to quantitative analysis of shape variations. This approach is applied to analysis of variation of the skull shape in the sample of 18 muroid genera belonging to the families Cricetidae, Arvicolidae and Gerbillidae. The skull shape is described by landmarks, and skulls are compared by resistant fit and superimposition methods. Under consideration is relation of skull shape to the trophic specialization, to family belonging, and to body size. Axial skull reveals more conspicuous relation to each of the factor analyzed as compared to mandible. Zygomatic region and tooth raw are most differentiable, while change of auditory bulla is of secondary effect. Transition from omnivorous through granivorous to grass-eating specialization involves the same trend in each of the family studied in the case of axial skull but not of mandible. Most dependent of trophic specialization appeared to be shape of axial skull rather then of mandible. Arvicolines are most specific in respect to the skull shape. Shape to size relation, although rather slight, also involves the same zygomatic-dental region. The results obtained indicate probably that geometric morphometrics does reveal variations in the skull shape that are free of the size effect.     

Aquatic adaptations in a Neotropical coral snake: A study of morphological convergence

Micrurus surinamensis is an aquatic member of the genus Micrurus. This species is known for its highly specialized venom and distinctive diet, mostly made of aquatic vertebrates. Here, we explore both external (head and body) and skull shape morphologies in M. surinamensis, comparing it with two terrestrial species of the genus (M. lemniscatus and M. spixii) and to aquatic and terrestrial species of distantly related groups. We use both traditional and geometric morphometrics to determine whether the presence of similar traits in head shape morphology is rather the result of adaptive convergences between M. surinamensis and other aquatic species, or whether it is the product of phylogenetic conservatism within the genus. Results from both traditional and geometric morphometrics show that M. surinamensis can be considered convergent with aquatic species, mainly in the skull shape. Micrurus surinamensis differs from the two terrestrial species of Micrurus by having a wider head, smaller distance between nostrils, and a long tail. Geometric morphometric analysis shows that despite having an extremely conserved skull and mandible shape, M. surinamensis shows a longer supratemporal and quadrate bones than in terrestrial Micrurus, indicating a larger gape for this species. A more kinetic skull combined with a larger gape would allow M. surinamensis to feed on fish, which represent larger and wider prey that contrast with the elongate prey, which compose the main diet of species in the genus Micrurus. Our results illustrate the importance of both phylogenetic conservatism and adaptation in shaping species morphology.     

Evolution of skull shape in carnivores 2. Additional modern carnivores

Fifteen functionally significant aspects of skull morphology were measured on skulls of 36 additional species of carnivores to complete a survey of skull shape in modern fissiped (land) carnivores that includes most of the living genera. The measurements were transformed to dimensionless variables based on the residuals from allometric equations, and were analysed singly and in a 10 variable principal components analysis. An initial study of 62 species of viverrids, canids, mustelids and felids had shown those families to be distinguished from each other by the functionally significant measurements. However, among the additional 36 species, some procyonids, ursids and mustelids display a range of diversity of skull morphology that overlaps that of the other families and diminishes the potential value of the measurements as taxonomic characters. Intraspecific variation is presented for 12 species, and is low enough to allow use of some features as species level diagnostic characters. The lack of correlation between diet and functionally significant aspects of skull morphology among omnivorous carnivores, and the absence of certain skull shapes among carnivores are discussed.     

A New Species of Allosathes Gaud, Atyeo & Berla (Astigmata: Crypturoptidae) on Feathers of the Chilean Tinamou, Nothoprocta perdicaria (Tinamiformes: Tinamidae) in Chile

Allosathes anitae n. sp. Casanueva & González-Acu?a (Astigmata: Crypturoptidae), collected on the Chilean tinamou Nothoprocta perdicaria (Tinamiformes: Tinamidae) from different localities of the ?uble Province, Chile, is described. Comments on its affinities and differences with Allosathes anepiandrius Gaud, Atyeo & Berla are also included.     

Morphometric analysis of six Gerbillus species (Rodentia,Gerbillinae) from Tunisia

Size and shape changes in the skull of the genus Gerbillus were investigated using geometric morphometrics. Six species from Tunisia were studied (G. gerbillus, G. campestris, G. nanus, G. tarabuli, G. simoni and G. latastei). Statistical analyses of shape variability allowed us to discriminate three morphological groups which are congruent with the three groups suggested by previous morphological and molecular studies. However, our results contrast with previous molecular investigations. In fact, according to results obtained by the use of principal component analysis, canonical variate analysis and UPGMA, we found a higher degree of divergence between the subgenus Dipodillus and the other two subgenera Gerbillus and Hendecapleura. This fact suggests that the morphometric differences observed among species within the genus Gerbillus are not mainly related to phylogeny. To reconciliate the molecular and morphological approaches, we propose a hypothesis of differential rates of phenotypic evolution in the genus Gerbillus. In this view, the species belonging to the subgenus Dipodillus evolved apomorphic features of the skull likely related to a higher degree of habitat specialization. By contrast, the more generalist Gerbillus and Hendecapleura subgenera show less differentiated plesiomorphic morphology.     

Variation in Discosauriscus austriacus (Makowsky, 1876) from the Lower Permian of the Boskovice Furrow (Czech Republic)

Two genera and four species of discosauriscids, Discosauriscus potamites, D. pulcherrimus, Letoverpeton austriacum and L. moravicum , have previously been described from the Boskovice Furrow in Moravia. These laxa were differentiated primarily on the basis of proportional differences of the skull together with other morphological characters.
Excavations at two localities in the Boskovice Furrow have produced new, relatively abundant, skeletal material. The specimens are preserved three-dimensionally. In order to evaluate the differences in skull proportions, 17 variable characters on 70 skulls were measured. Statistical analysis demonstrates the existence of several morphological trends in the shape of the skull. However, no trend permits the division of this set of specimens into two clearly differentiated groups. The morphological differences used previously to define genera and species seem to be incorrect. Pending the discovery of distinct morphological differences, the Moravian discosauriscids can be considered as belonging to only one taxon: Discosauriscus austriacus (Makowsky, 1876).     

Variation in the skull morphometry of four taxonomic units of Thrichomys (Rodentia: Echimyidae), from different Neotropical biomes

The echimyid rodents of the genus Thrichomys vary considerably in their behavior and feeding ecology, reflecting their occurrence in environments as different as the Caatinga, Cerrado, Pantanal, and Chaco biomes. While the genus was originally classified as monospecific, a number of Thrichomys species have been recognized in recent decades, based on morphometric, cytogenetic, and molecular analyses. While Thrichomys is well studied, the variation found in its cranial morphology is poorly understood, given the taxonomic and ecological complexities of the genus. Using a geometric morphometric approach, we characterized the differences found in the cranial morphology of four Thrichomys taxonomic units, including three established species, Thrichomys apereoides, Thrichomys fosteri, and Thrichomys laurentius, and one operational taxonomic unit (OTU), Thrichomys aff. laurentius. No significant differences were found among these units in cranium size, but significant variation was found in skull shape. The Procrustes distances provided a quantification of the differences in the shape of the skull, with the largest distances being found between T. aff. laurentius and T. fosteri in the dorsal view, and between T. aff. laurentius and T. apereoides in the ventral view. A Discriminant Function Analysis (DFA) with cross-validation determined that the pairings with the highest correct classification were T. aff. laurentius vs. T. apereoides and T. aff. laurentius vs. T. fosteri, in both views. The principal variation in skull shape was found in the posterior region and the zygomatic arch, which may be related to differences in diet.     

Testing the success of palaeontological methods in the delimitation of clam shrimp (Crustacea,Branchiopoda) on extant species

Fossil spinicaudatan taxonomy heavily relies on carapace features (size, shape, ornamentation) and palaeontologists have greatly refined methods to study and describe carapace variability. Whether carapace features alone are sufficient for distinguishing between species of a single genus has remained untested. In our study, we tested common palaeontological methods on 481 individuals of the extant Australian genus Ozestheria that have been previously assigned to ten species based on genetic analysis. All species are morphologically distinct based on geometric morphometrics (p ≤ 0.001), but they occupy overlapping regions in Ozestheria morphospace. Linear discriminant analysis of Fourier shape coefficients reaches a mean model performance of 93.8% correctly classified individuals over all possible 45 pairwise species comparisons. This can be further increased by combining the size and shape datasets. Nine of the ten examined species are clearly sexually dimorphic but male and female morphologies strongly overlap within species with little influence on model performance. Ornamentation is commonly species-diagnostic; seven ornamentation types are distinguished of which six are species-specific while one is shared by four species. A transformation of main ornamental features (e.g. from punctate to smooth) can occur among closely related species suggesting short evolutionary timescales. Our overall results support the taxonomic value of carapace features, which should also receive greater attention in the taxonomy of extant species. The extensive variation in carapace shape and ornamentation is noteworthy and several species would probably have been assigned to different genera or families if these had been fossils, bearing implications for the systematics of fossil Spinicaudata.     

23种伞形科植物果实形态及其分类学意义

该实验采用徒手切片法对伞形科17属23种植物的果实进行了外部形态和解剖特征的观察,结果表明:伞形科果实有背腹压扁、两侧压扁和不压扁3种类型;侧棱有宽、有窄;油管的分布有棱槽单油管型和棱槽多油管型;花柱基多数为圆锥状,少数种的花柱基为扁平垫状;萼齿明显或不明显。通过进一步对属间和属内果实解剖特征的比较得出:(1)果实表面被钩刺或刚毛及果棱特征在属间差异明显,在属内表现出一致性,可作为伞形科属间分类的依据。(2)果实横切面的形状、胚乳腹面的凹凸以及萼齿的形态特征在属内种间的分类研究中有重要的意义。基于果实形态特征,编制了17属23种植物的分种检索表。     

The Impact of Tooth Wear on Occlusal Shape and the Identification of Fossils of New World Porcupines (Rodentia: Erethizontidae)

Although isolated dental remains are a central source of data in mammalian paleontology, the teeth of many taxa remain unidentifiable at low taxonomic levels because of intraspecific variation. Extant New World porcupines, Erethizontinae, illustrate this problem. Since being declared unusually variable over 50 years ago, molar occlusal morphology has been neglected as a potential source of diagnostic traits. Here, we use geometric morphometrics to re-evaluate the occlusal morphology of the third lower molar of erethizontines as the basis for identification of genera while accounting for variation, particularly changes in shape caused by progressive tooth wear. We model the impact and predictability of this source of variation by using CT-based digital sectioning. We use modelled occlusal shapes to analytically search for traits that can be used to improve identification of isolated dental remains, as well as training machine-learning models to assign occlusal surfaces to genus. Nearly half of all shape variation can be explained by variables such as wear and size. Resulting models of shape predict six univariate traits that may diagnose the two extant genera, especially when specimens are compared at the same wear stage. However, high levels of variation previously noted qualitatively also influence quantitative analyses, rendering machine learning models unreliable. Overall, although the identity of some specimens that are highly worn or contain certain combinations of traits will remain ambiguous, large segments of the gnathic erethizontine fossil record have renewed utility when analyzed using these methods.

     

The pace of morphological change: historical transformation of skull shape in St Bernard dogs

Owing to the great morphological diversity of domestic dogs (Canis familiaris), the study of historical shape change in dog skulls provides an excellent opportunity for investigating the dynamics of morphological evolution. Breed standards make known which features were selected by breeders. Here we use the methods of geometric morphometrics to study change of skull shape in a series of purebred St Bernard dogs spanning nearly 120 years. A regression of shape on time was highly significant and revealed a consistent trend of shape change that corresponded to the features deemed desirable by the breed standard. Historical shape change in St Bernards involves a broadening of the skull and a tilting of the palate and upper jaw relative to the rest of the skull. This trend appears to be linear throughout the entire period and appears to be continuing. Allometry was ruled out as a contributing factor to this change because there was no consistent trend of historical change in skull size and because neither the patterns of static nor ontogenetic allometry corresponded to the historical shape change. The dramatic modification of the St Bernard skull demonstrates that selection can achieve sustained and substantial change and can completely overcome constraints such as allometry.     

Identification of asymmetrically winged samaras from the Western Hemisphere

A key is presented for use in identifying asymmetrically winged fruits (samaras) with either proximal or distal locules. It aids identification based on dispersed fuit morphology and can be used to identify undetermined extant herbarium specimens or fossil fruits to the correct extant family and genus. The 39 genera from 11 families (Aceraceae, Anacardiaceae, Fabaceae, Malpighiaceae, Phytolaccaceae, Polygalaceae, Polygonaceae, Rutaceae, Sapindaceae, Trigoniaceae, Ulmaceae) are distinguished on the basis of wing venation, size of fruit, presence and position of attachment surface, presence and type of subsidiary wings on the ovary wall. ornamentation, size and shape of the ovary, locule position, shape of locule cross section, style position and ornamentation, distinction between ovary wall and wing, and angle of attachment between individual samaras. The developmental origins of some of these features are discussed.     

大蹄蝠头骨形态地理变化

本文采用几何形态测量法对中国大蹄蝠9个不同地理种群头骨形态变化进行研究。结果表明,不同地理种群的头骨大小及形状存在显著差异,其中云南思茅种群与海南陵水种群差异最大。回归分析表明头骨形态的地理变化与气候因素相关。随着年均温度、年均湿度的升高以及年均降水量的增多,大蹄蝠头骨变小,上颌、齿、咬肌附着部分以及耳蜗部分的形状发生变化。此外,头骨大小与海拔高度呈正相关,头骨形状变化与纬度显著相关。本研究表明对栖息地生态条件的适应是中国大蹄蝠头骨形态地理变化的重要原因。     

Heterochronic mechanisms of morphological diversification and evolutionary change in the neotropical salamander,Bolitoglossa occidentalis (Amphibia: Plethodontidae)

Bolitoglossa occidentalis, a lowland salamander of Mexico and Guatemala, has a highly derived morphology. The features that are derived with respect to the condition in generalized members of the genus include the following: (1) small body size; (2) short tail; (3) fully webbed hands and feet; (4) reduction and loss of certain phalangeal elements; (5) fusion of carpals and tarsals; (6) absence of prefrontal bones; and (7) reduced skull ossification. The ontogeny of this species was analyzed quantitatively and compared with the patterns of growth and differentiation encountered in two morphologically generalized members of the genus, B. rostrata and B. subpalmata. Most of the derived features can be explained by invoking a single heterochronic process: truncation of development at a small size (most likely the product of early maturation). Therefore, B. occidentalis is a paedomorphic species whose morphology has been attained through the process of progenesis. This result supports Alberch's ('80a) prediction, based on functional analysis, that the principle adaptation to arboreality in B. occidentalis is small size; other derived morphological features are associated with the organism's truncated development and may have no adaptive significance. However, patterns of dissociation are found within this overall progenetic process. Some of these include the following: (1) accelerated growth rates of the metatarsals and first phalanges, and retarded growth rates of the second and third phalangeal elements; (2) dissociation between rates of ossification of the skull and the autopodial elements; and (3) dissociation between the timing of termination of the process of shape change during the ontogeny of the foot (the product of differential growth between digital and interdigital areas) and termination of growth in overall foot size (foot surface area). This later result illustrates the independence of morphogenetic phenomena (shape change) from processes of growth (size increase).     

Patterns of morphospace occupation and mechanical performance in extant crocodilian skulls: A combined geometric morphometric and finite element modeling approach

Extant and fossil crocodilians have long been divided into taxonomic and/or ecological groups based on broad patterns of skull shape, particularly the relative length and width of the snout. However, these patterns have not been quantitatively analyzed in detail, and their biomechanical and functional implications are similarly understudied. Here, we use geometric morphometrics and finite element analysis to explore the patterns of variation in crocodilian skull morphology and the functional implications of those patterns. Our results indicate that skull shape variation in extant crocodiles is much more complex than previously recognized. Differences in snout length and width are the main components of shape variation, but these differences are correlated with changes in other regions of the skull. Additionally, there is considerable disparity within general classes such as longirostrine and brevirostrine forms. For example, Gavialis and Tomistoma occupy different parts of morphospace implying a significant difference in skull shape, despite the fact that both are traditionally considered longirostrine. Skull length and width also strongly influence the mechanical performance of the skull; long and narrow morphotypes (e.g., Tomistoma) experience the highest amount of stress during biting, whereas short and broad morphotypes (e.g., Caiman latirostris) experience the least amount of stress. Biomechanical stress and the hydrodynamic properties of the skull show a strong relationship with the distribution of crocodilians in skull morphospace, whereas phylogeny and biogeography show weak or no correlation. Therefore, ecological specializations related to feeding and foraging likely have the greatest influence on crocodilian skull shape. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Bentho-Pelagic Divergence of Cichlid Feeding Architecture Was Prodigious and Consistent during Multiple Adaptive Radiations within African Rift-Lakes

Background

How particular changes in functional morphology can repeatedly promote ecological diversification is an active area of evolutionary investigation. The African rift-lake cichlids offer a calibrated time series of the most dramatic adaptive radiations of vertebrate trophic morphology yet described, and the replicate nature of these events provides a unique opportunity to test whether common changes in functional morphology have repeatedly facilitated their ecological success.

Methodology/Principal Findings

Specimens from 87 genera of cichlid fishes endemic to Lakes Tanganyka, Malawi and Victoria were dissected in order to examine the functional morphology of cichlid feeding. We quantified shape using geometric morphometrics and compared patterns of morphological diversity using a series of analytical tests. The primary axes of divergence were conserved among all three radiations, and the most prevalent changes involved the size of the preorbital region of the skull. Even the fishes from the youngest of these lakes (Victoria), which exhibit the lowest amount of skull shape disparity, have undergone extensive preorbital evolution relative to other craniofacial traits. Such changes have large effects on feeding biomechanics, and can promote expansion into a wide array of niches along a bentho-pelagic ecomorphological axis.

Conclusions/Significance

Here we show that specific changes in trophic anatomy have evolved repeatedly in the African rift lakes, and our results suggest that simple morphological alterations that have large ecological consequences are likely to constitute critical components of adaptive radiations in functional morphology. Such shifts may precede more complex shape changes as lineages diversify into unoccupied niches. The data presented here, combined with observations of other fish lineages, suggest that the preorbital region represents an evolutionary module that can respond quickly to natural selection when fishes colonize new lakes. Characterizing the changes in cichlid trophic morphology that have contributed to their extraordinary adaptive radiations has broad evolutionary implications, and such studies are necessary for directing future investigations into the proximate mechanisms that have shaped these spectacular phenomena.