Convergence and divergence in the evolution of cat skulls: temporal and spatial patterns of morphological diversity

Background

Studies of biological shape evolution are greatly enhanced when framed in a phylogenetic perspective. Inclusion of fossils amplifies the scope of macroevolutionary research, offers a deep-time perspective on tempo and mode of radiations, and elucidates life-trait changes. We explore the evolution of skull shape in felids (cats) through morphometric analyses of linear variables, phylogenetic comparative methods, and a new cladistic study of saber-toothed cats.

Methodology/Principal Findings

A new phylogenetic analysis supports the monophyly of saber-toothed cats (Machairodontinae) exclusive of Felinae and some basal felids, but does not support the monophyly of various saber-toothed tribes and genera. We quantified skull shape variation in 34 extant and 18 extinct species using size-adjusted linear variables. These distinguish taxonomic group membership with high accuracy. Patterns of morphospace occupation are consistent with previous analyses, for example, in showing a size gradient along the primary axis of shape variation and a separation between large and small-medium cats. By combining the new phylogeny with a molecular tree of extant Felinae, we built a chronophylomorphospace (a phylogeny superimposed onto a two-dimensional morphospace through time). The evolutionary history of cats was characterized by two major episodes of morphological divergence, one marking the separation between saber-toothed and modern cats, the other marking the split between large and small-medium cats.

Conclusions/Significance

Ancestors of large cats in the ‘Panthera’ lineage tend to occupy, at a much later stage, morphospace regions previously occupied by saber-toothed cats. The latter radiated out into new morphospace regions peripheral to those of extant large cats. The separation between large and small-medium cats was marked by considerable morphologically divergent trajectories early in feline evolution. A chronophylomorphospace has wider applications in reconstructing temporal transitions across two-dimensional trait spaces, can be used in ecophenotypical and functional diversity studies, and may reveal novel patterns of morphospace occupation.     

Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates

The fossil record is our only direct means for evaluating shifts in biodiversity through Earth''s history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic–Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent ‘Lagerstätten effects’. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous–Palaeogene extinction.     

A procolophonoid reptile with temporal fenestration from the Middle Triassic of Brazil

The small tetrapod Candelaria barbouri, from the Middle Triassic of southern Brazil, is the first example of an owenettid procolophonoid outside Africa and Madagascar. Candelaria barbouri was originally described as a primitive procolophonid; however, a re-examination of the holotype, as well as new material, reveals that C. barbouri is in fact the youngest member of the Owenettidae, extending the chronological range of the group by more than 10 million years. The recognition of C. barbouri as an owenettid points to a broader diversity and distribution for owenettids than hitherto thought. In addition, C. barbouri is the first member of the Owenettidae to exhibit temporal fenestrae, a discovery that draws attention to the significance of this feature in 'anapsid' reptiles.     

Morphological patterns in the skulls of cats

Measurements on the skull and dentition of 159 specimens of 22 species of recent felid were analysed using multivariate statistical methods (principal components analysis and correspondence analysis). The resulting patterns were considered in the light of recent work on the systematics of felids (large cats and small cats were clearly differentiated). Within each group there are subgroups. In the small cat group one subgroup was found to consist of the species Lynx lynx, L. rufus, L. pardina, L. canadensis and possibly Felis manul. Another subgroup consisted of the species F. bengalensis, F. rubiginosa, F. planiceps and F. viverrina. These two subgroups agree well with other work on felid systematics. Subgroups which do not agree so well with other work were also found, and the reasons for this are considered. Within the large cats there are two species, F. concolor and Neofelis nebulosa , which are clearly deviant from the norm. The reasons for this deviation are also considered.     

Patterns and processes in the early evolution of the tetrapod ear

This article reviews some of the latest information on the evolution of the tetrapod ear region as seen in the fossil record. It looks at the changes that can be documented across the fish-tetrapod transition, the patterns that they show and what can be inferred of the processes that brought some of them about. These processes include an increased role for neural crest, and heterochronic processes such as pedomorphosis. The earliest tetrapods show a common pattern of a short stout stapes with a large stapedial foramen, that primitively contacted the palatal bones and probably supported the braincase. Modifications to this pattern can be seen in tandem with changes to the occiput and are bound up with changes to jaw and breathing mechanisms. By the Late Carboniferous, tetrapods had diversified into a range of groups showing a wide variety of otic morphologies, some of which were probably tympanic, while others were not, and some which are very different from those found in extant tetrapods. In amniotes, the evolution of a tympanic ear appears to correlate with consolidation and integration of the occiput to the skull roof. Competing phylogenies suggest different numbers of iterations for the origin of a tympanic ear, but a minimum of four separate occasions is implied.     

Development of limbs in urodeles and the origin of tetrapod limbs

On the basis of studies on serial sections of larval Ranodon sibiricus limbs and published data, the hypothesis of the origin of tetrapod limbs from the biserial archipterygium is proposed. The mesomeres of the central axis of the biserial fin correspond (in proximodistal direction) to the humerus, ulna, ulnare, all carpalia distalia, metacarpale 1, and phalanges of the first digit in the forelimb of caudate amphibians and to the femur, fibula, fibulare, tarsalia distalia, metatarsale 1, and phalanges of the first digit in the hind limb. The preaxial elements of the zygopodium and autopodium, which are positioned proximal to the digital arch, correspond to the preaxial rays of the biserial fin, and digits 2–5 correspond to its postaxial rays. As the fin transformed into the limb, the central axis curved preaxially, forming the digital arch and resulting in partial reduction and fusion of preaxial rays.     

Structure of bone in the skulls of Neanderthal fossils

Close inspection of surface bone in skulls of Neanderthal man reveals weathering cracks extensive enough in one specimen, La Chapelle-aux-Saints, to allow preliminary analysis of major patterns of orientation and to make inferences about functional relationships of structures. The fine structure of the bone of the brow ridges is very different from the rest of the skull in the two adults examined, having a peculiar vermiculate surface pattern. Weathering cracks do not appear in this region. This indicates that Neanderthal brow ridges are not closely related to normal mechanical forces such as chewing exertion. It may, however, give further support to theories of Neanderthal brow ridges as protection for the eyes. The localized structure of bone often differs from region to region, and offers new possibilities for the analysis of both contemporary and fossil forms.     

Evolution of the tetrapod ear: an analysis and reinterpretation

The dominant view of tetrapod otic evolution–the “standard view”–holds that the tympanum developed very early in tetrapod history and is homologous in all tetrapods and that the opercular process of the rhipidistian hyomandibula is homologous to the tympanic process of the stapes in lower tetrapods. Under that view, the labyrinthodont amphibians of the Paleozoic are usually considered ancestral to reptiles, and thus the “otic notch” of labyrinthodonts and the tympanum it presumably contained form the starting-point for middle ear evolution in reptiles. Four problems have classically been identified with the standard view: the differing relationships of the internal mandibular branch of N. VII (chorda tympani) to the processes of the stapes in amniotes and anurans; the differing orientations of the stapes in key fossil and living groups; the location of the tympanum in early fossil reptiles; and the transferral of the tympanum, during the origin of mammals, from the stapes to the articular bone of the lower jaw. An examination of these problems and of the solutions proposed under the standard view reveals the ad hoc, and therefore unsatisfactory, nature of the proposed solutions. To organize and review alternative hypotheses of otic evolution an analytical table is constructed, using three characters (tympanic process, Nerve VII, tympanum), each with two possible states. A total of eight hypotheses about middle ear evolution are possible under this system, one of which is the standard view. The seven “non-standard” hypotheses, only five of which have been argued in the literature, are briefly examined. Six of the “non-standard” hypotheses appear unattractive for various reasons, including reliance on ad hoc arguments. The seventh was first proposed by Gaupp in 1898. It is today almost universally ignored but apparently largely for historical rather than scientific reasons. This hypothesis, her called the “alternative view”, appears to rest on assumptions equally as plausible as those of the standard view. Moreover, it offers a solution of the problems associated with the standard view without, apparently, raising any similarly serious problems. This paper compares the standard and alternative views of middle ear evolution in detail. Comparison proceeds on two levels. On one level, they are compared in terms of the hypotheses of phyletic tetrapod relationships each promotes and how strongly each supports its hypothesis. Both views promote the same hypothesis of tetrapod relationships. The alternative view is the more parsimonious, but the difference is not considered sufficient to provide a choice. On another level, the two views are compared in terms of their implications for: (1) the evolution of relative and absolute auditory perceptive ability; (2) the origin of reptiles; (3) the evolution of the suspensorium and cranial kinesis; and (4) the origin and evolution of recent amphibians. The nature of the data required for a test of the implications of the two views is specified in each case. Where data are available. the alternative view is consistent and the standard view is inconsistent with these data. We conclude that the alternative view is the preferable hypothesis of middle-ear evolution. This conclusion implies the following: the tympanic membranes and the tympanic processes of the stapes in recent mammals, reptiles + birds. and frogs. are not homologous; the evolution of “special periotic systems” in the ancestors of amphibians and amniotes were independent events and preceded the evolution of tympanic membranes; the amphibian tympanic membrane. probably including that of labyrinthodonts. is not ancestral to that of amniotes. and that labyiinthodonts with an otic notch are not suitable as amniote ancestors; the stapes of early reptiles functioned primarily as part of the jaw suspension rather than in hearing; the mechanisms and abilities of sound perception in recent tetrapods are likely to be diverse rather than forming parts of a cline; and the lack of a tympanum in Gymnophiona and Caudata may be a retention of a primitive condition.     

Adaptive radiation through phenological shift: the importance of the temporal niche in species diversification

Abstract 1. Phenological shift in oviposition in seed predators may be a key factor for adaptive radiation if temporal differences lead to less intense competition. 2. This hypothesis was tested at two sites in the French Alps in three sympatric species of larch cone flies grouped into two phenological groups (early and late) differing in adult emergence and oviposition timing by approximately 2 weeks. The present study assessed the intensity of competition within and between groups by measuring four larval traits. Cone traits were measured, and the impact of early species parasitism on cone development was assessed. 3. The occupation of the central axis of a developing cone by one early larva has a strong detrimental effect on cone growth and seed production. However, there was almost no correlation between the variables measured on the cones and on the larvae, suggesting that the resources available were not limiting. 4. Inter‐group competition had no significant effect on early larvae. In contrast, both inter‐ and intra‐group competition had a significant negative effect on late larvae length (–11% and –16% respectively), dry mass (–8% and –23%), and lipid mass (–15% and –26%). The intensity of competition was stronger among larvae in the same phenological group, which is consistent with the hypothesis that shifts in oviposition promote adaptive radiation in larch cone flies by reducing competition among larvae.     

四足动物起源研究的新进展

四足动物的起源是生物进化过程中的关键问题之一。2006年,美国学者Daeschler E B和Shubin N H在加拿大努纳武特地区南部的埃勒斯米尔岛上发掘了一系列距今3.75亿年的Tiktaalik鱼化石。该鱼与希望螈（Elpisto-stegalian）一样均没有背鳍、没有鳃盖、子鳃盖、外肩胛骨;而具有宽大的背腹性明显的扁平颅骨,以及位于颅骨背面的眼睛,还具有成对的额骨、微鼻孔和位于末端的口,并具备了四足动物的一些特征,诸如较大的通气孔、可活动的颈部、伏瓦状排列的肋骨,胸鳍出现了挠骨的分化,并有骨质关节。这一发现填补了从海洋动物到四足动物的空白。     

Technical possibilities of experimentation, registration and surgery for solving chronobiological problems in animal experiments

Common possibilities of keeping and registering in chronobiological experiments and special techniques of surgical operations (pinealectomy, thyroidectomy, and exstirpation of the ganglia cervicalia superiora) are presented with a special chronoendocrinological conception for an experiment to investigate the role of the pineal gland.     

Evolution of the Permian and Triassic tetrapod communities of Eastern Europe

The Permo-Triassic terrestrial and freshwater tetrapod communities of Eastern Europe are reconstructed as food-webs. The Late Permian theriodont-dinocephalian community (Ocher, Mezen, Isheyevo) changes to a latest Permian theriodont-pareiasaur community (North Dvina, Vyazniki). After a major extinction, the Triassic thecodontian-dicynodont communities appear, a lystrosaurid one in the Early Triassic (Lower and ?Upper Vetluga), and a kannemeyerid one in the later Early Triassic (?Yarenga) and the Mid Triassic (Donguz, Bukobay). Similar stages are represented in the evolution of aquatic communities: the Late Permian temnospondyl community (Ocher, Isheyevo), the latest Permian chroniosuchian one (North Dvina, Vyazniki), the Lower and Middle Triassic new temnospondyl one (from Vetluga to Bukobay). The faunal changes in Eastern Europe are mirrored in other parts of the world, although there are some endemic Russian forms.     

Environmental influences in the evolution of tetrapod hearing sensitivity and middle ear tuning

Vertebrates inhabit and communicate acoustically in most natural environments. We review the influence of environmental factors on the hearing sensitivity of terrestrial vertebrates, and on the anatomy and mechanics of the middle ears. Evidence suggests that both biotic and abiotic environmental factors affect the evolution of bandwidth and frequency of peak sensitivity of the hearing spectrum. Relevant abiotic factors include medium type, temperature, and noise produced by nonliving sources. Biotic factors include heterospecific, conspecific, or self-produced sounds that animals are selected to recognize, and acoustic interference by sounds that other animals generate. Within each class of tetrapods, the size of the middle ear structures correlates directly to body size and inversely to frequency of peak sensitivity. Adaptation to the underwater medium in cetaceans involved reorganization of the middle ear for novel acoustic pathways, whereas adaptation to subterranean life in several mammals resulted in hypertrophy of the middle ear ossicles to enhance their inertial mass for detection of seismic vibrations. The comparative approach has revealed a number of generalities about the effect of environmental factors on hearing performance and middle ear structure across species. The current taxonomic sampling of the major tetrapod groups is still highly unbalanced and incomplete. Future expansion of the comparative evidence should continue to reveal general patterns and novel mechanisms.