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.     

Basicranial axis length v. skull length in analysis of carnivore skull shape

Skull length is the measurement most commonly used as a standard against which other aspects of cranial morphology are compared to derive an index of relative size or proportions. However, skull length is composed of two different functional components, facial skull and cerebral skull, which vary independently and have different scaling relationships with body size. An analysis of carnivore skull shape with measurements standardized against basicranium length produced very different results than an analysis using skull length as the standard. For example, expressions of relative size of cranial measurements were reduced by 13% in mustelids and increased by 20% in canids, reflecting removal of jaw length (short in mustelids and long in canids) from the comparative standard (basicranial axis length). Cranial measurements scale with higher allometric exponents against basicranial axis length than against skull length.     

Canine tooth strength and killing behaviour in large carnivores

Bending strength of upper canine teeth is examined among living canids, felids, hyaenids and several extinct species including sabretooth cats, borophagine dogs and the dire wolf, Canis dirus. The tooth is modelled as a cantilever with an elliptical cross-section. Using beam theory, the bending strength of the upper canine is calculated given a constant force applied to the canine tip. Results indicate that felids and hyaenids have relatively stronger canines than canids, particularly in bending about the anteroposterior (AP) rather than the mediolateral axis. It is suggested that canine shape reflects the forces produced during killing and feeding. As shown by an analysis of jaw muscle moment arms, felids and hyaenids have relatively stronger bites than canids. Moreover, the canines of hyaenids and felids are perhaps more likely to contact bone during feeding and killing and consequently may be subjected to larger and more frequent bending moments about the AP axis. The canines of sabretooth cats are shown to be more similar in shape and strength characteristics to those of living canids than felids, whereas those of the borophagine dogs and the dire wolf are closer to modern hyaenas.     

Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.     

Mid-Pliocene Carnivora from the Woranso-Mille Area,Afar Region,Ethiopia

The Woranso-Mille paleontological study area, located in the central Afar region of Ethiopia, is one of the most important Pliocene sites in eastern Africa. Since the Woranso-Mille Research Project began its investigation in 2005, more than 7,500 mammalian fossils, including hominins, have been collected from 80 vertebrate localities. This paper provides a preliminary assessment of the Woranso-Mille carnivoran record, a record that is of great interest given the high level of species richness of African carnivorans during the middle Pliocene. Craniodental and postcranial material of canids, lutrine mustelids, viverrids, herpestids, machairodontine and feline felids, and hyaenids has been recovered. Thus, this diverse fauna includes not only the largest carnivorans from this time period (e.g., Enhydriodon and Homotherium), but also some of the smallest, including mongooses, civets, genets, and felids, some of which represent new species. However, the diversity of small taxa does not yet approach that found in the roughly contemporaneous Upper Laetolil Beds of Tanzania. In contrast, lutrine mustelids are better represented at Woranso-Mille than at Kanapoi (Kenya) or Laetoli (Tanzania), which is to be expected given the diversity of habitats represented at these sites. While more material from these sites and others are necessary to truly understand the increased diversity within the early to middle Pliocene eastern African carnivoran guild, it is clear that the material from Woranso-Mille has the potential to fill many of the gaps in our knowledge of carnivorans during this time period.     

Paragonimus westermani in Malaysian Felidae and Viverridae: probable modes of transmission in relation to host feeding habits

Examination of naturally infected felids and viverrids in Malaysia confirmed previously published records which indicated that P. westermani occurred only in felid cats. Felis planiceps and F. temnickli were reported as new host records. Analysis of stomach contents revealed no crab remains in either family of cats, but confirmed that felids were strictly carnivorous while viverrids were often omnivorous. In feeding experiments, only viverrids ate the host crabs Potomiscus johorensis and Parathelphusa maculata. The probable transmission of P. westermani to felids via paratenic hosts was discussed.     

Sexual dimorphism in the canines and skulls of carnivores: effects of size, phylogency, and behavioural ecology

Sexual dimorphism in craniodental features is investigated in a sample of 45 carnivore species in relation to allometry, phylogeny, and behavioural ecology. Dimorphism is more pronounced in both upper and lower canine size and strength than in carnassial size, skull dimensions and biomechanical features, but all dimorphism indices covary. As with most morphological characters, differences in canine sexual dimorphism are significantly related to phylogeny, estimated from either taxonomic rankings or a limited matrix of molecular distances; in particular, mustelids, felids and procyonids are more dimorphic than other carnivore families. Thus, because of problems related to species dependence in comparative data, remaining analyses are based on phylogenetically transformed values using a spatial autoregressive method.
In contrast to other mammals, sexual dimorphism in carnivore canines is not correlated with differences in body weight, skull length or basicranial axis length. Nor is it correlated with categorical variables of activity pattern, habitat, or diet. In our Carnivore sample, canine dimorphism is related only to breeding system: uni-male, group-living (harem) species have significantly greater canine dimorphism than multi-male, multi-female groups and monogamous pair-bonding species. By contrast, dimorphism in carnassial size is related to dietary differences, specifically greater dimorphism in meat-eating species, and not breeding patterns. Dimorphism in estimates of jaw muscle size suggest functional demands from both diet and breeding type. It is concluded that, befitting patterns of heterodont dentition, sexual selection influences variation in canine dimorphism while feeding ecology is related to carnassial dimorphism.     

Body size and species-richness in carnivores and primates.

We use complete species-level phylogenies of extant Carnivora and Primates to perform the first thorough phylogenetic tests, in mammals, of the hypothesis that small body size is associated with species-richness. Our overall results, based on comparisons between sister clades, indicate a weak tendency for lineages with smaller bodies to contain more species. The tendency is much stronger within caniform carnivores (canids, procyonids, pinnipeds, ursids and mustelids), perhaps relating to the dietary flexibility and hence lower extinction rates in small, meat-eating species. We find significant heterogeneity in the size-diversity relationship within and among carnivore families. There is no significant association between body mass and species-richness in primates or feliform carnivores. Although body size is implicated as a correlate of species-richness in mammals, much of the variation in diversity cannot be attributed to size differences.     

Elbow-joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores

Among the hunting strategies employed by members of the order Carnivora (Mammalia), two, stalk and ambush and sustained pursuit, are particularly prevalent among larger species of the order. It has been difficult to identify morphological traits that support this distinction and ecological observations have shown that most carnivorans adopt a continuum of strategies, depending on available habitat and prey. In this paper, the shape of the distal humerus articulation is analysed, with the aim of exploring the use of the forelimb in prey procurement, and as a guide to such behaviour among fossil carnivorans. The results suggest that manual manipulation and locomotion are conflicting functions. Elbow‐joint morphology supports a division between grapplers (i.e. ambushers) and nongrapplers (i.e. pursuers). Joints of the former are characterized by being relatively wide and the latter, by being relatively narrow and box‐like with pronounced stabilizing features. At intermediate and large body sizes, carnivorans show a pattern suggesting mutually exclusive feeding strategies that involve either grappling with prey or sustained pursuit. The former allows for large body sizes, such as pantherine felids and ursids; the latter includes species of only moderate size, such as hyenids and canids. Elbow‐joint morphology is closely linked to phylogeny, but the morphology of the cheetah converges with that of nongrapplers, showing that strong selective forces may override the phylogenetic component. Two taxa of giant mustelids from the Miocene were analysed to test whether this sort of analysis is applicable to carnivorans of the past. The African Late Miocene species Ekorus ekakeran has a joint morphology comparable to that of modern‐day nongrapplers. Two joint morphologies were found in the North American Late Oligocene‐Early Miocene Megalictis ferox. The first morphology is comparable to that of modern pantherine cats and the second forms an intermediate between grapplers and nongrapplers that is not present in the modern carnivoran fauna. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 91–104.     

Incus facet morphology in carnivorous mammals from different ecosystems: Taxonomy vs. habitat

This study is prompted by the discovery of an incus of Hyaenodon, the first known auditory ossicle of this genus and thus of any hyaenodont mammal so far. A large set of incudes of recent Carnivora, including felids, hyaenids, viverrids, herpestids, nandiniid and canids of different ecosystems, was set up for morphological comparison. This study examines especially the incudo-mallear facet. Typically, the incudo-mallear facet is composed of: (1) three articular surfaces in felids, (2) a U-shaped surface in hyaenids and (3) four surfaces in canids. Both taxonomy (on family level) and habitat (open, closed or mixed habitat preference) might have an impact on the morphology of the incus facets, the former having a major impact in our sample. The Hyaenodon incus is small, delicate and possesses an incudo-mallear facet of a general saddle-shape with two articulation facets, a large superior articulation area and a circular, inferior articulation area. Herein, its general morphology and facet shape is most similar to the felid incus morphology.     

Carnivore olfactory bulb size: allometry, phylogeny and ecology

Olfactory bulb size was measured in 146 species of Carnivora in order to examine whether recently observed functional patterns for overall brain size were similar for component parts of the brain. Comparative measures were analysed in relation to various allometric characters (body, brain and skull size), phylogeny, behaviour and ecology. Olfactory bulbs are significantly and positively correlated with all allometric variables, but indices of skull size correlate slightly more closely than other variables. This probably relates to functional aspects of skull size, facial proportions, and anterior elements of the brain. Phylogenetic associations were examined by two comparative methods: the method of independent contrasts and phylogenetic autoregression. Both revealed similar phylogenetic correlation at generic and familial levels. Using calculated values from either method, relative olfactory bulb size only correlates with zonation among seven behavioural and ecological variables; aquatic otters have smaller bulb sizes than carnivores of other zonal types. This agrees with discussion about the diminution of olfactory communication in aquatic environments. Also, olfactory bulb size correlates with home range size, which is consistent with a recent model on the use of olfaction for foraging in designated home ranges. Generally, comparative differences in olfactory bulb size in carnivores do not associate with functional variables found in other comparative studies. Nevertheless, future analyses of specific brain components in mammals may be more useful than overall brain size for testing evolutionary hypotheses of mammalian brain size.     

Convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint?

Since the canids and felids diverged in the mid‐Eocene or earlier, each family has developed a suite of morphological and behavioural adaptations for obtaining and consuming prey. We here distinguish between prey taxa captured and eaten as a result of these phylogenetic adaptations, and those because they are fortuitously encountered, and argue that such supplementary prey, often opportunistically caught, create a buffer between sympatric, and potentially competitive, canids and felids and thus enhance coexistence. We base our analysis on dietary data derived from the stomach contents of four sympatric canid and felid species in the Free State Province, South Africa (canids: Cape fox Vulpes chama and black‐backed jackal Canis mesomelas; felids: African wild cat Felis silvestris lybica and caracal Caracal caracal), and from results of studies on these species elsewhere in southern Africa. The two canid species preyed heavily on invertebrates, and thus opportunistically, while the felids (especially the caracal) concentrated on mammals, prey they are phylogenetically adapted to capture. Only three species of mammalian prey are shared by the four species. The ratio of opportunistically‐to‐phylogenetically mediated prey taxa used (the O/P ratio) differ between the species, with the black‐backed jackal having the most opportunistically caught taxa in its diet, and the caracal the least. As predicted, a comparison of this data with those from dietary studies of the same species carried out elsewhere indicates that the number of opportunistically obtained prey taxa varies more than those resulting from phylogenetic adaptations. The largest canid had the widest food spectrum (35 prey taxa) while the smallest felid had the most restricted one (11 prey taxa). We argue that using the O/P distinction allows a better understanding of changes in food niche breadth of particular species, especially in xeric areas, and gives a better indication of possible exploitative competition for food by sympatric carnivores than when regarding all prey taxa as actively pursued. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 527–538.     

Cephalisation bei Caniden

Cephalisation in Canidae Studied were the intraspecific and interspecific relationships between brain weight and body weight in Canidae. Inclination values of the allometric lines: intraspecific – 0,25, interspecific 0,571. Most of the studied species have nearly the same degree of cephalisation, deviations from the interspecific allometric line are small; the cephalisation of Nyctereutes and Otocyon is less developed than in the other canid species. There is no difference in cephalisation between most canids and felids. Mustelids show a high variability in cephalisation. Criticized were the data of Bauchot (1985) and Gittleman (1986) on brain weight and body weight.     

Allometry and performance: the evolution of skull form and function in felids

Allometric patterns of skull‐shape variation can have significant impacts on cranial mechanics and feeding performance, but have received little attention in previous studies. Here, we examine the impacts of allometric skull‐shape variation on feeding capabilities in the cat family (Felidae) with linear morphometrics and finite element analysis. Our results reveal that relative bite force diminishes slightly with increasing skull size, and that the skulls of the smallest species undergo the least strain during biting. However, larger felids are able to produce greater gapes for a given angle of jaw opening, and they have overall stronger skulls. The two large felids in this study achieved increased cranial strength by increasing skull bone volume relative to surface area. Allometry of skull geometry in large felids reflects a trade‐off between the need to increase gape to access larger prey while maintaining the ability to resist unpredictable loading when taking large, struggling prey.     

Phylogenetic relationships within caniform carnivores based on analyses of the mitochondrial 12S rRNA gene

The complete 12S rRNA gene of 32 carnivore species, including four feliforms and 28 caniforms, was sequenced. The sequences were aligned on the basis of their secondary structures and used in phylogenetic analyses that addressed several evolutionary relationships within the Caniformia. The analyses showed an unresolved polytomy of the basic caniform clades; pinnipeds, mustelids, procyonids, skunks,Ailurus (lesser panda), ursids, and canids. The polytomy indicates a major diversification of caniforms during a relatively short period of time. The lesser panda was distinct from other caniforms, suggesting its inclusion in a monotypic family, Ailuridae. The giant panda and the bears were joined on the same branch. The skunks are traditionally included in the family Mustelidae. The present analysis, however, showed a less close molecular relationship between the skunks and the remaining Mustelidae (sensu stricto) than between Mustelidae (sensu stricto) and Procyonidae, making Mustelidae (sensu lato) paraphyletic. The results suggest that the skunks should be included in a separate family, Mephitidae. Within the Pinnipedia, the grouping of walrus, sea lions, and fur seals was strongly supported. Analyses of a combined set of 12S rRNA and cytochromeb data were generally consistent with the findings based on each gene.     

Creodonts and carnivores from the Middle Miocene Muruyur Formation at Kipsaraman and Cheparawa,Baringo District,Kenya

The Middle Miocene Muruyur Formation (ca 14.5 Ma), Tugen Hills, Kenya, has yielded a huge creodont and a variety of carnivores ranging in size from mongoose-sized viverrids and herpestids to lion-sized amphicyonids. The fauna partly fills what used to be a major gap in our knowledge of Neogene African carnivores, spanning the period between the better known Early Miocene assemblages of western Kenya and eastern Uganda, and the Late Miocene and Plio–Pleistocene faunas of East Africa. Present in the deposits are Megistotherium, two species of Hecubides, one species of Agnotherium, Herpestes, Vishnuictis, and one or two undetermined felids.     

Variation of cranial and dental measurements and dental correlations in the pampean fox (Dusicyon gymnocercus)

The knowledge of the craniodental variability of South American canids is mainly anecdotal, and the variability and dental correlation of living canids have only been studied in several species of holarctic foxes ( Vulpes vulpes , Vulpes lagopus and Urocyon cinereoargeteus ) and Canis lupus . These studies have shown that the last molars, canines and incisors are the most variable and the central molars and premolars are the least variable, and that correlations are highest among the posterior premolars. This pattern has been accounted for by developmental factors, functional utility and tooth complexity. In this paper the pattern of dental correlation and variability of craniodental measurements in the South American fox Dusicyon gymnocercus is studied, with the aim of checking the extension of the pattern observed in other species and contrasting previous hypotheses. The most variable in D. gymnocercus measurements are some cranial widths, width of the incisors, lower canine and lower last molar, whereas others such as width of the braincase, length of the upper carnassials, and first and second lower and upper molars have lower variability. The premolar region is the most correlated, in terms of values and number of significant correlation coefficients. The results show that measurement error is negatively correlated with size of the variables, which could explain the bias found in this index as earlier authors have suggested, but the pattern captured by this index cannot be completely considered to be an artefact. The observed results confirm that the patterns of variability and correlation found in holarctic foxes are widespread in the family Canidae, and suggest the presence of an evolutionary constraint.     

The structure of the mandibular corpus and its relationship to feeding behaviours in extant carnivorans

Assuming some optimization of bone structure to applied mechanical loadings in vivo , different killing and feeding behaviours in carnivores should be reflected in observed differences in cross-sectional shape of their mandibular corpora. Section moduli are used to gauge the magnitudes of bending moments in the mandibular corpus and, when dentary length is controlled, the magnitudes of forces applied to the corpus. Comparisons are made of section moduli at the P₃P₄ and P₄M₁ interdental gaps among canids, felids and hyaenids; in canids only, the M₁M₂ interdental gap was also studied. Local variations in loadings are identified by comparing the section moduli at adjacent loci along the corpus within each family.
The findings of this study show that the precarnassial corpora of canids and hyaenids have greater strength in bending than the corpora of felids of similar body weight. This is taken to reflect relatively greater bending moments under loading in the corpora of canids and hyaenids due, in part, to their elongate dentaries (relative to body weight). Relative to dentary length , however, the precarnassial corpora of felids and hyaenids have much greater strength in bending than the corpora of canids. These scaling relationships appear to reflect the high customary forces (i.e. not moments) applied to the precarnassial corpora of felids and hyaenids with sustained canine killing bites and with bone ingestion using the premolars, respectively. An increase in bending strength of the corpus caudal to the camassial blade in canids is interpreted to be an adaptation for bone-crushing with the postcarnassial molars.     

甘肃临夏盆地晚中新世鬣狗类群头骨的几何形态测量学及生态形态学分析(英文)

在晚新生代大型食肉目动物中,鬣狗科动物地史分布广、种类和数量多,在中国出现于中中新世至更新世晚期的地层中。形态功能学家将鬣狗科的60多个化石种分为似豺、似狼和似灵猫等生态类群,这些生态类群和现生仅存的3个食骨和1个食虫类群形成鲜明的对比。生态形态学是基于生物形态与生态环境的密切关系研究不同物种形态所代表的生态位的一种方法;研究者依靠化石鬣狗类与现生种类牙齿和头骨整体形态的相似度进行了生态形态(ecomorphology)的分类,但尚未对这些已定的生态形态进行过多变量的头骨形状分析。本文拟采用平面几何形态测量学的方法对晚中新世临夏盆地的鬣狗类进行研究,并探讨以头骨形状划分其生态形态类型的可行性。几何形态测量法是以生物形态的轮廓作为数据的计算方法,虽然目前尚未被国内古生物研究者广泛采用,但自20世纪30年代多变量统计学的理论成熟以来,这一方法的理论基础和软件算法在国外得到迅速发展,至今已成为近代生物学(neontology)和古生物学形态研究领域的一个常用工具。三维几何形态测量分析也因为三维激光扫描仪的普及而变得更简单和直观。然而,化石标本由于埋藏和保存的关系通常会出现变形或残缺不全,使目前几何形态测量方法在古生物学的应用仍主要以平面数据分析为主。平面数据采集的第一步是从与标本某一平面相垂直的角度拍摄数码照片;虽然拍摄平面的选择通常由研究者自定,但一般会选择待研究类群形状变异最具代表性的一面(如,鱼类一般选取侧视)。第二步是形态的数字化,即使用某一种形态测量软件来标出地标点(landmarks;或称界标点、标志点、标点)。地标点的选择对于分析的结果有直接而且重要的影响,一般会选择Bookstein分类中的I型或II型作为地标点的标准。这两种地标点的共同点是可以从解剖学特征上准确地在每一个标本上找到,而无不明确或复现困难的可能性。数字化后的数据在软件中用数学算法叠加起来(superimposition,或称叠合,重叠),以去除原始图片数据中标本位置、角度及其他与几何形态无关的冗余信息。所有的数字化数据叠加起来后,利用软件计算所有标本之间的几何距离。现在常用的软件中都会以平均几何形状作为中心,而且利用类似主成分分析(principal components analysis)的多变量计算来呈现相对扭曲(或称相对反卷)轴(relative warp axis),后者即是可以用来把形态差异视觉化的多变量数据。这些数据可以用平面坐标图来看不同形态的空间分布,也可以用所谓的薄板样条曲线图(thin-plate spline grid)来表示相对形态上的变化。相对扭曲轴所代表的形状数据还经常被拿来与代表几何物体大小的距心值(或称重心距离,centroid size)做回归曲线分析,以便发现种群中某些形态的异速生长模式(allometry)。本文采用几何形态测量的方法对产自和政地区的鬣狗科Hyaenictitherium,Ictitherium,Adcrocuta和旁鬣狗科Dinocrocuta的材料进行研究,分析了这4个属保存完好的头骨标本的侧面形态。与东非大草原现生食肉动物的头骨整体形态分布的比较和分析表明,和政的鼬鬣狗(Ictitherium)和鬣型鼬鬣狗(Hyaenictitherium)的头骨形状分布介于现生斑鬣狗(Crocuta crocuta)、犬科猎狗(Lycaon pictus)和金豺(Canis aureus)之间,为二者似豺生态形态的解释提供了几何形态测量证据。再者,上述两属化石鬣狗的形状分布与现生斑鬣狗的幼年个体形状重叠,表明现生斑鬣狗头骨的发育机制可能是在鼬鬣狗祖先类型的异速生长规律基础上的持续发育,进而演化出现有的粗壮形态。此外,巨鬣狗(Dinocrocuta)和副鬣狗(Adcrocuta)的头骨形状与现生的斑鬣狗在几何形态测量空间内有普遍重叠的现象,指示了这些异时出现的种类具有相似的生态形态,因而可能占据相近的生态位。结果还显示巨鬣狗和斑鬣狗的幼年个体形状相近,以及两者从幼年到成年发育的形状变化过程也具有相似的规律。因而,巨鬣狗和斑鬣狗之间的趋同演化不仅表现在成年头骨的粗壮程度上,而且在幼年发育模式中也存在平行演化现象。现生发育学与行为生态学已经证实,相对其他大型食肉动物,现生斑鬣狗发育粗壮头骨形态的机制不是以增速生长,而是以延长发育期来实现的。由此推断,巨鬣狗的发育期有可能和现生鬣狗相当(35个月),也可能由于具有相对粗壮和巨大的头骨形态,其发育期会延长些。当然,这个新解释仍需要更多的化石数据和发育研究来证实。     

Evolution of community composition in several carnivore palaeoguilds from the European Pleistocene: the role of interspecific competition

This study focuses on ecological processes such as competition or predation from an evolutionary perspective. First, we attempt to test the idea that species with similar feeding requirements tend to coexist by separating morphologically or behaviourally. Then, the Barton–David test was applied to several carnivore communities (felids and canids) separated in time. Although the preservation bias of the fossil record renders our conclusions tentative, the general equal size–ratio pattern in most of the guilds examined indicates that inter-specific competition for prey species seems to be a good candidate to explain the evolution of guild composition and morphological traits throughout the Pleistocene for the two groups considered, felids and canids.