Megafaunal extinctions and the disappearance of a specialized wolf ecomorph

The gray wolf (Canis lupus) is one of the few large predators to survive the Late Pleistocene megafaunal extinctions [1]. Nevertheless, wolves disappeared from northern North America in the Late Pleistocene, suggesting they were affected by factors that eliminated other species. Using skeletal material collected from Pleistocene permafrost deposits of eastern Beringia, we present a comprehensive analysis of an extinct vertebrate by exploring genetic (mtDNA), morphologic, and isotopic (delta(13)C, delta(15)N) data to reveal the evolutionary relationships, as well as diet and feeding behavior, of ancient wolves. Remarkably, the Late Pleistocene wolves are genetically unique and morphologically distinct. None of the 16 mtDNA haplotypes recovered from a sample of 20 Pleistocene eastern-Beringian wolves was shared with any modern wolf, and instead they appear most closely related to Late Pleistocene wolves of Eurasia. Moreover, skull shape, tooth wear, and isotopic data suggest that eastern-Beringian wolves were specialized hunters and scavengers of extinct megafauna. Thus, a previously unrecognized, uniquely adapted, and genetically distinct wolf ecomorph suffered extinction in the Late Pleistocene, along with other megafauna. Consequently, the survival of the species in North America depended on the presence of more generalized forms elsewhere.     

泥河湾山神庙嘴化石点直隶狼埋藏学研究

长期以来,泥河湾盆地的脊椎动物化石研究聚焦于类群演化和生物地层,缺乏对化石埋藏过程和居群生态的专门研究。近年来,山神庙嘴化石点的正规发掘及丰富的化石发现为引入化石埋藏学方法探讨上述问题提供了可能。本文对山神庙嘴化石点直隶狼化石的骨骼部位组成、居群死亡年龄和生物成层过程进行了研究。骨骼部位组成的分析显示直隶狼的不同骨骼部位存在明显的差异保存,但在居群层面可能仍具有相当的可信度。本文借鉴现生灰狼中常用的牙齿磨耗法、犬齿髓腔闭合率法对山神庙嘴直隶狼的死亡年龄进行了判断。直隶狼的死亡年龄以老年为主,死亡方式为衰老、疾病导致的磨耗型死亡,可能对应了较小的生态压力。直隶狼的埋藏方式为快速的原地埋藏,在被埋藏前受到其他食肉动物和湖滨水流的的改造。     

Functional morphology of the feeding apparatus of ten species of Pacific butterflyfishes (Perciformes,Chaetodontidae): an ecomorphological approach

Synopsis The functional morphology of the feeding apparatus and the feeding ecology of an assemblage of ten species of butterflyfishes was investigated using a comparative ecomorphological approach. Behavioral observations in situ and in vivo, morphological measurements of fresh-killed specimens, scanning electron microscopy, and kinematic modeling were utilized. The fishes show varying degrees of morphological and behavioral specialization and generalization. The more specialized species group according to how they feed, rather than what they feed on. The feeding guild concept is therefore not very instructive in understanding the functional significance of the feeding apparatus. Many of the morphologically specialized butterflyfishes demonstrate evolutionary convergence in feeding morphology. Whereas the more morphologically specialized species do generally have more ecologically specialized diets, these data do not particularly support the ecomorphological hypothesis in that similar diets do not correspond to similar morphologies.     

Phylogeny of the large extinct South American Canids (Mammalia,Carnivora, Canidae) using a “total evidence” approach

South America currently possesses a high diversity of canids, comprising mainly small to medium‐sized omnivorous species, but in the Pleistocene there were large hypercarnivorous taxa that were assigned to Protocyon spp., Theriodictis spp., Canis gezi, Canis nehringi and Canis dirus. These fossils have never been included in phylogenies based on quantitative cladistics, but hand‐constructed cladograms published in the 1980s included some of them in the South American canine clade and others in the Canis clade. In this work, the phylogenetic position of the large extinct South American canids was studied using a large sample of living and extinct canids, as well as different sources of characters (e.g. DNA and 133 osteological characters). The phylogenetic analysis corroborates the inclusion of Theriodictis and Protocyon in the “South American clade”, where C. gezi is also included. In addition, the position of C. dirus as a highly derived Canis species is confirmed. The simultaneous analysis supports hypercarnivory having arisen at least three times in Caninae and once in the “South American clade”. The combination of the phylogenetic analyses, the fossil record and divergence dates estimated in previous works suggests that at least three or four independent lineages of the “South American clade” invaded South America after the establishment of the Panama bridge around 3 million years ago, plus other events corresponding to the immigration of Urocyon and Canis dirus.
© The Willi Hennig Society 2009.     

Understanding the ecology of mammalian carnivorans and herbivores from Valdegoba cave (Burgos, northern Spain) through stable isotope analysis

Stable carbon and oxygen isotope analyses of mammalian carnivoran and herbivore species from the late Pleistocene Valdegoba cave site in northern Spain imply competition and partitioning in resource use. In general, the data support the previously recognized ecology for the analyzed species. δ¹³C values show that the ecosystem around the cave was dominated by C₃ plants. The observed δ¹⁸O values are similar to what is found in modern environments. The analyzed bovids, Bos primigenius, Capra pyrenaica, and Rupicapra rupicapra, showed the most positive δ¹³C values. Bos primigenius had the most positive mean carbon isotope value and is suggested to feed on grasses in open environments. Values for Capra pyrenaica primarily indicate grass feeding, while Rupicapra rupicapra had the widest diet for the analyzed species, likely including grass and browse. Cervus elaphus, Equus ferus, Equus hydruntinus, and Stephanorhinus hemitoechus displayed more negative δ¹³C values indicating the use of similar resources. The smallest species analyzed, Castor fiber, displayed the most negative δ¹³C and δ¹⁸O values, implying a preference for eating C₃ plants and being semi-aquatic. The canids, Canis lupus and Vulpes vulpes, displayed the most positive δ¹³C and δ¹⁸O values, and overlap many of the sampled ungulate species. Positive δ¹⁸O values in canids implies that this group obtains much of its water from its prey, uses a different water source, or has physiological differences from the other carnivorans that influence oxygen isotope values. Lynx pardinus had values similar to the canids. Crocuta crocuta had δ¹³C values more negative than expected for a generalist predator. These values are likely due to concentration of diet on taxa from more forested environments. The most negative δ¹³C values are observed in the bears, Ursus arctos and Ursus spelaeus. These values are the likely result of hibernation and the inclusion of significant vegetation in bear diets.     

Canid Coprolites from the Late Pleistocene of Hidalgo,Central Mexico: Importance for the Carnivore Record of North America

We present the first reported occurrence of canid coprolites from the late Pleistocene of central Mexico. The sample consists of five associated coprolites recovered from Quaternary deposits that crop out in southeastern Hidalgo, central Mexico. The material shows several of the typical features of canid feces, including elongated cylindrical shape, uncommon and nondistinct constrictions, and one tapered end. The coprolites are similar in size to feces of North American foxes, such as Vulpes macrotis, V. lagopus, V. vulpes, and Urocyon cinereoargenteus; likewise, their size might be comparable to those of the Pleistocene canid Canis cedazoensis. The content of the studied coprolites consists of numerous teeth and bone fragments referable to the pocket gophers Pappogeomys or Cratogeomys. The coprolite content is related to a mostly carnivorous diet comparable to that of C. cedazoensis, wolves (Canis lupus, C. rufus, and C. dirus), and some foxes (V. macrotis and V. velox). As a result, it is proposed that the potential trace maker of the coprolites could be a form similar to a carnivorous fox or a small dog. These trace fossils represent the first indirect evidence of canids in the late Pleistocene of Hidalgo and provide additional information regarding the geographic distribution of the Canidae in temperate areas of North America that now are part of central Mexico during the second half of the Pleistocene.     

Phylogeny of the Caninae (Carnivora): Combining morphology,behaviour, genes and fossils

Phylogenetic relationships among 36 Recent and 42 extinct species of the Caninae (Canidae) were analysed, based on 360 morphological, developmental, ecological, behavioural and cytogenetic characters and 24 mitochondrial and nuclear markers. Primary phylogenetic analyses were accompanied by experimental analyses based on various combinations of data partitions and taxon samples. Leptocyon was recovered as a paraphyletic stem lineage of the Caninae; monophyly/paraphyly of the fox‐like canids (Vulpini) remains uncertain; Urocyon and Metalopex form a clade, possibly sister to all non‐Leptocyon canids; Otocyon, Nyctereutes and Nurocyon form a clade; dog‐like canids (Canini) are monophyletic (with South American Cerdocyonina and Afro‐Holarctic Canina); all South American hypercarnivores (Canis gezi, Protocyon, Speothos, Theriodictis) form a clade, close to Chrysocyon and Dusicyon; Canis arnensis, C. ferox, C. thooides, C. lepophagus and Eucyon spp. are basal to the Canina; Lycaon is an isolated African hypercarnivore; Cuon and its relatives (Xenocyon, possibly also Canis antonii, C. falconeri and Cynotherium) form a clade close to Canis s. str.; C. edwardii–C. etruscus–C. mosbachensis–C. palmidens–C. variabilis and hypercarnivorous Canis armbrusteri–C. dirus clades belong to Canis s. str. As the highly homoplastic morphological characters connected to dietary biology are the prominent characters available for the key fossil species, we conclude that macroevolutionary and palaeoecological analyses of the extinct and extant Caninae were to some extent compromised by the phylogenies used.     

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.     

The predatory behaviour of the thylacine: Tasmanian tiger or marsupial wolf?

The extinct thylacine (Thylacinus cynocephalus) and the extant grey wolf (Canis lupus) are textbook examples of convergence between marsupials and placentals. Craniodental studies confirm the thylacine's carnivorous diet, but little attention has been paid to its postcranial skeleton, which would confirm or refute rare eyewitness reports of a more ambushing predatory mode than the pack-hunting pursuit mode of wolves and other large canids. Here we show that thylacines had the elbow morphology typical of an ambush predator, and propose that the 'Tasmanian tiger' vernacular name might be more apt than the 'marsupial wolf'. The 'niche overlap hypothesis' with dingoes (Canis lupus dingo) as a main cause of thylacine extinction in mainland Australia is discussed in the light of this new information.     

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.     

Effects of diet on female fecundity and larval development in the carrion beetle Necrophila japonica

The carrion beetle subfamily Silphinae (Coleoptera: Silphidae) contains dominant macroinvertebrates of soil ecosystems in temperate zones. However, their feeding habits, which determine the role of each species in the ecosystem, have not been sufficiently studied. Moreover, although a diet shift from necrophagy on vertebrate carcasses to predatory feeding on invertebrates is known to occur in this subfamily, the processes and mechanisms of this shift have also been inadequately addressed. We examined female fecundity and larval development on various diets in a Silphinae species, Necrophila (Eusilpha) japonica (Motschulsky). The experimental diets included a meat diet and various invertebrate diets, which reflect the ‘ancestral’ feeding habit in Silphinae, necrophagy, and the ‘derivative’ feeding habit, predatory feeding. Female fecundity was significantly higher on the meat diet (minced beef) than on an insect larvae diet (mealworms and dipteran larvae) but did not significantly differ from that on an earthworm diet. Larval developmental performance was significantly higher on the earthworm diet than on the meat and insect larvae diets. Our results for larval development were consistent with those of previous stable isotope analyses of the same species, in which isotopic values of larval samples agree with those of hypothetical consumers that utilize earthworms. The consistency of results among different methods indicates that N. japonica larvae are most likely earthworm feeders. In contrast, our results for the female fecundity experiment differed from those of previous stable isotope analyses, in which vertebrate carcasses unlikely serve as the staple diet of adults in the field; thus, the feeding habits of N. japonica adults remain unresolved. Our observations that females and larvae performed best on the meat and earthworm diets, respectively, may indicate that, in Silphinae, the diet shift from necrophagy to predatory habits occurs earlier in larvae than in adults.     

Biotic responses of canids to the terminal Pleistocene megafauna extinction

Trophic downgrading is a major concern for conservation scientists. The largest consumers in many ecosystems have become either rare or extirpated, leading to worry over the loss of their ecosystem function. However, trophic downgrading is not a uniquely modern phenomenon. The extinction of 34 genera of megafauna from North America ～13 000 yr ago must have led to widespread changes in terrestrial ecosystem function. Studies that have examined the event address impacts on vegetative structure, small mammal communities, nutrient cycling, and fire regimes. Relatively little attention has been paid to community changes at the top of the food chain. Here, we examine the response of carnivores in North America to the Pleistocene extinction. We employ fossil data to model the climatic niche of endemic canids, including the extinct dire wolf Canis dirus, over the last 20 000 yr. Quantifying the abiotic niche allows us to account for expected changes due to climate fluctuations over the Late Quaternary; deviations from expected responses likely reveal influences of competition and/or resource availability. We quantify the degree of niche conservatism and interspecific overlap to assess species and community responses among canids. We also include in our analyses a novel introduced predator, the domestic dog Canis lupus familiaris, which accompanied humans into the New World. We find that endemic canid species display low fidelity to their climatic niche through time, We find that survivors increasingly partition their climatic niche throughout the Holocene and, surprisingly, do not expand into niche space presumably vacated by the extinction of very large carnivores. These results suggest that loss of megaherbivores and competition with humans likely outweighed advantages conferred from the loss of very large predators. We also find that wolves and dogs decrease their niche overlap throughout the Holocene, suggesting a distinctive relationship between dogs and man.     

Assessing the prevalence of hybridization between sympatric Canis species surrounding the red wolf (Canis rufus) recovery area in North Carolina

Predicting spatial patterns of hybridization is important for evolutionary and conservation biology yet are hampered by poor understanding of how hybridizing species can interact. This is especially pertinent in contact zones where hybridizing populations are sympatric. In this study, we examined the extent of red wolf (Canis rufus) colonization and introgression where the species contacts a coyote (C. latrans) population in North Carolina, USA. We surveyed 22,000km(2) in the winter of 2008 for scat and identified individual canids through genetic analysis. Of 614 collected scats, 250 were assigned to canids by mitochondrial DNA (mtDNA) sequencing. Canid samples were genotyped at 6-17 microsatellite loci (nDNA) and assigned to species using three admixture criteria implemented in two Bayesian clustering programs. We genotyped 82 individuals but none were identified as red wolves. Two individuals had red wolf mtDNA but no significant red wolf nDNA ancestry. One individual possessed significant red wolf nDNA ancestry (approximately 30%) using all criteria, although seven other individuals showed evidence of red wolf ancestry (11-21%) using the relaxed criterion. Overall, seven individuals were classified as hybrids using the conservative criteria and 37 using the relaxed criterion. We found evidence of dog (C. familiaris) and gray wolf (C. lupus) introgression into the coyote population. We compared the performance of different methods and criteria by analyzing known red wolves and hybrids. These results suggest that red wolf colonization and introgression in North Carolina is minimal and provide insights into the utility of Bayesian clustering methods to detect hybridization.     

Ecological Changes in Coyotes (Canis latrans) in Response to the Ice Age Megafaunal Extinctions

Coyotes (Canis latrans) are an important species in human-inhabited areas. They control pests and are the apex predators in many ecosystems. Because of their importance it is imperative to understand how environmental change will affect this species. The end of the Pleistocene Ice Age brought with it many ecological changes for coyotes and here we statistically determine the changes that occurred in coyotes, when these changes occurred, and what the ecological consequences were of these changes. We examined the mandibles of three coyote populations: Pleistocene Rancho La Brean (13–29 Ka), earliest Holocene Rancho La Brean (8–10 Ka), and Recent from North America, using 2D geometric morphometrics to determine the morphological differences among them. Our results show that these three populations were morphologically distinct. The Pleistocene coyotes had an overall robust mandible with an increased shearing arcade and a decreased grinding arcade, adapted for carnivory and killing larger prey; whereas the modern populations show a gracile morphology with a tendency toward omnivory or grinding. The earliest Holocene populations are intermediate in morphology and smallest in size. These findings indicate that a niche shift occurred in coyotes at the Pleistocene/Holocene boundary – from a hunter of large prey to a small prey/more omnivorous animal. Species interactions between Canis were the most likely cause of this transition. This study shows that the Pleistocene extinction event affected species that did not go extinct as well as those that did.     

The cryptic African wolf: Canis aureus lupaster is not a golden jackal and is not endemic to Egypt

The Egyptian jackal (Canis aureus lupaster) has hitherto been considered a large, rare subspecies of the golden jackal (C. aureus). It has maintained its taxonomical status to date, despite studies demonstrating morphological similarities to the grey wolf (C. lupus). We have analyzed 2055 bp of mitochondrial DNA from C. a. lupaster and investigated the similarity to C. aureus and C. lupus. Through phylogenetic comparison with all wild wolf-like canids (based on 726 bp of the Cytochrome b gene) we conclusively (100% bootstrap support) place the Egyptian jackal within the grey wolf species complex, together with the Holarctic wolf, the Indian wolf and the Himalayan wolf. Like the two latter taxa, C. a. lupaster seems to represent an ancient wolf lineage which most likely colonized Africa prior to the northern hemisphere radiation. We thus refer to C. a. lupaster as the African wolf. Furthermore, we have detected C. a. lupaster individuals at two localities in the Ethiopian highlands, extending the distribution by at least 2,500 km southeast. The only grey wolf species to inhabit the African continent is a cryptic species for which the conservation status urgently needs assessment.     

Ecomorphological patterns of aerial feeding in oscines (Passeriformes: Passeri)

The hindlimb (myology and osteology) of swallows (Hirundinidae) is studied and compared with that of seven other passerine families to identify ecomorphological patterns. Muscular and osteological differences are found among swallow species and associations between morphology and foraging technique are examined. We explain morphological differences found in hirundinids as adaptations favouring flexion and adduction of the legs in these aerial foragers, which devote very little time to cursorial locomotion. This adaptive hypothesis is tested using a phylogenetic approach on the basis of an available molecular phylogenetic hypothesis. A clear ecomorphological pattern emerges relating foraging behaviour and pelvic morphology in hirundinids: aerial feeding technique is correlated with short distal leg segments, a large pelvis, a medial insertion of M. iliotibialis cranialis, an absence of pars accessoria of M. flexor cruris lateralis and a fused M. pubo-ischiofemoralis.     

Convergence in trophic morphology and feeding performance among piscivorous natricine snakes

Piscivory has independently evolved numerous times amongst snakes, and therefore these animals provide a powerful opportunity to test for convergent evolution in a vertebrate feeding system. In this study, we integrate performance trials with comparative methods to test the hypothesis that piscivory drives convergence in trophic morphology and feeding performance among natricine snakes. Within and across species, increasing the relative length of the quadrate bone in the skull is positively and strongly linked to a reduction in the time needed to swallow large fish prey. Thus, our feeding experiments suggest that a longer quadrate bone enables snakes to better conform their head shape to the shape of the prey during swallowing. Ancestral diet reconstructions and phylogenetically corrected multiple regression analyses further reveal that evolutionary increases in piscivory are coupled to the evolution of relatively longer quadrates, and hence improved feeding performance on fish prey in these animals. The exploitation of similar dietary niches drives the evolution of convergent trophic morphologies and feeding performances in natricine snakes.     

History of canids in Chile and impacts on prey adaptations

Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe), are reported to be very susceptible to predation by pack‐hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack‐hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies, and hunting behavior. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviors of guanacos: predator inspection of ambush predators, for example, Puma concolor, and rushing at and kicking smaller cursorial predators, for example, Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group‐hunting canids east of the Andes, there were no native species creating group‐hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group‐hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.