Trends in Proboscidean Diversity in the African Cenozoic

The evolution of the Order Proboscidea has been proposed as a series of adaptive shifts in response to changing environments and evolutionary trends in skull and dentition that were linked to increasing dietary specialization. The assumption that climate and environmental change are the driving forces behind evolution is prevalent in studies of adaptation and diversification. However, there are other forces at work that may be equally responsible for evolutionary change, those involving biotic interaction and competition. In this paper, a summary of the African proboscidean record is presented as a means to examine overall trends in diversity and adaptation. Specimens from 175 localities in East, Central, and Southern Africa were entered into a locality database. Proboscideans from these localities comprise 11 families, 27 genera, and 53 species. First Appearance Datum (FAD) and Last Appearance Datum (LAD) for each species were plotted in 1-million-year intervals. Three periods of diversification were noted, the first in the Oligocene, the second in the early-middle Miocene, and the last in the Pliocene. Three ecomorphological categories based on dental functional morphology were identified in each of the three periods, suggesting that these are stable categories through time. As a result, it is proposed that biotic competition (resulting in resource partitioning) played a more prominent role in proboscidean evolutionary change and diversity than previously thought.     

Review of Japanese Cenozoic (Miocene–Modern) Vertebrate Tracks

One hundred and seventy three Cenozoic vertebrate track sites from Miocene to 1600 A.D have been reported in Japan. Three ichnofaunas can be recognized: a perissodactyl and artiodactyl ichnofauna in the Miocene, an artiodactyl and proboscidean ichnofauna in the Plio-Pleistocene, and human ichnofauna from about 900–800 B.C. to about 1400–1600 A.D. Track data indicate that a predominance of large vertebrates in fluvio-lacustrine environment in lowland changed from perissodactyls to proboscidean through Miocene to Plio-Pleistocene, and ancient people then occupied lowlands instead of large animals. Pes length of proboscidean tracks revealed temporal variation, and the relationship between proboscidean body sizes and tracks was observed. The Cenozoic Japanese proboscidean trackways can be distinguished on the basis of trackway width, as narrow- and wide-gauge, but the difference between of those narrow- and wide-gauge trackways probably indicates generic level differences. The Cenozoic Japanese bird tracks can be identified as four types: ?crane (Family Gruidae?), ?heron (Family Ardeidae?), ?stork (Family Ciconiidae?), and ?shorebird tracks.     

甘肃东乡几种早中新世哺乳动物化石

本文记述了在甘肃东乡椒子沟发现的几种早中新世哺乳动物化石:Gomphotherium sp、Dzungariotheriura orgosense、Rhinocerotidae gen. indet.和Paraentelodon macrognathus sp. nov.,着重讨论了全北区猪齿兽类的系统关系。根据化石的性质判断,东乡椒子沟化石点的地质时代应与欧洲Burdigalian早期,亦即MN_3相当或稍早。笔者将原定为上新世临夏组的第一、二岩性段划出,命名为椒子沟组,其时代为早中新世。     

Valsequillo biostratigraphy. IV: Proboscidean ecospecies in Paleoindian sites

Five proboscidean species have been found in Paleoindian sites from North to South America: two open-country adaptations, Mammuthus columbi and Cuvieronius tarijensis, two woodland and riparian forms, Mammut americanum and Mammuthus jeffersonii, and one tropical savanna species, Haplomastodon chimborazi. Their value in biostratigraphy and as ecological index fossils is discussed with particular emphasis on the Central Mexican Paleoindian sites.     

Enamel Structure of Cuvieronius hyodon (Proboscidea, Gomphotheriidae) with a Discussion on Enamel Evolution in Elephantoids

Dental material of the South American elephantoid Cuvieronius hyodon from the Late Pleistocene of the Tarija Basin, Bolivia was sampled for a comprehensive analysis of the microstructure of the enamel. To examine variability at the dentition level, enamel samples of the upper incisor, second deciduous premolar, and molars were sectioned. The incisor and cheek teeth enamel is compared to that of other proboscideans in order to reveal phylogenetically and functional informative features useful to reconstruct the evolution of elephantoid enamel. Studies of the adaptations and evolution of proboscidean enamel have focused so far on molars. Nevertheless, given the possibility of an independent evolution of the enamel at different tooth positions, the variation of the enamel throughout the dentition needs to be taken into consideration when using enamel microstructural characters to infer proboscidean diversity and phylogeny. The results obtained from this study demonstrate the generality, among elephantoids, of the basic microstructural features of Cuvieronius hyodon enamel, allowing the characterization of the Elephantoid Enamel (EE). The differentiation between incisor and molar enamel seen in elephantoids is shown to represent a primitive elephantiform trait, as it also occurs in Phiomia. The three-layered enamel of the cheek teeth appears as the sole synapomorphy of the Elephantoidea, though the character might be homoplastic within the Proboscidea. Characters of the prisms cross-section might be used, on the other hand, to define less inclusive clades within the Elephantoidea.     

Cochlea anatomy of Numidotherium koholense: auditory acuity in the oldest known proboscidean

Court, N. 1992 04 15: Cochlea anatomy of Numidotherium koholense: auditory acuity in the oldest known proboscidean. Lethaia , Vol. 25, pp. 211–215. Oslo. ISSN 0024–1164.
A natural cast of the spiral cochlea space from the inner ear of the oldest known proboscidean, Numidotherium koholense , is described. The cast was sectioned to reveal internal structures. Cochlea structure in numiodtheres, and by inference auditory acuity, is shown to have been very different from that of living proboscideans. The cochlea of Numidotherium is deemed to reflect tuning to relatively high frequencies and contrasts with the low-frequency specializations seen in the inner ear of living proboscideans. It is suggested that evolution of the elephant otic region might be explained by a shift in developmental timing such that features occurring earlier in ontogeny are characteristic of the adult stage. * Proboscidea, Numidotherium, cochlea, natural cast, auditory acuity .     

甘肃临夏盆地的渐新世巨犀化石

描述了 2种巨犀化石 :霍尔果斯准噶尔巨犀 (Dzungariotheriumorgosense)和牙沟副巨犀(新种 ) (Paraceratheriumyagouensesp .nov.)。新种的主要特征是 :个体小 ;上颊齿齿冠高 ,有薄层白垩质覆盖 ;DP2～M1反前刺大 ,原尖后收缩沟明显 ,次尖有深的前收缩沟 ,中谷和后凹中常有附属小柱等。巨犀的演化历史可能比过去想象的更复杂。Indricotherium、Dzungariotherium和Paraceratherium为代表不同进化水平和支系的 3个属 ,不应合并为Paraceratherium一属。牙沟含巨犀化石层位的地质时代为晚渐新世。     

Reinforcement of mate preference among hybridizing Heliconius butterflies

Recent models of mate preference evolution suggest that direct selection on alleles at preference loci and correlated evolution of preference with locally adapted mating cues are more likely to drive the evolution of assortative mate preference than reinforcement. Mate preference evolution in mimetic Heliconius butterflies has been attributed to all three forms of selection, but here we show that reinforcement has been critical. By examining geographical variation in assortative mating and male mate preference among seven populations of three hybridizing Heliconius species from Costa Rica, we found pronounced character displacement of preference such that sexual isolation was enhanced in areas of interspecific contact. Of the different explanations for the evolution of assortative mate preference, only reinforcement is dependent on interspecific contact in this system. Thus, the observed pattern of reproductive character displacement of mate preference is best explained as a product of indirect selection generated by natural selection against nonmimetic hybrids.     

中国北方上中新统的早期玛姆象属(Mammut)及其在玛姆象科(Mammutidae)分化和演化中的意义

玛姆象属是长鼻类玛姆象科这一重要类群的最终成员.虽然这一属在上新世的欧亚大陆和更新世的北美大陆广泛分布,它早期的进化历史却鲜为人知.报道了中国北方上中新统发现的斜脊玛姆象(相似种)(Mammut cf.M.obliquelophus)的新材料,包括一个几乎完整的幼年头骨,这些材料显示了玛姆象科的许多原始特征,因此很好地解释了玛姆象属形态特征的形成过程.斜脊玛姆象(相似种)具有强烈向两侧扩展的枕部,在门齿窝的基部具有收缩,这些特征与莫罗托始轭齿象(Eozygodon morotoensis)和广河豕脊齿象(Choerolophodon guangheensis)均具有相似性,后两者分别为玛姆象科与豕脊齿象科的早期代表.因此,玛姆象科与豕脊齿象科(Choerolophodontidae)具有近的亲缘关系,二者同位于象形类(Elephantimorpha)系统发育中的基部.支序分析支持了这一结论.     

贵州毕节麻窝口洞与智人伴生的长鼻类化石

最近在贵州毕节麻窝口洞发现了3枚古人类牙齿化石和伴生的哺乳动物群。其中,古人类牙齿经初步研究可归入解剖学上的现代人,而与古人类相伴的大、小哺乳动物化石,经初步鉴定共计8目20科43属53种。本文系统记述了该动物群中大哺乳动物的典型代表——长鼻类化石,共2属2种:东方剑齿象(Stegodon orientalis)和亚洲象(Elephas maximus)。麻窝口洞的长鼻类缺失我国南方早更新世的典型种类——中华乳齿象(Sinomastodon)和华南剑齿象(Stegodon huananensis),具有从典型的中更新世大熊猫-剑齿象动物群(Ailuropoda-Stegodon fauna)向晚更新世亚洲象动物群(Asian elephant fauna)过渡的特征。依动物群的性质和地貌地层的特征,毕节麻窝口洞的智人及伴生动物群的地质时代很可能为中更新世晚期或晚更新世早期,这与堆积物的光释光年代测定的初步结果(距今约11.2-17.8万年)基本吻合。麻窝口洞东方剑齿象与亚洲象的组合明显具有东洋界亚热带动物群的特点,指示温暖潮湿的气候,这些长鼻类与智人等生存于近水的森林和灌丛中,并镶嵌了一些草地。     

Evolution of specialization in resource utilization in structured metapopulations

We study the evolution of resource utilization in a structured discrete-time metapopulation model with an infinite number of patches, prone to local catastrophes. The consumer faces a trade-off in the abilities to consume two resources available in different amounts in each patch. We analyse how the evolution of specialization in the utilization of the resources is affected by different ecological factors: migration, local growth, local catastrophes, forms of the trade-off and distribution of the resources in the patches. Our modelling approach offers a natural way to include more than two patch types into the models. This has not been usually possible in the previous spatially heterogeneous models focusing on the evolution of specialization.     

Evolution of specialization in resource utilization in structured metapopulations

We study the evolution of resource utilization in a structured discrete-time metapopulation model with an infinite number of patches, prone to local catastrophes. The consumer faces a trade-off in the abilities to consume two resources available in different amounts in each patch. We analyse how the evolution of specialization in the utilization of the resources is affected by different ecological factors: migration, local growth, local catastrophes, forms of the trade-off and distribution of the resources in the patches. Our modelling approach offers a natural way to include more than two patch types into the models. This has not been usually possible in the previous spatially heterogeneous models focusing on the evolution of specialization.     

Proboscideans from the upper Miocene localities of Thermopigi,Neokaisareia and Platania (Northern Greece)

In the present article, we study the proboscidean remains from three upper Miocene localities of Northern Greece: Thermopigi (Serres), Neokaisareia (Pieria) and Platania (Drama). The material from the Turolian locality of Thermopigi includes only postcranial specimens. The morphological features of the scapula indicate the presence of the deinotheriid Deinotherium sp., whereas the rest of the specimens are morphologically distinct from Deinotherium and can be referred to Elephantimorpha indet. The material from Neokaisareia consists of a partial skeleton of a single individual and is attributed to the mammutid Mammut sp. (M. obliquelophus?). This taxon is known in Greece from the early–middle Turolian. The Platania proboscidean belongs to the tetralophodont amebelodontid Konobelodon cf. atticus. The genus Konobelodon was already present during the Vallesian of the wider area, but the lower tusk of the Platania shovel-tusker presents some morphological and metrical differences from the Vallesian representative, yet it has also smaller dimensions in its deciduous dentition than the morphologically similar Turolian specimens. The type locality of K. atticus is Pikermi (Attica, Greece), correlated to the middle Turolian, but the known biostratigraphic range of this species covers the entire Turolian. Platania is possibly correlated close to the Vallesian/Turolian boundary and the possible record of this species could document one of its earliest occurrences.     

Histogenesis of the unique morphology of proboscidean ivory

The chequered pattern (often called Schreger pattern), which can be seen by unaided eye on transverse profiles of several proboscidean tusks and which can be emphasized by the spreading pattern of the cracks or by mineral discoloration, is an autapomorph feature of the clade Elephantoidea. The pattern differs among proboscidean taxa; therefore, it allows the coarse differentiation of elephants, mammoths, and some other basal groups. Such identification methods could facilitate efforts concerned with protection of the remaining elephant populations through ivory trade restrictions, since the tooth dentine from extinct Mammuthusprimigenius and from extant Loxodontaafricana and Elephasmaximus are the most common raw materials of the ivory carvings. The aim of this study was to show the internal structure of proboscidean ivory and to revise the existing theories on the aforementioned pattern of the elephantoids with reflections on the events which lead to the development of this microstructure. Thin sections and natural crack surfaces with various orientations of M.primigenius, Elephasantiquus, Prodeinotherium, and Deinotherium tusk fragments were used to produce a three‐dimensional model which explains the features on all profiles. The “phase shift” model is introduced, which assumes a sinusoid undulation of the dentinal tubules in radial profiles in the case of elephantoids. The model was confirmed by photomicrographs, scanning electron microscopic images, interpretation of natural crack surfaces, and radial displacement analysis of the dentinal tubules. The latter proved that the adjacent waves are not in the same phase. Several new nondestructive distinguishing methods are described here on the basis of the correlation between some microscopic and macroscopic features related to the Schreger pattern. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Phylogeny and evo-devo: characters, homology, and the historical analysis of the evolution of development

The concept of homology continues to attract more and more commentary. In systematic and evolutionary biology the meaning of homology as synapomorphic similarity inherited from a common ancestor has gained wide acceptance over the last three or four decades. In recent years, however, developmental biologists, in particular, have argued for a new approach to, and new definition for, homology that revolves around the desire to make it more process-oriented and more mechanistic. These efforts raise questions about the relationship between developmental and evolutionary biology as well as how the evolution of development is to be studied. It is argued in this paper that this new approach to homology seemingly decouples developmental biology from the study of the evolution of development rather than to facilitate that study. In contrast, applying the notion of historical, phylogenetic homology to developmental data is inherently comparative and therefore evolutionary.     

Phylogenetics and the evolution of major structural characters in the giant genus Euphorbia L. (Euphorbiaceae)

Euphorbia is among the largest genera of angiosperms, with about 2000 species that are renowned for their remarkably diverse growth forms. To clarify phylogenetic relationships in the genus, we used maximum likelihood, bayesian, and parsimony analyses of DNA sequence data from 10 markers representing all three plant genomes, averaging more than 16kbp for each accession. Taxon sampling included 176 representatives from Euphorbioideae (including 161 of Euphorbia). Analyses of these data robustly resolve a backbone topology of four major, subgeneric clades--Esula, Rhizanthium, Euphorbia, and Chamaesyce--that are successively sister lineages. Ancestral state reconstructions of six reproductive and growth form characters indicate that the earliest Euphorbia species were likely woody, non-succulent plants with helically arranged leaves and 5-glanded cyathia in terminal inflorescences. The highly modified growth forms and reproductive features in Euphorbia have independent origins within the subgeneric clades. Examples of extreme parallelism in trait evolution include at least 14 origins of xeromorphic growth forms and at least 13 origins of seed caruncles. The evolution of growth form and inflorescence position are significantly correlated, and a pathway of evolutionary transitions is supported that has implications for the evolution of Euphorbia xerophytes of large stature. Such xerophytes total more than 400 species and are dominants of vegetation types throughout much of arid Africa and Madagascar.     

Convergent evolution of gigantism in the flora of an isolated archipelago

Animals often evolve conspicuous differences in body size after colonising isolated islands. However, far less is known about repeated patterns in the evolution of plant size on islands. We tested for convergent evolution of leaf area, seed size and plant stature in the flora of the Chatham Islands (i.e. Rēkohu), which are located 700 km off the east coast of New Zealand. First, we measured leaf area, seed size and plant stature from populations of 22 plant taxa on the Chatham Islands. These data were then compared to analogous measurements from their sister taxa on the mainland to test for evidence of island gigantism, and whether size changes differ between leaves, seeds and stature. Second, we collated data from the literature to test whether size changes in endemic taxa differed among plant growth forms or were correlated with divergence times. Results showed that all three plant traits tended to increase in size on the Chatham Islands. However, field data showed that size increases tended to be more consistent in seeds and leaves than in stature. Data from the literature indicated that size increases also differed among growth forms. Herbaceous species showed the strongest evidence of gigantism, followed by woody plants, while graminoids showed a weak trend towards dwarfism. Insular size increases in seeds and leaves were also positively related to divergence times, indicating that taxa which have resided on the Chatham Islands for longer periods show stronger evidence of gigantism than taxa which arrived more recently. Overall results illustrate that gigantism is a hallmark of the Chatham Island flora, providing a remarkable example of convergent evolution.     

Anatomy and phylogenetic value of the mandibular and coronoid canals and their associated foramina in proboscideans (Mammalia)

Characters associated with the mandibular canal are differently distributed amongst proboscidean lineages and provide useful information on the systematics and relationships of proboscideans. The aim of this paper is to describe the pattern of the mandibular canal and its associated foramina in proboscideans in order to fully appreciate the extent of interspecific variation of these structures within the group and to discuss its systematic and phylogenetic value. Outgroup comparison indicates that the condition presented by the basal proboscidean Phosphatherium is morphotypic for proboscideans. Primitive proboscidean characters are: the low position of the mandibular foramen, and its crescent‐shaped outline, the occurrence of a coronoid foramen (canal), the occurrence of two lateral mental foramina, the posterior one at the level of (or slightly behind) the posterior margin of the symphysis, the anterior one in a more distal position, the absence of a medial mental foramen (MMF), the mandibular canal set just below the tooth row. The occurrence of a single lateral mental foramen may represent a shared derived character of Daouitherium, Numidotherium, and Barytherium. A unique derived feature of the Elephantinae mandible is the occurrence of a medial mental foramen on the medial side of the incisive part of the mandible. MMFs have never been observed in other proboscideans excluding elephantines. The very high frequency of MMFs observed in Mammuthus meridionalis–Mammuthus trogontherii–Mammuthus primigenius (>93 per cent of the studied specimens) could be considered a synapomorphy of this group. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 391–413.     

记同心铲齿象一幼年头骨化石

本文记述了一个采自宁夏同心丁家二沟的同心铲齿象幼年头骨。通过与PL.danovi.Pl.grangeri比较,进一步证明了Pl.tongxinensis的进化水平介于二者之间,并与后者的关系更为密切。笔者选择Phiomia和现生Elephas maximus为参照分析了同心头骨的进化特征。尽管三者不属于同一枝系,但表现在它们身上的性状变化仍可大体反映出长鼻类的进化趋向。     

Extremely intron-rich genes in the alveolate ancestors inferred with a flexible maximum-likelihood approach

Chromalveolates are a large, diverse supergroup of unicellulareukaryotes that includes Apicomplexa, dinoflagellates, ciliates(three lineages that form the alveolate branch), heterokonts,haptophytes, and cryptomonads (three lineages comprising thechromist branch). All sequenced genomes of chromalveolates haverelatively low intron density in protein-coding genes, and fewintron positions are shared between chromalveolate lineages.In contrast, genes of different chromalveolates share many intronpositions with orthologous genes from other eukaryotic supergroups,in particular, the intron-rich orthologs from animals and plants.Reconstruction of the history of intron gain and loss duringthe evolution of chromalveolates using a general and flexiblemaximum-likelihood approach indicates that genes of the ancestorsof chromalveolates and, particularly, alveolates had unexpectedlyhigh intron densities. It is estimated that the chromalveolateancestor had, approximately, two-third of the human intron density,whereas the intron density in the genes of the alveolate ancestoris estimated to be slightly greater than the human intron density.Accordingly, it is inferred that the evolution of chromalveolateswas dominated by intron loss. The conclusion that ancestralchromalveolate forms had high intron densities is unexpectedbecause all extant unicellular eukaryotes have relatively fewintrons and are thought to be unable to maintain numerous intronsdue to intense purifying selection in their, typically, largepopulations. It is suggested that, at early stages of evolution,chromalveolates went through major population bottlenecks thatwere accompanied by intron invasion.