Evidence of predation damage in Pliocene Apletosia maxima (Brachiopoda)

Abstract:  Little is known about predation of Mesozoic and Cenozoic articulated brachiopods, but it is far from clear whether this is because they suffered very little predation pressure or because there have been few attempts to search for evidence of it. A study of 248 museum specimens of the large Pliocene terebratulid Apletosia maxima from the Coralline Crag (UK) has revealed that more than 16 per cent of them show evidence of having been attacked by predators. The styles of damage can be attributed to drilling muricid gastropods (most of which were successful) and failed crushing attacks probably by decapods. Brachiopods are usually thought to offer a poor tissue yield to potential predators, but in this instance it appears that A. maxima was attractive to predators even though they were living with a rich molluscan fauna. It is suggested that the mass of adductor and diductor muscles (likely to be spicule-free) of these particularly large brachiopods may have made them profitable. Further studies of post-Palaeozoic brachiopod faunas are required, particularly those from mixed shallow-water communities, before it can be established whether articulated brachiopods have or have not been driven into refugia by increasing predation pressure.     

Uncovering the holes and cracks: from anecdote to testable hypotheses in predation studies

Biological interactions between organisms have long been believed to be very important in structuring communities and, when scaled up over geological time, in the evolution of organisms. Investigations of palaeontological evidence for predator–prey interactions have been popular pursuits, and a number of attractive hypotheses have been proposed which link increased predation pressure with a wide range of morphological and ecological changes which are apparent over the course of the Phanerozoic. In particular studies of fossil drill holes and repair scars in shelly prey have been common targets for research. However, the nature of some of our data has been rather anecdotal and restricted in range. Perhaps we should be more concerned that we are not picking up the true range of natural variability. This paper aims to highlight the sources of variability in our data and, going forward, to urge the collection of quantitative data from many more samples and (palaeo)environmental settings in order that we might properly be able to separate the intrinsic natural variability in our data from robust temporal or spatial trends.     

Causes of damage to brachiopods from the Middle Pennsylvanian Naco Formation, central Arizona

The Kohl Ranch locality in the Naco Formation of central Arizona has yielded an abundant and diverse fauna including eight genera of brachiopods and four of sharks including Cochliodus. The most abundant brachiopod, Composita subtilita , was analyzed for predation damage and of 935 specimens 503 (53.7%) were found to show the type of unrepaired crushing previously attributed to shark predation. Reconstruction of the jaw of Cochliodus , however, suggests that it may have been a selective feeder similar to the modern Heterodontus and unlikely therefore to have left so many damaged but uningested specimens. Evidence from specimens collected in situ suggests that the damage mostly occurred post-mortem due to the compaction of the sediments and was not due to the sharks who would have comminuted any prey items taken.     

A note on the genus Artemia in Algeria

Five new locality records for Artemia in Algeria are given. Some physical and chemical characteristics of their environment are recorded. Predation on Artemia by the fish Aphanius fasciatus occurs where the river Khrouf debouches in Chott Merouan but not elsewhere.     

A new species of Castanopsis (Fagaceae) from the upper Pliocene of West Yunnan,China and its biogeographical implications

Two fossil leaves identified as Castanopsis presclerophylla n. sp. collected from a diatomite bed in the upper Pliocene (3.3–2.8 Ma) in Tengchong County, Yunnan Province, Southwest China, are characterized by elliptic laminar shapes with acuminate apex, convex base, simple trichome bases and cyclocytic stomata. The cuticles of extant leaves from six selected Castanopsis species that are similar to the Pliocene fossil leaves in laminar shape were examined in detail for the comparison. Based on the analysis of leaf architecture and cuticular characteristics, the present fossil leaves most closely resemble the extant leaves of Castanopsis sclerophylla (Lindl.) Schott., which prefers a subtropical climate. The published fossil records of Castanopsis also indicate that it was once widely distributed in North America, Europe and Asia before the Pliocene, and then rapidly retreated to the East and Southeast Asia as a result of the Pleistocene glaciation.     

Upper Pliocene bivalve shell concentrations from the Lower Chelif basin (NW Algeria): Systematics,sedimentologic and taphonomic framework

Sedimentologic and palaeontological investigation of the Upper Pliocene Slama Formation in the Lower Chelif Basin (NW Algeria) led us to collect important bivalve assemblages for taxonomic and taphonomic purposes. A rather comprehensive inventory list of Upper Pliocene bivalves from northwestern Algeria is now available and consists of 30 species, 17 of which are extinct ones. Four principal taphonomic attributes were analysed: bioerosion, encrustation, fragmentation, and abrasion. Physical and biogenic sedimentary structures are used for palaeoenvironmental interpretations. The taphonomic, sedimentologic and ichnological characteristics of most of the deposits suggests they originated from discontinuous processes of winnowing and bypassing of sediments, probably due to the action of storms in shallow waters, mainly in the shoreface depositional environment. The bivalve assemblage is dominated by disarticulated valves and displays significant taphonomic alteration in the shells. Sclerobionts traces in shells particulary affect the oyster shells. Bioerosion traces are predominately those of clionid sponges (Entobia isp.), polychaetes (Maeandropolydora isp. and Caulostrepsis isp.), bivalves (Gastrochaenolites isp.), and of predatory gastropods (Oichnus isp.). Among the sclerobionts, the identified encrusters were juvenile oyster recruits, barnacles, polychaetes (serpulid tubeworms), bryozoans (Microporella sp. and Acanthodesia sp.), and vermetid gastropods (Petaloconchus intortus).     

A fossil Dendrocygninae (Aves,Anatidae) from the Early Pliocene of the Argentine Pampas and its paleobiogeographical implications

In the present paper, the distal end of a humerus referable to a Dendrocygninae anseriform bird is reported. The specimen was collected at the Monte Hermoso Formation (early Pliocene) of the Farola Monte Hermoso locality (Buenos Aires Province, Argentina). This record constitutes one of the oldest for the group in South America and elsewhere. The overview of the anseriform record in South America indicates a similar pattern to that of other localities across the world. Paleogene localities are dominated by non-Anatinae taxa, whereas more recent faunas are dominated by anatines. Following this pattern, in South America the Anatinae appears in the fossil record as recently as in the Pleistocene, probably arriving from North America during the Great American Biotic Interchange. Pleistocene and Recent anseriform avifaunas are dominated by Anatinae taxa, and this dominance is probably due to some “key characters” regarding reproductive biology. In fact, the anatines exhibit a high reproductive success probably due to the capability of having two molts per year, and that the parental care of the downy young is only conducted by the female. The combination of characters may have allowed anatines to dominate most aquatic environments, surpassing in diversity more plesiomorphic taxa.     

Post‐metamorphic allometry in the earliest acrotretoid brachiopods from the lower Cambrian (Series 2) of South China,and its implications

The earliest growth of post‐metamorphic (post‐larval) shells in two species of Eohadrotreta is described from the Cambrian Shuijingtuo Formation of South China. Two different growth patterns can be observed by quantifying developmental variations in size and shape of successive stages of post‐metamorphic shell growth (including the pedicle foramen forming stage, pedicle foramen enclosing stage and intertrough increasing stage) of Eohadrotreta zhenbaensis and Eohadrotreta ? zhujiahensis . The pedicle foramen is never enclosed within the metamorphic shell of E. zhenbaensis , while the enclosed pedicle foramen of E .? zhujiahensis is located directly outside the metamorphic shell after the pedicle foramen enclosing stage. A strongly allometric growth pattern of E. zhenbaensis is demonstrated by the early enclosure of the pedicle foramen; an accelerated lengthening of the ventral intertrough is associated with the development of a more complex dorsal median septum during the intertrough increasing stage. By contrast, E .? zhujiahensis demonstrates possible paedomorphic development by delayed enclosure of pedicle foramen and an associated decreased lengthening of ventral intertrough during the intertrough increasing stage. This ontogenetic developmental sequence represents the marginal accretionary formation and growth of the pedicle foramen, which resembles that of linguloid brachiopods. Furthermore, the developmental process of the pedicle foramen of Eohadrotreta seems to recapitulate the likely evolutionary transition from the Botsfordiidae, with open delthyrium, to the Acrotheloidea, with an enclosed foramen. This study provides a unique opportunity to obtain a complete understanding of the ontogenetic development of the earliest acrotretoids, and casts new light on the phylogeny of lingulate brachiopods.     

Orthid brachiopods from the Lower and Middle Devonian of northeastern Russia

Brachiopods of the order Orthida from the Lower and Middle Devonian of northeastern Russia: Skenidium diversus Baranov (family Skenidiidae); Datnia gen. nov., with the type species D. asiatica sp. nov. (subfamily Prokopiinae); Janzhinshinia gen. nov., with the type species J. datnensis sp. nov. (subfamily Isorthinae); Cortezorthis chobotchalensis (Alekseeva) (subfamily Cortezorthinae); Elenia gen. nov., with the type species E. gerensis sp. nov. (family Dicoelosiidae); Seimtchania communis (family Seimtchaniidae); Simakovia gen. nov., with the type species S. rara sp. nov. (family Draboviidae); Schizophoria distenta sp. nov., S. grande Baranov, S. striatula (Schlotheim) (family Schizophoriinae); and Hypsomionia sinsera Baranov (family Hypsomioniidae) are described.     

The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its palaeobiological implications

The Pliocene Beds of the Pisco Formation on the southern coast of Peru have yielded three new skulls of the walrus-like odontocete, Odobenocetops (Delphinoidea, Cetacea). Two of the skulls are from a slightly younger horizon than the holotype of O. peruvianus and belong to a different species, O. leptodon . The holotype of O. leptodon bears a 135 cm long needle-like right tusk and a small 25 cm long left tusk, the apex of which was erupted. The third skull, referred to a female of O. peruvianus , bears two small tusks similar in size to the small left tusk of the holotype. The new specimens include periotic, tympanic and ear ossicles, which confirm referral to the Order Cetacea and the morphology of the humerus reinforces affinities of odobenocetopsids to monodontids. Because the anterodorsal edge of its orbit is slightly concave O. leptodon had reduced anterodorsal binocular vision, a condition compensated for by the probable presence of a small melon (and inferred echolocation). The head was bent ventrally when swimming in such a way that the long tusk was approximately parallel to the axis of the body. The extremely salient occipital condyles of Odobenocetops are indicative of great mobility of the neck, probably related to bottom-feeding. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 423–452.     

云南宜良上新世栎属植物研究及其古环境指示意义

栎属(Quercus)植物叶片化石在新生代地层中分布普遍, 对亚热带常绿阔叶林具有重要的指示意义。本文对采自云南省宜良县上新统茨营组中的栎属植物叶片化石进行角质层结构分析, 结合叶形态特征, 系统描述了4种栎属植物: 阔叶栎(Quercus latifolia)、线叶栎(Q. scottii)、楔基栎(Q. simulata)和黄毛青冈相似种(Q. cf. delavayi)。通过对比化石和现生植物的叶形态和角质层特征, 本文认为叶片形状、叶基和叶尖特征、二级脉数量、表皮细胞和垂周壁特征、气孔大小和形状、气孔密度、毛基类型、形态和分布情况可以作为区分不同种类的特征, 但是不能只考虑其中的一个因素, 应该综合这些特征作为判断依据。结合宜良植物群中其他化石资料认为, 包括壳斗科柯属(Lithocarpus)、栲属(Castanopsis)以及樟科、榆科、木兰科等在内的植物, 代表了以栎属青冈组为优势树种的半湿润常绿阔叶林。宜良植物群与同时期相近纬度的植物群相比, 植被类型相似, 但是落叶成分较少。宜良植物群中的线叶栎、楔基栎和阔叶栎同晚始新世的线叶栎以及早中新世的楔基栎和阔叶栎相比, 形态并未发生大的变化, 与现生植物也非常相似, 进一步说明云南现代常绿阔叶林的主要成分来自于古老植物的承袭, 没有发生较大的改变。     

New Carboniferous brachiopods from the eastern Great Basin, Nevada, USA: implications for loop ontogeny and evolution in Late Palaeozoic terebratuloids

Terebratuloid brachiopods from two localities in the eastern Great Basin, Nevada, USA, include the new taxa Cryptacanthia savagei sp. nov., Fletcherithyris infrequens sp. nov., Cryptonella simplex sp. nov., and Albelenina alvarezi gen. et sp. nov. The faunas are considered to be mid Desmoinesian (late Moscovian) in age. The brachiopods were found associated in clusters, and many of the specimens are well preserved. Records of Pennsylvanian terebratuloid brachiopods are uncommon, and these faunas have enabled an understanding of internal features previously unknown or poorly understood. Systematic analyses of the faunas have generated some new concepts on the ontogeny and evolution of the loop in Late Palaeozoic terebratuloids. From ontogenetic analyses, it is inferred that the stage of loop development may be the most important aspect in taxonomic classification. Analyses of loop evolution suggest the possibility of a transition from a teloform stage to long-flanged deltiform and deltiform stages. These transitions may be possible through reduction in length of the descending lamellae of the crura by resorption of calcium carbonate or posteroventral folding of flanges. Subsequently, these changes in loop morphology are retained by paedomorphosis within long-term evolutionary processes.     

The early ontogeny of Jurassic thecideoid brachiopods and its contribution to the understanding of thecideoid ancestry

Abstract: A collection of very early brephic juveniles from the Upper Aalenian of the Cotswolds, England, has provided the first opportunity to study the ontogeny of thecideoids prior to the recognizable development of a median septum. Traced towards their origin, the enduring characters of thecideides appear to be cementation, a delthyrium closed by a pseudodeltidium, an almost circular dorsal valve with a prominent erect bilobed cardinal process, well‐defined inner socket ridges, widely spaced lateral adductor muscle fields, tubercles, fibrous secondary shell lining both valves and probably endopunctae. The subperipheral rim and median septum so characteristic of adult thecideoids are undeveloped in the earliest juveniles. Initiation of the development of the free ventral wall in the ventral valve is identified as an important event in thecideoid ontogeny. Further important discoveries arising from the study are that the crura‐like outgrowths which form the brachial bridge in thecideides are not structurally homologous with the crura of spiriferides or terebratulides and that during their earliest ontogenetic development thecidellinids and thecideids are indistinguishable. Also, the identification of morphological characters correlative with both palaeontological and neontological approaches to thecideoid phylogeny has important implications for thecideide taxonomy. Interpretation of the morphology exhibited by the Cotswold specimens introduces the probability that during their earliest ontogeny moorellinin, thecidein and lacazellin dorsal valves follow the same development pattern and the described ontogenetic sequence has been corroborated by evidence from Jurassic rocks of Argentina, North America and France, from Cretaceous rocks of England, Central Europe and Czechoslovakia, also from extant species from the Atlantic, Indian and Pacific oceans, and to an extent which suggests the pattern of ontogenetic development revealed has been typical of thecideoids throughout their history.     

Caradoc strophomenoid and plectambonitoid brachiopods from Wales and the Welsh Borderland

Abstract: Ordovician strophomenoidean and plectambonitoidean brachiopods are reviewed and partially redescribed from the Caradoc (Sandbian to Early Katian) age rocks of Wales and the Welsh Borderland of England, then forming part of the Avalonia Terrane. There are 51 nominal species available, of which 12 are synonymised here and many transferred to different genera. Strophomenoids occurred sporadically, often rarely but occasionally abundantly, particularly in mid‐shelf benthic assemblages, and 21 named species and subspecies are noted, including the new Kiaeromena (Kiaeromena) harperi, as well as six in open nomenclature. The common genus Kjaerina, with six species known from the area, is revised in detail. In contrast to the strophomenoids, plectambonitoids were more common in shallow and mid‐shelf assemblages, including Eoplectodonta (Eoplectodonta) abigailae sp. nov., with Sowerbyella in particular dominating many bedding planes, and 17 named species are reviewed, as well as three in open nomenclature. The distinctive Gunningblandella, hitherto known only from Australia and Kazakhstan, is recorded for the first time from Europe. The Avalonian strophomenides are compared with those from neighbouring terranes, particularly Laurentia and Baltica, during a time when the seas between those terranes were narrowing, and the faunas from the three terranes are found to be largely similar at the generic level, although there are few species in common. In contrast, there are few genera in common between those three terranes and the Mediterranean Province of Northwestern Gondwana.     

A hominoid humeral fragment from the Pliocene of Kenya

The purpose of this communication is to describe and provide a preliminary interpretation of a hominoid proximal humeral fragment recovered from sediments more than 4.0 m.y. old in the Baringo District, Kenya. The geological and chronological context of the fossil is examined and associated fauna is analyzed.     

Conifer Woods of the Pliocene Age from Yunnan, China

The Tertiary floras play an integral role in understanding the biodiversity and interactions between climate and vegetation in Yunnan, China. The fossil spores, pollen grains, and leaves in this region have been investiaged intensively. In comparison, the woods have been studied relatively little. A large number of Pliocene wood specimens was collected from the opencast lignite coal mine of Hongxing situated in Changning County of Yunnan Province. Among the collection, Tsuga cf. dumosa (D. Don) Eichler and Pinus cf. armandii Franchet were identified based on wood structures. The subtle feature oftori extensions is reported in the wood of T. cf. dumosa. Considering the climatic requirements of modern T. dumosa and P.armandii, the two species of conifer described from Pliocene sediments probably grew in mountainous terrain at an elevation of approximately 2 300 m, in a cool and humid environment.     

Conifer Woods of the Pliocene Age from Yunnan, China

The Tertiary floras play an integral role in understanding the biodiversity and interactions between climate and vegetation in Yunnan, China. The fossil spores, pollen grains, and leaves in this region have been investiagedintensively. In comparison, the woods have been studied relatively little. A large number of Pliocene wood specimens was collected from the opencast lignite coal mine of Hongxing situated in Changning County of Yunnan Province. Among the collection, Tsuga cf. dumosa (D. Don) Eichler and Pinus cf. armandii Franchet were identified based on wood structures. The subtle feature of tori extensions is reported in the wood of T. cf. dumosa. Considering the climatic requirements of modern T. dumosa and P.armandii, the two species of conifer described from Pliocene sediments probably grew in mountainous terrain at an elevation of approximately 2 300 m, in a cool and humid environment.     

Reappraising the early evidence of durophagy and drilling predation in the fossil record: implications for escalation and the Cambrian Explosion

The Cambrian Explosion is arguably the most extreme example of a biological radiation preserved in the fossil record, and studies of Cambrian Lagerstätten have facilitated the exploration of many facets of this key evolutionary event. As predation was a major ecological driver behind the Explosion – particularly the radiation of biomineralising metazoans – the evidence for shell crushing (durophagy), drilling and puncturing predation in the Cambrian (and possibly the Ediacaran) is considered. Examples of durophagous predation on biomineralised taxa other than trilobites are apparently rare, reflecting predator preference, taphonomic and sampling biases, or simply lack of documentation. The oldest known example of durophagy is shell damage on the problematic taxon Mobergella holsti from the early Cambrian (possibly Terreneuvian) of Sweden. Using functional morphology to identify (or perhaps misidentify) durophagous predators is discussed, with emphasis on the toolkit used by Cambrian arthropods, specifically the radiodontan oral cone and the frontal and gnathobasic appendages of various taxa. Records of drill holes and possible puncture holes in Cambrian shells are mostly on brachiopods, but the lack of prey diversity may represent either a true biological signal or a result of various biases. The oldest drilled Cambrian shells occur in a variety of Terreneuvian‐aged taxa, but specimens of the ubiquitous Ediacaran shelly fossil Cloudina also show putative drilling traces. Knowledge on Cambrian shell drillers is sorely lacking and there is little evidence or consensus concerning the taxonomic groups that made the holes, which often leads to the suggestion of an unknown ‘soft bodied driller’. Useful methodologies for deciphering the identities and capabilities of shell drillers are outlined. Evidence for puncture holes in Cambrian shelly taxa is rare. Such holes are more jagged than drill holes and possibly made by a Cambrian ‘puncher’. The Cambrian arthropod Yohoia may have used its frontal appendages in a jack‐knifing manner, similar to Recent stomatopod crustaceans, to strike and puncture shells rapidly. Finally, Cambrian durophagous and shell‐drilling predation is considered in the context of escalation – an evolutionary process that, amongst other scenarios, involves predators (and other ‘enemies’) as the predominant agents of natural selection. The rapid increase in diversity and abundance of biomineralised shells during the early Cambrian is often attributed to escalation: enemies placed selective pressure on prey, forcing phenotypic responses in prey and, by extension, in predator groups over time. Unfortunately, few case studies illustrate long‐term patterns in shelly fossil morphologies that may reflect the influence of predation throughout the Cambrian. More studies on phenotypic change in hard‐shelled lineages are needed to convincingly illustrate escalation and the responses of prey during the Cambrian.     

Early Carboniferous spiriferoid brachiopods from the Qaidam Basin,Northwest China: Taxonomy,biostratigraphy and biogeography

Six Early Carboniferous brachiopod species in four genera of the Superfamily Spiriferoidea are described from the Qaidam Basin, northwestern China, including a new genus, Qaidamospirifer, and two new species: Grandispirifer qaidamensis and Qaidamospirifer elongatus. Additionally, a new genus, Triangulospirifer, is also proposed to replace Triangularia Poletaev, 2001 that was preoccupied by a Devonian molluscan genus.On the basis of the new material as well as published information, we have reviewed the taxonomic composition and the stratigraphic and palaeobiogeographic distributions of the three previously established genera from the viewpoint of palaeobiogeography. The study reveals that Grandispirifer has a relatively long stratigraphic range from the late Tournaisian to Serpukhovian. During this interval, the genus attained a wide geographical distribution, reaching Northwest China, western Yunnan of West China, Japan, as well as Iran and North Africa. Angiospirifer first occurred in western Europe in the Viséan, and later migrated to North Africa during the late Viséan. In the Serpukhovian, it migrated eastward, reaching the Donets Basin of Ukraine and the Qaidam Basin in Northwest China. Anthracothyrina evolved from Brachythyrina in North Africa in late Viséan, then dispersed north-westward to western and eastern Europe and, further eastward to the Qaidam Basin during the Serpukhovian.