Fragilicetus velponi: a new mysticete genus and species and its implications for the origin of Balaenopteridae (Mammalia,Cetacea, Mysticeti)

A new extinct genus, F ragilicetus gen. nov. , is described here based on a partial skull of a baleen‐bearing whale from the Early Pliocene of the North Sea. Its type species is F ragilicetus velponi sp. nov. This new whale shows a mix of morphological characters that is intermediate between those of Eschrichtiidae and those of Balaenopteridae. A phylogenetic analysis supported this view and provided insights into some of the morphological transformations that occurred in the process leading to the origin of Balaenopteridae. Balaenopterid whales show specialized feeding behaviour that allows them to catch enormous amounts of prey. This behaviour is possible because of the presence of specialized anatomical features in the supraorbital process of the frontal, temporal fossa, glenoid fossa of the squamosal, and dentary. F ragilicetus velponi gen. et sp. nov. shares the shape of the supraorbital process of the frontal and significant details of the temporal fossa with Balaenopteridae but maintains an eschrichtiid‐ and cetotheriid‐like squamosal bulge and posteriorly protruded exoccipital. The character combination exhibited by this cetacean provides important information about the assembly of the specialized morphological features responsible for the highly efficient prey capture mechanics of Balaenopteridae. © 2015 The Linnean Society of London     

Phylogeny of mysticete whales based on mitochondrial and nuclear data

Mysticetes or baleen whales are comprised of four groups: Eschrichtiidae, Neobalaenidae, Balaenidae, and Balaenopteridae. Various phylogenetic hypotheses among these four groups have been proposed. Previous studies have not satisfactorily determined relationships among the four groups with a high degree of confidence. The objective of this study is to determine the relationships among the mysticete whales. Mitochondrial and nuclear DNA were sequenced for phylogenetic analysis. Most species relationships determined using these data were well resolved and congruent. Balaenidae is the most basal group and Neobalaenidae is the second most basal and sister group to the balaenopterid-eschrichtiid clade. In this phylogenetic study, the resolution of Eschrichtiidae with two main lineages of Balaenopteridae was problematic. Some data partitions placed this group within the balaenopterids, and other partitions placed it as a sister taxon to the balaenopterids. An additive likelihood approach was used to determine the most optimal trees. Although it was not found in the combined phylogenetic analyses, the "best" tree found under the additive likelihood approach was one with a monophyletic Balaenopteridae.     

A NEW BASAL BALAENOPTERID WHALE FROM THE PLIOCENE OF NORTHERN ITALY

Abstract:  A new basal balaenopterid genus and species, Archaebalaenoptera castriarquati , is described and compared with all the living and fossil members of the family Balaenopteridae and related fossil rorqual-like taxa. It was found in the Lower Pliocene of northern Italy, and is characterized by a supraoccipital with a transversely compressed anterior process, the zygomatic process of the squamosal diverging from the longitudinal axis of the skull, very long nasal bones, and subtle exposition of the parietal on the dorsal wall of the skull. It is primitive in having a maxilla with a long ascending process that is posteriorly unexpanded and round, and a dentary that is straight and not bowed outward, unlike that of living Balaenopteridae. In particular, the discovery of this new genus suggests that, among the early members of Balaenopteridae, the acquisition of the typical sutural pattern shown by maxilla, frontal, parietal and supraoccipital preceded the acquisition of the feeding-related traits that are characteristic of the family. The primitive morphology of the feeding-related structures of A. castriarquati (i.e. the straight dentary and the flat glenoid fossa of the squamosal) suggests that this whale was unable to undertake the intermittent ram feeding typical of Balaenopteridae as efficiently as living members of the family.     

First porcupine fossils (Mammalia,Rodentia) from the late Miocene of NW Iran,with notes on late Miocene–Pliocene dispersal of porcupines

This paper describes the first fossil porcupine remains from Iran. Four upper cheek teeth and two fragmentary lower incisors present sufficient characters for identification as Hystrix aryanensis, a species previously known from the late Miocene locality of Molayan (Afghanistan) estimated at ca. 7–8 Ma. The dental features of porcupines are discussed to show their systematic value and highlight evolutionary trends in late Miocene and Pliocene porcupines. This study also discusses the dispersal history of fossil porcupines in relation to paleobiogeographic provinces and environmental changes during late Miocene to late Pliocene time.     

Proterozoic stromatolites: The first marine evolutionary biota

A biometric analysis of the morphology of a late Pliocene planktic diatom lineage, Rhizosolenia praebergonii Mukhina indicates that it appeared abruptly in the fossil record in the Indian Ocean at 2.9 Ma after which it remained virtually unchanged. The observed pattern can be explained by “local”; evolution from the ancestral form Rhizosolenia bergonii, Peragallo or by migration from thecentral Pacific Ocean, where it originated gradually, in conjunction with an accelerated rate of evolution. At present time, it is not possible to conclude which one of the two hypotheses is more likely.     

The oldest marine vertebrate fossil from the volcanic island of Iceland: a partial right whale skull from the high latitude Pliocene Tjörnes Formation

Extant baleen whales (Cetacea, Mysticeti) are a disparate and species‐rich group, but little is known about their fossil record in the northernmost Atlantic Ocean, a region that supports considerable extant cetacean diversity. Iceland's geographical setting, dividing North Atlantic and Arctic waters, renders it ideally situated to shed light on cetacean evolution in this region. However, as a volcanic island, Iceland exhibits very little marine sedimentary exposure, and fossil whales from Iceland older than the late Pleistocene are virtually unknown. Here, we present the first fossil whale found in situ from the Pliocene Tjörnes Formation (c. 4.5 Ma), Iceland's only substantial marine sedimentary outcrop. The specimen is diagnosed as a partial skull from a large right whale (Mysticeti, Balaenidae). This discovery highlights the Tjörnes Formation as a potentially productive fossil vertebrate locality. Additionally, this find indicates that right whales (Eubalaena) and bowhead whales (Balaena) were sympatric, with broadly overlapping latitudinal ranges in the Pliocene, in contrast to the modern latitudinal separation of their living counterparts.     

Origins, phytogeny and taxonomy of the gadfly petrels Pterodroma spp

Taxonomic characters of gadfly petrels (Procellariidae: Pterodroma spp.) are reviewed and the genus is redefined. The structure of the upper intestines, which have become helicoidally twisted to varying degrees in most species, is an important character not hitherto used. Results of a phylogenetic study of the genus based mainly on the development of helicoidal intestines agree substantially with findings from studies of the Mallophaga parasitizing these petrels. One species, the Kerguelen Petrel Lugensa brevirostris, previously classified in Pterodroma, is shown to have closer affinity with some fulmar genera. The Tahiti Petrel Pseudobulweria rostrata and its rare or extinct relatives, formerly placed in a superspecies of Pterodroma, are more closely related to the Procellaria and Bulweria petrels, and may be derived from the link between the subfamilies Procellariinae and Fulmarinae. From consideration of the fossil record, anatomical characters and Mallophaga, a phylogeny of the Procellariidae is proposed which supports its monophyletic origin. Gadfly petrels apparently descended from the fulmars through the ancestral lines of Snow Petrels Pagodroma, the Kerguelen Petrel and finally the Blue Petrel Halobaena, which may have given rise separately both to gadfly petrels and to the prions Pachyptila. A late Pliocene origin of Pterodroma in the vicinity of southern New Zealand is possible. The genus Pterodroma is divided into four subgenera, representing four connected radiations, and 29 species are recognised. In probable chronological order, Proaestrelata (new subgenus) contains five species and is restricted to the Pacific Ocean, Cookilaria comprises six species restricted as breeders to the South Pacific, Hallstroma has seven species mainly in the tropics of the Pacific, Indian and South Atlantic Oceans, whereas Pterodroma includes 11 species which extend as breeders from the southwest Pacific west to the South Atlantic and into the North Atlantic. Ecological, physical and physiological adaptations in the evolution of gadfly petrels are discussed.     

Revisiting the Great American Biotic Interchange through analyses of amphitropical bees

The Great American Biotic Interchange (GABI) is zoogeographic event characterized by the exchange of taxa between North and South America, typically associated with the rise of the Isthmus of Panama in the late Pliocene. Recent geologic evidence suggests the connections between North and South America may be much older, and that the interchange of organisms between the two continents could have therefore happened much earlier than 3 Ma. Most of the research investigating the GABI has come from tropical vertebrate taxa; little work has been done on invertebrates or on non‐tropical species. To investigate how the GABI shaped the distribution of arid‐adapted species, particularly those with amphitropical distributions (i.e. taxa found in South and North American xeric regions yet absent from the tropics), we examine the historical biogeography of the bee genus Diadasia using a hypothesis of Diadasia phylogenetic relationships. Nuclear and mitochondrial genetic loci are used to reconstruct a phylogeny of Diadasia, which is then used to estimate divergence dates and reconstruct ancestral area relationships. Our analyses suggest the divergence between North and South American Diadasia species occurred between 20.5 and 15 Ma, long before the formation of the Isthmus of Panama. This study is the first to show a Miocene connection for an amphitropically‐distributed insect group. It suggests that the biotic connection between continents is more complicated than previously thought and may have initiated long before the late Pliocene.     

RECENT RADIATION OF THE PALMARIACEAE (RHODOPHYTA)1

Molecular phylogenetic studies on the evolution of the red algae indicate that this ancient division has many lineages that have recently undergone radiations. One such example is the cold–temperate family Palmariaceae. In this study, sequences from the ribosomal DNA internal transcribed spacer regions were compared among ten species in the Palmariaceae from both Atlantic and Pacific sites, Phylogenetic analyses of sequence data, in which Rhodophysema georgii Batters was used as outgroup and root, indicate a radiation into four clades, three of which contain species of “Palmaria” and the fourth species of Halosaccion. Palmaria palmata (L.) Kuntze, the type and only North Atlantic species in the genus, stands apart from all remaining species in the family and terminates the most basal branch in the rooted tree. The three more derived clades have radiated mainly in the North Pacific. Southern Ocean Palmaria and North Atlantic Devaleraea are hypothesized to have invaded from separate but closely related North Pacific ancestors. The ease with which sequences could be aligned combined with an unsaturated transition: transversion ratio and modest divergence involving predominantly point mutations suggests that the initial radiation is relatively recent (late Miocene–Pliocene) and that the Devaleraea–Palmaria clade is even more recent (late Pliocene–Pleistocene).     

Globorotalia puncticulata: Population divergence, dispersal and extinction related to Pliocene–Quaternary water masses

The isolating effect of water mass partitioning of populations on the morphology, stratigraphic distribution and extinction of planktonic foraminifera is assessed from the Pliocene–Quaternary record of Globorotalia puncticulata. Southern Hemisphere, Mediterranean and North Atlantic data on these aspects of its history are examined and appear consistent with a limited dispersal biogeographic model wherein populations are largely confined by hydrographic barriers.Earliest populations appeared during the latest Miocene in Southern Hemisphere middle latitude water masses. However, morphometric analysis shows that significant differentiation in the axial shape of shells had developed by 4 Ma between Southwest Pacific populations from ODP Site 1123 (temperate water) and ODP Site 1119 (subantarctic water). These sites are in close proximity but separated by the Subtropical Front. At Site 1123 inflation of late-formed chambers and reduction in the number included in the outer whorl created shell profiles that anticipated the globose form of Globorotalia inflata. The latter's gradual evolution from G. puncticulata s.s. took place in this temperate water mass, with the earliest morphotypes with three chambers in the outer whorl present by 4.1 Ma. In contrast, subantarctic populations at Site 1119 retained four chambers but their axial shape was modified. The development of a large, highly arched aperture and increase in the number of chambers in the outer whorl in Mediterranean–North Atlantic Globorotalia puncticulata bononiensis is an example of population differentiation later in the Pliocene.Chronostratigraphy shows that the northward expansion of central temperate water populations commenced with their occupation of Southwest Pacific subtropical water about 4.8 Ma. The rather abrupt entry of substantial populations into Mediterranean and North Atlantic water at 4.5 Ma marked a major biogeographic expansion and established G. puncticulata as a bipolar species. It was widely distributed about 3 Ma, with major populations in several water masses during a period of middle Pliocene warmth.After this acme North Atlantic and Mediterranean G. puncticulata bononiensis populations collapsed as late Pliocene Northern Hemisphere glacials intensified. They were extinguished in MIS 96 (2.4 Ma). Concurrently, G. puncticulata s.s became extinct in the warm subtropical Southwest Pacific. Subantarctic populations persisted but in turn were decimated in severe glacials during the Middle Pleistocene Transition. Most had disappeared by MIS 16 (0.66 Ma). However, at Sites 594 and 1119 there was a small Lazarus-like revival in MIS 11 (0.41 Ma). The highest known occurrence is in MIS 9 (0.33 Ma) at Site 1119. Confinement of the species to subantarctic water in the Pleistocene may have raised its vulnerability to extinction.While stable isotope data indicate that the lineage's evolution is related to depth habitat selection about the thermocline, its biogeography suggests that hydrographic barriers significantly isolated populations and probably facilitated speciation. Eddies such as the North Brazil Current rings provide conduits for inter-water mass transfer of populations but the history of G. puncticulata suggests that such mechanisms seldom operated successfully.The morphology of the lectotype of G. puncticulata s.s., from beach sand at Rimini, Italy, is consistent with a lower Pliocene source. Reports of living occurrences are poorly documented and the species is considered to be extinct.     

Proceedings of the SMBE Tri-National Young Investigators' Workshop 2005. Baleen whale phylogeny and a past extensive radiation event revealed by SINE insertion analysis

Baleen whales (suborder Mysticeti) comprise 11 extant species that are classified into four families. Although several phylogenetic hypotheses about these taxa have been proposed, their phylogenetic relationships remain confused. We addressed this problem using short interspersed repetitive element (SINE) insertion data, which now are regarded as almost ideal shared, derived characters at the molecular level. We reconstructed the phylogenetic relationships of baleen whales by characterizing 36 informative SINE loci. One of the intriguing conclusions is that balaenopterids and eschrichtiids radiated very rapidly during a very short evolutionary period. During this period, speciation occurred in balaenopterids and eschrichtiids while newly inserted SINE loci remains polymorphic. Later on, these SINEs were sorted incompletely into each lineage. Thus, there are now inconsistencies among species regarding the presence or absence of a given SINE. This is in sharp contrast to the phylogeny of toothed whales, for which no SINE inconsistencies have been found. Furthermore, we found monophyletic groupings between humpback and fin whales as well as between (sei+Bryde's) whales and blue whales, both of which have not previously been recognized. The comprehensive SINE insertion data, together with the mitochondrial DNA phylogeny that was recently completed (Sasaki, T., M. Nikaido, H. Healy et al. 2005. Mitochondrial phylogenetics and evolution of mysticete whales. Syst. Biol. 56:77-90; Rychel, A. L., T. W. Reeder, and A. Berta. 2004. Phylogeny of mysticete whales based on mitochondrial and nuclear data. Mol. Phylogenet. Evol. 32:892-901), provide a nearly complete picture of the evolutionary history of baleen whales.     

Molecular lineage diversity and inter‐oceanic biogeographical history in Hiatella (Mollusca,Bivalvia)

Hiatella is one of the most widespread marine bivalve genera, occurring in diverse habitats from the temperate to polar latitudes in both hemispheres, and in fossil strata since almost 150 Myr ago. Despite variation in some biological and morphological traits, characters to resolve the current systematic structure consistently across the range of the genus are not known: all samples are often referred to a single species, Hiatella arctica (L.). Exploring the systematics of Hiatella using partial sequences of three genes (mitochondrial COI, and the nuclear ANT and 28S rRNA), we find high diversity of deep lineages (11–22% p‐distance in COI), and identify at least 13 distinct taxa both by heuristic criteria (congruence of the nuclear and mtDNA data) and by coalescence‐based analyses. At several localities, two or three of these cryptic species were found in sympatry. In the framework of previous fossil evidence and of hypotheses of paleoceanographical connections, scenarios of the phylogeny and biogeographical history of the identified species at a range of different time scales are outlined. The distinction between the main North Pacific and North Atlantic Hiatella clades and systematic diversification within each of them seem to have followed a Miocene trans‐Panamanian invasion. Apart from such earlier intra‐basin diversification, the data suggest that three successive counter‐invasions from the Pacific to the Atlantic via the Arctic Ocean route have later contributed to the current North Atlantic Hiatella diversity. These invasions probably took place in connection with (i) the Great Trans‐Arctic Biotic Interchange in the Pliocene, (ii) the last interglacial period c. 120 kya and (iii) the Holocene, postdating the last glaciation. This sequence of trans‐Arctic invasions is largely analogous to that hypothesized for some other boreal‐arctic bivalves.     

Towards a panbiogeography of the seas

A contrast is drawn between the concept of speciation favoured in the Darwin–Wallace biogeographic paradigm (founder dispersal from a centre of origin) and in panbiogeography (vicariance or allopatry). Ordinary ecological dispersal is distinguished from founder dispersal. A survey of recent literature indicates that ideas on many aspects of marine biology are converging on a panbiogeographic view. Panbiogeographic conclusions supported in recent work include the following observations: fossils give minimum ages for groups and most taxa are considerably older than their earliest known fossil; Pacific/Atlantic divergence calibrations based on the rise of the Isthmus of Panama at 3 Ma are flawed; for these two reasons most molecular clock calibrations for marine groups are also flawed; the means of dispersal of taxa do not correlate with their actual distributions; populations of marine species may be closed systems because of self‐recruitment; most marine taxa show at least some degree of vicariant differentiation and vicariance is surprisingly common among what were previously assumed to be uniform, widespread taxa; mangrove and seagrass biogeography and migration patterns in marine taxa are best explained by vicariance; the Indian Ocean and the Pacific Ocean represent major biogeographic regions and diversity in the Indo‐Australian Archipelago is related to Indian Ocean/Pacific Ocean vicariance; distribution in the Pacific is not the result of founder dispersal; distribution in the south‐west Pacific is accounted for by accretion tectonics which bring about distribution by accumulation and juxtaposition of communities; tectonic uplift and subsidence can directly affect vertical distribution of marine communities; substantial parallels exist between the biogeography of terrestrial and marine taxa; biogeographically and geologically composite areas are tractable using panbiogeographic analysis; metapopulation models are more realistic than the mainland/island dispersal models used in the equilibrium theory of island biogeography; and regional biogeography is a major determinant of local community composition. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 675–723.     

A rapidly deposited pennate diatom ooze in Upper Miocene-Lower Pliocene sediment beneath the North Pacific polar front

Rapidly deposited Thalassionema-Thalassiothrix pennate diatom oozes previously have been described in Upper Miocene-Lower Pliocene sediment beneath the frontal boundary of the eastern equatorial Pacific. Here we document a new occurrence of Thalassionema-Thalassiothrix ooze in Upper Miocene-Lower Pliocene sediment beneath the frontal boundary of the subarctic North Pacific. The ooze is a 6 m interval of siliceous sediment at Ocean Drilling Program (ODP) sites 885/886 that was rapidly deposited between approximately 5.0 and 5.9 Ma. Bulk sediment in this interval may contain greater than 85% pennate diatom tests. There are also abundant laminae and pockets that are composed entirely of Thalassionema and Thalassiothrix diatoms. The presence of a rapidly deposited ooze dominated by pennate diatoms indicates unusual past conditions in the overlying surface waters. Time coincident deposition of such oozes at two distinct frontal boundary locations of the Pacific suggests that the unusual surface water conditions were causally linked to large-scale oceanographic change. This same oceanographic change most likely involved (1) addition of nutrients to the ocean, or (2) redistribution of nutrients within the ocean. The occurrence and origin of pennate diatom oozes may be a key component to an integrative understanding of late Neogene paleoceanography and biogeochemical cycling.     

The systematics of right whales (Mysticeti: Balaenidae)

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.     

Phylogeny and biogeography of tetralophodont rodents of the tribe Oryzomyini (Cricetidae: Sigmodontinae)

Oryzomyini is the richest tribe among the Sigmodontine rodents, encompassing 32 living and extinct genera and including an increasing number of recently described species and genera. Some Oryzomyini are tetralophodont showing a reduction in the number of molar folds to four, while most taxa in this tribe retain the plesiomorphic pentalophodont state. We applied phylogenetic methods, molecular dating techniques and ancestral area analyses to members of an oryzomyini clade informally named ‘D’ in former studies and included related fossil tetralophodont forms. Based on 98 morphological characters and sequences of five gene fragments, we found that the tetralophodont condition is paraphyletic. Among living taxa, Pseudoryzomys is sister to Holochilus, and Lundomys is derived from a basal divergence. A clade formed by living Holochilus and the fossils Noronhomys and Carletonomys is sister to Holochilus primigenus, making Holochilus paraphyletic. Therefore, we describe a new genus that accommodates the fossil H. primigenus. Because trans‐Andean taxa currently share a common ancestor with taxa of cis‐Adean distribution, the northern Andes uplift may have worked as a postdispersal barrier. The tetralophodont lineages diverged during the Pliocene from a cis‐Andean ancestor, and the Great Plains in South America may have favoured the diversification of tetralophodont forms adapted to open habitats during the Pliocene.     

A scolopocryptopid centipede (Chilopoda: Scolopendromorpha) from Mexican amber: synchrotron microtomography and phylogenetic placement using a combined morphological and molecular data set

The first scolopocryptopid centipede known from the fossil record is a specimen of the subfamily Scolopocryptopinae in Miocene amber from Chiapas, southern Mexico. It is described here as Scolopocryptops simojovelensis sp. nov. , displaying a distinct combination of morphological characters compared to extant congeners. Anatomical details of the fossil specimen were acquired by non‐invasive 3D synchrotron microtomography using X‐ray phase contrast. The phylogenetic position of the new species is inferred based on a combination of morphological data with sequences for six genes (nuclear 18S and 28S rRNA, nuclear protein‐coding histone H3, and mitochondrial 12S rRNA, 16S rRNA, and protein‐coding cytochrome c oxidase subunit I) for extant Scolopendromorpha. The data set includes eight extant species of Scolopocryptops and Dinocryptops from North America, east Asia, and the Pacific, rooted with novel sequence data for other blind scolopendromorphs. The molecular and combined data sets, analysed in a parsimony/direct optimization framework, identified a stable pattern of two main clades within Scolopocryptopinae. North American and Asian species of Scolopocryptops are united as a clade supported by both morphological and molecular characters. Its sister group is a Neotropical clade in which the type species of Dinocryptops is nested within a paraphyletic assemblage of Scolopocryptops species; Dinocryptops is placed in synonymy with Scolopocryptops. The strength of support for the relationships of extant taxa from the molecular data allow the Chiapas fossil to be assigned with precision, despite ambiguity in the morphological data; the fossil is resolved as sister species to the extant Laurasian clade. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 768–786.