‘After Africa’: the evolutionary history and systematics of the genus Charaxes Ochsenheimer (Lepidoptera: Nymphalidae) in the Indo‐Pacific region

The predominantly Afrotropical genus Charaxes is represented by 31 known species outside of Africa (excluding subgenus Polyura Billberg). We explored the biogeographic history of the genus using every known non‐African species, with several African species as outgroup taxa. A phylogenetic hypothesis is proposed, based on molecular characters of the mitochondrial genes cytochrome oxidase subunit I (COI) and NADH dehydrogenase 5 (ND5), and the nuclear wingless gene. Phylogenetic analyses based on maximum parsimony and Bayesian inference of the combined dataset implies that the Indo‐Pacific Charaxes form a monophyletic assemblage, with the exception of Charaxes solon Fabricius. Eight major lineages are recognized in the Indo‐Pacific, here designated the solon (+African), elwesi, harmodius, amycus, mars, eurialus, latona, nitebis, and bernardus clades. Species group relationships are concordant with morphology and, based on the phylogeny, we present the first systematic appraisal and classification of all non‐African species. A biogeographical analysis reveals that, after the genus originated in Africa, the evolutionary history of Charaxes in the Indo‐Pacific, in particular Wallacea, may be correlated with the inferred geological and climatic history of the region. We propose that Wallacea was the area of origin of all Charaxes (excluding C. solon) occurring to the east of Wallace's [1863] Line. The earliest Indo‐Pacific lineages appear to have diverged subsequent to the initial fragmentation of a palaeo‐continent approximately 13 million years ago. Further diversification in Indo‐Pacific Charaxes appears primarily related to climatic changes during the Pliocene and possibly as recently as the Pleistocene. Although both dispersal and vicariance have played important roles in the evolution of the genus within the region, the latter has been particularly responsible for diversification of Charaxes in Wallacea. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 457–481.     

High frequency of drill holes in brachiopods from the Pliocene of Algeria and its ecological implications

The fossil record holds a wealth of ecological data, including data on biotic interactions. For example, holes in the skeletons of invertebrates produced by drilling activities of their enemies are widely used for exploring the intensity of such interactions through time because they are common and easily distinguished from non-biotic holes or holes produced by other types of interactions. Such drill holes have been described in numerous studies of Palaeozoic brachiopods but rarely in those focusing on brachiopods of the post-Palaeozoic, a striking pattern given that in the late Mesozoic and Cenozoic drilling gastropods diversified and frequencies of drilled molluscs increased dramatically. During the past several years, however, drilled brachiopods were reported in several studies of the Mesozoic and Cenozoic, suggesting that this phenomenon may be more common than has been previously assumed. Here we report on drilled brachiopods from a Pliocene locality in Algeria where 90 of 261 (34.5%) specimens of Megerlia truncata show evidence of predatory drilling. These data confirm that Cenozoic drilling frequencies of brachiopods may be locally high and, when taken together with other published data, that drilling frequencies are highly heterogeneous in space and time.     

Pleistocene fish otoliths from the Mediterranean Basin: a synthesis

An overview of upper pPliocene and pleistocene otolith assemblages is compiled on the basis of both literature data and newly collected material from several sections located mainly in southern Italy. One hundred and five taxa are listed. Additional comments are provided for taxa subject to discussion. The composition and affinities of the Mediterranean Pleistocene otolith associations (consisting mainly of deep sea fishes) is checked against the available data for Pre-Messinian, Pliocene, and Recent Mediterranean fishes. This analysis is based on the recorded nominal species, but some taxa that could be identified at the generic level only are also included when they are relevant from a biogeographic point of view. Some new data on Piacenzian (middle Pliocene) and Gelasian (upper Pliocene) otoliths are also provided because this time interval still constitutes a major gap in the knowledge of the stratigraphic range of Plio-Pleistocene fish taxa. The Pleistocene deepwater fish fauna of the Mediterranean shows a markedly more oceanic character than the present-day one, a tendency that was all ready observed from early Oligocene till Pliocene times. Starting from the late Piacenzian, an increasing number of Atlantic taxa progressively invaded the Mediterranean including several subpolar and temperate forms. This invasion became stronger in the Gelasian. From this period up to the middle and late Pleistocene, the Mediterranean deepwater fish fauna (mesopelagic and benthopelagic) is enriched by taxa which today are typical for the north Atlantic cold deep water. This faunal shift seems to be correlated with the evolution of the Plio-Pleistocene Mediterranean paleoceanographic setting as well as with the global climatic deterioration.     

In situ occurrence of a gall midge (Insecta,Diptera, Cecidomyiidae) on fossilized angiosperm leaf cuticle fragments from the Pliocene sediments of eastern India

In situ preservation of fossil insect damage in plant fossils is an excellent tool to study the coevolution of flora and fauna through geological time, but finding both damage and the insect causing that damage in the same specimen is a very rare phenomenon. Galling is a common form of angiosperm leaf damage, which can be regarded as a kind of extended phenotype of the causal insects, essentially the gall midges, but galls usually lack remains of the insects themselves. Here we report the in situ occurrence of a gall midge (Insecta, Diptera, Cecidomyiidae) as well as its pupal exuviae on the abaxial cuticular surface of fossilized leaf cuticle fragments of Fabaceae leaves (cf. Albizia) that also bear galls, recovered from the latest Neogene (Rajdanda Formation, Pliocene) sediments of the Chotonagpur Plateau, Jharkhand, northeastern India. This Pliocene gall midge features well-preserved legs, segmented antenna with distinct and enlarged scape, elongate curved setae, and longer than broad terminal plate of the ovipositor lamellae. The in situ presence of a gall midge on a host fabaceous leaf cuticle indicates the existence of a host-ectoparasite relationship in the ancient warm and humid tropical monsoon-influenced forests of eastern India during the Pliocene. This is the first authentic fossil record of an in situ phytophagous insect of Cecidomyiidae from India, as well as southeast Asia. Although the identification of the recovered phytophagous insect associated with the fossil leaf cuticle is only possible to family level, this find reveals that such plant-insect relationships existed in the Pliocene of eastern India.     

Pre-Quaternary hominin settlements in Asia: Archaeology,bio-lithostratigraphy and magnetostratigraphy evidences at Masol,Siwaliks, Northwestern India

The National Museum of Natural History, Paris, the National Centre for Scientific Research (CNRS/UMR 7194 and 8148) and the Society for Archaeological and Anthropological Research, Chandigarh, conduct paleontological and archaeological surveys in a sub-Himalayan territory well known for its Upper Pliocene fossiliferous deposits and named the Quranwala Zone. Since 2007, the team is investigating this formation in the Masol anticline and has collected four fossils with marks of butchery as well as numerous lithic artifacts. In this paper we show the first chopper extracted in stratigraphy from the deepest fossiliferous layer at more than 2.6 Ma as shown by the undisputable magnetostratigraphy, we describe the lithostratigraphy of the excavation, the techno-typology of the chopper and compare it with the collection of its locality and two other major paleonto-archaeological sites of Masol. The characteristics of this Masol lithic assemblage do not match those of the early Lower Pleistocene African Oldowan (2.55 Ma), but they are similar to those of the Chinese lithic industry of Longgupo (2.5 Ma). These activities open new perspectives regarding the phylogenetic and geographical origins of these Asian hominins as well as on their descendants, and justify the pursuit of surveys and test pits in the Quranwala zone.     

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).     

Quantitative calcareous nannofossil biostratigraphy of the subsurface Pliocene sequence,north-east Nile Delta,Egypt

Detailed quantitative calcareous nannofossil biostratigraphy was carried out on a subsurface lower–upper Pliocene succession (Kafr El Sheikh Formation) in three offshore wells (Ras El Barr-1, Bougaz-1 and Bougaz East-1), north-east Nile Delta area, Egypt. The standard zonations are easily applied in these sediments and additional biohorizons have been adopted in this study. For early to late Pliocene interval, the following five bioevents are considered reliable: last occurrences (LOs) of Discoaster surculus, D. tamalis, Reticulofenestra pseudoumbilicus and Amaurolithus tricorniculatus and first occurrence of D. asymmetricus. The LO of R. pseudoumbilicus was used to detect the early/late Pliocene boundary in the three studied wells. In Bougaz-1-Well, the LO of D. intercalcaris was used to approximate the Miocene/Pliocene boundary. The calcareous nannofossils are moderate to good preserved. Lower–upper Pliocene sediments are dominated by small-sized reticulofenestrids (R. minuta), Pseudoemiliania lacunosa, Helicosphaera carteri and H. kamptneri. Large size (>7 μm) R. pseudoumbilicus is recorded. Discoaster species is rarely represented. No nannofossils were found in the top part of the Kafr El Sheikh and the overlying Wastani Formations.     

SKULL MORPHOLOGY AND PHYLOGENETIC RELATIONSHIPS OF A NEW DIMINUTIVE BALAENID FROM THE LOWER PLIOCENE OF BELGIUM

Abstract:  A new small balaenid is described and compared to all fossil and living balaenid taxa. The specimen represents a new genus and species and is named Balaenella brachyrhynus . It was discovered in the Lower Pliocene of Kallo (north-west Antwerp, Belgium) and adds new information on the diversity and evolution of Balaenidae. Based on both comparative morphology and phylogenetic analysis, Balaenella brachyrhynus is morphologically closer to the genus Balaena , including the living Greenland Bowhead whale ( B. mysticetus ), and two Pliocene species ( B. montalionis and B. ricei ) from central Italy and the eastern USA. Balaenella brachyrhynus has very short nasals, a short rostrum relative to the total skull length and a horizontal supraoccipital. A cladistic treatment of 81 morphological character states scored for 10 balaenids and nine non-balaenid cetaceans revealed that the other small balaenids generally included in the genus Balaenula (including Balaenula astensis, B. balaenopsis and a Pliocene Balaenula sp. from Japan) are closer to the living genus Eubalaena (the Right whale). As the new skull is so different from the nominal Balaenula species, and as it is more closely related to Balaena than to Eubalaena , it is concluded that a small body size was a common condition in different Balaenidae clades.     

Late Miocene vegetation and climate reconstruction based on pollen data from the Sofia Basin (West Bulgaria)

The analysis of fossil palynomorh assemblages in the Late Miocene freshwater sediments of the Sofia Basin (West Bulgaria) was done to collect data on the vegetation and climate dynamics during the Late Miocene. On the basis of pollen data, we described the main palaeocomunities developed in the region. The mixed mesophytic forests dominated the vegetation in which species of Quercus, Ulmus, Zelkova, Fagus, Carpinus, Betula, Castanea, Corylus, Pterocarya, Carya, Juglans, and Eucommia played important roles. Swamp forests were also recorded, including Taxodiaceae, Alnus, Glyptostrobus, Nyssa, and Myrica. Herbaceous vegetation was distributed in the middle part of the section, with a maximum of 35.5%. The vegetation dynamic passes through several phases, which were associated with changes in paleoclimate and palaeoecological conditions. Coexistence Approach (CA) was applied to palynological data to calculate four climatic parameters. The values of coexistence intervals for mean annual temperatures are 13.6–16.6 °C, with winter temperatures being 3.7–6.6 °C and summer temperatures being 23.6–27.8 °C. Mean annual precipitation ranged most frequently between 828 and 1308 mm. The palaeoclimatic reconstruction illustrates existence of a warm-temperate and relatively humid climate with higher mean annual temperature than the present day climate.     

Pleistocene genetic connectivity in a widespread,open‐habitat‐adapted mosquito in the Indo‐Oriental region

Aim The environmental effect of Pleistocene climatic change in the Indo‐Oriental region has resulted in allopatric fragmentation and the generation of diversity in forest‐associated species. The aim of this study was to determine the extent to which Pleistocene climatic change has resulted in the fragmentation and speciation of an open‐habitat‐adapted mosquito, Anopheles vagus s.l., across its range. Location Anopheles vagus s.l. was sampled across the Indo‐Oriental region. Methods We generated 116 mitochondrial cytochrome c oxidase subunit I (COI) and 121 nuclear internal transcribed spacer 2 (ITS2) DNA sequences from 18 populations. Relationships between mitochondrial haplotypes were reconstructed using minimum spanning networks, and population structure was examined using analyses of molecular variance. The population history, including lineage divergence times, population expansion and gene flow, was inferred using beast and the isolation with migration (IM) model. Results There was no evidence to support the presence of the endemic Philippines species, A. limosus; instead, Philippine populations were closely related to, and derived from, A. vagus on the eastern Southeast Asian mainland. The most distinct populations were those from Java and East Timor, which differed from all other populations by all individuals having a 4‐bp insertion in the ITS2 sequence. The corresponding mitochondrial haplotypes had an estimated divergence time of 2.6 Ma [95% confidence interval (CI) 1.9–3.6 Ma]. Haplotype networks and analysis of molecular variance for COI supported western (Sri Lanka, India and Myanmar) and eastern (Thailand, Singapore, Cambodia, Vietnam and the Philippines) population groupings. This grouping structure results from the divergence of an eastern and a western mitochondrial lineage, estimated to have occurred 0.37 Ma (95% CI 0.26–0.55 Ma). Subsequent migration from the east to the west (0.16 Ma) is inferred to have created an admixture zone in Myanmar and Thailand. Main conclusions With the possible exception of populations from Java and East Timor, A. vagus appears to be one widespread genetically diverse taxon across its extensive range. The abundance of grassland during long interglacial periods may have facilitated population connectivity and range expansion across the Oriental and western Australasian regions.