Early Miocene subtropical water temperatures in the southeast Pacific

Cenozoic climate of western South America is strongly controlled by features like Andean uplift and the Humboldt Current. The first strontium isotope age data from central and southern Chile provide a latest Oligocene to late early Miocene age for classic warm-water mollusk faunas reaching as far south as 45°S. Comparison with the biogeography of congeneric living species indicates that sea surface temperatures off central and southern Chile during that time were at least 5 °C higher than today; i.e., minimum annual mean sea surface temperatures for Darwin's Navidad fauna at 34°S are estimated as 20 °C. As expected, the number of tropical taxa decreases towards the south but several are still present as far south as 45°S. The ages scatter relatively broadly between ~ 24 and ~ 16 Ma, partly even within individual localities. Shallow-water and deeper-water faunas are revealed to have similar ages. When considered in light of convincing micropaleontological evidence for late Miocene to early Pliocene depositional ages, the Sr isotope data support a hypothesis that the mollusk fauna is reworked.     

MIOCENE COMMUNITIES AND PALEOENVIRONMENTS OF SOUTHERN COSTA RICA

Three Tertiary microfossil floras from northern Latin America are considered with reference to paleoenvironments and paleoenvironmental trends, especially paleotemperatures: Gatuncillo (middle (?) to late Eocene, Panama), Uscari (early Miocene, Costa Rica), and Paraje Solo (late Miocene, Veracruz, Mexico). The composition of the floras reflect lower temperatures in the late Miocene for terrestrial environments at ca. 18°N latitude, and temperatures comparable to the present for the middle (?) to late Eocene and early Miocene at ca. 9°N latitude. These values are consistent with Tertiary paleotemperature curves derived from ¹⁸O and ¹³C isotope studies of marine invertebrates, and show that the global drop in temperature at the end of the Miocene affected terrestrial biotas at least as far south as 18°N latitude. Comparison of the paleobotanical data with paleotemperature curves further reveals that the late Miocene was not only a likely time for the introduction of northern temperate elements into the Latin American biota (viz., establishment of the floristic and faunal relationship between eastern United States and eastern Mexico), but that earlier times were progressively less favorable because of generally rising temperatures and more tropical conditions. Information on the Tertiary history of vegetation in the Gulf/Caribbean region is also being monitored with reference to the effect of global sea level changes, although the specific influence of these fluctuations cannot as yet be detected.     

Phytoextraction potential of the nickel hyperaccumulators Leptoplax emarginata and Bornmuellera tymphaea

Leptoplax emarginata and Bornmuellera tymphaea are nickel hyperaccumulators of the Brassicaceae family endemic to serpentine soils in Greece. The aims of this work were to compare the growth and uptake behavior of these plants with the Ni hyperaccumulator species Thlaspi caerulescens and Alyssum murale, and to evaluate their effect on soil Ni availability. Plants were grown for 3 mo on three soils that differ in Ni availability. Ni availability in soils was measuredby isotopic exchange kinetics and DTPA-TEA extractions. Results showed that L. emarginata produced significantly more biomass than other plants. On the serpentine soil, B. tymphaea showed the highest Ni concentration in shoots. However, Niphytoextraction on the three soils was maximal with L. emarginata. The high initial Ni availability of soil Serp (470.5 mg kg(-1)) was the main explanation for the high Ni concentrations measured in plant shoots grown on this soil, compared to those grown on soils Calc and Silt A. murale was the least efficient in reducing Ni availability on the serpentine soil L. emarginata appeared as the most efficient species for Ni phytoextraction and decrease of the Ni available pool.     

Scorched mussels (Brachidontes spp., Bivalvia: Mytilidae) from the tropical and warm‐temperate southwestern Atlantic: the role of the Amazon River in their speciation

Antitropicality is a distribution pattern where closely related taxa are separated by an intertropical latitudinal gap. Two potential examples include Brachidontes darwinianus (south eastern Brazil to Uruguay), considered by some authors as a synonym of B. exustus (Gulf of Mexico and the Caribbean), and B. solisianus, distributed along the Brazilian coast with dubious records north of the intertropical zone. Using two nuclear (18S and 28S rDNA) and one mitochondrial gene (mtDNA COI), we aimed to elucidate the phylogeographic and phylogenetic relationships among the scorched mussels present in the warm‐temperate region of the southwest Atlantic. We evaluated a divergence process mediated by the tropical zone over alternative phylogeographic hypotheses. Brachidontes solisianus was closely related to B. exustus I, a species with which it exhibits an antitropical distribution. Their divergence time was approximately 2.6 Ma, consistent with the intensification of Amazon River flow. Brachidontes darwinianus, an estuarine species is shown here not to be related to this B. exustus complex. We suspect ancestral forms may have dispersed from the Caribbean to the Atlantic coast via the Trans‐Amazonian seaway (Miocene). The third species, B rodriguezii is presumed to have a long history in the region with related fossil forms going back to the Miocene. Although scorched mussels are very similar in appearance, their evolutionary histories are very different, involving major historical contingencies as the formation of the Amazon River, the Panama Isthmus, and the last marine transgression.     

Patterns of population differentiation in annual killifishes from the Paraná–Uruguay–La Plata Basin: the role of vicariance and dispersal

Aim To elucidate the role of vicariance versus dispersal at the microevolutionary scale in annual killifish populations belonging to the Austrolebias bellottii species complex (Rivulidae). Within this complex, A. bellottii and A. apaii have low vagility and occur widely within the study area, making them excellent models for testing biogeographic hypotheses of differentiation. Location South America, in the Paraná–Uruguay–La Plata river basin. Methods Molecular data and morphometric analyses were used to reconstruct the phylogeographic history and morphological variation of 24 populations of two taxa of the A. bellottii species complex. Phylogenetic analyses using maximum likelihood (ML) and Bayesian inference (BI) model‐based methods, estimates of clade divergence times implemented in beast , non‐metric multidimensional scaling, analysis of molecular variance results, and morphological analyses elucidated the role of vicariance versus dispersal hypotheses in population differentiation in the aforementioned river basin. Results In the A. bellottii species complex from the Paraná–Uruguay–La Plata river basin, past allopatric fragmentation from vicariance events seems to be the most plausible scenario for diversification since the Late Miocene and more recently since the Plio‐Pleistocene. The Plio‐Pleistocene vicariance produced the differentiation of three major clades in A. bellottii populations. One clade from the eastern Uruguay River drainage was separated from another in western Uruguay and the Paraná–La Plata River drainages. A later vicariance event split populations to the south (lower Paraná–La Plata Basin) and north (middle Paraná) of the western Paraná River drainage. However, our results do not exclude the possibility of dispersal events among A. bellottii populations from both the Uruguay and Paraná river drainages, which could occur in these river basins during hypothesized connectivity cycles of the Late Pliocene and Pleistocene. Main conclusions Past allopatric fragmentation caused by different vicariance events seems to be the main driver of diversification in the A. bellottii species complex since the Plio‐Pleistocene. However, the current molecular data suggest that populations from both drainages of the Paraná–Uruguay rivers may have experienced cycles of connectivity during the Pleistocene, perhaps including multiple vicariance or dispersal events from populations located in the western lower Uruguay River drainage, which encompassed climatic and geological changes in the Paraná–Uruguay–La Plata Basin.     

HISTORICAL ALLOPATRY AND THE BIOGEOGRAPHY OF SPECIATION IN THE PROSOBRANCH SNAIL GENUS NUCELLA

Two recently diverged northeastern Pacific sibling snail species, Nucella ostrina and N. emarginata, currently inhabit adjacent zoogeographic provinces. Their distributions overlap in central California to the north of a major faunal boundary at Point Conception, California (PC). To test the hypothesis that modern sympatry is due to a recent northward range expansion by N. emarginata, I analyzed the population structures of both species with nuclear (allozyme) and mitochondrial DNA (mtDNA) markers. Populations of N. emarginata in the region of overlap exhibit significantly lower heterozygosity and allelic diversity than either populations to the south of PC or populations of N. ostrina. A single mtDNA haplotype characterizes all but one population of N. emarginata sampled in this region, but no haplotype to the south of PC is found at more than one locality. MtDNA haplotypes and allozyme allele frequencies also indicate monophyly of central California populations of N. emarginata. Sharp differences in allelic diversity over small geographic distances may reflect the action of natural selection, but because both nuclear and mtDNA markers display concordant patterns, a range expansion across PC best explains patterns of genetic variation in N. emarginata. Allozymes and mtDNA also reveal that the geologically older N. ostrina is paraphyletic with respect to N. emarginata. This pattern is consistent with, but not indicative of, a peripheral isolation model of speciation. Low genetic diversity is also expected if a significant bottleneck occurred at speciation. However, low allelic diversity is not universal throughout the geographic range of N. emarginata; high allelic diversity at the southern end of the distribution of N. emarginata suggests that in the past N. emarginata has been geographically restricted much further south than PC. A northward range expansion across PC by N. emarginata may thus represent only the most recent postglacial movement by the species. The thermal and oceanographic discontinuities found at PC may not have been directly involved in geographic isolation if N. emarginata originated much further south of this modern boundary. Despite uncertainty regarding the exact spatial distribution of populations at speciation, genetic data indicate that even though N. ostrina and N. emarginata currently exhibit a broad range of geographic overlap, speciation was likely allopatric and was initiated by physical isolation of populations in different zoogeographic provinces.     

Late Neogene nannofossil biostratigraphy and paleoceanography of the northeastern Gulf of Mexico and adjacent areas

Two 305 m cored sections from the northwest Florida continental shelf contain a nearly complete record of late Neogene hemipelagic sedimentation. One of the sites, south and east of De Soto Canyon, is isolated from terrigenous sediment except for sediment transported in suspension. This site contains a continuous record from the late Miocene to the Recent. The second site, on the western rim of De Soto Canyon, is more expanded and continuous from the late Pliocene to the Recent. A hiatus separates the late Pliocene from the middle Miocene. Six prominent nannofossil biohorizons were recognized within the Pleistocene, seven within the Pliocene, and three within the Miocene; in addition one biohorizon marks the base of the Pleistocene and another the base of the Pliocene.Nearly all carbonate in the sediment is pelagic. Terrigenous detrital sedimentation was controlled by glacioeustatic sea level fluctuations during the Pleistocene, and sea level changes are probably responsible for fluctuations in the ratio of pelagic carbonate to clayey detritus in pre-Pleistocene sediments also. Carbonate content, coarse fraction percent, and relative abundances of environmentally sensitive nannoplankton species suggest important paleoceanographic changes in the northeastern Gulf of Mexico and adjacent areas. Fluctuations in the relative abundance of the solution-resistant coccoliths of the genusCyclococcolithus indicate that waters at a depth of 600–1000 m were more corrosive during the late Miocene than they are today. The decrease in carbonate dissolution during the late Miocene probably was a response to gradual constriction of the Central American passage and the consequent restriction of flow of corrosive water from the Pacific. Short term fluctuations in dissolution during the Pliocene and Pleistocene are related to climatic cycles.Productivity variations in the surface waters, recorded mainly by the relative abundance of small and large morphotypes of closely related coccolith species, indicate that productivity increased during the Pliocene, but the most dramatic change — a major oceanwide increase in productivity — occurred during the Pleistocene, during and just prior to the Jaramillo magnetic event about 0.9 m.y. ago. Surprisingly the late Miocene Messinian event did not leave a significant imprint in the northeastern Gulf of Mexico.     

Some observations on shell shape variation in Pacific Nucella

This paper considers the patterns of shell shape variation shown by Nucella canalicuata, N. emarginata and N. lamellosa from two areas of the Pacific Northwest: the shores near Friday Harbour on San Juan Island and near Bamfield on the west coast of Vancouver Island. No clear pattern of variation in association with changes in exposure was seen in either N. canaliculata or N. lamellosa . It appears that genetic influences are more important controls of shell shape than environmental selection in both these species. Nucella emarginata shows the nearest approximation to the pattern shown by the Atlantic species, N. lapillus , but only at the exposed end of the wave-action gradient. On those shores, enclaves from the most surf-washed open coast headlands have shells with proportionally larger apertures (and thus a shorter, squatter form) than their equivalents in local shelter. But, unlike in N. lapillus , the trend does not continue onto genuinely sheltered shores. Under these circumstances the species is generally rare and, where enclaves do occur, their shells are of much the same shape (although of a much larger size) as in more exposed situations.     

Pleistocene and Holocene rhinocerotids (Mammalia,Perissodactyla) from the Indochinese Peninsula

Rhinocerotids were abundant and diverse in southern Asia during the Pleistocene and the Holocene epochs, as shown by palaeontological and archaeological discoveries published throughout the last century, whereas the only living rhinoceros in the Indochinese Peninsula is Rhinoceros sondaicus (Cat Loc Reserve, Vietnam). The Pleistocene-Holocene Indochinese rhinocerotid record consists of the extinct species Dicerorhinus gwebinensis (Early Pleistocene, Myanmar) and representatives of the Recent Asian Species Rhinoceros unicornis (Middle-Late Pleistocene), R. sondaicus (Middle Pleistocene-Recent), and Dicerorhinus sumatrensis (Middle Pleistocene-Holocene). This fossil record is synthesized, mapped for Early/Middle/Late Pleistocene and Holocene/Recent times, and then compared with coeval rhinocerotid assemblages from the adjacent areas (South China), subregions (Indian, Sundaic, Philippine, and Wallacean), and region (Palearctic), from a biochronological and biogeographical perspective.     

Effects of temperature on the respiration rates and the kinetics of citrate synthase in two species of Idotea (Isopoda, Crustacea)

The two species of isopods, Idotea baltica (Pallas) and Idotea emarginata (Fabricius), co-occur frequently near Helgoland, North Sea, occupying different ecological niches. Respiration rates and kinetic properties of citrate synthase (CS) were compared in these species in order to identify possible mechanisms of temperature adaptation. Specimens were acclimated to 5 and 15 degrees C prior to further investigations. Respiration rates were measured under normoxic conditions at 5, 10 and 15 degrees C. CS was partly purified chromatographically and influences of temperature, pH, substrate saturation and ATP-concentration on enzyme activity were examined. In both species, rising temperatures led to linearly increasing oxygen consumption, with estimated Q10 values between 3.2 and 4.2. Only I. baltica showed an effect of short term acclimation: warm adapted animals had always higher respiration rates than cold adapted ones. In I. emarginata, the acclimation temperature had no effect on oxygen consumption. Furthermore, its CS slightly indicates higher affinity to oxaloacetic acid when specimens were adapted to 15 degrees C compared to those maintained at 5 degrees C. Any effect of the experimental temperature on CS in I. baltica was negligible. The results are discussed in view of the different habitats occupied by the species compared.     

Sea-surface temperature changes in the North Pacific during the Late Miocene

Twenty-seven radiolarian species and species groups are identified and used in this study. Their stratigraphic ranges span from Late Miocene to modern times and they appear to have distributions which have changed little over this interval of time. Census data for these 27 taxa in surface sediment samples are used to define six assemblages (Q-mode factors). The distribution of these assemblages are similar to modern water-mass distributions and can be statistically related to modern sea-surface temperatures.Census data on these same 27 taxa have been collected in the Upper Miocene intervals of six Pacific sites. These faunal data are described in terms of the modern radiolarian assemblages; and based on the multiple regression equations relating the modern assemblages to modern temperatures, estimates of Late Miocene temperatures are made. The estimated temperatures indicate that there is an overall cooling trend in the later part of the Miocene (magnetic Chron 8 to Chron 5), punctuated by several distinct cooling events. In August the low-latitude sites appear to be slightly cooler, and the mid-latitude sites warmer than in modern times. In February both the low- and mid-latitude sites appear to be warmer than modern temperatures at the same locations. The relatively warm temperatures near the base of the interval studied may be associated with the passage of warm tropical Atlantic waters into the eastern Pacific via the Isthmus of Panama. The general cooling trend seen in these records is thought to be associated with the gradual closure of these straits and the increased influence of the high southern latitudes.     

Rainforest air-conditioning: the moderating influence of epiphytes on the microclimate in tropical tree crowns

Epiphytes are often assumed to influence the microclimatic conditions of the tree crowns that they inhabit. In order to quantify this notion, we measured the parameters "temperature" (of the substrate surface and the boundary layer of air above it), "evaporative drying rate" and "evapotranspiration" at various locations within tree crowns with differing epiphyte assemblages. The host tree species was Annona glabra, which was either populated by one of three epiphyte species (Dimerandra emarginata, Tillandsia fasciculata, or Vriesea sanguinolenta) or was epiphyte-free. We found that during the hottest and driest time of day, microsites in the immediate proximity of epiphytes had significantly lower temperatures than epiphyte-bare locations within the same tree crown, even though the latter were also shaded by host tree foliage or branches. Moreover, water loss through evaporative drying at microsites adjacent to epiphytes was almost 20% lower than at exposed microsites. We also found that, over the course of several weeks, the evapotranspiration in tree crowns bearing epiphytes was significantly lower than in trees without epiphytes. Although the influence of epiphytes on temperature extremes and evaporation rates is relatively subtle, their mitigating effect could be of importance for small animals like arthropods inhabiting an environment as harsh and extreme as the tropical forest canopy.     

The Mediterranean hydrologic budget from a Late Miocene global climate simulation

During the Late Miocene the Mediterranean experienced a period of extreme salinity fluctuations known as the Messinian Salinity Crisis (MSC). The causes of these high amplitude changes in salinity are not fully understood but are thought to be the result of restriction of flow between the Mediterranean and Atlantic, eustatic sea level change and climate. Results from a new Atmospheric General Circulation Model (AGCM) simulation of Late Miocene climate for the Mediterranean and adjacent regions are presented here. The model, HadAM3, was forced by a Late Miocene global palaeogeography, higher CO₂ concentrations and prescribed sea surface temperatures. The results show that fluvial freshwater fluxes to the Mediterranean in the Late Miocene were around 3 times greater than for the present day. Most of this water was derived from North African rivers, which fed the Eastern Mediterranean. This increase in runoff arises from a northward shift in the intertropical convergence zone caused by a reduced latitudinal gradient in global sea surface temperatures. The northwards drainage of the Late Miocene Chad Basin also contributes. Numerical models designed to explore Late Miocene salt precipitation regimes in the Mediterranean, which typically make use of river discharge fluxes within a few tens of percent of present-day values, may therefore be grossly underestimating these fluxes.Although the AGCM simulated Late Miocene river discharge is high, the model predicts a smaller net hydrologic budget (river discharge plus precipitation minus evaporation) than for present day. We discuss a possible mechanism by which this change in the hydrologic budget, coupled with a reduced connection between the Mediterranean and the global ocean, could cause the salinity fluctuations of the MSC.     

Preliminary investigation of dental microstructure in the Yuanmou hominoid (Lufengpithecus hudienensis), Yunnan Province,China

Fieldwork in the Yuanmou Basin of southern China has uncovered a large assemblage of late Miocene hominoid fossils assigned to Lufengpithecus hudienensis. Two mandibular first molars from this species were made available for histological analysis as part of a larger ongoing study on the ontogeny of dental development in Miocene to Recent hominoids. Results are compared with published and unpublished data on tooth growth in a wide range of extant and extinct hominoids. The Yuanmou molars are smaller than those of Lufengpithecus lufengensis and have markedly shorter crown formation times, overlapping slightly with Pan, but most similar to Proconsul and Dryopithecus. In other aspects of molar development (including enamel extension rates and enamel thickness), L. hudienensis shows similarities with all extant hominoids, in particular, Pongo. Ultimately, charting the ontogeny of molar crown formation may help shed light on the relationship of Lufengpithecus hudienensis to orangutans, and other Miocene to Recent hominoids.     

How caterpillars avoid overheating: behavioral and phenotypic plasticity of pipevine swallowtail larvae

We tested the hypothesis that larvae of the pipevine swallowtail butterfly, Battus philenor, employ behavioral and phenotypic plasticity as thermoregulatory strategies. These larvae are phenotypically varied across their range with predominantly black larvae (southeastern USA and California) and red larvae (western Texas, Arizona) occurring in different regions. Two years of field observations in south Texas indicate that the proportion of red larvae increases with increasing daily temperatures as the growing season progresses. Larvae were also observed to shift their microhabitats by climbing on non-host vegetation and avoided excessive heat in their feeding microhabitat. Larvae of ten half-sib families from populations in south Texas and California, reared under different temperature regimes in common garden experiments, exhibited plasticity in larval phenotype, with larvae from both populations producing the red phenotype at temperatures greater than 30°C and maintaining the black phenotype at cooler temperatures. However, larvae from Texas were more tolerant of higher temperatures, showing no decrease in growth rate in the highest temperature (maximum seasonal temperature) treatment, compared to the California population. In a field experiment, black larvae were found to have higher body temperatures when exposed to sunlight compared to red larvae. These results suggest that microhabitat shifts and the color polyphenism observed in pipevine swallowtail larvae may be the adaptive strategies that enable larvae to avoid critical thermal maximum temperatures.     

Un nouveau gisement miocène à ichthyofaune au sud de la chaîne des Chotts (Tunisie méridionale). Paléoenvironnement et paléobiogéographie

A new fish-bearing Miocene site, south of the ‘Chaîne des Chotts’ (southern Tunisia). Palaeoenvironment and palaeobiogeography. A systematic excavation in the Miocene continental sands of the Béglia Formation (southwestern Tunisia), south of the ‘Chaîne des Chotts’, yielded a rich fossil-fish assemblage. These sands, with soft rollers and oblique stratifications are in unconformity on the Cretaceous and are covered by Miocene clays. The organisation and the nature of the facies, as well as the palaeocurrent figures testify to a proximal fluvial-deltaic regime. It is SW–NE prograding. The three fish taxa (polypterid, clariid, latid) are typical of modern Nilo-Sudanese assemblages, but are absent from Maghreb. The regional palaeobiogeographic relations are discussed. To cite this article: B. Mannaï-Tayech, O. Otero, C. R. Palevol 4 (2005).     

Late Miocene paleoclimatology: Subantarctic water mass,Southwest Pacific

The Late Miocene—Early Pliocene paleoclimatic history has been evaluated for a deep drilled sediment sequence at Deep Sea Drilling Project Site 281 and a shallow water marine sediment sequence at Blind River, New ealand, both of which lay within the Subantarctic water mass during the Late Miocene.A major, faunally determined, cooling event within the latest Miocene at Site 281 and Blind River coincides with oxygen isotopic changes in benthonic foraminiferal composition at DSDP Site 284 considered by Shackleton and Kennett (1975) to indicate a significant increase in Antarctic ice sheet volume. However, at Site 281 benthonic foraminiferal oxygen isotopic changes do not record such a large increase in Antarctic ice volume. It is possible that the critical interval is within an unsampled section (no recovery) in the latest Miocene.Two benthonic oxygen isotopic events in the Late Miocene (0.5‰ and 1‰ in the light direction) may be useful as time-stratigraphic markers. A permanent, negative, carbon isotopic shift at both Site 281 and Blind River allows precise correlations to be made between the two sections and to other sites in the Pacific region. Close interval sampling below the carbon shift at Site 281 revealed dramatic fluctuations in surface-water temperatures prior to a latest Miocene interval of refrigeration (Kapitean) and a strong pulse of dissolution between 6.6 and 6.2 ± 0.1 m.y. which may be related to a fundamental geochemical change in the oceans at the time of the carbon shift (6.3?6.2 m.y.). No similar close interval sampling at Blind River was possible because of a lack of outcrop over the critical interval.Paleoclimatic histories from the two sections are very similar. Surface water temperatures and Antarctic ice-cap volume appear to have been relatively stable during the late Middle—early Late Miocene (early—late Tongaporutuan). By 6.4 m.y. cooler conditions prevailed at Site 281. Between 6.3 and 6.2 ± 0.1 m.y. the carbon isotopic shift occurred followed, within 100,000 yr, by a distinct shallowing of water depths at Blind River. The earliest Pliocene (Opoitian) is marked by increasing surface-water temperatures.     

New fossil anthropoids from the middle Miocene of East Africa and their bearing on the origin of the oreopithecidae

Recent paleontological collections at the middle Miocene locality of Maboko Island in Kenya, dated at 15-16 million years, have yielded numerous new specimens belonging to at least five species of fossil anthropoids. The most common species of ape at the site, a medium-sized primate with a very distinctive dental morphology, clearly represents a previously undescribed taxon. When compared with other Miocene anthropoids from East Africa, it has its closest affinities with the poorly known species Rangwapithecus vancouveringi from the early Miocene locality of Rusinga Island. The species from Maboko Island is described here as belonging to a new genus of fossil anthropoid, to which "Rangwapithecus" vancouveringi is also referred. The new genus has a highly distinctive suite of derived characters of its molars and premolars, which it shares with Oreopithecus bambolii from the late Miocene of Europe. These synapomorphies indicate a close phyletic relationship between the East African species and Oreopithecus and form the basis for the inclusion of these taxa in a single family, the Oreopithecidae Schwalbe, 1915. In many respects, however, the East African forms are more conservative than Oreopithecus, and in a general sense they can be regarded as an intermediate grade between Oreopithecus and the more generalized early Miocene catarrhines, the proconsuloids. There is, therefore, good fossil evidence to indicate that the origins of the Oreopithecidae can be traced back to the early Miocene of Africa.     

An alpha 2-macroglobulinlike activity in the blood of chelicerate and mandibulate arthropods

The alpha 2-macroglobulins are large molecular weight proteinase-binding proteins that inhibit the ability of proteinases to hydrolyze protein substrates without suppressing activity against amide or ester substrates. They are also able to protect the active site of bound proteinases from active site inhibitors of suitably high molecular weight. The ability to protect the amidolytic activity of trypsin from the macromolecular active site inhibitor, soybean trypsin inhibitor, was used to demonstrate an alpha 2-macroglobulinlike activity in the blood of the horseshoe crab, Limulus polyphemus and the crustaceans Libinia emarginata (the spider crab) and Cancer borealis (the Jonah crab). The alpha 2-macroglobulinlike activities of L. polyphemus and L. emarginata are sensitive to methylamine, but that of C. borealis is relatively insensitive. The molecular weights (mw) of the trypsin-protecting proteins in L. emarginata and C. borealis, estimated from gelfiltration studies, are, respectively, 480 X 10(3) and 460 X 10(3), and are significantly smaller than that of L. polyphemus (Mr = 570 X 10(3)).     

Possible Ways of Spreading and Evolution of Alcids

Based on the published data, a hypothesis is proposed on possible ways of spreading of the Alcids from the center of their primary radiation located in the Pacific. The Alcids colonized the Atlantic first in the Early–Middle Miocene. They rounded North America from the south: at that time, the strait between the Americas was open, while the Bering Strain was closed. Later, in the Late Miocene, Late Pliocene, and Pleistocene, exchange of species between the Atlantic and Pacific took place through the periodically opening Bering Strait.