Correlations between baltic amber and Pinus resins

The composition of the acid fractions of four pine resins has been studied before and after an ‘ageing’ process. Correlations are established between Baltic amber and ‘aged’ Pinus halepensis resin.     

13C NMR and IR analyses of structure,aging and botanical origin of Dominican and Mexican ambers

¹³C NMR analysis of Oligocene amber from the Dominican Republic, Oligo-Miocene amber from Mexico and trunk resins from certain extant species of Hymenaea, suggests a polylabdane structure for all and gives insights into structural changes during aging. Additionally, IR spectra, some aspects of the ¹³C NMR spectra and paleobotanical data suggest a close relationship between Dominican amber and H. verrucosa. By contrast, spectral evidence shows that Mexican amber differs, not only from the former polymers, but from the H. courbaril polymer.     

Sciadopitys cladodes from Eocene Baltic amber

The Baltic amber deposit represents the largest accumulation of any fossil resin worldwide and hundreds of thousands of entrapped arthropods have been recovered so far. The source plants of Baltic amber, however, are still controversial, and the botanical composition of the ‘Baltic amber forest’ remains poorly studied. Here, we provide the first unequivocal Baltic amber inclusions of the umbrella pine Sciadopitys (Sciadopityaceae), a genus that has been suggested as the source of succinite (the main variety of Baltic amber) based on chemical analyses. As previously suggested sciadopitoid inclusions must be reconsidered as being notional, representing angiosperm leaves instead, the new fossils are the first unambiguous macrofossil evidence of Sciadopitys from the ‘Baltic amber forest’, and the first pre‐Oligocene macrofossil record of Sciadopitys from Europe. The fossil Sciadopitys cladodes provide new insights into the conifer diversity of the ‘Baltic amber forest’ and broaden the picture of its palaeoecology, indicating the presence of humid swamp to raised bog habitats. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 258–268.     

Chemical preservation of plants and insects in natural resins

The morphological preservation of fossils in amber is remarkable, but their chemical composition is largely unknown. The likelihood of DNA preservation in amber has been questioned but, surprisingly, the fate of more decay-resistant macromolecules such as ligno-cellulose in plants or the chitin–protein complex in insect cuticle has not been investigated. Here we report the results of investigations using pyrolysis-gas chromatography/mass spectrometry (py-GC/MS) of the tissues of insects and the plant Hymenaea from ancient and sub-fossil resins (2–20 ka) from Kenya, and from Dominican amber (25–30 Ma). The volatile components of the resin have penetrated even the internal tissues, resulting in the exceptional three-dimensional preservation of amber inclusions. Chitin is preserved in the bee and ligno-cellulose in the Hymenaea leaf from the Kenyan resins. There was no trace, however, of these macromolecules in tissues in Dominican amber. The presence of aliphatic polymer and sulphur-containing moieties in these tissues indicates that they have undergone diagenetic alteration; in view of this, the preservation in Dominican amber of a macromolecule as labile as DNA would be extraordinary.     

Fossil evidence of insect pathogens

The present report describes fossil evidence of insect pathogens, heretofore, almost non-existent, from six samples of amber ranging in age from 15 to 100 million years. They include a cytoplasmic polyhedrosis virus and trypanosomatid infection in an adult biting midge (Diptera: Ceratopogonidae), and a nuclear polyhedrosis virus in an adult sand fly (Diptera: Phlebotomidae), both from Early Cretaceous Burmese amber, several types of fungal thalli on the cuticle of an adult mosquito (Culicidae: Diptera), as well as a fungal growth on the prothorax of a fungus gnat (Mycetophilidae: Diptera) in Dominican amber and large tumors in the body cavity of a caterpillar (Lepidoptera) in Mexican amber. These discoveries suggest that insect polyhedrosis viruses were present 100 million years ago and present the possibility that vertebrate arboviruses (especially those in the family Reoviridae) could have evolved from cytoplasmic polyhedrosis viruses infecting biting insects. The flagellates in the Early Cretaceous biting midge represent the first fossil record of monogenetic trypanosomatid infections of arthropods.     

A new proposal concerning the botanical origin of Baltic amber

Baltic amber constitutes the largest known deposit of fossil plant resin and the richest repository of fossil insects of any age. Despite a remarkable legacy of archaeological, geochemical and palaeobiological investigation, the botanical origin of this exceptional resource remains controversial. Here, we use taxonomically explicit applications of solid-state Fourier-transform infrared (FTIR) microspectroscopy, coupled with multivariate clustering and palaeobotanical observations, to propose that conifers of the family Sciadopityaceae, closely allied to the sole extant representative, Sciadopitys verticillata, were involved in the genesis of Baltic amber. The fidelity of FTIR-based chemotaxonomic inferences is upheld by modern–fossil comparisons of resins from additional conifer families and genera (Cupressaceae: Metasequoia; Pinaceae: Pinus and Pseudolarix). Our conclusions challenge hypotheses advocating members of either of the families Araucariaceae or Pinaceae as the primary amber-producing trees and correlate favourably with the progressive demise of subtropical forest biomes from northern Europe as palaeotemperatures cooled following the Eocene climate optimum.     

The role of acid catalysis in the genesis of amber

Analysis of the hydrocarbon fraction from baltic amber is described. Transformations which have occurred in resins during the formation of amber are discussed on the grounds of acid-catalysed reactions undergone by 7,13-abietadiene and sclareol.     

Burmese amber: evidence of Gondwanan origin and Cretaceous dispersion

Abstract

Burmese amber is an extremely important source of mid-Cretaceous plant and animal remains with over 870 species of organisms, ranging from protozoa to vertebrates, described from this source. The amber mines are located on the West Burma Block that according to geologists was originally part of Gondwana. The present study introduces some angiosperms and insects in Burmese amber whose closest extant relatives have a Gondwanan distribution and there is no previous evidence of a Laurasian distribution. Based on this evidence, it is proposed that organisms in Burmese amber represent a selection of tropical to subtropical life forms that inhabited the interconnected continents of Gondwana in the Early Cretaceous. Based on the fossil record of angiosperms and their diversity in Burmese amber, the West Burma Block could not have rafted from Gondwana to SE Asia before the Early Cretaceous.     

Protist-like inclusions in amber, as evidenced by Charentes amber

The mid-Cretaceous amber of France contains thousands of protist-like inclusions similar in shape to some ciliates, flagellates and amoebae. The sheer abundance of these inclusions and their size variation within a single amber piece are not concordant with true fossil protists. French amber is coniferous in origin, which generally does not preserve well protists without cell walls. Thus, it would be surprising if French Cretaceous amber had preserved millions of protists. Here, we present a survey of the protist-like inclusions from French amber and attempt to elucidate their origins.Diverse Cretaceous ambers (from Spain, Germany and Lebanon), also derived from conifer resins, contain thousands of protist-like inclusions. In contrast, Tertiary ambers and modern resins are poor in protist-like fossils. This suggests these inclusions originated from early Cretaceous plant resins, probably secreted with the resin by trees that did not survive after the Cretaceous (such as the Cheirolepidiaceae). A review of the recent literature on amber microfossils indicates several protist-like inclusions that are unlikely to have a biological origin have already been described as real fossil protists. This is problematic in that it will bias our understanding of protist evolution.     

Palaeoecological perspectives in Dominican amber

Palaeoecological and palaeobiogeographical aspects of Dominican amber are discussed based on the known insect fauna. Topics covered are examples of speciation, extinctions, longevity as well as associations between insects and plants, insects and vertebrates and various arthropod groups. Examples of camouflage, oviposition and predator-prey associations are presented. Ecological and medical implications from recently discovered vector-parasite associations (malaria, trypanosomiasis and leishmaniasis) are discussed. Paleosymbiotic associations, with examples of phoresis, mutualism, parasitism and pathogens, are examined. Insects in Dominican amber, together with their cohabitants, can assist in determining specific habitats and reconstructing ancient landscapes.     

Diverse fossil amoebae in German Mesozoic amber

Fossil amoebae are very rare, although their evolutionary history extends back into the Proterozoic. The Cenomanian amber of Schliersee (southern Germany) is very rich in micro-organisms and contains the highest diversity of fossil freshwater rhizopods (Gymnamoebia and Testacealobosia) yet discovered. Altogether seven testate amoebae and one gymnamoebian species are recorded from this Mesozoic amber. The four newly discovered taxa described in this paper can be assigned to the extant species Centropyxis delicatula , Centropyxis hirsuta , Phryganella acropodia and Phryganella paradoxa . Over 200 individuals of Phryganella paradoxa are preserved. Together with their syninclusions, the amoebae are species of limnetic or limnetic–terrestrial microcoenoses. The presence of 100-myr-old fossils with extant representatives suggests evolutionary stasis of these freshwater amoebae. However, not all modern testacean families have been recorded from Mesozoic limnetic habitats. Our experimental studies verify that naked and testate amoebae can be embedded in resins.     

The isotopic footprint of irrigation in the western Mediterranean basin during the Bronze Age: the settlement of Terlinques,southeast Iberian Peninsula

The isotopic composition of the remains of cereals and pine has been studied, from three different chronological phases from 2140 to 1500 cal bc at the Terlinques site, southeast Iberian Peninsula. The δ¹³C values range between ?24.91 and ?21.19 ‰ (V-PDB), with an average of ?23.05 ‰ (STD = 0.69). The archaeological cereals show an average isotopic discrimination (Δ¹³C) with the past atmospheric CO₂ of 16.96 ‰, which is much greater than the average Δ¹³C of 13.89 ‰ of the rainfed Triticeae (wheat and barley) in modern times. However, considering the effect of the atmospheric CO₂ concentration, which is included in the WUEi (intrinsic water-use efficiency), this difference is even greater, 77 for archaeological samples versus 144 for present-day rainfed cereals. This could represent some of the earliest evidence of the use of irrigation techniques in Europe. Modern cereals which have been irrigated show a general Δ¹³C average of 17.17 ‰, very similar to those of the middle Holocene. However, when the WUEi is calculated, the value of 108 indicates that present-day irrigated cereals are more stressed than the archaeological samples. For comparison, we have included pine trees, since these have an extensive root development which is capable of reaching the water table. In the past, both cereals and pine present similar WUEi values (77 vs. 72), however at present only irrigated cereals show similar WUEi values to pine (108 vs. 107). This again suggests irrigation of cereals in the past. The processes of climatic degradation towards drier conditions which started in the middle Holocene could be responsible for the use of land near water sources, on riverbanks and near shallow lakes. According to the isotopic and plant macrofossil data, irrigation or water management techniques were used at the Terlinques site, located close to the Laguna de Villena, a lake which has now dried out.     

A new genus of sphaeropsocid bark lice from the Early Cretaceous amber of Lebanon (Psocodea: Sphaeropsocidae)

A new genus and species of sphaeropsocid bark louse is described and figured from a single individual in Early Cretaceous amber from Hammana, central Lebanon. Asphaeropsocites neli gen. n., sp. n. is the second sphaeropsocid described from Lebanese amber. Like Sphaeropsocites lebanensis Grimaldi & Engel 2006, it has a basal phylogenetic position within Sphaeropsocidae, and adds evidence that these insects were once widespread and global, in the past. The new species is distinguished from related taxa, and a discussion and checklist of sphaeropsocids are provided.     

Arthropods in contemporary and some fossil resins

More than 4800 arthropod inclusions were isolated and identified from resin of various contemporary conifer trees in various parts of northern Eurasia. Their composition is compared with that in representative collections of Baltic and Rovno ambers (Upper Eocene) and with that in Dominican amber (Lower Miocene). The original composition of inclusions of Dominican amber is reconstructed for the first time using a procedure intended to reduce the effect of human bias. Taphonomical characteristics of resins and their effects on the composition of inclusions are studied. The actuapaleontological approach reveals a trend towards a decrease in the relative abundance of arboreal springtails and nematoceran dipterans and an increase in that of the true bugs, beetles, lepidopterans, and hymenopterans (especially ants) between the Eocene and the present. Relative abundances of spiders and mites show no clear trend. The available data on other arthropods are still insufficient for elucidating evolutionary trends. Surprisingly, a small contemporary sample from Taimyr (N. Siberia) was inexplicably more similar to the Eocene amber than to other contemporary resins. No other significant differences in composition of inclusions, compared across different conifer genera or geographic areas, have been revealed. A more detailed comparison between contemporary and fossil hymenopteran and beetle inclusions reveals correlations with both age (= evolutionary change) and geography. The absolute dominance of ants, particularly Formicinae and Myrmicinae, and, among solitary hymenopterans, Ichneumonidae, Braconidae, and Pteromalidae, and a corresponding decline in the abundance of Scelionidae and Dolichoderinae in contemporary resins compared to amber reflect evolutionary changes. In contrast, the overwhelming abundance of Formicinae and consistent occurrence of sawflies in contemporary resins of northern Eurasia appear to be explained by geography. The Eocene assemblages of beetle inclusions are characterized by a wider and more variable set of dominant families, in sharp contrast to contemporary resins, which are uniformly dominated by Curculionidae, Chrysomelidae, and Staphylinidae. Additional analyses are needed to explain this difference.     

Using fossils to determine an amber source: aphids and crane flies in Chinese or Baltic amber?

Aphids and crane flies in amber purported to have originated in China were examined. Six aphid specimens were characterised and determined to be conspecific with Germaraphis defuncta Heie, originally described from Baltic amber. Two crane flies from the same collection were identified as Cheilotrichia minuta and Rhabdomastix brevis, species also previously described from Baltic amber. This study shows that when the locality of an amber source is ambiguous, fossils can be used to determine the true province.     

Palaeobiology of tanaidaceans (Crustacea: Peracarida) from Cretaceous ambers: extending the scarce fossil record of a diverse peracarid group

Diverse assemblages of tanaidacean peracarid crustaceans from western Tethyan continental deposits suggest that the group was relatively common in or around ancient resin‐producing forests. Here we report the results of an examination of 13 tanaidacean specimens from three Cretaceous (Albian to Turonian) French amber deposits. Two new species of the fossil family Alavatanaidae are placed in the previously described Early Cretaceous genus Eurotanais: Eurotanais pyrenaensis sp. nov. from Cenomanian Pyrenean amber (Fourtou, Aude) and Eurotanais seilacheri sp. nov. from Turonian Vendean amber (La Garnache, Vendée). The remaining specimens are placed in three newly erected genera and species (but family incertae sedis): Arcantitanais turpis gen. et sp. nov. from Albian–Cenomanian Charentese amber (Archingeay, Charente‐Maritime), and Tytthotanais tenvis gen. et sp. nov. and Armadillopsis rara gen. et sp. nov. from Pyrenean amber. These are the first formally described fossils that might be related to the paratanaoidean families Nototanaidae and Paratanaidae, sharing with these some putatively derived features and providing possible evidence for the antiquity and morphological stability of these families and the suborder Tanaidomorpha. The distinctive features and character combinations of these fossil taxa are discussed in connection with possible relationships to the living lineages of tanaidaceans. Propagation phase‐contrast X‐ray synchrotron microtomography was used to obtain high‐quality 3D images for some fossils. A discussion is provided on the putative palaeobiology of tanaidaceans and the French resiniferous forest ecosystem. The discovery of these new tanaidaceans extends the palaeogeographical distribution and stratigraphical range of the family Alavatanaidae and sheds new light on the palaeoecology and diversity of tanaidaceans in pre‐angiospermous woodlands.     

How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)

Six new fossils of Micromalthus (Coleoptera: Archostemata) from Dominican amber are compared with extant and previously described fossil micromalthid beetles. The amber inclusions are well preserved and all important morphological features are visible. Taking into account the morphological variability of the extant species Micromalthus debilis LeConte, 1878 , it is not possible to find any morphological features that distinguish the fossils from the extant species. This also applies to the Dominican amber inclusion described as Micromalthus anasi Perkovsky, 2008, which therefore is considered a junior synonym of M. debilis. The lack of morphological change in M. debilis over time might possibly be explained by unusually stable environmental conditions, as this species occupies a very specialized ecological niche in decaying timber. A general survey of fossil insects indistinguishable from extant species is presented. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 300–311.     

Anti-bacterial function in the sexually dimorphic pollinator rewards of Clusia grandiflora (Clusiaceae)

Many species of the dioecious, neo-tropical plant genus Clusia secrete a viscous, hydrophobic resin from glandular tissues in both male and female flowers. This substance is readily gathered by meliponine and euglossine bees for whom it most often serves as the sole pollinator reward. Bees use Clusia resin as a nest-building material. As such, resin clearly serves an indispensable mechanical function. However, resins with antimicrobial properties may also serve to reduce the risk of pathogenesis in the nest. If resin-gathering apids benefit from antimicrobial properties in nesting materials and are able to discern these characteristics in the forage they gather, one might predict that the resin reward presented in Clusia could have evolved under selection for both mechanical and antimicrobial properties. In dioecious species, where females and males each present a resin reward, selection regimes may differ between the sexes with the result that resin form and function diverge. We investigated both the form and function of the male and female pollinator reward resins of Clusia grandiflora. Using thin-layer chromatography (TLC), we compared the chemical compositions of floral resins from five widely separated populations of this species growing in southeastern Venezuela. We found that male and female resins exhibited a marked chemical dimorphism, with females having two major TLC-resolvable fractions and males having seven. This dimorphism was stable: there were no component differences between populations in either sex. Using a disk-diffusion technique, we surveyed the same resins for antimicrobial activity using assay microorganisms isolated from eusocial meliponine bees. Both male and female Clusia grandiflora resins had pronounced but relatively directed antimicrobial activity: both were toxic to 10 of 11 Gram-positive bacteria, 7 of 15 Gram-negative or variably-staining bacteria, 0 of 3 yeasts, and 0 of 3 filamentous fungi. Again with the disk-diffusion technique, we performed more detailed tests of resin bioactivity using two Gram-positive honeybee associates, Paenibacillus larvae and P. alvei, as model pathogens. Both male and female C. grandiflora resins were highly toxic to these honeybee pathogens. Female resin, however, produced zones of inhibition with more than twice the mean diameter of those produced by the male resin. This divergence in form and function of the C. grandiflora pollinator reward resins could be in response to different selective regimes as mediated by the pollinating insects. Received: 19 October 1998 / Accepted: 9 February 1999     

When phylogeny meets geology and chemistry: doubts on the dating of Ethiopian amber

The first fossil species of a Melissotarsus ant is described (Melissotarsus ethiopiensis n. sp.), suggesting that Ethiopian amber could be cenozoic rather than cretaceous. This hypothesis is supported by a Fourier transform infrared spectroscopy (FTIR) analysis of the amber itself, suggesting a weak maturation of the fossil resin, and a revision of the pyrolysis–gas chromatography–mass spectrometry (PY-GC-MS) analysis done by Schmidt and his collaborators in 2010, in an article in which the deposit was described for the first time.