On the other hand, the preservation of internal structures i. It has to be noted that the presence of a white aureole surrounding the body of inclusions, considered to be a foam of microscopic bubbles by Mierzejewski  and Weitschat and Wichard  can also help in some cases to reveal which insects where trapped alive or dead in the resin  , as it is possibly the result of an early diagenesis reaction between fluids, produced by decay and decomposition of labile tissues, with sugar and terpenes in the resin.
However, such foams are rare in Mexican and Dominican ambers. In the present case, no foam is visible around our specimens to help us in our analysis. The two homogenous density groups are not distinguishable under optic microscope. Some preserved internal organs are visible, but the level of preservation of the cuticles is difficult to evaluate. One particularly odd case is the Nasutitermes with the damaged gaster Na3 as this gaster is obviously empty of any kind of internal structures, while CT-scan images show that its whole body is denser than the amber.
The fact that the damaged part of the gaster exhibits a bite mark caused by an ant, and is entirely covered by what seems to be the edge of an air bubble Figure 2B strongly suggests that this specimen was dead before entombment in fresh resin. In fact, regarding what we mentioned above, if the two closely contiguous Nasutitermes Na3 and Na4 are likely to have been both dead before entombment in the resin, the similarity of their density levels may not have the same taphonomical origin, and remain to be elucidated for specimen Na3 Nasutitermes.
Biostratinomic processes the period between the moment when resin is exuded from the tree to the moment when it is buried in sediment could also be responsible for these differences in matter density, as great differences can occur between flows in term of duration of the flowing event viscosity , degree of humidity of the air, level and time of exposure to air and to UV, etc.
Such variations can therefore create disparities in the taphonomic process between inclusions present in different flows. If we therefore cross-compare the density distribution between specimens with the different flows in which they were embedded, our amber piece shows that the two density groups almost fit with the distribution of the specimens inside the flows, except for the closely contiguous Nasutitermes soldier and the Nasutitermes worker with a damaged gaster, as they share the same density level as the two Azteca ants that belong to a different flow.
TREATISE ON A NALYSlS Volume This is Volume in PURE AND APPLIED MATHEMATICS A series of Monographs and Tex. Treatise on Analysis: TREATISE ON A NALYSlS Volume This is Volume in PURE AND APPLIED MATHEMATICS A series of Monographs and.
Regarding what has been mentioned above, this result can be due to differences between the insects themselves before entombment in the resin. Our study provides evidence that some degree of relationship between Azteca ants and Nasutitermes termites might have already existed in Central America during the late Oligocene-middle Miocene period, together with the predation of army ants on other eusocial insects in the same community.
However, the condition that led to the apparition of such interactions and their stability through time are still to be elucidated. The main question to further address is to know in which measure both flow structures of the amber pieces and physical density variations of insect bodies can help to further reconstruct necrolysis, biostratinomic and diagenetic processes that occurred in the amber and its inclusions.
Movie of the syninclusion, showing the different flows of the resin that comprise the amber piece.
We thank the two anonymous referees for their useful comments on the first version of the paper. DC thanks his friends, Manuel Ramirez and his son Heriberto, owners and miners of Salt River Mine, and their family, for their great help in his field research. Browse Subject Areas?
Click through the PLOS taxonomy to find articles in your field. Abstract We describe here a co-occurrence i. Introduction Ants and termites represent ecologically critical organisms in intertropical and subtropical ecosystems, impacting by their abundance, organization and variety of occupied niches the availability of nutrients as well as the composition of soils  — . Download: PPT. Material, Locality and Method This amber piece was discovered in a batch of crude amber acquired by one of us DC from locals exploiting the Totolapa amber deposit Salt River Mine.
No permits were required for the described study. Systematic palaeontology Identifications of the specimens were possible at the generic level but not at the specific level, for the reasons indicated below.
General description of the amber piece Our amber piece is 1. CT-scan results X-ray tomographic analysis revealed that our amber piece is in fact made of eight distinct layers corresponding to different flows, and that the distribution of the insects does not reflect a synchronous event.
Figure 3. Virtual slicing from CT scan analysis showing flow boundaries. Discussion Ants-termites interactions — palaeoecological interpretation of the syninclusion In the Neotropics, Nasutitermes is often involved in relationships with ants, possibly in relation with the fact that it is the termite genus with the highest number of species building conspicuous nests  , . Interpretation of CT-scan results Structure of the amber. Differences of density between specimens. Conclusion Our study provides evidence that some degree of relationship between Azteca ants and Nasutitermes termites might have already existed in Central America during the late Oligocene-middle Miocene period, together with the predation of army ants on other eusocial insects in the same community.
Supporting Information. Figure S1. PDF 3D of the syninclusion. Figure S2. Acknowledgments We thank the two anonymous referees for their useful comments on the first version of the paper. References 1. Belknap Harvard University Press, Springer publ. Kaspari M Taxonomic level, trophic biology, and the regulation of local abundance.
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