Problematic fossils, extinct taxa of enigmatic morphology that cannot be assigned to a known major group, were once a major issue in palaeontology. A long-favoured solution to the 'problem of the problematica', particularly the 'weird wonders' of the Cambrian Burgess Shale, was to consider them representatives of extinct phyla. A combination of new evidence and modern approaches to phylogenetic analysis has now resolved the affinities of most of these forms. Perhaps the most notable exception is Tullimonstrum gregarium, popularly known as the Tully monster, a large soft-bodied organism from the late Carboniferous Mazon Creek biota (approximately 309-307 million years ago) of Illinois, USA, which was designated the official state fossil of Illinois in 1989. Its phylogenetic position has remained uncertain and it has been compared with nemerteans, polychaetes, gastropods, conodonts, and the stem arthropod Opabinia. Here we review the morphology of Tullimonstrum based on an analysis of more than 1,200 specimens. We find that the anterior proboscis ends in a buccal apparatus containing teeth, the eyes project laterally on a long rigid bar, and the elongate segmented body bears a caudal fin with dorsal and ventral lobes. We describe new evidence for a notochord, cartilaginous arcualia, gill pouches, articulations within the proboscis, and multiple tooth rows adjacent to the mouth. This combination of characters, supported by phylogenetic analysis, identifies Tullimonstrum as a vertebrate, and places it on the stem lineage to lampreys (Petromyzontida). In addition to increasing the known morphological disparity of extinct lampreys, a chordate affinity for T. gregarium resolves the nature of a soft-bodied fossil which has been debated for more than 50 years.
Within modern gymnosperms, conifers and Ginkgo are exclusively wind pollinated whereas many gnetaleans and cycads are insect pollinated. For cycads, thrips are specialized pollinators. We report such a specialized pollination mode from Early Cretaceous amber of Spain, wherein four female thrips representing a genus and two species in the family Melanthripidae were covered by abundant Cycadopites pollen grains. These females bear unique ring setae interpreted as specialized structures for pollen grain collection, functionally equivalent to the hook-tipped sensilla and plumose setae on the bodies of bees. The most parsimonious explanation for this structure is parental food provisioning for larvae, indicating subsociality. This association provides direct evidence of specialized collection and transportation of pollen grains and likely gymnosperm pollination by 110-105 million years ago, possibly considerably earlier.paleoecology | paleoethology | coevolution | plant-insect interactions
El Soplao outcrop, an Early Cretaceous amber deposit recently discovered in northern Spain (Cantabria), has been shown to be the largest site of amber with arthropod inclusions that has been found in Spain so far. Relevant data provided herein for biogeochemistry of the amber, palynology, taphonomy and arthropod bioinclusions complement those previously published. This set of data suggests at least two botanical sources for the amber of El Soplao deposit. The ñrst (type A amber) strongly supports a source related to Cheirolepidiaceae, and the second (type B amber) shows non‐specific conifer biomarkers. Comparison of molecular composition of type A amber with Frenelopsis leaves (Cheirolepidiaceae) strongly suggests a biochemical affinity and a common botanical origin. A preliminary palynologlcal study indicates a regional high taxonomical diversity, mainly of pteridophyte spores and gymnosperm pollen grains. According to the preliminary palynologlcal data, the region was inhabited by conifer forests adapted to a dry season under a subtropical climate. The abundant charcoalified wood associated with the amber in the same beds is evidence of paleofires that most likely promoted both the resin production and an intensive erosion of the litter, and subsequent great accumulation of amber plus plant cuticles. In addition, for the first time in the fossil record, charcoalified plant fibers as bioinclusions in amber are reported. Other relevant taphonomic data are the exceptional presence of serpulids and bryozoans on the surfaces of some amber pieces indicating both a long exposure on marine or brackish‐water and a mixed assemblage of amber. Lastly, new findings of insect bioinclusions, some of them uncommon in the fossil record or showing remarkable adaptations, are reported. In conclusion, a documented scenario for the origin of the El Soplao amber outcrop is provided.
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