Jorge A. Santiago-Blay

A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies

Long-Lost Pollinators The rise of angiosperms in the Early Cretaceous (∼140 million years ago) was accompanied by coevolution of a variety of insects, including flies, bees, and wasps required for pollination. Ren et al. (p. 840; see the Perspective by Ollerton and Coulthard) show that three families of scorpionflies had already evolved specialized mouth parts for feeding on the nectar of gymnosperms, as early as the Middle Jurassic (∼170 million years ago). The diversity and specialization of these insects and related plant structures suggests that they were also involved in pollination. These families died out later in the Cretaceous as angiosperms began to dominate. Prior to the coevolution of angiosperms and pollinating insects, scorpionflies may have been pollinating gymnosperms. The head and mouthpart structures of 11 species of Eurasian scorpionflies represent three extinct and closely related families during a 62-million-year interval from the late Middle Jurassic to the late Early Cretaceous. These taxa had elongate, siphonate (tubular) proboscides and fed on ovular secretions of extinct gymnosperms. Five potential ovulate host-plant taxa co-occur with these insects: a seed fern, conifer, ginkgoopsid, pentoxylalean, and gnetalean. The presence of scorpionfly taxa suggests that siphonate proboscides fed on gymnosperm pollination drops and likely engaged in pollination mutualisms with gymnosperms during the mid-Mesozoic, long before the similar and independent coevolution of nectar-feeding flies, moths, and beetles on angiosperms. All three scorpionfly families became extinct during the later Early Cretaceous, coincident with global gymnosperm-to-angiosperm turnover.

Chemical signatures of fossilized resins and recent plant exudates.

Amber is one of the few gemstones based on an organic structure. Found over most of the world, it is the fossil form of sticky plant exudates called resins. Investigation of amber by modern analytical techniques provides structural information and insight into the identity of the ancient plants that produced the source resin. Mass spectrometric analysis of materials separated by gas chromatography has identified specific compounds that are the basis of a reliable classification of the different types of amber. NMR spectroscopy of bulk, solid amber provides a complementary classification. NMR spectroscopy also can be used to characterize modern resins as well as other types of plant exudates such as gums, gum resins, and kinos, which strongly resemble resins in appearance but have very different molecular constitutions.

Paleontology of leaf beetles

The fossil record of the Chrysomelidae can be tentatively traced back to the late Paleozoic to early Mesozoic (Triassic). Mesozoic records (at least 9 subfamilies, 19 genera, and 35 species), are represented by the Sagrinae, the exclusively Mesozoic Protoscelinae, Clytrinae, Cryptocephalinae, Eumolpinae, Chrysomelinae, Galerucinae, Alticinae, and Cassidinae. Cenozoic records (at least 12 subfamilies — 63% of the extant — 121 genera, and 325 species), include the same extant subfamilies as well as the Donaciinae, Zeugophorinae, Criocerinae, and Hispinae and can be frequently identified to genus, especially if preserved in amber. Quaternary records are often identified to extant species. In total, at least 131 genera (about 4% of total extant), and 357 species (< 1%) have been reported. At least, 24 genera (< 1% of the extant) seem to be extinct.

New fossil Lepidoptera (Insecta: Amphiesmenoptera) from the Middle Jurassic Jiulongshan Formation of Northeastern China.

Background The early history of the Lepidoptera is poorly known, a feature attributable to an inadequate preservational potential and an exceptionally low occurrence of moth fossils in relevant mid-Mesozoic deposits. In this study, we examine a particularly rich assemblage of morphologically basal moths that contribute significantly toward the understanding of early lepidopteran biodiversity. Methodology/Principal Findings Our documentation of early fossil moths involved light- and scanning electron microscopic examination of specimens, supported by various illumination and specimen contrast techniques. A total of 20 moths were collected from the late Middle Jurassic Jiulongshan Formation in Northeastern China. Our principal results were the recognition and description of seven new genera and seven new species assigned to the Eolepidopterigidae; one new genus with four new species assigned to the Mesokristenseniidae; three new genera with three new species assigned to the Ascololepidopterigidae fam. nov.; and one specimen unassigned to family. Lepidopteran assignment of these taxa is supported by apomorphies of extant lineages, including the M1 vein, after separation from the M2 vein, subtending an angle greater than 60 degrees that is sharply angulate at the junction with the r–m crossvein (variable in Trichoptera); presence of a foretibial epiphysis; the forewing M vein often bearing three branches; and the presence of piliform scales along wing veins. Conclusions/Significance The diversity of these late Middle Jurassic lepidopterans supports a conclusion that the Lepidoptera–Trichoptera divergence occurred by the Early Jurassic.

The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies

Mid-Mesozoic kalligrammatid lacewings (Neuroptera) entered the fossil record 165 million years ago (Ma) and disappeared 45 Ma later. Extant papilionoid butterflies (Lepidoptera) probably originated 80–70 Ma, long after kalligrammatids became extinct. Although poor preservation of kalligrammatid fossils previously prevented their detailed morphological and ecological characterization, we examine new, well-preserved, kalligrammatid fossils from Middle Jurassic and Early Cretaceous sites in northeastern China to unravel a surprising array of similar morphological and ecological features in these two, unrelated clades. We used polarized light and epifluorescence photography, SEM imaging, energy dispersive spectrometry and time-of-flight secondary ion mass spectrometry to examine kalligrammatid fossils and their environment. We mapped the evolution of specific traits onto a kalligrammatid phylogeny and discovered that these extinct lacewings convergently evolved wing eyespots that possibly contained melanin, and wing scales, elongate tubular proboscides, similar feeding styles, and seed–plant associations, similar to butterflies. Long-proboscid kalligrammatid lacewings lived in ecosystems with gymnosperm–insect relationships and likely accessed bennettitalean pollination drops and pollen. This system later was replaced by mid-Cretaceous angiosperms and their insect pollinators.

Characterization of Plant Exudates by Principal-Component and Cluster Analyses with Nuclear Magnetic Resonance Variables

Principal-component and cluster analyses have been applied to nuclear magnetic resonance data for exudates derived from both conifers and angiosperms in order to classify these materials on the basis of molecular structure. The method succeeds in distinguishing resins produced by the conifer families Araucariaceae, Cupressaceae, and Pinaceae from each other and from resins produced by the angiosperm family Fabaceae. Other exudate types, including gums, gum resins, and kinos, also are distinguished from each other and from the resins.

Examination of amber and related materials by NMR spectroscopy

Examination of the solid‐state 13C and solution 1H NMR spectra of fossilized resins (ambers) has generated five groupings of materials based on spectral characteristics. The worldwide Group A is associated with the botanical family of the Araucariaceae. The worldwide Group B is associated with the Dipterocarpaceae. Baltic amber or succinite (Group C) is related to Group A but with a disputed conifer source. Amber from Latin America, the Caribbean, and Africa is associated with the Fabaceae, the genus Hymenaea in particular. The minor Group E contains the rare fossil polystyrene. The spectra of jet indicate that it is a coal‐like material with a rank between lignite and sub‐bituminous coal. Copyright

Characterization of Eucalyptus and Chemically Related Exudates by Nuclear Magnetic Resonance Spectroscopy

Exudates from six species of the genus Eucalyptus and one of the genus Corymbia (formerly Eucalyptus), from the family Myrtaceae, have been characterized by solid-state 13C and solution 1H NMR spectroscopy for the first time. Although these eucalypt kinos, as these exudates often are called, resemble resin (terpenoid) and gum (carbohydrate) exudates in physical appearance, their NMR spectra are dramatically different. In addition to lacking the characteristic terpene saturated resonances, they exhibit strong unsaturated resonances, which are weak for resins and absent for gums. We additionally report that exudates from genera of several other families of flowering plants (Amyris, Centrolobium, Guaiacum, Liquidambar, and Prosopis) also exhibit part or all of this kino spectroscopic signature.

Life habits, hox genes, and affinities of a 311 million-year-old holometabolan larva

BackgroundHolometabolous insects are the most diverse, speciose and ubiquitous group of multicellular organisms in terrestrial and freshwater ecosystems. The enormous evolutionary and ecological success of Holometabola has been attributed to their unique postembryonic life phases in which nonreproductive and wingless larvae differ significantly in morphology and life habits from their reproductive and mostly winged adults, separated by a resting stage, the pupa. Little is known of the evolutionary developmental mechanisms that produced the holometabolous larval condition and their Paleozoic origin based on fossils and phylogeny.ResultsWe provide a detailed anatomic description of a 311 million-year-old specimen, the oldest known holometabolous larva, from the Mazon Creek deposits of Illinois, U.S.A. The head is ovoidal, downwardly oriented, broadly attached to the anterior thorax, and bears possible simple eyes and antennae with insertions encircled by molting sutures; other sutures are present but often indistinct. Mouthparts are generalized, consisting of five recognizable segments: a clypeo-labral complex, mandibles, possible hypopharynx, a maxilla bearing indistinct palp-like appendages, and labium. Distinctive mandibles are robust, triangular, and dicondylic. The thorax is delineated into three, nonoverlapping regions of distinctive surface texture, each with legs of seven elements, the terminal-most bearing paired claws. The abdomen has ten segments deployed in register with overlapping tergites; the penultimate segment bears a paired, cercus-like structure. The anterior eight segments bear clawless leglets more diminutive than the thoracic legs in length and cross-sectional diameter, and inserted more ventrolaterally than ventrally on the abdominal sidewall.ConclusionsSrokalarva berthei occurred in an evolutionary developmental context likely responsible for the early macroevolutionary success of holometabolous insects. Srokalarva berthei bore head and prothoracic structures, leglet series on successive abdominal segments – in addition to comparable features on a second taxon eight million-years-younger – that indicates Hox-gene regulation of segmental and appendage patterning among earliest Holometabola. Srokalarva berthei body features suggest a caterpillar-like body plan and head structures indicating herbivory consistent with known, contemporaneous insect feeding damage on seed plants. Taxonomic resolution places Srokalarva berthei as an extinct lineage, apparently possessing features closer to neuropteroid than other holometabolous lineages.

A fossil scorpion Tityus geratus new species (Scorpiones: Buthidae) from Dominican Amber

A fossil scorpion, Tityus geratus (Scorpioncs: Buthidae) n. sp., is described from Dominican amber. This is the third described amber Tityus sp. and the third Dominican amber scorpion. On the basis of the amber finds and available knowledge on the present distribution of Tityus spp. in the Antilles, the vicariance model of Caribbean biogeography can be evoked to best explain the present distribution of this genus.