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A larval brush-footed butterfly (Lepidoptera: Nymphalidae) in Dominican amber, with a summary of fossil Nymphalidae
Abstract
The first fossil member of the family Nymphalidae (Lepidoptera) in Dominican amber and the first fossil nymphalid caterpillar is placed near the genus Smyrna of the tribe Coloburini (Nymphalinae) on the basis of the morphology, position and arrangement of its spines. The larva is regarded as prototypic ancestor near the Pycina-Vanessa lineage and its closest descendants probably occur in Central and South America, thus illustrating yet another case of disjunct distribution in the Dominican amber fauna. This represents the second family of butterflies reported from Dominican amber (previously only the Riodinidae). A summary of the confirmed cases of fossil Nymphalidae is presented.
... 06. Specimen from Hammond & Poinar (1998), Nymphalidae, image flipped. 07. ...
... No scale is available for the specimen. Hammond & Poinar (1998) reported a heavily armoured caterpillar from Dominican amber interpreted as a larva of the group Nymphalidae (specimen 06; Fig. 1). The authors provided micrographs in dorsal view (p. ...
... 58 fig. 53B) of specimen 06, i.e. the heavier armoured one reported by Hammond & Poinar (1998). Also, the fourth one is an already known specimen, specimen 01, i.e. the one reported by Poinar (1993). ...
... age that is considered to belong to the Urticaceae-feeding tribe Nymphalini (Fig. 13). Today caterpillars of Vanessa Fabricius and the related Hypanartia Hübner are the only butterflies that feed on Urticaceae in Hispaniola (Hammond and Poinar 1998). ...
... The fossil shown in Fig. 12, although not so well-preserved, is similar enough to be worthy of note. Caterpillars of Vanessa Fabricius and the related Hypanartia Hübner are the only butterflies that feed on Urticaceae in Hispaniola today (Hammond and Poinar, 1998). Scale bar = 2.3 mm. ...
The first fossil flowers of Neotropical Urticaceae (Boehmerieae) are described from the Dominican Republic and Mexico as belonging to a new genus, Ekrixanthera. Ekrixanthera hispaniolae sp. nov. from Dominican amber has pentamerous staminate flowers on short pedicels with a pilose pistillode and heteromorphic pilose tepals: two are clavate and three linear. Ekrixanthera ehecatli sp. nov. has pentamerous staminate flowers lacking pedicels, a pistillode with greatly reduced pilosity, glabrous and heteromorphic tepals with two linear and three wedge-shaped with truncate tips. The presence or absence of a pedicel, heterotrophic condition of the tepals, and the presence or absence of pilosity of the pistillode and tepals separate the two species. Those characters, together with the pentamerous flowers separate both fossil species from extant genera. The floral structures indicate explosive pollen release and pollination by wind (anemophily). Pistillate flowers have not been found for this usually dioecious tribe. Lepidopteran herbivory is suggested by a damaged stipule in one specimen and a nymphalid butterfly (Vanessa-like) caterpillar that may have used Ekrixanthera as a food plant is illustrated. The fossils establish an early lineage of Boehmerieae with characteristic explosive pollen release and perhaps associated herbivorous insects in the West Indies and North America during the mid-Tertiary.
... For Miocene Dominican amber, the list of fossilized Lepidoptera includes other families: Blastobasidae, Cosmopterygidae, Gelechiidae, Noctuidae, Tineidae and Tortricidae (Poinar[1992]), Tortricidae (Poinar and Brown[1993]), and Oecophoridae (Kristensen and Skalski[1999]).Grimaldi and Engel (2005)figured a caterpillar and two adults of the family Geometridae, an adult of the genus Acrolophus (Acrolophidae), and other adults of Tortricidae and Gelechioidea; an interesting tineoid moth figured by these authors is a case with the caterpillar inside. The scarce butterflies found in Dominican amber have been studied in detail, and are represented as caterpillars of the families Nymphalidae and Riodinidae (DeVries and Poinar, 1997;Hammond and Poinar, 1998) and adults of Riodinidae (Poinar, 1992;Grimaldi, 1996;DeVries, 1997;Hall et al., 2004). Here, we report two additional adult butterfly specimens of the families Riodinidae and Nymphalidae as inclusions in Miocene Dominican amber; the nymphalid specimen is the first adult known of this family preserved in amber. ...
... 4) because the wings are incomplete, overlapping, and touching. This specimen belongs to the genus Dynamine due to the combination of the following characters: presence of a white median band on fore-and hind wings, absence of ocelli on the forewing, two ocelli under the hind wing restricted toThe family Nymphalidae has been found previously in Dominican amber, represented by a caterpillar placed near the genus Smyrna of the subfamily Nymphalinae (Hammond and Poinar, 1998), and thus without any relationship to the new fossil nymphalid specimen. COMPARISON WITH EXTANT DYNAMINE: The genus Dynamine has very characteristic wing color patterns, especially that of the underside, and for this reason it can be confused only with the genus Lucinia. ...
A new, virtually complete and well-preserved female specimen of Voltinia dramba Hall, Robbins, and Harvey, 2004 (Lepidoptera: Riodinidae) provides new data on this fossil species, and a new fossil species of the Recent genus of Nymphalidae Dynamine Hübner, 1819 (Lepidoptera: Nymphalidae) is described as Dynamine alexae n.sp., on the basis of a male specimen. The two species are preserved in Miocene amber from the Dominican Republic. Dynamine alexae n.sp. represents the first adult nymphalid butterfly found as a fossil in amber. The four taxa of butterflies found up to the present in Dominican amber indicate post-Miocene extinctions in Hispaniola, probably caused by insularization. The butterflies found in Dominican amber do not support a hypothesis of a Gondwanan origin for many butterfly tribes and subfamilies as previously proposed; we conclude that this hypothesis is implausible based on the age of the butterflies as inferred from the fossil record. Some palaeoecologic and taphonomic questions are discussed.
... A specimen reported by DeVries & Poinar [85] (their figure 1) (re-figured in [86] (their figure 107, colour plates 107, p. 101-102,) and also refigured in Poinar [87] (his figure 48)) has short, stubby, and balloon-like setae at the head-trunk transition and softer-appearing setae on the trunk. A more heavily armoured specimen in Dominican amber was reported by Poinar & Hammond [88] (their figures 1-3) (re-figured in [86] (their figure 53B p. 58,) and Poinar et al. [83] (their figure 13)). The spines are not very long, but they are prominent, carrying secondary spinules. ...
Resin is a plastic-like product of trees. Older occurrences of such resin are referred to as amber and are considered fossil resin. Younger resins are termed copals. Even younger ones have been dubbed defaunation resins. Non-fossil resins remain in a terminological limbo, often referred to as “sub-fossils”. We report two lepidopteran caterpillars preserved in non-fossil resin: one from Madagascar, one from Brazil. Prominent hairs (=setae) and spines (=spine-like setae) of the specimens make it likely that they represent larvae of Erebidae (e.g., tussock moths and others). So far, most known caterpillars preserved in resins are either “naked” or bear protective cases; only few are armoured with spines or hairs. In particular, long-haired caterpillars such as the ones reported here are so far almost absent. Only one specimen with comparable setae has been reported from 15-million-year-old Dominican amber, but no significant details of this specimen are accessible. We briefly also review the record of caterpillars known from the Holocene, recognising that it is very sparse. The new specimens demonstrate that very hairy caterpillars can readily be preserved in resins in fine detail. Furthermore, the specimens increase the known size range of caterpillars preserved in resins, with one measuring more than 12 mm.
... phone SCUDDER (Nymphalidae) découvert dans les couches de l " Oligocène à Florissant , Colorado (USA) et dont l " état de conservation est absolument exceptionnel (BROSIUS,1994), peu de fossiles de Lépidoptères sont connus et surtout rarement en très bon état. Du Jurassique supérieur des montagnes de Shara-Teg en Mongolie sont connus de très nombreux insectes dont plusieurs Lépidoptères (KUPRIJANOV, 1998), quelques rares chenilles, chrysalides et imagos du Stampien vers Aix-en-Provence (NEL & NEL, 1985, 1986 PFRETZSCHNER, 1998) et de l'ambre du Crétacé inférieur du Nord de l " Espagne à Alava (ORTUÑO & ARILLO, 1998 ) et de la République Dominicaine (HAMMOND & POINAR, 1998). Les rares découvertes de papillons fossiles sont résumées ci-après : ...
The authors relate in the present paper the occurrence of several fossils dis-covered in the Upper Miocene diatomite deposit of Saint-Bauzile (F-07). This site shows an ex-ceptional richness with some fossils particularly well preserved. During one research, a lepidop-teron fossil was unexpectedly discovered. Ac-cording to the estimated age of the members of the subfamily, the known larval food plants and others criteria, this moth appears to belonging to the subfamily of the Ennominae (Geometridae). The authors propose to name this new lepidop-teron fossil Problongos baudiliensis gen. & spec. nov.
... In a piece of amber containing the haematophagous Triatoma dominicana Poinar, 2005, bat hairs indicated the probable host (Poinar, 2005). Bat hairs were also found adjacent to a larval brush-footed butterfly (Lepidoptera: Nymphalidae) entrapped in Dominican amber (Hammond & Poinar, 1998). According to Wenzel & Peterson (1981), whereas streblids can occur on colonial, cave-dwelling bats, most New World streblids occur on bats that roost in forest habitats. ...
The first fossil streblid, Enischnomyia stegosoma n. g., n. sp. (Diptera: Hippoboscoidea: Streblidae), is described from Dominican amber. Placed in the subfamily Nycterophiliinae Wenzel, 1966, which includes two New World extant genera, Nycterophilia Ferris, 1916 and Phalconomus Wenzel, 1984 (=Phalcophila Wenzel, 1976), the male specimen of E. stegosoma is characterised by the following features: a laterally compressed body, well-developed two-segmented antennae with the scape fused with the head, a tubular pedicel with an annulated basal portion and swollen apical portion bearing setae and bristles, a distinct flagellum with a dorsal boss bearing microsetae and a subterminal pectinate arista, a large tubular labium (proboscis) with the tip held upwards, eyes reduced to three facets, an expanded and flattened profemur, an anteriorly curved protarsus, and a well-developed wing with an entire distal margin. The possession of wings separates E. stegosoma from the species of Phalconomus, and the wing outline and venation, as well as the structure of the antennae and palps, distinguish it from species of Nycterophilia.
... The minimum age of the split between Dynamine and its sister group was constrained to be 20 Ma based on the Dominican amber fossil of Dynamine alexae, which clearly belongs to this extant genus (Peñ alver & Grimaldi 2006). The minimum age of the split between Smyrna and its sister group was constrained to be 20 Ma based on the Dominican amber fossil larva of Smyrna (Hammond & Poinar 1998). The minimum age of the split between Lethe and its sister group was constrained to be 25 Ma based on the fossil Lethe corbieri found in the Oligocene deposits of southeast France (Nel et al. 1993). ...
The butterfly family Nymphalidae contains some of the most important non-drosophilid insect model systems for evolutionary and ecological studies, yet the evolutionary history of the group has remained shrouded in mystery. We have inferred a robust phylogenetic hypothesis based on sequences of 10 genes and 235 morphological characters for exemplars of 400 of the 540 valid nymphalid genera representing all major lineages of the family. By dating the branching events, we infer that Nymphalidae originated in the Cretaceous at 90 Ma, but that the ancestors of 10-12 lineages survived the end-Cretaceous catastrophe in the Neotropical and Oriental regions. Patterns of diversification suggest extinction of lineages at the Cretaceous/Tertiary boundary (65 Ma) and subsequent elevated speciation rates in the Tertiary.
Caterpillars are an omnipresent component in modern terrestrial faunas. Not surprisingly, they represent an important food source for larger animals. The oldest fossils of caterpillars are known from the Cretaceous. Yet, so far, only nine possible Cretaceous specimens have been reported. Here we expand the known record by four new specimens of caterpillars preserved in 99 million-year-old Kachin amber from Myanmar. The specimens cannot easily be interpreted in a taxonomic or phylogenetic frame. A simple morphometric comparison reveals that the new specimens differ in their relative body dimensions from those of the previously known specimens, expanding the morphological diversity of Cretaceous caterpillars. All caterpillars from the Cretaceous of which sizes are known are rather small, the largest definite one being only about 5 mm. Comparison to younger ambers reveals no clear directed preservation bias towards preserving only small caterpillars. In addition, the sizes of known adult lepidopterans from the Cretaceous are compatible with rather small caterpillars, although possibly slightly larger than the known ones. While small, the observed size range appears still to be within the size range of food items known to be consumed by modern birds. It therefore seems likely that also in the Cretaceous caterpillars were adequate food items for early birds.
In this catalog, we attempt to assemble all fossil records of Lepidoptera described formally or informally in the world literature. A total of 667 records dealing with at least 4,568 specimens have been compiled. They include descriptions of 131 fossil genera and 229 fossil species, as well as 72 extant genera and 21 extant species to which some of these fossils supposedly belong or show superficial similarity. Replacement names of two fossil genera are proposed to avoid homonymy: Baltopsyche Sohn, gen. nov. for Palaeopsyche Sobczyk and Kobbert, 2009 and Netoxena Sohn, gen. nov. for Xena Martins-Neto, 1999. New generic combinations are proposed for: Tortrix? destructus Cockerell, 1916, Tortrix florissantanus Cockerell, 1907, and Tortrix sp. sensu Gravenhorst (1835), all three to Tortricites Kozlov, 1988; Pterophorus oligocenicus Bigot, Nel and Nel, 1986, to Merrifieldia Tutt, 1905; Aporia sp. sensu Branscheid (1969) to Pierites Heer, 1849; Noctua spp. sensu Hope (1836) and Lomnicki (1894), both to Noctuites Heer, 1849. Eleven names improperly proposed for lepidopteran fossils are invalidated: Baltonides roeselliformis Skalski in Kosmowska-Ceranowicz and Popiolek, 1981; Baltodines Kupryjanowicz, 2001; Barbarothea Scudder, 1890; Lepidopterites Piton, 1936; Palaeozygaena Reiss, 1936; Psamateia calipsa Martins-Neto, 2002; Saxibatinca meyi Skalski in Kristensen and Skalski, 1998; Spatalistiforma submerga Skalski, 1976; Thanatites juvenalis Scudder, 1875; Tortricibaltia diakonoffi Skalski, 1976; and Zygaenites Reiss, 1936. An unnecessary subsequent type designation for Pierites Heer, 1849, is discussed. A total of 129 records include lepidopteran fossils which cannot be placed in any taxonomic rank. There also exist at least 25 fossil records which lack any evidence of the supposed lepidopteran association. Misidentified specimens, including 18 fossil genera, 29 fossil species and 12 unnamed fossils, are excluded from Lepidoptera. All the known lepidopteran fossils are annotated by fossil type, specimen deposition, excavation locality, association with plants when present, and geological age. A bibliographic list of lepidopteran fossils is provided.
This work is a first attempt to integrate into one list and quantify all the known species of Hispaniolan arthropods. It includes all the terrestrial and surrounding marine arthropod species (plus those of Tardigrada and Onychophora) known to me to be reported for the island of Hispaniola until the end of 2007, as well as 158 species that are reported here as new records for the Dominican Republic and the island. A total of 8,237 valid species (6,833 extant and 1,404 fossils) are listed, of which the largest component are the insects (5,676 extant and 824 fossil species). Preliminarily, 2,521 arthropod species (36.9%) are considered to be endemic or unique to Hispaniola. Also 84 species are recognized as introduced. The bibliography complements the taxonomic information and includes over 4,000 titles. Brief annotations are also given on the history of entomology in Hispaniola.
The age and depositional history of Dominican amber-bearing deposits have not been well constrained. Resinites of different
ages exist in Hispaniola, but all of the main amberiferous deposits in the Dominican Republic (including those famous for
yielding biological inclusions) were formed in a single sedimentary basin during the late Early Miocene through early Middle
Miocene (15 to 20 million years ago), according to available biostratigraphic and paleogeographic data. There is little evidence
for extensive reworking or redeposition, in either time or space. The brevity of the depositional interval (less than 5 million
years) provides a temporal benchmark that can be used to calibrate rates of molecular evolution in amber taxa.
Although symbiotic association with ants is pervasive in the butterfly families Lycaenidae and Riodinidae the age of these symbioses has never been estimated explicitly. Here we report the first known fossil riodinid caterpillar. This fossil can be dated minimally between 15 and 20 Ma old, and confidently placed in the extant genus Theope. Differing little from modern day Theope, this fossil from Dominican amber provides direct evidence that secretory and acoustical organs used by modern caterpillars to mediate symbioses with ants have been highly developed at least since the Miocene. This fossil therefore becomes the point of reference for future studies using molecular clock methods for dating these symbioses within the riodinid butterflies. Modern evidence, and the abundance of dolichoderine ants in Dominican amber (now extinct in the West Indies) imply that specialized symbiotic relationships between Theope caterpillars and these ants were likely in existence at least 15 Ma ago. The current distribution of neotropical riodinid butterfly and ant faunas indicates the extinction in the West Indies of at least two unrelated taxa that formed a tightly linked symbiotic association, which persisted to the present elsewhere.
DNA has been successfully isolated from both fossilized plant and animal tissues. The oldest material, dated as 25-40 million years old (Tertiary), was obtained from amber-entombed bees and termites. Tissues from both these insects yielded DNA of good quality, which could be amplified by the polymerase chain reaction (PCR) and subsequently sequenced, including the genes encoding 18S ribosomal RNA and 16S rRNA. We report here the extraction of DNA from a 120-135-million-year-old weevil (Nemonychidae, Coleoptera) found in Lebanese amber, PCR amplification of segments of the 18S rRNA gene and the internal transcribed spacer, and the corresponding nucleotide sequences of their 315- and 226-base-pair fragments, respectively. These sequences were used for preliminary phylogenetic analysis of the nemonychid's sequence with three extant coleopterans: Lecontellus pinicola (Nemonychidae), Hypera brunneipennis (Curculionidae) and the mealworm Tenebrio molitor (Tenebrionidae), and two extant dipterans: the fruitfly Drosophila melanogaster (Drosophilidae) and mosquito Aedes albopictus (Culicidae) for the purpose of ascertaining the origin of the extracted and amplified DNA. The results revealed that the PCR-amplified material is that of the extinct nemonychid weevil. This represents the oldest fossil DNA ever extracted and sequenced, extending by 80 million years the age of any previously reported DNA.
It was not at first intended to do any work at Florissant during the summer of 1909, but two of my students, Messrs. Rusk and Duce, having become interested in the study of fossils, arranged to go there for a short time. Their collections include the usual percentage of insects, several of which are new and are herewith described.
Hymenaea protera is described from amber originating from La Toca mine in the Dominican Republic. The fossil species is characterized by the presence of reduced petals as well as distinctly clawed showy petals with cordate to reniform bases and a glabrous, verrucose ovary with long hirsute hairs at its base and along one margin. The fossil species most closely resembles the extantH. verrucosa Gaertner which occurs in East Africa and adjacent islands. These findings are interpreted as supporting a hypothesis that the genusHymenaea arose in the late Cretaceous on the combined South American-African continents and that Cretaceous and early Tertiary landmass movements were significant in determining the present amphi-Atlantic distribution of the genus. It is proposed that at least the majority, if not all, of the amber recovered from La Toca mine and other mines in the vicinity with similar-aged deposits originated fromH. protera.