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Poneromorph ants (Hymenoptera, Formicidae: Amblyoponinae, Ectatomminae, Ponerinae) of Grube Messel, Germany: Diversification during the Eocene
Abstract
Ants are a very successful group among insects, but the course of evolution of their biodiversity is still unclear. In this study, light is shed on the ant diversification during the Eocene. The analysis of the ant taphocoenosis of the fossil site Grube Messel, Germany, 47 Ma, yielded three poneromorph subfamilies and 22 new species in six genera of which four genera are new (Pseudectatomma gen. nov., Cephalopone gen. nov., Cyrtopone gen. nov., Messelepone gen. nov.). Only one extant genus, Pachycondyla, is present in the taphocoenosis from Messel. The high diversity of poneromorph ants from Messel is very conspicuous in comparison with the middle to late Eocene European ambers. In the ambers, a significant lower portion of species could be assigned to poneromorph ants, and even compared in absolute numbers, fewer poneromorph species are known from European ambers than from Messel. A possible gradual decline of the diversity of poneromorphs from the Eocene to the Miocene seems to be detectable worldwide. These insights are discussed in the context of the morphology and ecology of Poneromorpha and Formicomorpha. The proportion of ant castes in amber seems to indicate that already during the Eocene, poneromorphs inhabited preferably litter and soil, whereas formicomorphs preferred the arboreal realm. The “ponerine paradox” of having only a primitive social organization, yet being an old phylogenetic line with global distribution is discussed with emphasis on the paleontological data basis, but still remains unsolved. The evolutionary history of Myrmicinae is discussed. With the new available paleontological data, the timing and the dynamics of predominance of different subgroups of the ants can be traced more precisely than before.
... Regarding their biodiversity and ecological position, the success story of ants has been elaborated by Wilson (1990, 2010) and since then repeatedly mentioned by many other authors (e.g. Dlussky and Wedmann 2012;LaPolla et al. 2013). According to AntCat (Bolton 2019), there are 13,575 extant valid species described. ...
... At that time, myrmicine ants were very likely arboreal dwellers or at least foraging on trees, because the number of workers found in amber is relatively high, compared to the number of workers belonging to real epigaeic living taxa. Although the proportion of myrmicine specimens in European Eocene amber is low (Dlussky and Rasnitsyn 2009), Radchenko and Dlussky (2013) stress the increase of new myrmicine genera in late Eocene amber as possibly being the cause of the decrease of poneroid taxa in amber, compared to their high number found in mid-Eocene Messel deposits (Dlussky and Wedmann 2012). As most of the Eocene myrmicine genera are extinct, the authors suppose that they could not compete with other myrmicine taxa or with taxa from other subfamilies invading Europe. ...
... Only during the Late Oligocene to mid Miocene do the number of myrmicine ants specimens increase to contemporary percentages, i.e. Kleinkems 36% (early Oligocene), Enspel 36.6% (Upper Oligocene MP 28, this paper), Rott 50% (late Oligocene, MP 30), Radoboj 22% (early Miocene) and Vishnevaya balka, Stavropol 40% (mid Miocene) (Dlussky and Wedmann 2012). These findings, however, are in accordance with earlier statements by Ward (2000), according to whom the numbers of fossil myrmicine ant specimens became similar to the modern ones (about 74%) by the mid Miocene. ...
New species of the subfamily Myrmicinae are described from the fossil volcanogenic Lake Enspel, Westerwald, Germany. This Fossil-Lagerstätte comprises lake sediments, which essentially consist of intercalated “oilshales” and volcaniclastics. The sediments were dated to be of late Oligocene age. Out of the 287 fossil ants from Enspel examined, 96.5% (n = 277) belong to the formicoid clade (Formicinae, Dolichoderinae, Myrmicinae), and only 3.5% (n = 10) to the poneroid clade (Ponerinae, Agroecomyrmecinae). The high proportion of Myrmicinae (36.6%) indicates that this subfamily was already strongly represented in Central Europe in the late Oligocene. However, it should be noted that the large number of specimens are alate reproductives. The number of workers is low: n = 19, or 6.1% of 312 specimens when 25 isolated wings are included. Ten new species, belonging to the genera Paraphaenogaster, Aphaenogaster, Goniomma and Myrmica are named. Taxonomic questions regarding the separation of the two genera Aphaenogaster and Paraphaenogaster as well as aspects of biogeography, palaeoenvironment and palaeobiology in Enspel are discussed.
... Species included. Casaleia inversa (Dlussky, 1981) (Chon-Tuz, Kirgizia, Middle Miocene), C. longiventris (Heer, 1849) (Radoboj, Croatia, Early Miocene), and C. eocenica Dlussky et Wedmann, 2012 (Messel, Germany, Middle Eocene) [Heer, 1849;Dlussky, 1981;Dlussky, Wedmann, 2012]. ...
... Described as entire faunas are also the one from the Middle Miocene of Stavropol (Cis-Caucasian [Dlussky, 1981]), Late Eocene (Priabonian) of the Bembridge Marls (north in Isle of Wight, UK [Antropov et al., 2014]) and the Early Miocene of Radoboj in Croatia [Dlussky, Putyatina, 2014]. Ants of the Middle Eocene of Grube Messel and Eckfeld Maar in Germany are described only in part [Lutz, 1986;Dlussky et al., 2008Dlussky, 2012;Dlussky, Wedmann, 2012]. However, complete lists of ants identified up to subfamily are available for these fossil sites. ...
... In temperate forests, 80% of myrmicine ants are foraging within the leaf litter, soil and/or lower herb layer. As to Poneromorpha, it was only mid-Eocene when these ants override Myrmicinae in species and specimen diversity in Europe [Dlussky, Wedmann, 2012]. In the Late Eocene, abundance of myrmicine ants surpassed that of poneromorphs but left low yet until grew considerably in majority of the Oligocene and Miocene assemblages (Radoboj excluded). ...
The ant fossils of the Bol’shaya Svetlovodnaya (Late Eocene of Sikhote-Alin, Russian Far East) are studied. A total of 80 specimens are identified as Formicidae. The location is quite diverse in composition of ant: described 15 genera and 9 subfamilies. A generic level comparison reveals Svetlovodnaya ants as more similar to European rather than North American ant fauna. Unusually high proportion Myrmicinae (21.4%) for the known deposits of Europe, where these ants become abundant only in the Miocene. A distinctive feature of the Svetlovodnaya ant assemblage under comparison with the European and North American ones is the lack of highly dominanting species. Presences of strictly tropical and temperate taxa support the equable climate concept as applicable to the Late Eocene climate of the Sikhote-Alin area. New genera and species are described from the Bol’shaya Svetlovodnaya: Aneuretinae – Paraneuretus dubovikoffi sp. n., Dolichoderinae – Dolichoderus pinguis sp. n., Emplastus macrops sp. n., Emplastus elongates sp. n., Emplastus dubius sp. n., Liometopum incognitum sp. n., Formicinae – Formica biamoensis sp. n., Formica paleosibirica sp. n. Gesomyrmex magnus sp. n., Gesomyrmex macrops sp. n., Gesomyrmex incertus sp. n., Myrmeciinae – Ypresiomyrma orientalis sp. n., Amblyoponinae – Casaleia orientalis sp. n., Ponerinae – Pachycondyla oligocenica sp. n., Pachycondyla aberrans sp. n., Proceratiinae – Proceratium petrosum sp. n., Myrmicinae – Agastomyrma laticeps sp. n., Biamomyrma zherikhini sp. n., Biamomyrma lata sp. n., Biamomyrma rugosa sp. n., Myrmecites sibiricus sp. n., Myrmecites dubius sp. n., Myrmecites major sp. n. , also at least two species of Lasius and one species of Cerapachyinae.
... Today, the "big four" subfamilies Dolichoderinae, Formicinae, Myrmicinae, and Ponerinae comprise the vast majority of the biomass and taxonomic diversity in most ecosystems (WILSON & HÖLLDOBLER 2005a). Aside from the Messel Formation where formiciines are highly abundant ( DLUSSKY & WEDMANN 2012), dominance of the "big four" is largely consistent into the Eocene. Green River (~ 51 Ma;SMITH & al. 2008), perhaps best known for its wealth of well-preserved fish fossils, has long been a source for ant fossils (SCUDDER 1877b), although they are not as abundant or well preserved as other approximately contemporaneous fossil beds. ...
... ( GRIMALDI & AGOSTI 2000, DLUSSKY & RASNITSYN 2002, LAPOLLA & al. 2013), although it has been noted that the faunal composition preserved in the fossil record may not accurately reflect that of ancient ecosystems (e.g., SOLÓRZANO KRAEMER & al. 2015). There are some anomalies, but the general trend is as follows: Ants never make up more than 1.5% of insects in fossil localities during the Cretaceous, in the Cenozoic prevalence increases to as high as 13.1% in the Messel Formation during the Eocene (recall that reproductive formiciines are responsible for half of this abundance; DLUSSKY & RASNITSYN 2002, DLUSSKY & WEDMANN 2012) and 20% in the Eocene-aged Florissant Formation ( CARPENTER 1930). During the Miocene, in Dominican amber, ant prevalence reaches ~ 27-36% ( , DLUSSKY & WEDMANN 2012). ...
... There are some anomalies, but the general trend is as follows: Ants never make up more than 1.5% of insects in fossil localities during the Cretaceous, in the Cenozoic prevalence increases to as high as 13.1% in the Messel Formation during the Eocene (recall that reproductive formiciines are responsible for half of this abundance; DLUSSKY & RASNITSYN 2002, DLUSSKY & WEDMANN 2012) and 20% in the Eocene-aged Florissant Formation ( CARPENTER 1930). During the Miocene, in Dominican amber, ant prevalence reaches ~ 27-36% ( , DLUSSKY & WEDMANN 2012). Today, highly dominant species such as members of Formica in parts of Europe and Iridomyrmex in some landscapes of Australia ( HÖLLDOBLER & WILSON 1990) make up the majority of ant biomass. ...
The ant fossil record is summarized with special reference to the earliest ants, first occurrences of modern lineages, and the utility of paleontological data in reconstructing evolutionary history. During the Cretaceous, from approximately 100 to 78 million years ago, only two species are definitively assignable to extant subfamilies – all putative crown group ants from this period are discussed. Among the earliest ants known are unexpectedly diverse and highly social stemgroup lineages, however these stem ants do not persist into the Cenozoic. Following the Cretaceous-Paleogene boundary, all well preserved ants are assignable to crown Formicidae; the appearance of crown ants in the fossil record is summarized at the subfamilial and generic level. Generally, the taxonomic composition of Cenozoic ant fossil communities mirrors Recent ecosystems with the "big four" subfamilies Dolichoderinae, Formicinae, Myrmicinae, and Ponerinae comprising most faunal abundance. As reviewed by other authors, ants increase in abundance dramatically from the Eocene through the Miocene. Proximate drivers relating to the "rise of the ants" are discussed, as the majority of this increase is due to a handful of highly dominant species. In addition, instances of congruence and conflict with molecularbased divergence estimates are noted, and distinct "ghost" lineages are interpreted. The ant fossil record is a valuable resource comparable to other groups with extensive fossil species: There are approximately as many described fossil ant species as there are fossil dinosaurs. The incorporation of paleontological data into neontological inquiries can only seek to improve the accuracy and scale of generated hypotheses.
... These included ten ants (1.2% of the total number of insects), based on which Dlussky (1988) described five extinct genera and seven species, assigned to the extant subfamilies Ponerinae (Protoo pone Dlussky), Aneuretinae (Aneuretellus Dlussky), Dolichoderinae (Eotapinoma Dlussky, Zherichinius Dlussky), and Formicinae (Chimaeromyrma Dlussky). It is interesting that representatives of the two genera of Dolichoderinae described from the Sakhalin amber were later found in the much older deposits of Alberta (Canada, Upper Cretaceous, Campanian, 78-79 Ma) (Eotapinoma macalpini Dlussky) ( Dlussky, 1999) and in a younger Late Eocene Bitterfeld (=Saxonian) amber (35-37 Ma) (as yet undescribed species of Zherichinius) ( Dlussky and Rasnitsyn, 2009;Duboo vikoff, 2012), whereas seven new species of the genus Protopone were described from the Middle Eocene (Lutetian) shale in Germany (Grube Messel, 47 Ma) ( Dlussky and Wedmann, 2012). The age of the Sakhalin amber was debated for a long time. ...
... Originally, these beds were dated as Late Eocene-Early Oligocene ( Viana et al., 1957), but, later, these deposits were dated as Late Eocene ( Rossi de Garcia, 1983), or were dated within a wide range from the Late Paleocene to Early Oligocene ( Petrulevv icius, 1999); Dlussky previously considered this spee cies Paleocene ( Dlussky and Fedoseeva, 1988). Finally, in the last decade, the Ventana Formation was dated as Middle Eocene (Early Lutetian, 47-48 Ma) ( Wilf et al., 2005;Dlussky, 2012). Previously, the Ølst Formation (Denmark) was dated Upper PaleoceneLower Eocene ( Dlussky and Rasnitsyn, 2007) with numerous remains of Ypresiomyrma rebekkae ( Rust et Andersen), but now it is clear that these remains belong to the very beginning of the Ypresian (Lower Eocene, ca. ...
... It is noteworthy that, for a long time, Irii domyrmex (Dolichoderinae) was thought to be the earr liest occurrence of an extant genus, while Eocenidris Wilson was thought to be the earliest member of the subfamily Myrmicinae. The extant genera Gesomyrmex Mayr, Oecophylla F. Smith (Formicinae), and Pachycondyla (Ponerii nae) as well as a number of extinct genera from the extant subfamilies Dolichoderinae, Amblyoponinae, Ectatomminae, Ponerinae, Myrmeciinae, Pseudomyr mecinae, Cerapachyinae, and Myrmicinae, and the extinct subfamily Formiciinae were found in the Midd dle Eocene beds of Germany (Messel, 47 Ma; Eckk feld, 44 Ma) ( Lutz, 1986;Dlussky et al., 2008Dlussky, 2012;Dlussky and Wedmann, 2012;LaPolla et al, 2013). It should be stressed that material from these Lagerstätten is only partly studied and no generic level identification is referred to the subfamii lies Dolichoderinae, Pseudomyrmecinae, Cerapachyy inae, and Myrmicinae from these localities in available publications. ...
The earliest member of an extant ant genus of the subfamily Myrmicinae, Aphaenogaster dlusskyana sp. nov., from the Sakhalin amber (Russia, Middle Eocene, 43–47 Ma) is described. Fossil representatives of the genus Aphaenogaster are critically analyzed and it is proposed to transfer many of these to the fossil morphotaxon Paraphaenogaster. The morphology, distribution, and possible evolutionary trends of the genus Aphaenogaster are reviewed and it is suggested that this genus appeared in the territory currently occupied by the Palearctic not later than the Early Eocene (over 50 Ma). Aphaenogaster dlusskyana can be considered the oldest described representative of an extant genus of the subfamily Myrmicinae, although earlier, as yet undescribed, records of extant genera of Myrmicinae belong to the Early Eocene.
... Work on fossil ants has mainly focused on taxonomic analysis of individual specimens (e.g. Baroni Urbani, 1980;Perrichot et al., 2008a;Dlussky & Wedmann, 2012). Only a few synthetic analyses exist on particular deposits with a rich ant palaeocommunity, such as Baltic and Dominican ambers (Wheeler, 1915;Dlussky & Putyatina, 2014). ...
... However, these authors verbally described patterns but did not test them quantitatively. Furthermore, as those original studies, subsequent work has enriched our understanding of both the fossil and modern ant faunal distributions, with studies on new deposits available (Europe: Dlussky & Wedmann, 2012;Dominican amber: Wilson, 1985;Baroni Urbani, 1995;Chiapas: Sol orzano Kraemer, 2007), and improvements in taxonomy of both modern and fossil taxa (Ward, 2007). The aggregation of the most recent data for both the fossil and modern ant distributions (Gu enard et al., 2010;Perrichot, 2014) allow now for more rigorous comparison of the affinities between those different assemblages. ...
... These numbers would undoubtedly increase significantly with the inclusion of genera only known from the fossil record, which were excluded in our analysis. For example, 6 extinct genera of the subfamily Aneuretinae are present in European Tertiary fossil deposits but the subfamily is currently restricted to Sri Lanka (Dlussky, 2012). Similarly, extinct ant genera of the subfamily Myrmeciinae are present in Nearctic and European fossil faunas (Archibald et al., 2006), but the subfamily is now restricted to Australasia. ...
We investigate the dynamics of ant biodiversity patterns and community structure from the Eocene until the present. Our goal is to empirically test hypotheses regarding the similarities in composition and structure of fossil and modern ant faunas to understand the historical processes influencing modern global biodiversity patterns.LocationGlobal.Methods
We integrated two recently developed databases of the geographical distributions of fossil and modern ant genera and analysed the evolution of diversity and composition since the Eocene, c. 55 Ma. We assembled a third new database on community structure of ants, and used it to compare community structure of fossil assemblages with modern communities in different bioregions.ResultsThe analyses of generic composition and community structure were congruent, supporting a strong affinity between the Western Palaearctic ant fauna and modern Indomalayan and Australasian assemblages, and of a widespread Holarctic ant palaeofauna, and affinity between fossil Caribbean and modern Neotropical faunas. In addition, neither generic composition nor community structure of fossil assemblages showed evidence of taphonomic bias towards arboreal taxa.Main Conclusions
The aggregated fossil record reveals the dynamic nature of macroecological patterns in ant biodiversity during the Cenozoic, with continental-scale generic extinctions common and important in shaping modern ant assemblages. Our results suggest major compositional changes for the Western Palaearctic bioregion and the Caribbean bioregions. The ant palaeofauna of the Western Palaearctic bioregion was then very similar in composition and structure to the one observed now in the Indomalayan bioregion, supporting the notion that modern-day ‘tropical’ biomes were historically much more widespread, while temperate biotas are more recently evolved. Our results underscore the importance of a palaeogeographical perspective in understanding modern-day macroecological patterns.
... Species included. Casaleia inversa (Dlussky, 1981) (Chon-Tuz, Kirgizia, Middle Miocene), C. longiventris (Heer, 1849) (Radoboj, Croatia, Early Miocene), and C. eocenica Dlussky et Wedmann, 2012 (Messel, Germany, Middle Eocene) [Heer, 1849;Dlussky, 1981;Dlussky, Wedmann, 2012]. ...
... Described as entire faunas are also the one from the Middle Miocene of Stavropol (Cis-Caucasian [Dlussky, 1981]), Late Eocene (Priabonian) of the Bembridge Marls (north in Isle of Wight, UK [Antropov et al., 2014]) and the Early Miocene of Radoboj in Croatia [Dlussky, Putyatina, 2014]. Ants of the Middle Eocene of Grube Messel and Eckfeld Maar in Germany are described only in part [Lutz, 1986;Dlussky et al., 2008Dlussky, 2012;Dlussky, Wedmann, 2012]. However, complete lists of ants identified up to subfamily are available for these fossil sites. ...
... In temperate forests, 80% of myrmicine ants are foraging within the leaf litter, soil and/or lower herb layer. As to Poneromorpha, it was only mid-Eocene when these ants override Myrmicinae in species and specimen diversity in Europe [Dlussky, Wedmann, 2012]. In the Late Eocene, abundance of myrmicine ants surpassed that of poneromorphs but left low yet until grew considerably in majority of the Oligocene and Miocene assemblages (Radoboj excluded). ...
The ant fossils of the Bol’shaya Svetlovodnaya (Late Eocene of Sikhote-Alin, Russian Far East) are studied. A total of 80 specimens are identified as Formicidae. The location is quite diverse in composition of ant: described 15 genera and 9 subfamilies. A generic level comparison reveals Svetlovodnaya ants as more similar to European rather than North American ant fauna. Unusually high proportion Myrmicinae (21.4%) for the known deposits of Europe, where these ants become abundant only in the Miocene. A distinctive feature of the Svetlovodnaya ant assemblage under comparison with the European and North American ones is the lack of highly dominanting species. Presences of strictly tropical and temperate taxa support the equable climate concept as applicable to the Late Eocene climate of the Sikhote-Alin area. New genera and species are described from the Bol’shaya Svetlovodnaya: Aneuretinae – Paraneuretus dubovikoffi sp. n., Dolichoderinae – Dolichoderus pinguis sp. n., Emplastus macrops sp. n., Emplastus elongates sp. n., Emplastus dubius sp. n., Liometopum incognitum sp. n., Formicinae – Formica biamoensis sp. n., Formica paleosibirica sp. n. Gesomyrmex magnus sp. n., Gesomyrmex macrops sp. n., Gesomyrmex incertus sp. n., Myrmeciinae – Ypresiomyrma orientalis sp. n., Amblyoponinae – Casaleia orientalis sp. n., Ponerinae – Pachycondyla oligocenica sp. n., Pachycondyla aberrans sp. n., Proceratiinae – Proceratium petrosum sp. n., Myrmicinae – Agastomyrma laticeps sp. n., Biamomyrma zherikhini sp. n., Biamomyrma lata sp. n., Biamomyrma rugosa sp. n., Myrmecites sibiricus sp. n., Myrmecites dubius sp. n., Myrmecites major sp. n. , also at least two species of Lasius and one species of Cerapachyinae.
... [5,16,17]. Recently, the discovery of both a new basal lineage of subterranean, predatory ants [16], and new fossil deposits [11,18,19] have re-ignited focus on the ecology of the earliest ant lineages. These new findings have prompted further discussion of the habitat strata in which early ants lived, including speculation about possible arboreality in some of the earliest ant fossils [20]. ...
... It was based on early attempts to date ant origins [3] as well as morphology of the oldest ant and stem-group ant fossils, about which much more is known today [34][35][36][37][38]. Recently, the age of the earliest ants and the subsequent diversification of the family were estimated with molecular and fossil evidence [5,9,15]. The earliest ants, fossils from the Cretaceous, are indeed dissimilar to the extant lineages known now to be sister to all the remaining ants [11,18,19]. The earliest fossils representing the presumed sister-group to ants, the extinct subfamily Sphecomyrminae, are known from workers and have large eyes, suggesting they were not soildwelling, but rather epigaeic or arboreal [39]. ...
... These factors, combined with other hints about early ant lifestyles, make it tempting but difficult to draw clear conclusions about habitat use based on early ant fossils. For example, Poneromorphs are represented in amber mostly by alates [18,36,37], indicating that workers were soil dwelling, whereas the workers of surfacedwelling species (Formicinae and Dolichoderinae) are primarily preserved in the amber fossil record and thought to be arboreal [38]. Finally, the subterranean habitat is frequented by the descendants of the lineage from which early ants evolved, the aculeate Hymenoptera. ...
Formicidae) is increasingly well-understood due to recent phylogenetic analyses, along with estimates of divergence times and diversification rates. Yet, leading hypotheses regarding the ancestral habitat of ants conflict with new findings that early ant lineages are cryptic and subterranean. Where the ants evolved, in respect to habitat, and how habitat shifts took place over time have not been formally tested. Here, we reconstruct the habitat transitions of crown-group ants through time, focusing on where they nest and forage (in the canopy, litter, or soil). Based on ancestral character reconstructions, we show that in contrast to the current consensus based on verbal arguments that ants evolved in tropical leaf litter, the soil is supported as the ancestral stratum of all ants. We also find subsequent movements up into the litter and, in some cases, into the canopy. Given the global importance of ants, because of their diversity, ecological influence and status as the most successful eusocial lineage on Earth, understanding the early evolution of this lineage provides insight into the factors that made this group so successful today.
... Work on fossil ants has mainly focused on taxonomic analysis of individual specimens (e.g. Baroni Urbani, 1980;Perrichot et al., 2008a;Dlussky & Wedmann, 2012). Only a few synthetic analyses exist on particular deposits with a rich ant palaeocommunity, such as Baltic and Dominican ambers (Wheeler, 1915;Dlussky & Putyatina, 2014). ...
... However, these authors verbally described patterns but did not test them quantitatively. Furthermore, as those original studies, subsequent work has enriched our understanding of both the fossil and modern ant faunal distributions, with studies on new deposits available (Europe: Dlussky & Wedmann, 2012;Dominican amber: Wilson, 1985;Baroni Urbani, 1995;Chiapas: Sol orzano Kraemer, 2007), and improvements in taxonomy of both modern and fossil taxa (Ward, 2007). The aggregation of the most recent data for both the fossil and modern ant distributions (Gu enard et al., 2010;Perrichot, 2014) allow now for more rigorous comparison of the affinities between those different assemblages. ...
... These numbers would undoubtedly increase significantly with the inclusion of genera only known from the fossil record, which were excluded in our analysis. For example, extinct genera of the subfamily Aneuretinae are present in European Tertiary fossil deposits but the subfamily is currently restricted to Sri Lanka (Dlussky, 2012). Similarly, extinct ant genera of the subfamily Myrmeciinae are present in Nearctic and European fossil faunas (Archibald et al., 2006), but the subfamily is now restricted to Australasia. ...
The dominance of ants in the terrestrial biosphere has few equals among animals today, but this was not always the case. The oldest ants appear in the fossil record 100 million years ago, but given the scarcity of their fossils, it is presumed they were relatively minor components of Mesozoic insect life. The ant fossil record consists of two primary types of fossils, each with inherent biases: as imprints in rock and as inclusions in fossilized resins (amber). New imaging technology allows ancient ant fossils to be examined in ways never before possible. This is particularly helpful because it can be difficult to distinguish true ants from non-ants in Mesozoic fossils. Fossil discoveries continue to inform our understanding of ancient ant morphological diversity, as well as provide insights into their paleobiology.
... épigéique. Et bien que la proportion de myrmicines trouvées dans l'ambre éocène d'Europe soit relativement faible (Dlussky & Rasnitsyn, 2009), ces derniers auteurs soulignent également que l'apparition de nouvelles lignées 13 de myrmicines à la fin de l'Éocène pourrait expliquer la diminution de la présence de taxons ponéroïdes, alors que leur proportion était encore forte comme le montrent notamment les dépôts de Messel datés du milieu de l'Éocène (Lutétien, (Dlussky & Wedmann, 2012). De nombreuses lignées de myrmicines apparues à l'Éocène, devaient sans doute être en compétition entre elles (ainsi qu'avec d'autres sous-familles) pour les mêmes niches écologiques, et le nombre important de genres typiques de cette époque (ex : Boltonidris, Eocenomyrma, Fallomyrma...) ayant disparu avant l'Oligocène-Miocène, s'expliquerait par une valeur fitness moins importante et témoignerait de cette compétition écologique. ...
... L'âge de calibration donne les limites inférieure et supérieure de l'âge attribué au taxon, dont les limites sont intégrées dans l'onglet Priors du logiciel sous le module Sampled Ancestors (distribution uniforme) et CladeAge (distribution automatique). Les valeurs de calibration sous une distribution en log-normale utilisées pour l'analyse FBD (FBD-SD) sont en grande partie issues de Brady et al.Basée sur Pseudectatomma eocenica et P. striatula décrites dans les schistes bitumineux de Grube Messel (Allemagne) parDlussky & Wedmann (2012). Celles-ci sont rattachées à la sous-famille Ectatomminae. ...
Avec plus de 7000 espèces, les fourmis Myrmicinae constituent l’un des plus grands succès écologiques de l’histoire. Pourtant, leur histoire évolutive reste mal comprise. Le présent travail tente de retracer l’évolution du groupe, par l’étude taxonomique des récentes découvertes de myrmicines fossiles (ambre éocène de l’Oise, ambre miocène de Zhangpu et d’Éthiopie), pour les utiliser comme nouveaux points de calibration. L’approche combine les plus récents outils permettant de mieux considérer les données paléontologiques (taux de diversification, d’échantillonnage, etc.), et leur intégration dans l’analyse phylogénétique (modèle FBD, CladeAge). L’effet des diverses modalités de calibration (calibration à la racine, du groupe-couronne ou à l’origine, modèles de distribution, node-dating vs. tip-dating) sur les estimations des temps de divergence est également testé et discuté. Enfin, l’histoire biogéographique est revue à l’aune des nouvelles occurrences et des résultats phylogénétiques. Le groupe serait apparu dans le Nouveau Monde au Crétacé Supérieur (85-95 Ma), sans toutefois montrer une appartenance plus marquée au Néarctique ou au Néotropique. Les grandes lignées se seraient ensuite rapidement dispersées, en particulier à l’Éocène à travers l’Antarctique, la Béringie et le Greenland. L’extension des latitudes tropicales à la suite d’évènements hyperthermiques (ETM, MECO) auraient permis des dispersions successives entre Nouveau et Ancien Monde, et expliquent la disparité des distributions actuelles, où les lignées basales sont respectivement restreintes au Néarctique-Paléarctique et au Nouveau Monde, tandis que les lignées plus dérivées montrent des distributions plus larges mais plus hétérogènes.
... Best known are the vertebrates, but ecologically just as important is the insect fauna. A wide range of different groups of insects has been documented from Messel [29], including e.g. the first fossil leaf insect and a very diverse hymenopteran fauna [30][31][32][33]. Permission for digging for fossils in the Messel Pit Fossil Site was issued to the Senckenberg Research Institute Frankfurt by the following authority: Archaeologische und palaeontologische Denkmalpflege, hessenArchaeologie, Schloss Biebrich/Ostfluegel, 65203 Wiesbaden, Germany. ...
Newly discovered fossil bugs (Insecta: Hemiptera: Heteroptera:
Pentatomidae) from the Eocene of Messel (Germany) and
Green River (North America) exhibit an exaggerated
morphology including prominent spiny humeral and
anterolateral angles of the pronotum and a spiny lateral
abdominal margin. Especially the humeral angles are unique;
they consist of expansive, rounded projections with strong
spines, which is a rare trait among pentatomids. A hypothesis
for the function of this extreme morphology is defence against
small vertebrate predators, such as birds or reptiles. The same
protuberances also produce a disruptive effect camouflaging
the specimen in its environment and provide additional
protection. Therefore, the extreme morphology provides
primary as well as secondary anti-predator defence. The
morphology of Eospinosus peterkulkai gen. et sp. nov. and
E. greenriverensis sp. nov. resembles that of Triplatygini, which
today occur exclusively in Madagascar, as well as that of
Discocephalinae or Cyrtocorinae, which today occur in the
Neotropics. Due to a lack of conclusive characters, it cannot be
excluded that the fossil species may represent a case of
remarkable convergence and are not related to either taxon.
Phylogenetic analyses using parsimony as well as Bayesian
algorithms confirmed that the new genus is a member of
Pentatomidae, but could not solve its phylogenetic relationships
within Pentatomidae.
... comm., 2013) (Table 1) interesting that at least for two amber faunas with the same (Danish, 1.6) or very similar (Rovno, 1.9) ratios were calculated for biting midges (Ceratopogonidae): predominance of Holarctic biting midges in the representative collections from other ambers is even higher than for Holarctic ants (Perkovsky, 2017). In the early Eocene at Messel and middle Eocene at Eckfeld, tropical taxa absolutely dominate, nearly as much as in tropical climate Oligocene Sicilian amber (Emery, 1891;Wappler, 2003;Dlussky et al., 2008Archibald et al., 2011;Dlussky, 2012;Dlussky, Wedmann, 2012). The prevalence of Holarctic species in all representative amber ant collections is associated with decreasing of temperature in the late Eocene (see below). ...
... The scarcity of outcrops with well-preserved fossil insects from the Late Cretaceous to the Eocene is the reason that only a few studies compare insect assemblages from different ages. One study, based on Middle Eocene ants (Formicidae) from the Messel maar showed that the dominance and diversity of different subgroups of ants changed substantially from the Middle to the Late Eocene ( Dlussky and Wedmann, 2012). ...
The Paleocene is an important epoch in the development of the flora and fauna in the Cenozoic,but relatively few terrestrial fossil sites of this age are known in Europe. For many groups, like plants and mammals, the catastrophic events at the Cretaceous–Paleogene (K/Pg, formerly K/T) boundary had ushered in significant changes, and our understanding of western European flora and fauna during this period is still very incomplete. The Paleocene Konservat- Lagerstätte Menat (Puy-de-Dôme, France) is an archive of an ecosystem that can provide insights into this crucial epoch. The sediments of Menat were deposited in a former lake, probably a maar lake, and excavations during the last century have yielded an
extensive flora and fauna. An overview of the current state of paleontological investigations is given and new results based on recent excavations are presented. We find that the preservation of organic matter
strongly differs between different excavation sites, probably influenced by weathering processes. Recently our team collected many impression fossils, those in more bituminous layers being exceptionally wellpreserved, including three-dimensional plant remains. The occurrence of charcoal in many horizons at Menat signifies the occurrence of paleowildfires during the Paleocene in the vicinity of Paleolake Menat.
These results demonstrate the high potential for further scientific studies at Menat.
... The scarcity of outcrops with well-preserved fossil insects from the Late Cretaceous to the Eocene is the reason that only a few studies compare insect assemblages from different ages. One study, based on Middle Eocene ants (Formicidae) from the Messel maar showed that the dominance and diversity of different subgroups of ants changed substantially from the Middle to the Late Eocene (Dlussky and Wedmann, 2012). ...
The Konservat-Lagerstatte Menat (Puy-de-Dome, France) is an outstanding archive of a Paleocene ecosystem, which was deposited in a former maar lake. Excavations during the last century have yielded an extensive flora and fauna record, therefore an overview of the current state of paleontological investigations is given in this paper. Additionally, new results based on excavations from the years 2012 to 2014 are presented. The preservation of organic matter differed strongly between excavation sites, probably influenced by weathering processes. The stratigraphic succession consists mostly of organic-rich clays, intercalated with hard, silicified claystones. In 2013 and 2014 both impression and compression fossils were collected from different outcrops. Compression fossils from organic-rich clays were exceptionally well-preserved and included three-dimensional plant remains. A new database on insect paleobiodiversity was compiled. The occurrence of charcoal in almost all horizons investigated suggests that paleowildfires were frequent during the Paleocene in the vicinity of the paleolake. The results confirm the high potential of the Konservat-Lagerstatte Menat for future paleontological research.
... While molecular analyses indicate that many higher-level ant taxa were established in the Cretaceous, only a few are confirmed by fossils Moreau et al. 2006;LaPolla et al. 2013). Most recent hypotheses on the diversification of ants use "diversification" to mean divergence of their higher-level sub-taxa, although a few concern increase of richness at lower levels, which we mean here (e.g., see differing meanings by Leston 1973;Wilson and Hölldobler 2005;Moreau et al. 2006;Dunn et al. 2007;Dlussky and Wedmann 2012;Lucky et al. 2013;Ward 2014). The Cretaceous fossil record suggests a low community presence and impact of small numbers of individuals and species (Zherikhin 2002;Barden and Grimaldi 2016). ...
Most major modern families of Hymenoptera were established in the Mesozoic, but the diversifications within ecologically key trophic guilds and lineages that significantly influence the character of modern terrestrial ecosystems – bees (Apiformes), ants (Formicidae), social Vespidae, parasitoids (Ichneumonidae), and phytophagous Tenthredinoidea – were previously known to occur mostly in the middle to late Eocene. We find these changes earlier, seen here in the early Eocene Okanagan Highlands fossil deposits of western North America. Some of these may have occurred even earlier, but have been obscured by taphonomic processes. We provide an overview of the Okanagan Highlands Hymenoptera to family level and in some cases below that, with a minimum of 25 named families and at least 30 when those tentatively assigned or distinct at family level, but not named are included. Some are poorly known as fossils (Trigonalidae, Siricidae, Peradeniidae, Monomachidae), and some represent the oldest confirmed occurrences (Trigonalidae, Pompilidae, Sphecidae sensu stricto , Peradeniidae, Monomachidae, and possibly Halictidae). Some taxa previously thought to be relictual or extinct by the end of the Cretaceous (Angarosphecidae, Archaeoscoliinae, some Diapriidae) are present and sometimes abundant in the early Eocene. Living relatives of some taxa are now present in different climate regimes or on different continents.
... The site is famous for its extraordinarily well preserved fossils of early mammals (e. g. Franzen 2007, Franzen et al. 2009), but has also supplied a huge number of fossil insect specimens (e. g. lutz 1990, Wedmann 2005, dluSSky & Wedmann 2012. Presently, the Senckenberg collection of Messel insects comprises over 16,500 fossils, some of them unusually well preserved, e. g. jewel beetles and moths, with more or less original structural colours (HörnSCHemeyer & Wedmann 1994, mCnamara 2013 and the first known fossil leaf insect (Wedmann et al. 2007). ...
Six fossil specimens of Plecia Wiedemann, 1828 were found in Grube Messel, a widely known Eocene locality. A male and a female were identified as Plecia hoffeinsorum Skartveit, 2009, a species originally described from Baltic amber, which is the first record of a Baltic amber species in Grube Messel. Another female is either close to or conspecific with Plecia acourti Cockerell, 1921, a species previously known from the late Eocene of Isle of Wight (United Kingdom). The remaining three specimens are liekly to belong to an undescribed species, which we hesitate to name here as our material is in too poor a condition.
... Th e main diff erence between these faunas and those of Priabonian ambers is the dominance of Formica and Lasius in the latter. Th e Messel fauna is dominated (49 % of all ants) by extremely thermophilic Formiciini (not known in Priabonian) of the genus Titanomyrma (Archibald et al., 2011), poneromorph ants are quite large in number and extremely diverse (22 species, Dlussky, Wedmann, 2012), whereas Formicinae are apparently 12 species out of 60, and 4 of those are assigned to the tropical genera Gesomyrmex and Oecophylla . Conversely, there are at least 18 known species of Holarctic Formicini and Lasius in Priabonian ambers, including 15 in Baltic amber. ...
Based on representative collections, the ratio of tropical and Holarctic ant species in Priabonian (Late Eocene) Baltic, Bitterfeld (Saxonian), Danish and Rovno ambers is analyzed for the first time. In surveyed representative collections of Baltic amber, the ratios of Holarctic and tropical ant species are from 1.1 to 1.5; with 10 Holarctic and 9 tropical species (out of 31) in the PIN-964 collection, and 9 and 5 species (out of 29) in the Giecewicz collection; the ratio in the representative collection of Saxonian amber is 0.9, 11 Holarctic species vs. 12 tropical species (out of 55); in the representative collection of Rovno amber it is 0.65, 15 vs. 23 species (out of 79); and in the representative collection of Danish amber it is 0.64, 7 vs. 11 species (out of 36). Hence, in representative collections of Baltic amber, Holarctic species clearly prevail not just in terms of the share of their specimens (by 9.8 to 19.6 times), but also by the number of species. In Bitterfeld amber, Holarctic species are somewhat less numerous than tropical ones, but their specimens are 6 times greater. In representative collections of Rovno and Danish ambers, the number of Holarctic species is 1.5 to 1.7 times smaller than that of tropical species, but the number of their specimens is 4.9 to 6.9 times greater. The numbers of tropical and Holarctic species represented by more than one specimen is similar in Priabonian ambers, 25 versus 22, but Holarctic species include four dominants or subdominants. The abundance of temperate elements in the Priabonian amber ant fauna along with the relatively small number of tropical elements greatly distinguishes it from the Middle European Lutetian ant faunas of Messel and Eckfeld in shale, which do not have temperate elements at all. Formica phaethusa Wheeler, Glaphyromyrmex oligocenicus Wheeler, Plagiolepis squamifera Mayr, Proceratium eocenicum Dlussky, Hypoponera atavia (Mayr), Ponera lobulifera Dlussky, Aphaenogaster mersa Wheeler, and Ennaemerus reticulatus Mayr are new records for Rovno amber, and Formica gustawi Dlussky and Gnamptogenys europaea (Mayr) for Danish amber.
... The ongoing excavations have yielded a multitude of insect fossils that comprise a very diverse fauna (e.g . Lutz 1990;Wedmann 2005;Wedmann et al. 2007Wedmann et al. , 2009Wedmann et al. , 2011McNamara et al. 2011;Dlussky & Wedmann 2012). Odonata are rather scarce, generally preserved as isolated wings, unlike the other insects (Lutz 1987;Wedmann 2005). ...
The study of a new specimen of Petrolestes hendersoni from the Eocene Green Formation allows a more precise description of the enigmatic damselfly and the diagnosis of the Petrolestini. Petrolestes messelensis sp. nov. is described from the Eocene Messel Formation in Germany, extending the distribution of the Petrolestini to the European Eocene. The new damsel-dragonfly family Pseudostenolestidae is described for the new genus and species Pseudostenolestes bechlyi, from the Eocene Messel Formation. It is the first Cenozoic representative of the Mesozoic clade Isophlebioptera.
... The ongoing excavations have yielded a multitude of insect fossils which comprise a very diverse fauna (e. g . Lutz 1990;Wedmann 2005;Wedmann et al. 2007, McNamara et al. 2011, Dlussky & Wedmann 2012. ...
The phylogenetic placement of siricid hymenopterans and especially the relationships among Siricidae are not resolved, and are in need of further investigation. The fossil record helps to illuminate the evolutionary history of this group. In this paper, Xoanon? eocenicus sp. nov. is newly described from the Eocene Fossillagerstätte Grube Messel (Germany). Although several characters of the wing venation of this fossil are similar to the extant genus Xoanon, there remain some doubts concerning the generic placement. A newly found fossil is attributed to Urocerus ligniticus (Piton, 1940) from the Paloecene Fossillagerstätte Menat (France) and provides more morphological information on this species. An overview of the fossil record of Siricidae in the strict sense is given and the state of knowledge on the different fossils is reviewed.
... These genera are known only from a handful of fossils from Dominican amber and dating no older than the Early Miocene (summarized in Table 1), but these records and their surprisingly modern character (rather than belonging to stemgroup Odontomachiti) serve to demonstrate that this clade is considerably older. Indeed, Ponerinae as a whole extends well into the Mesozoic (e.g., Grimaldi et al. 1997; (1994) and Grimaldi and Engel 2005;LaPolla et al. 2013) and has experienced a rich history of continuous diversification throughout the Tertiary (e.g., Dlussky and Wedmann 2012;LaPolla et al. 2013). This impressive diversification resulted in a collection of taxa so divergent from one another that some have at times been placed in distinctive tribes (Ouellette et al. 2006;Fisher and Smith 2008;Santos et al. 2010). ...
Odontomachus paleomyagra sp. nov. is described from the Early Miocene of the Most Basin (Czech Republic) on the basis of a single-winged female, representing one of the rare reports of fossil Odontomachini. The new species is separated easily from other trap-jaw ant species groups by differences in mandibular morphology (without denticles on the inner side) and distributional occurrence. The evolutionary and biogeographic history of the Odontomachini is briefly discussed.
Parasitoid wasps of the family Ichneumonidae are one of the most diverse and species-rich groups of organisms with a worldwide distribution. We here describe seven new ichneumonid fossil species and two new genera from a remarkable insect fossil site, the Eocene Messel Pit in Germany (~47Ma). The unique fossil preservation allows us to place five out of the seven new species unequivocally in extant subfamilies and genera. For the first time, lobed claws which are a clear synapomorphy for the subfamily Pimplinae, are observed in a fossil, making the newly described Scambus fossilobus sp. nov. the oldest unequivocal representative of the group. We also describe a fossil of Labeninae (Trigonator macrocheirus gen. et sp. nov.), an ichneumonid subfamily that was until now believed to be an exclusively Gondwanan element. Furthermore, the newly described Rhyssella vera sp. nov., Xanthopimpla messelensis sp. nov., and X. praeclara sp. nov. provide evidence that these extant genera date back as far as the Early/Middle Eocene. In contrast to the clear placement of most of the newly described species, we were unable to place Polyhelictes bipolarus gen. et sp. nov. and Mesornatus markovici gen. et sp. nov. in an ichneumonid subfamily, mostly due to the high levels of homoplasy found in this group. These findings on the one hand demonstrate the need for a more rigorous approach in the taxonomic placement of fossil ichneumonids, and on the other hand provide more precise minimum ages for several ichneumonid genera and subfamilies.
The fossil record of Australasian Formicidae is extremely sparse. It currently comprises two ants in the subfamilies Ponerinae and Dolichoderinae from Plio/Pleistocene strata in Victoria, Australia, 14 as-yet undescribed ants from Cape York amber, and one ant in the subfamily Amblyoponinae from the early Miocene Foulden Maar in southern New Zealand. Here, we report on a diverse myrmecofauna preserved as compression fossils from Foulden Maar and describe Amblyoponinae gen. et sp. indet.,
Rhytidoponera waipiata
n. sp.,
Rhytidoponera gibsoni
n. sp.,
Myrmecorhynchus novaeseelandiae
n. sp., and
Austroponera schneideri
n. sp. Further isolated wings are designated as Formicidae sp. A, B, and C, the former resembling a member of subfamily Dolichoderinae. Fossils of
Austroponera
and
Myrmecorhynchus
are reported for the first time, whereas
Rhytidoponera waipiata
n. sp. and
R
.
gibsoni
n. sp. are the first Southern Hemisphere fossil records of this genus.
The fossil taxa from Foulden Maar establish the subfamilies Ectatomminae, Formicinae, Ponerinae and, possibly, Dolichoderinae in the Australasian region in the early Miocene and provide evidence that the few native ants in the extant New Zealand fauna are the surviving remnant of taxonomically different, possibly more diverse, warm-temperate to subtropical myrmecofauna.
Problems of the formation of a complex of diagnostic characters that determine taxa of the subfamily Myrmeciinae are discussed. A comparative analysis of morphological data on the recent and extinct Myrmeciinae was performed. The wings of recent and Paleogene representatives of Myrmeciinae have different complexes of diagnostic characters. The wings of extinct Myrmeciinae have intermediate features of venation and demonstrate more primitive states of some characters as compared to those of recent poneromorphs.
Problem is how the complex of diagnostic characters of subfamily forms in evolution: gradually or in one step? Imperfection of the paleontological data, parallel morphological evolution seriously complicates the problem. Problems of the formation and distinguishing of a complex of diagnostic characters that determine taxonomic groups of the subfamily Myrmeciinae are discussed. We attempt to investigate the process using morphological data of recent and extinct Myrmeciinae. At the first stage of the research we examined wing morphology basing on our pervious findings. We demonstrated that recent Myrmeciinae had a diagnostic complex of wing characters, which are distinct from that of extinct myrmeciines and recent poneromorphs. Wings of extinct Myrmeciinae has intermediate features of venation and demonstrate more primitive state of some features than recent poneromorphs. However differences between complexes of diagnostic characters recent and extinct myrmeciines are size of cell 1+2r and position of 1RS, 1M. Poneromorph wings differ from extinct myrmeciines by changes in position of m-cu, rs-m.
Kaulfuss, U., Harris, A.C., Conran J.G. & Lee, D.E., 2014. An early Miocene ant (subfam. Amblyoponinae) from Foulden Maar: the first fossil Hymenoptera from New Zealand. Alcheringa 38, 568–574. ISSN 0311-5518.The ant subfamily Amblyoponinae is presently represented in New Zealand by one endemic species in the cosmopolitan genus Stigmatomma and an introduced Australian species of Amblyopone. The fossil record of the group is restricted to two species of Stigmatomma from late Eocene Baltic Amber. Here, we describe the third fossil record, an Amblyopone-like specimen from the early Miocene of Otago, southern New Zealand, based on a winged male that resembles the extant A. australis Erichson in size, general habitus and characters of wing venation, but also shares features with the African amblyoponine genus Zymmer. This represents the first fossil record of Amblyoponinae from the Southern Hemisphere and the first example of Hymenoptera among the few pre-Quaternary insect fossils known from New Zealand. It suggests a long history of Amblyoponinae in New Zealand and Australia.Uwe Kaulfuss [uwe.kaulfuss@otago.ac.nz] and Daphne E. Lee [daphne.lee@otago.ac.nz], Department of Geology, University of Otago, PO Box 56, Dunedin 9016, New Zealand; Anthony C. Harris [anthony.harris@otagomuseum.govt.nz], Otago Museum, PO Box 6202, Dunedin 9059, New Zealand; John G. Conran [john.conran@adelaide.edu.au], ACEBB & SGC, School of Earth and Environmental Sciences, The University of Adelaide, Benham Bldg, DX 650 312, Adelaide SA 5005, Australia. Received 18.3.2014; revised 15.5.2014; accepted 23.5.2014.
A new fossil ant genus, Bilobomyrma new genus, and two new species are described based on males from the late Eocene Rovno (B. ukrainica n. sp.) and Baltic (B. baltica n. sp.) ambers. We tentatively place this genus in the myrmicine tribe Formicoxenini. Bilobomyrma is characterized by its 13-segmented antennae without an apical club; by the short scape, which is subequal to the length of the first and second funicular segments together; by the shape of the second funicular segment, which is distinctly longer than the any other funicular segment except for the apical one; by the presence of notauli on the scutum; by the absence of spurs on the middle and hind tibiae. At the same time, Bilobomyrma differs from other myrmicine genera by the peculiar shape of its clypeus, having a strongly incised medially, bilobed anterior margin, and its forewing venation: the wings have three closed cells-mcu, 1+2r and 3r; the cell 3r is very short, only twice as long as its width; the distal section of veins RS and M diverge from the cell 1+2r separately.
The ant assemblage of Radoboj (Croatia) described by Heer (1849, 1867) is considered the richest known Miocene assemblage of Europe. However, Heer's data can no longer be used for analysis of the palaeontological history of ants, because they are strongly outdated and require a revision. Such a revision was the purpose of our study. We found in collections of three museums of Austria (Universalmuseum Joanneum in Graz, Geologische Bundesanstalt, and Naturhistoricshes Museum in Wien) a total of 537 compression fossils of ants from Radoboj, 459 of which were identified earlier by Heer. We designated the holotypes, lectotypes and neotypes for 54 of the 62 species described by Heer, and subsequently compared the other specimens to these types. As a result, we have identified 350 specimens to subfamily and 309 specimens to species. We re-described 23 species originally described by Heer (1849, 1867) and two species described by Mayr (1867). One genus and eight species are described as new; 27 species and varieties described by Heer are synonymized. The taxonomic placement of eight species described by Heer remains unclear. As a result of our revision, the known assemblage of Radoboj includes 33 species of 15 genera and five subfamilies. The assemblage of Radoboj is especially similar at the subfamily level to the assemblage of Bembridge, UK (Late Eocene) and Stavropol, RF (Middle Miocene).
The Messel Pit near Darmstadt (Hesse, Germany) is worldwide one of the most important localities for fossils of animals and plants. For this reason the UNESCO declared the Messel Pit a World Heritage Site in 1995; it is the first and until now sole natural heritage in Germany. "Oil shale" has been mined in that pit since the middle of the 19th century. The "oil shale" is a very dark, finely layered clay rock, rich in organic substances; it was deposited during the Paleogene (middle Eocene). Although much is known about this oil shale pit, the genesis of the Messel structure was mostly unknown. Geophysical studies and a cored borehole in autumn 2001 clarified how the basin originated 49 million years ago.
Molecular data have revolutionized our understanding of ant evolution and systematics by making available large quantities of data and providing an independent character set to infer phylogenetic relationships. Although the first ant phylogeny from molecular sequence data was published less than 15 years ago. the field has grown rapidly with implications for all levels of ant biology. Not only has molecular data helped resolve the phylogenetic relationships of many ant groups. but with well-resolved phylogenies inferences about the evolution of morphology. ages of clades. diversification. associations with mutualists. behavior. development. and other areas of research in ant biology have benefited from this active avenue of research. Questions of speciation. population genetics. phylogeography. systematics. biogeo- graphy. and many other active lines of ant research are now routinely addressed with molecular data. With the ease of generating large amounts of sequence data increasing and costs decreasing the next frontier in myrmecology will be to insure that future ant biologists are able to take advantage of this important tool while still being trained in basic ant biology and taxonomy.
. The higher phylogeny of the Formicidae was analysed using 68 characters and 19 taxa: the 14 currently recognized ant subfamilies plus 5 potentially critical infrasubfamilial taxa. The results justified the recognition of 3 additional subfamilies: Aenictogitoninae Ashmead (new status), Apomyrminae Dlussky & Fedoseeva (new status), and Leptanilloidinae Bolton (new subfamily). A second analysis on these better delimited 17 subfamilies resulted in 24 equally most parsimonious trees. All trees showed a basal division of extant Formicidae into two groups, the first containing (Myrmicinae, Pseudomyrmecinae, Nothomyrmeciinae, Myrmeciinae, Formicinae, Dolichoderinae, Aneuretinae) and the second the remaining subfamilies. Clades appearing within these groups included the Cerapachyinae plus ‘army ants’, the Nothomyrmeciinae plus Myrmeciinae, the ‘formicoid’ subfamilies (Aneuretinae + Dolichoderinae + Formicinae), and the Old World army ants (Aenictinae + Aenictogitoninae + Doryline), but relationships within the last two groups were not resolved, and the relative positions of the Apomyrminae, Leptanillinae and Ponerinae remained ambiguous. Moreover, a bootstrap analysis produced a consensus tree in which all branches were represented in proportions much lower than 95%. A reconstruction of the ground plan of the Formicidae indicated that the most specialized of all recent ants are the members of the subfamily Dorylinae and the least specialized ones are the monotypic Apomyrminae.
Described is the first syninclusion of dolichoderine ant Ctenobethylus goepperti (Mayr) (two workers) and scale insect Matsucoccus (male and fe- male) in Rovno amber. РЕЗЮМЕ. Описан первый сининклюз долиходе- рины Ctenobethylus goepperti (Mayr) (двое рабочих) и кокциды Matsucoccus (самец и самка) в ровенс- ком янтаре.
Fossil representatives of the hydrophilid genera Hydrochara Berthold, 1827, Hydrobiomorpha Blackburn, 1888 and Hydrophilus Geoffroy, 1762 were recorded at the lower Middle Eocene locality Grube Messel in Germany. Four morphospecies were recognised, including Hydrobiomorpha eopalpalis, sp. nov. showing sexually dimorphic maxillary palpomere 2 unknown in any recent or fossil species of the genus. These fossils are the oldest known records of the mentioned genera and indicate a minimum age of 47million years for the divergence of the Hydrobiomorpha and Hydrophilus clades. Based on these data, we assume that the diversification of the 'greater hydrophilines' clade predated the lower Middle Eocene. The fossil record of the subtribe Hydrophilina is briefly reviewed, the reasons of the scarcity or absence of some genera in the fossil record are discussed, and the paleoenviromental significance of the presented fossils is discussed.
The Middle Eocene (Lutetian) bee faunas of Eckfeld and Messel, Germany are revised. In addition to the previously known Electrapis electrapoides (Lutz), five additional species are recognized. Four new species of the Electrapini (Apidae: Apinae) are described: Electrapis micheneri Wappler and Engel, E. prolata Engel and Wappler, Protobombus pristinus Wappler and Engel, and P. messelensis Engel and Wappler. In addition, the new genus Pygomelissa Engel and Wappler is proposed for Pygomelissa lutetia Engel and Wappler new species, which cannot presently be classified into any tribe of the Apidae. The tribe Megachilini (Megachilidae: Megachilinae) is also recorded from Eckfeld but in the absence of any body fossil. Megachilines include the leaf-cutter bees (Megachile) and from the occurrence of the distinctive semicircular damage they produce in leaves, we conclude that such bees were also present in the fauna. The bee fauna is compared with that of the contemporaneous Baltic amber. As with Baltic amber, the majority of bee specimens are from the advanced eusocial lineages of the corbiculate Apinae. Lastly, comments are made on the phylogenetic and paleobiological significance of the faunas.
Lace bugs (Heteroptera: Tingidae) comprise over 2.100 species classified in two subfamilies. So far, nearly 36 fossil species of Tingidae have been described, most of Cenozoic age. The present paper reports on three new species genera of Tingidae from the late Middle Eocene Grube Messel (Germany), inscribed on the UNESCO World Heritage List, describing Exmesselensis disspinosus gen. et sp. n. (Canatacaderinae: Phatnomini), Chorotingiotes prisca gen. et sp. n. (Tinginae: Ypsotingini), and Oblongomorpha lutetia gen. et sp. n. (Tinginae: Litadeini). The specimens from Messel show once more that Cantacaderinae, especially of the tribe Phatnomini were widespread and highly diverse in Europe during the Paleogene than today. Oblongomorpha n. gen. and Chorotingioites n. gen. demonstrate that even the differentiation of the Tinginae in the main tribes has taken place in this period or earlier. A taxonomic catalog of all described, fossil lace bugs is appended.
The Late Cretaceous Grassy Lake and Cedar Lake amber deposits of western Canada are among North America’s most famous amber-producing localities. Although it has been suggested for over a century that Cedar Lake amber from western Manitoba may be a secondary deposit having originated from strata in Alberta, this hypothesis has not been tested explicitly using geochemical fingerprinting coupled to comparative analyses of arthropod faunal content. Although there are many amber-containing horizons associated with Cretaceous coals throughout Alberta, most are thermally mature and brittle, thus lacking the resilience to survive long distance transport while preserving intact biotic inclusions. One of the few exceptions is the amber found in situ at Grassy Lake. We present a suite of new analyses from these and other Late Cretaceous ambers from western Canada, including stable isotopes (H and C), Fourier transform infrared (FTIR) spectra, and an updated faunal compendium for the Grassy and Cedar lakes arthropod assemblages. When combined with amber’s physical properties and stratigraphic constraints, the results of these analyses confirm that Cedar Lake amber is derived directly from the Grassy Lake amber deposit or an immediate correlative equivalent. This enables the palaeoenvironmental context of Grassy Lake amber to be extended to the Cedar Lake deposit, making possible a more inclusive survey of Cretaceous arthropod faunas.
The ant fauna of a rain forest in Sabah, Malaysia was sampled by using different collecting methods in three strata. In total, 524 morphospecies of ants could be distinguished. They belong to seven subfamilies and 73 genera. So far, the ant community described is the most species rich published for a primary tropical rain forest. Regarding the stratification in the forest, the leaf litter community comprised as many ant species as the lower vegetation or canopy. Furthermore the litter stratum had the highest generic diversity. The stratification of ants in rain forests seems to be a very strict one with the majority of species (75%) being related to only one stratum. This is in contrast to findings on the stratification of beetles in rain forests. The stratification and a radiation of some groups into vegetation and canopy, where a broad spectrum of permanent habitats exist, is responsible for the high diversity of ants in tropical rain forests.
The formicid genus Gesomyrmex is reviewed and several new species are described from the middle Eocene (about 47 Ma) of Grube Messel, Germany, and from the middle Eocene (about 43 Ma) of Eckfeld maar, Germany. The new taxa are Gesomyrmex curiosus n. sp., Gesomyrmex breviceps n. sp., and Gesomyrmex pulcher n. sp. from Messel, and Gesomyrmex flavescens n. sp., and Gesomyrmex germanicus n. sp. from Eckfeld maar. Two previosly described Oligocene species must be excluded from Gesomyrmex. Former G. expectans Théobald, 1937 is transferred to Eoformica expectans (Théobald, 1937) (comb. nov.), and former G. miegi Théobald, 1937 has to be considered as Formicidae incertae sedis (comb. nov.). A key to the living and fossil reproductive female caste (gyne) of the genus Gesomyrmex is provided. Given the fossil records of Gesomyrmex hoernesi Mayr, 1868 from different European amber deposits the presence of this genus in Europe during the Eocene is well established. Both extant and fossil Gesomyrmex species have an arboreal mode of life. The comparison of arboreal ant faunas from Eocene to Recent times shows that their community structure apparently changed considerably during this period. We infer that Gesomyrmex, along with other genera, was most prosperous during the middle Eocene of Europe, and today has a relict distribution in southern Asia.
S. 2008. New middle Eocene formicid species from Germany and the evolu− tion of weaver ants. Acta Palaeontologica Polonica 53 (4): 615–626. Two new species of weaver ant are described from the Eocene of Germany. Males and gynes of Oecophylla longiceps Dlussky sp. nov. are found in the middle Eocene (about 47 Ma) of Grube Messel, Germany. Males, gynes and two work− ers of O. eckfeldiana Dlussky sp. nov. are recorded from the middle Eocene (about 43 Ma) of Eckfeld maar, Germany. The two new species are among the oldest records of the extant genus Oecophylla. First adaptations for moving in the for− est canopy are present in the workers of O. eckfeldiana. Even more specialized adaptations for arboreal life are found in the workers of O. brischkei from Baltic amber. The coexistence of two species in Baltic amber and in the Bembridge marls suggests that in these cases different ecological niches were realised by sympatric species. Comparisons of the queens from different fossil and extant species indicate morphological trends, partly connected with increasing fertility. Most likely Oecophylla originated in the early Paleogene in the Palaearctic realm, radiating strongly during the climatic changes of the Eocene–Oligocene transition. Gennady M. Dlussky [dlusskye@mail.ru], Moskovskij gosudarstvennyj universitet imeni M.V. Lomonosova; Biolo− gicheskij fakul'tet Vorobjovy gory, 119992, Moskva, Russia; Torsten Wappler [twappler@uni−bonn.
The formicid genus Gesomyrmex is reviewed and several new species are described from the middle Eocene (about 47 Ma) of Grube Messel, Germany, and from the middle Eocene (about 43 Ma) of Eckfeld maar, Germany. The new taxa are Gesomyrmex curiosus n. sp., Gesomyrmex breviceps n. sp., and Gesomyrmex pulcher n. sp. from Messel, and Gesomyrmex flavescens n. sp., and Gesomyrmex germanicus n. sp. from Eckfeld maar. Two previosly described Oligocene species must be excluded from Gesomyrmex. Former G. expectans Théobald, 1937 is transferred to Eoformica expectans (Théobald, 1937) (comb. nov.), and former G. miegi Théobald, 1937 has to be considered as Formicidae incertae sedis (comb. nov.). A key to the living and fossil reproductive female caste (gyne) of the genus Gesomyrmex is provided. Given the fossil records of Gesomyrmex hoernesi Mayr, 1868 from different European amber deposits the presence of this genus in Europe during the Eocene is well established. Both extant and fossil Gesomyrmex species have an arboreal mode of life. The comparison of arboreal ant faunas from Eocene to Recent times shows that their community structure apparently changed considerably during this period. We infer that Gesomyrmex, along with other genera, was most prosperous during the middle Eocene of Europe, and today has a relict distribution in southern Asia.
A key to workers of all the genera of ants known from Baltic amber is presented. For each of these genera data on the modern geographical distribution, the numbers of extant and extinct species, a list of species described from Baltic amber, and the main synonyms are given. A monotypic genus Ctenobethylus Brues for Iridomyrmex goepperti (Mayr) is resurrected from the synonymy.
Fallomyrma, a new monotypic ant genus from the Rovno, Saxonian, and Danish ambers (Late Eocene), and a new species, F. transversa, are described. The taxonomic position and morphological similarity of the new genus to other genera is discussed.
Although Messel Pit, as a UNESCO World Natural Heritage Site, is one of the most important fossil deposits in the world, the genesis of the deposit was disputed for a very long time. Only recently, in the year 2001, a research well in combination with geophysical and geological investigations allowed to clearly prove the origin of the deposit. The drilling project "Forschungsbohrung Messel 2001" was carried out in cooperation between the Senckenberg Research Institute (Forschungsinstitut Senckenberg, Frankfurt a.M.), the Leibniz Institute for Applied Geosciences (Institut für Geowissenschaftliche Gemeinschaftsaufgaben, Hannover) and the Hessian Agency for Environment and Geology (Hessisches Landesamt für Umwelt und Geologie, Wiesbaden). The Messel 2001 well reached a depth of 433 m beneath the floor of the pit. A volcano-lacustrine sequence was found in the well, in which the deposits of the Eocene lake go down to a depth of about 240 m. From depths down to 373 m a tuff was drilled that consists of basaltic, vesicle-poor globular lapilli. Further below to the bottom of the well the tuff is underlain by a breccia. This lithology confirmed that the lake basin originated from phreatomagmatic eruptions about 50 million years before present and did not, as in previously favoured theories, form in a tectonic basin (graben). The sequence found in the well - albeit having some individual features correlates well with the maar lithozones introduced by Pirrung (1998). Also in 2001, the inclinometer well 28 was drilled for engineering purposes at the slope of the pit, about 50 m south of the northern basin margin. The sequence observed in the well of 94 m depth consists of 30 m thick sands and bituminous shales, underlain by breccias containing fragments of the surrounding wall rock as well as basaltic lapilli. The volcano-sedimentary fill of the Messel basin was not drilled through. This points to a steep dip angle (> 60°) of the basin margin. It is very likely that the oilshale deposit of Messel has never been overlain by a significant post-depositional cover, and neither have the deposits experienced a significant post-Eocene erosion, in contrast to those found in the Odenwald further south. Therefore the pre-eruptive surface is supposed to have lain only about 50-100 m above the present surface. This is why most of the sediments filling the former maar lake of Messel have been preserved.
The ants from Burmese amber were not known before. Three ants from this amber were found out in the collection of the Natural History of Museum in London. Burmomyrma rossi sp. nov. (fig. 3) differs from other known ants by the combination of reduced wing venation (absence of closed cells), the presence of sting, one segmented waist and absence of constriction between 1 and 2 gastral segments. The holotype of B. rossi has no head, so it is impossible to determine precisely taxonomic position of new genus. Haidomyrmex cerberus sp. nov. (fig. 1) is the extraordinary specialised Sphecomyrminae ant with L-shaped mandibles and unusual structure of cranio-mandibular system. All known ants with trap-like mandibles have increased back part of the head, while H. cerberus have extended genae. The fragments of other ant (gaster, legs and deformed petiolus and propodeum) are present in the same piece of amber (fig. 26). The poor preservation state of specimen does not permit to make species description. The presence of sting and absence of constriction between 1 and 2 gastral segments in this ant and in B. rossi definitely indicates that they belongs to subfamilies Aneuretinae, Nothomyrmeciinae, or Sphecomyrminae.
The Messel Pit Fossil Site (a UNESCO World Heritage Site) is located within the Sprendlingen Horst (SW Germany), a basement unit forming the northeastern shoulder of the Upper Rhine graben. The Messel Pit is interpreted as an isolated lake sediment occurrence created by phreatomagmatic activity. 40Ar/39Ar dating with stepwise heating via laser on a redeposited basalt fragment within a lapilli tuff recovered by the Messel research drilling of 2001 yields a plateau age of 47.8 ± 0.2 Ma (Middle Eocene). This age correlates with a major phase of volcanic activity in the Sprendlingen Horst during the short time span from 49 to 47 Ma. The 40Ar/39Ar date enables a numerical calibration of the Messel lake sedimentation. The contemporaneously deposited taphocoenosis corresponds to the mammalian biostratigraphic reference level Mammal Paleogene MP 11. For phreatomagmatic activity associated with the maar crater collapse and contemporaneous onset of lake sedimentation, an age of 48 Ma might be derived. Frequency analyses along with sedimentological criteria point to a subsequent phase of sedimentation lasting approximately 1 Ma. The age of the fossils excavated thus far from the uppermost part of the Messel sediments could thus be 47 Ma. Correspondingly, an estimate for the MP 11 time interval is 47.5 to 46.5 Ma. Since MP 11 represents the base of European Land Mammal Age (ELMA) Geiseltalian, the boundary with the older ELMA Grauvian can be estimated at 47.5 Ma.
A new genus and two new species, Eotapinoma macalpini sp. nov. (Dolichoderinae) and Canapone dentata gen. et sp. nov. (Ponerinae) from Canadian amber (Upper Cretaceous, Campanian; Medicine Hat, Alberta, Canada) are described.
A new fossil ant genus, Bilobomyrma new genus, and two new species are described based on males from the late Eocene Rovno (B. ukrainica n. sp.) and Baltic (B. baltica n. sp.) ambers. We tentatively place this genus in the myrmicine tribe Formicoxenini. Bilobomyrma is characterized by its 13-segmented antennae without an apical club; by the short scape, which is subequal to the length of the first and second funicular segments together; by the shape of the second funicular segment, which is distinctly longer than the any other funicular segment except for the apical one; by the presence of notauli on the scutum; by the absence of spurs on the middle and hind tibiae. At the same time, Bilobomyrma differs from other myrmicine genera by the peculiar shape of its clypeus, having a strongly incised medially, bilobed anterior margin, and its forewing venation: the wings have three closed cells-mcu, 1+2r and 3r; the cell 3r is very short, only twice as long as its width; the distal section of veins RS and M diverge from the cell 1+2r separately.
An annotated catalogue of the invertebrate animals from the Eocene deposit Grube Messel is provided here. Members of the following taxa are recorded: Dinoflagellata, Porifera, Gastropoda, Crustacea, Chelicerata, and Insecta. Several taxa are recorded for the first time, including Sternorrhyncha, Thysanoptera, Neuropterida, and Mecoptera. Both fossils and ichnofossils are listed.
The importance of litter in the total energy flow dynamics of a central Amazonian rain forest near Manaus, Brazil, is discussed. The study area is located in the hinterland of Manaus between the Rio Negro and the Amazon. Its substrate is Tertiary sediment. The area receives 1771 mm rainfall per year, and the soil is classified as yellow latosol. The forest comprises 93,780 dicotyledonous trees and palms per hectare reaching 38.10 meters in height. Over 500 species of palms and dicotyledonous trees above 1.5 m. in height are identified for a 2000 sq. m. plot. The estimate for fresh living dicotyledonous tree and palm biomass is 939.5 metric tons per hectare consisting of 1.9% leaves, 49.7% stems, 21.3% branches and twigs, and 27.1% roots. Lianas, vascular epiphytes, and parasites are estimated to comprise 46.2 mt/hectare in the fresh state. At the soil surface there are 59 mt/hectare of fresh litter. Living animal biomass is about 200 kg/hectare of which half is soil fauna. The high proportion of soil fauna, the type of humus, the decomposition of litter, the apparent dependence of soil fauna on fungi, and the low nutrient content of litter are all factors which strongly support a consumer food chain based almost entirely on dead organic matter. The fungi play a decisive role in concentrating the otherwise limited nutrient resources.
The librational modes of the water molecules in the isomorphic MX2·Y2O (M=Sr, Ba; X=Br, I; Y=H, D) are assigned to the wagging (Rγ), twisting (Rt), and rocking vibrations (Rr) by means of single crystal Raman technique. The order of the librations in Rγ > Rt > Rr in energy and Rt > Rr in intesity (Rγ not observable), as predicted for tetrahedrally coordinated water molecules in solids. The data obtained are discussed with respect to (i) the strength of the hydrogen bonds, (ii) repulsive effects of the lattice (owing to the different volume increments of the H2O molecules), and (iii) metal oxygen interactions (shown by the different enthalpies of hydration). The H2O twisting vibrations are only little affected by the hydrogen bonds present (opposite to the wagging and rocking modes), but very sensitive to both repulsive effects and metal oxygen interaction.
From collections of ants made by myself in the West Indies and Northern South America, incidental to studies on the biology of fungus-growing ants (Attini),1 the following remarkable new cerapachyines and ponerines are selected for description.2 All types, except that of Thaumatomyrmex manni, are in the author's collection. T. manni is in the U. S. National Museum.
The first fossil records of Holarctic representatives of the genus ComptosiaMacquart, 1840 from the middle Eocene Messel Pit, Germany, and the upper Eocene of Florissant, USA, are reported. The fossil from Messel, Comptosia pria sp. nov., is represented by a well-preserved wing, displaying characteristic wing venation, remains of the second wing and elements of the thorax. The Florissant fossil, C. miranda comb. nov., is preserved almost completely with both wings and most parts of the body visible. Extant members of the genus Comptosia are currently found only in Australia, and close relatives are known from southern South America. Previously this group of genera has been thought to be of Gondwanan origin, but these Eocene fossil representatives of Comptosia from the Holarctic region argue against this interpretation, and suggest that Comptosia and its relatives were more widespread in both Northern and Southern hemispheres during the Palaeogene. The fossil record of bee flies suggests a major radiation of bombyliids in the late Cretaceous or early Cenozoic. This assumption is supported by the comparatively young geological age of many bombyliid host taxa. A key factor driving the specialization of many bombyliid larvae on ground-dwelling hosts may have been avoidance of competition with tachinid flies and parasitic hymenopterans.