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Here we report fossil ants, including a new genus of Ponerinae, about 50 million years (Myr) older than the previous oldest specimens. These discoveries in amber from the Turonian stage (92 Myr ago) of New Jersey in the United States have important implications for estimates dating the origin of ants, and extend the age of an extant ant subfamily back about 50 Myr.
Nature © Macmillan Publishers Ltd 1998
Here we report fossil ants, including a
new genus of Ponerinae, about 50 mil-
lion years (Myr) older than the previous old-
est specimens. These discoveries in amber
from the Turonian stage (92 Myr ago) of
New Jersey in the United States have impor-
tant implications for estimates dating the
origin of ants, and extend the age of an
extant ant subfamily back about 50 Myr.
Until now, a specimen of Sphecomyrma
freyi in a piece of amber from Cliffwood
Beach, New Jersey, was one of the few non-
compression fossil ants from the Cretaceous
period showing most morphological details1.
The metapleural gland is the only morpho-
logical trait unique within the Hymenoptera
that distinguishes ants, and can be seen in
most ants embedded in amber. This gland
produces antibiotic-like substances2, neces-
sary to maintain nests underground or in
humid pieces of wood, where bacteria and
fungi would otherwise invade immobile
broods. The gland is seen in modern species,
opening above the hind coxae. Development
of the gland and eusociality were probably
correlated and are involved in the great eco-
logical success of the ants. In an Amazonian
rainforest, for example, ants make up more
than 25% of the total animal biomass3.
Wilson et al.1described an almost per-
fectly preserved worker of S. freyi, but the
debate remained whether this fossil is an ant
because the presence of the metapleural
gland was uncertain. Sphecomyrma was thus
excluded from a recent phylogenetic analysis
of the Formicidae4, and its relatively short
first antennal segment in workers was
assumed to prevent the basic social behav-
iour of trophallaxis5.
Our new specimens include three worker
and four male ants. One complete, well-
preserved worker was the second discovery
of S. freyi. Among other well-preserved traits
in this specimen are the external and even
part of the internal anatomy of the meta-
pleural gland. The gland has a circular open-
ing and a very large subcuticular atrium (Fig.
1a). Sphecomyrma is thus an ant.
Two of the males are a new species of the
Cretaceous genus Baikuris, known previous-
ly only from Upper Cretaceous (Santonian)
amber from Taymyr, northern Siberia. The
third male belongs to a genus as yet undeter-
mined. The fourth male is tentatively
assigned to Sphecomyrma and would be the
first known male of that genus.
Another worker represents a new genus,
distinguished by its clubbed antennae, pro-
portions of the antennal articles, thin man-
dibles that lack teeth and cross extensively
when closed, and a girdling constriction of
the gaster typical of the large modern sub-
family, the Ponerinae (Fig. 1b). Furthermore,
the broad attachment of the third and fourth
abdominal segments, and the apical denticles
on the genae, indicate a relationship with
ants of the ponerine tribe Amblyoponini.
Addition of these two new taxa confirms
the basal position of Sphecomyrma6, but as
part of a quadritomy. The position of the
new ponerine within one clade of the Poner-
inae demonstrates that one major lineage of
extant ants was established well before the
formation of amber from Sakhalin Island
(probably Palaeocene)7and of Eocene Baltic
amber8, which are about 50 Myr younger
than the New Jersey amber.
A reasonable estimate would thus place
the origin of the ants into the lowermost
Cretaceous (about 130 Myr ago), but proba-
bly no older. This conclusion is consistent
with the relationship of the Formicidae to
the wasp families Vespidae and Scoliidae9
and the phylogenetic position of the Creta-
ceous Vespidae10. On this basis, a hypothesis
of a Lower Jurassic origin of ants11, based on
estimated divergence time of mitochondrial
cytochrome bsequences, is unlikely.
Although ants originated in the Creta-
ceous, it was not until the Tertiary7period
that they became so dominant and diverse in
terrestrial ecosystems, as documented in
Baltic8and Dominican amber12. We are still
examining why the radiations were delayed
until the Tertiary.
Donat Agosti, David Grimaldi,
James M. Carpenter
Department of Entomology, American Museum of
Natural History, Central Park West at 79th Street,
New York, New York 10024-5192, USA
1. Wilson, E. O., Carpenter, F. M. & Brown, W. L. Jr. Psyche 74, 1–19
2. Beattie, A. J., Turnbull, C. L., Hough, T. & Knox, R. B. Ann.
Entomol. Soc. Am. 79, 448–450.
3. Fittkau, E. J. & Klinge, H. Biotropica 5, 2–14 (1973).
4. Baroni Urbani, C., Bolton, B. & Ward, P. S. Syst. Entomol. 17,
301–329 (1992).
5. Dlussky, G. M. Paleont. J. 1983 (3), 65–78 (1983).
6. Grimaldi, D. et al. J.M. Am. Mus. Novit. (in the press).
7. Dlussky, G. M. Paleont. J. 1988 (1), 50–61 (1988).
8. Wheeler, W. M. Schr. Phys.-Ökon. Ges. Königsb. 55, 1–142 (1915).
9. Brothers, D. J. & Carpenter, J. M. J. Hymenopt. Res. 2, 227–304
10. Carpenter, J. M. & Rasnitsyn, A. P. Psyche 97, 1–20 (1990).
11. Crozier, R., Jermiin, L. S. & Chiotis, M. Naturwissenschaften 84,
22–23 (1997).
12. Hölldobler, B. & Wilson, E. O. The Ants(Harvard Univ. Press,
Boston, 1990).
VOL 391
29 JANUARY 1997 447
scientific correspondence
Posterior end
0.5 mm
Figure 1 Photomicrographs of New Jersey ant
fossils. a, Alitrunk and portion of the gaster of
the new
S. freyi
worker with a drawing showing
the position and structure of its metapleural
gland (MGO). b, Lateral view of the new Creta-
ceous ponerine genus (m, mandible in frontal
view). A detailed description will be provided
Oldest known ant fossils discovered
tionship where a 1 °C fall in diurnal temper-
ature range increased Australian wheat yield
by 0.52 t ha11. This effect, taken with the
trend in diurnal range, accounted for 45% of
the yield increase between 1952 and 1992. In
an extended model with three climate vari-
ables, Nicholls found that changes in mini-
mum temperature had had most impact on
wheat yield and that rainfall change had con-
tributed little. In our view, Nicholls’s results
need qualification and should be interpreted
with caution. They are only estimates, and
do not include standard errors to indicate
their precision.
The analytical challenge is to separate the
effects of climate change from other factors
affecting Australian wheat yields. Non-
climate factors include total wheat area, the
Climate change and
Australian wheat yield
Nicholls1reported that 30–50% of the
increase in Australian wheat yields in the
period 1952–92 resulted from climate
change. He estimated a simple linear rela-
... One of the oldest unequivocal ant fossils is Sphecomyrma freyi (Fig. 3C) in New Jersey amber (dating 92 mya) (Wilson et al., 1967a). This was the first discovery of a Cretaceous fossil that undoubtedly had a metapleural gland (Agosti, Grimaldi, & Carpenter, 1998). The primitive characteristics of S. freyi, in addition to its remarkable combination of ant-like and wasp-like features, led to the establishment of the Sphecomyrminae subfamily (Wilson et al., 1967a;Wilson, Carpenter, & Brown, 1967b), considered by some myrmecologists to be the most basal subfamily in Formicidae and the closest relative to all extant ants (Wilson et al., 1967a). ...
... Following the discovery of the Cretaceous ant fossil S. freyi, further support for the early origin of wing polymorphism in ants came from the discovery of wingless females in numerous other Cretaceous fossils (Agosti et al., 1998;Barden & Grimaldi, 2014;Barden & Grimaldi, 2016;Wilson et al., 1967a). Wilson's (1987) analysis of various wingless ant fossils showed that they morphologically group with modern ants and hence are part of a worker caste as opposed to their grouping with solitary wingless female wasps (see Section 2). ...
... The metapleural gland, a defining feature of ants that is present in all early ant fossils (Agosti et al., 1998;Barden, 2016), may provide insight into the ancestral habitat as well as the relationship between wing polyphenism evolution and eusociality. In extant ants, the gland has been shown to secrete antibiotic substances essential to overcoming the pressures from microbes and parasite transmission associated with living in colonies as well as nesting under the soil or in decomposing leaf-litter or rotting wood (Boomsma, Schmid-Hempel, & Hughes, 2005;Holldobler & Wilson, 1990;Yek & Mueller, 2011). ...
The evolution of eusociality, where solitary individuals integrate into a single colony, is a major transition in individuality. In ants, the origin of eusociality coincided with the origin of a wing polyphenism approximately 160 million years ago, giving rise to colonies with winged queens and wingless workers. As a consequence, both eusociality and wing polyphenism are nearly universal features of all ants. Here, we synthesize fossil, ecological, developmental, and evolutionary data in an attempt to understand the factors that contributed to the origin of wing polyphenism in ants. We propose multiple models and hypotheses to explain how wing polyphenism is orchestrated at multiple levels, from environmental cues to gene networks. Furthermore, we argue that the origin of wing polyphenism enabled the subsequent evolution of morphological diversity across the ants. We finally conclude by outlining several outstanding questions for future work.
... erstmals weitere gesetzmäßige Besonderheiten im Trichterbau belegen: Nach dem Neubau eines Trichters (1) folgen tagesperiodisch (23:20±02:11 Std., n=96) Nachbauphasen, in denen im Erweiterungsbau (2) (Moreau & al. 2006). Der Beginn ihrer Entwicklung wird auf 130 Mio Jahre angesetzt (Agosti & al. 2008 (Yoshida & al. 2001), die Stabilität der Körpereiweiße bei Hochtemperaturen und die Aktivität der Verdauungsenzyme bei einem Temperaturoptimum von 59 °C (Bongers & Koch 1984). In der Namib-Wüste ergaben eigene Temperaturmessungen in den Trichtern mittags 12:00 Uhr Werte bis 62,8°C. ...
Antlion larvae and ants: Dimensions of predator-prey-relation in NorthEast-Germany (Euroleon nostras Fourcroy, Myrmeleontidae, Neuroptera) Six species of the genetic group Euroleon are widespread distributed form Korea to Spain. Reports about larvae of E. nostras of Northern Europe are scant; especially, it concerns reports about ecological conditions, prey profit in habitats, stipulation of biomass as well as survival strategies. Moreover, studies about Life history of larvae under intermitted access to prey will be under demand. This study deals with population, prey spectrum, prey diversity, pit design of antlions in habitats of southern Baltic Sea shores. It includes also abiotic characteristics as soil texture and water balance. Recent reports about evolution of antlion lacewings and simultaneous occurrence of Formicidae suggest existence of predator-prey relationships between them for around 100 million years. Earliest evidence appears in Baltic as well as Myanmar amber with ages of 50 and 100 million years, respectively. Populations were studied in sandy habitats (fine up to coarse sand). Sandy habitats allow for oxygen and also water exchange of larvae with surroundings. Dry sand comprises only 1 % of water. Therefore, antlions are dependent on rain and dew for stabilizing their fluid balance. On the other side, larvae dig pits for trapping of prey. It fails hugely when rain solidifies the sand design and living conditions for a population might become complicated. However, larvae are versatile in stabilizing water balance. Prey serves as prime source for fluid. Ants of species Formica, Porcellio (Isopoda), and Glomeris (Glomerida) comprise 76, 69 and 64 % of water, respectively. In contrast, water balance of antlion larvae is less stable it varies between 68 and 73 % of body mass. Ants, as prey, are not permanently available. Ants constitute around 60 to 80 % of prey in open-situated habitats. In forest-habitats, their portion shrinks to 5 %. Prey comes then chiefly from small animals of forest floor.-In summary, ambush predators live risky because they meet food unpredictably. Therefore, larvae must be equipped with low metabolic rates so that they survive for a long time without food. Let us have a quick glance to the ambush trap. The larva marks its site by a circular channel. Next, it crawls backwards in sand along the circular-spiral orbit. It pulls by aid of one front leg permanently the sand from inside out. The pit gets deeper and deeper when the larva moves along the spiral orbit. In fixing the open space and the depth of the pit it meets automatically the angle of repose of the sand. It has been found that larvae dig their pits in around 0.5 h. This is a very short time for constructing a pitfall trap in relation to the time at resting metabolic rate. This period of time does not depend on the larvae stage. The metabolism during this action exceeds the resting one by a factor of roughly nine (Example: L3, 22 mg; 5.3 µgC / h to 0.6 µgC / h). Therefore, the costs of the pit construction (volume, mass weight, energy) are of minor importance in relation to prey. The linear accession of pit traps diameter (benefit of prey catch) exponentially corresponds to ascent of costs. We collected data about transport of sand and time of design as well as related data by computer-aided data processing. The development cycle from ovi-positioning to pupa lasts for two years in NorthEast Germany. Interestingly, all three larvae stages exist in populations during certain periods, but with widely differing biomasses from 5 to 150 mg. This is chiefly caused by varying access to nourishment. * Meiner Frau Brigitte Meißner, geb. Schönermarck (15.07.1938-23.11.2019) gewidmet! Mitt. Dtsch. Ges. allg. angew. Ent. 22 Halle (Saale) 2020 52 We found in prey-poor habitats time restricted food access with waiting periods up to 24 days. It is interesting to note that single Formica may satisfy the resting metabolism over this very long period. Thus, difference in biomasses reflects prey-rich and prey-poor habitats. Antlions suck effectively their prey independently of their own mass. It has been observed, they gain from ants and larvae of Tenebrio 35 and 43 %, respectively. Lean larvae (30 mg) grow by 170 % of their mass whereas more heavy larvae (100 mg) bring it only to 40 % plus. This regime is important for their development towards pupa.
... The first unambiguous ant fossils date from the Lower Cretaceous [95,96], and molecular clock analyses have dated the ant crown group to 120 -160 million years ago (Ma) [97 -99]. Attine fungiculture is 50-60 million years old [100,101] and has a single evolutionary origin, but leafcutter ants appear to have evolved only 8-12 Ma [100]. ...
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This special feature results from the symposium 'Ants 2016: ant interactions with their biotic environments' held in Munich in May 2016 and deals with the interactions between ants and other insects, plants, microbes and fungi, studied at micro-and macroevolutionary levels with a wide range of approaches, from field ecology to next-generation sequencing, chemical ecology and molecular genetics. In this paper, we review key aspects of these biotic interactions to provide background information for the papers of this special feature. After listing the major types of biotic interactions that ants engage in, we present a brief overview of ant/ant communication, ant/plant interactions, ant/fungus symbioses, and recent insights about ants and their endosymbionts. Using a large molecular clock-dated Formi-cidae phylogeny, we map the evolutionary origins of different ant clades' interactions with plants, fungi and hemiptera. Ants' biotic interactions provide ideal systems to address fundamental ecological and evolutionary questions about mutualism, coevolution, adaptation and animal communication.
... The fragment of amber in which the fly is preserved is one of dozens such fragments, all from a large piece (AMNH NJ-90) that was found splintered. That piece yielded a remarkable array of inclusions, including the oldest fossil mushrooms (Hibbett et al., 1995;1997), several male ants (Grimaldi et al., 1997;Agosti et al., 1998), and many others. Both pieces with the flies were embedded in epoxy and trimmed. ...
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Sixty-five specimens representing 49 species in 37 genera and 12, possibly 13, families of brachycerous Diptera are described in-detail. Some genera are family incertae sedis. They are preserved in Cretaceous ambers from the following areas and ages (abbreviations after each are used to designate the following origins of the ambers): Manitoba and Alberta, Canada (C) (Campanian); central, New Jersey (NJ) (Turonian); and Lebanon (L) (Neocomian). All taxa described are new species and most genera are described as new, except where noted. The new taxa and their origins are the following: Tethepomyia thauma (NJ), an extremely apomorphic fly of probable nematocerous affinities. In RHAGIONIDAE: Paleochrysophilus hirsutus (L), Jersambromyia borodini (NJ), Mesobolbomyia acrai (L); and four additional genera (3 L, 1 NJ) that are described and illustrated but not named because of incomplete preservation. STRATIOMYIDAE: it new specimen of Cretaceogaster pygmaeus Teskey (C) is reported, showing newly observed structures that confirm its extremely primitive position in the family; in addition, in NJ amber an additional primitive genus is described but not named, with affinities in the Pachygastrinae, Chiromyzinae, or Beridinae. HILARIMORPHIDAE: Hilarimorphites superba, H. yeatesi, and H. longimedia, all in NJ amber, and the only fossil hilarimorphids. SCENOPINIDAE(?): Proratites simplex (NJ), probably a primitive (proratine) scenopinid, which would be the only Mesozoic fossil of the family. ASILIDAE: an incomplete, unnamed specimen in NJ amber, which is one of only two Cretaceous records, The most diverse and numerous brachycerans in Cretaceous ambers-are in the EMPIDOIDEA, With new taxa as follows. EMPIDINAE: Turonempis styx (NJ), Emplita casei (NJ). ATELESTINAE: Atelestites senectus (L). NEMEDINA GENUS GROUP: Cretodromia glaesa (C); Nemedromia campania (C), N, telescopica (C), N. turonia (NJ); Neoturanius asymmetrus (NJ), N. cretatus (NJ), and N, vetus (NJ, possibly also C); Phaetempis lebanensis (L), which is possibly a very plesiomorphic member of this group. The Nemedina group today is represented by a single extant species from Hungary. TACNYDROMIINAE: Cretoplatypalpus americanus (C), with Cretoplatypalpus Kovalev previously known from a species in Cenomanian amber from northern Siberia; and Mesoplatypalpus carpenteri (C). TRICHOPEZINAE: Apalocnemis canadambris (C), which is the only species studied here belonging to an extant genus, Apalocnemis Philippi (previously known only from extant species widespread in distribution). MICROPHORINAE: Microphorites similis and M. oculeus (L), two additional species of the extinct genus Microphorites Hennig, known only from Lebanese amber; Avenaphora hispida (L); Cretomicrophorus novemundus (NJ), the second species in the extinct genus Cretomicrophorus Negrobov, originally known from Cretaceous amber of Siberia; Archichrysotus incompletus (NJ) and A. manitobus (C), the genus also previously known from Siberian amber. DOLICHOPODIDAE: Sympycnites primaevus (L), which is the oldest definitive dolichopodid. Three new species are described in an unusual new genus, Chimeromyia, known only from Lebanese amber: C. intriguea, C. acuta, and C. reducta. Chimeromyia possesses features of Empidoidea and Cyclorrhapha. The few Cyclorrhapha in Cretaceous ambers are all very plesiomorphic. PLATYPEZIDAE: Electrosania cretica (NJ), the most plesiomorphic known platypezid. Lebambromyia acrai (L), formally unplaced to family, is a plesiomorphic phoroid closely resembling IRONOMYIIDAE (with one living species in Australia and Tasmania, and one extinct species previously described in Canadian amber). LONCHOPTERIDAE: Lonchopterites prisca (L) and Lonchopteromorpha asetocella (L), the only definitive fossils of this small, extant family, SCIADOCERIDAE: Archiphora pria (NJ); and Archisciada lebanensis (L), the oldest fossil of the family and perhaps the most plesiomorphic phoroid. In addition, two new species are described in the Mesozoic genus Prioriphora McAlpine and Martin, P. luzzii and P. casei (both NJ). This is the best represented brachyceran genus in the Cretaceous, although it might be a paraphyletic taxon. Three cyclorrhaphan larvae of uncertain family identities are described, all in NJ amber; one appears similar to Sciadoceridae. Phylogenetic significance of most of these fossils are discussed, as are certain characters of traditional importance in the higher classification of Brachycera, such as the number of aristal articles. The fossils are placed onto cladograms of the lower Brachycera, the Empidoidea, and basal Cyclorrhapha, and a chronology is proposed of the origins of brachyceran families. The Brachycera apparently originated in the Lower Jurassic, with the Asiloidea not diversifying until the Lower Cretaceous. The Eremoneura (Empidoidea + Cyclorrhapha), as expected, show later diversification, with subfamily-level radiations of empidoids in the Lower to mid-Cretaceous, and the most plesiomorphic families of Cyclorrhapha (e.g., Platypezoidea, Phoroidea, Lonchopteridae) appearing in the Lower to mid-Cretaceous. Origins and radiations of the Schizophora almost certainly are of more recent origin, in the mid to latest Cretaceous and especially the Cenozoic. The diversity and detailed preservation of these fossils contribute exceptional insight into the early evolution of the Brachycera and the Eremoneura in particular.
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In this study, somatic chromosome numbers and 2C nuclear DNA contents of some Polygala species which are Polygala supina, P. anatolica, P. papilionacea, P. inexpectata, P. hohenackeriana, P. vulgaris, P. alpestris, P. peshmenii, P. turcica and P. monspeliaca taxa were investigated. The taxon with the lowest chromosome number in terms of chromosome numbers of the specimens was determined as P. alpestris (2n= 24), and the taxon with the highest chromosome number was determined as P. vulgaris (2n= 68, 2n= 70). According to chromosome count studies, tetraploidy with 2n=4x=68 chromosome number was detected in P. vulgaris species. Chromosome numbers of P. papilionacea (2n=42), P. inexpectata (2n=34), P. peshmenii (2n=34) and P. turcica (2n=38) species were determined for the first time. P. alpestris, P. papilionacea, P. anatolica and P. vulgaris taxa differed from all other taxa in terms of somatic chromosome numbers. Flow cytometric analysis (or nuclear DNA content analysis) was carried out on 24 different populations and 66 different individuals for these species. Based on the results of flow cytometric analysis the lowest 2C nuclear DNA content (0.50 pg) was observed in P. monspeliaca while the highest 2C nuclear DNA content (2.42 pg) was observed in P. alpestris. P. monspeliaca was easily distinguished from other taxa as it had the most different nuclear DNA content. In this study, both the chromosome count in 3 Polygala species and the 2C nuclear DNA content in 7 Polygala species were determined for the first time and it was determined that some of the Polygala taxa we studied were different from each other in terms of these values.
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With a long history of more than 100 million years and about 14,000 described living species, ants are one of the most important and well-known groups of insects in the world. Ants are key elements in the structure and dynamics of terrestrial systems, especially in the tropics, as well as models in studies of evolution, ecology, and monitoring of disturbed ecosystems. For all this, it is essential to know their history and phylogeny, in order among other things, to have a solid base for their systematics and taxonomy. Within Aculeata ants appear to be the sister group of Apoidea, and the ancestor of the Formicidae may have inhabited the northern hemisphere during the early Cretaceous. Since then, and especially since the Eocene, ants have spread throughout the planet, occupying almost all environments from forests to deserts. The Neotropical region has 137 genera and around 3,100 species of ants. The Neotropics seem to have been the “cradle” and “museum” of the groups of ants, which would explain their great diversity and a high degree of endemisms. This review describes the current state of knowledge of ants in the Neotropical region from a systematic point of view, with a synopsis of all supraspecific taxa described to date. Critical genera, problems to be solved, and perspectives for the study of these insects are also presented.
The recent discovery of diverse fossil flowers and floral organs in Cretaceous strata has revealed astonishing details about the structural and systematic diversity of early angiosperms. Exploring the rich fossil record that has accumulated over the last three decades, this is a unique study of the evolutionary history of flowering plants from their earliest phases in obscurity to their dominance in modern vegetation. The discussion provides comprehensive biological and geological background information, before moving on to summarise the fossil record in detail. Including previously unpublished results based on research into Early and Late Cretaceous fossil floras from Europe and North America, the authors draw on direct palaeontological evidence of the pattern of angiosperm evolution through time. Synthesising palaeobotanical data with information from living plants, this unique book explores the latest research in the field, highlighting connections with phylogenetic systematics, structure and the biology of extant angiosperms.
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. 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.
This essay shows Alban Berg's intense interrelationship with some literary figures in his native city. Documentation from his life and letters indicate how the works of Karl Kraus and Peter Altenberg inspired the composer. In the process a cultural portrait of Vienna emerges with its culinary attractions, its literary coffee houses, musical offerings and the peculiar character traits of its inhabitants. But great sociological changes also take place in Alban Berg's lifetime: the mass immigration of Jews from the provinces, the politics of anti-Semitism and the incisive events of the war that mark the transition from Vienna, the glittering center of a rich Empire, to Vienna, the impoverished capital of a truncated Republic that nobody wanted.
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.
Secretions from the metapleural gland of the Australian ant Myrmecia nigriscapa Roger were bioassayed against the spores of soil fungi that commonly occur where this ant nests. Four of the seven fungal species were nonentomogenous: Cladosporium resinae (Lindau) DeVries, Colletotrichum gloeosporioides (Penz.) Sacc, Gliocladium roseum Bain, and Mucor plumbeus Bon. Three were entomogenous: Beauveria bassiana Bals. Vuill., Paecilomyces lilacinus (Thorn.) Samson, and Metarhizium brunneum Petch. Metapleural secretions significantly reduced germination of the spores of all fungal species except M. brunneum. In this species, experimental and control spores germinated at approximately equal rates, but the secretions significantly reduced subsequent hyphal growth. These data, together with previously published experiments that show suppression of mycelial growth in eight other soil fungi by metapleural secretions, are consistent with the hypothesis that the metapleural gland is a source of antibiotic. The other social Hymenoptera, the bees and wasps, protect their juvenile stages in antibiotic-impregnated brood cells. By contrast, ants appear to utilize antibiotic secretions from the metapleural gland.
  • E O Wilson
  • F M Carpenter
  • W L Brown
  • Jr
Wilson, E. O., Carpenter, F. M. & Brown, W. L. Jr. Psyche 74, 1–19 (1967).
  • Baroni Urbani
  • C Bolton
  • B Ward
Baroni Urbani, C., Bolton, B. & Ward, P. S. Syst. Entomol. 17, 301–329 (1992).