An opilioacarid mite in Cretaceous Burmese amber

A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opilioacarus groehni sp. nov. This ca. 99 Ma record (Upper Cretaceous: Cenomanian) represents only the third fossil example of this putatively basal mite lineage, the others originating from Eocene Baltic amber (ca. 44–49 Ma). Our new record is not only the oldest record of Opilioacarida, but it is also one of the oldest examples of the entire Parasitiformes clade. The presence of Opilioacarida—potentially Opiloacarus—in the Cretaceous of SE Asia suggests that some modern genus groups were formerly more widely distributed across the northern hemisphere, raising questions about previously suggested Gondwanan origins for these mites.     

Fossil evidence for the early ant evolution

Ants are one of the most studied insects in the world; and the literature devoted to their origin and evolution, systematics, ecology, or interactions with plants, fungi and other organisms is prolific. However, no consensus yet exists on the age estimate of the first Formicidae or on the origin of their eusociality. We review the fossil and biogeographical record of all known Cretaceous ants. We discuss the possible origin of the Formicidae with emphasis on the most primitive subfamily Sphecomyrminae according to its distribution and the Early Cretaceous palaeogeography. And we review the evidence of true castes and eusociality of the early ants regarding their morphological features and their manner of preservation in amber. The mid-Cretaceous amber forest from south-western France where some of the oldest known ants lived, corresponded to a moist tropical forest close to the shore with a dominance of gymnosperm trees but where angiosperms (flowering plants) were already diversified. This palaeoenvironmental reconstruction supports an initial radiation of ants in forest ground litter coincident with the rise of angiosperms, as recently proposed as an ecological explanation for their origin and successful evolution.     

The oldest micropepline beetle from Cretaceous Burmese amber and its phylogenetic implications (Coleoptera: Staphylinidae)

The staphylinid subfamily Micropeplinae includes small strongly sclerotized beetles with truncate elytra leaving the most part of abdomen exposed. Fossil micropeplines are rare and confined to Cenozoic representatives of extant genera. Here, we describe the oldest micropepline, Protopeplus cretaceus gen. and sp. n., from the Upper Cretaceous Burmese amber. Fluorescence microscope and confocal laser scanning microscopy (CLSM) were both used to reveal diagnostic features of Micropeplinae and some primitive traits that place Protopeplus very basally within Micropeplinae.     

Gamergates in the myrmicine genus <Emphasis Type="Italic">Metapone</Emphasis> (Hymenoptera:Formicidae)

Gamergates (i.e. mated reproductive workers) are mostly known from ant species within the Ponerinae. We report here the discovery of gamergates in two species of the subfamily Myrmicinae. Until now, mated reproductive females in colonies of myrmicine species have been considered morphologically distinct from the worker caste. However, in two species of the myrmicine Metapone (Metapone madagascarica and a newly discovered, not yet described Metapone species) all workers have six ovarioles and a spermatheca; and some of them are mated. There are no morphological differences between mated and non-mated workers. Field observations and laboratory studies indicate that colonies of the Metapone species can reproduce with gamergates only.     

A giant termite from the Late Miocene of Styria, Austria (Isoptera)

A giant termite is described and figured from the Late Miocene of the Styrian Basin in southeastern Austria. Gyatermes styriensis gen. n. et sp. n. is represented by a relatively complete forewing, with basal scale. The fossil approximates in size the largest of all termites today and is the largest fossil termite on record. The presence of this species in the Late Miocene fauna of Europe indicates that climatic conditions were appropriate for the persistence of species and colonies requiring relatively stable, warm conditions. The genus is primitive in overall features but shares some similarity with the dampwood termites.     

A unique strategy of host colony exploitation in a parasitic ant: workers of<Emphasis Type="Italic"> Polyrhachis lama</Emphasis> rear their brood in neighbouring host nests

The formicine ant Polyrhachis lama is a social parasite, exploiting its ponerine host ant species Diacamma sp. In most social parasitic associations, the parasitic species are closely related to their host species group, evolving directly from independent ancestors of the host species. However, in the Polyrhachis lama–Diacamma sp. association, the associated species belong to different ant subfamilies. Based on preliminary field surveys, we had presumed that P. lama might have given up its reproductive division of labour, i.e. workers would be able to produce males as well as workers and females parthenogenetically. In this study, this hypothesis was disproved: Polyrhachis lama workers cannot be fertilized and are only able to produce males. In the host–parasite association originating from nests possessing a P. lama queen, workers penetrate surrounding Diacamma sp. nests, carrying brood for rearing within these satellite nests. In this peculiar way, a single P. lama colony is able to exploit several Diacamma sp. colonies simultaneously.     

Fossil mesostigmatid mites (Mesostigmata: Gamasina, Microgyniina, Uropodina), associated with longhorn beetles (Coleoptera: Cerambycidae) in Baltic amber

Fossil mesostigmatid mites are extremely rare. Inclusions assignable to the tortoise mites (Mesostigmata, Uropodina) are described here for the first time from Eocene (ca. 44–49 Ma) Baltic amber. This is the oldest record of Uropodina and documents the first unequivocal amber examples potentially assignable to the extant genus Uroobovella Berlese, 1903 (Uropodoidea: Urodinychidae). Further mites in the same amber pieces are tentatively assigned to Microgynioidea (Microgyniina) and Ascidae (Gamasina), both potentially representing the oldest records of their respective superfamily and family groups. This new material also preserves behavioural ecology in the form of phoretic deutonymphs attached to their carriers via a characteristic anal pedicel. These deutonymphs in amber are intimately associated with longhorn beetles (Coleoptera: Cerambycidae), probably belonging to the extinct species Nothorhina granulicollis Zang, 1905. Modern uropodines have been recorded phoretic on species belonging to several beetle families, including records of living Uroobovella spp. occurring on longhorn beetles. Through these amber inclusions, a uropodine–cerambycid association can now be dated back to at least the Eocene.     

Hummingbird with modern feathering: an exceptionally well-preserved Oligocene fossil from southern France

Hummingbirds (Trochilidae) today have an exclusively New World distribution, but their pre-Pleistocene fossil record comes from Europe only. In this study, we describe an exceptionally preserved fossil hummingbird from the early Oligocene of southeastern France. The specimen is articulated, with a completely preserved beak and feathering. Osteological characters allow to identify it as Eurotrochilus sp. This genus is a stem group representative of Trochilidae and was recently described from the early Oligocene of southern Germany. The new fossil reveals that these European Trochilidae were remarkably modern in size, skeletal proportions and the shape of the wing, tail and beak and hyoid bones. These features confirm the early acquisition of the abilities of hovering and nectarivory in hummingbirds, probably before the Oligocene. In several morphological characteristics, they resemble members of the 'true hummingbirds' (subfamily Trochilinae) and differ from hermits (Phaethornithinae). These features, which include a short and square tail and a moderately long, almost straight beak, appear to be primitive within the family Trochilidae.     

A structural intermediate between triisodontids and mesonychians (Mammalia, Acreodi) from the earliest Eocene of Portugal

A new mammal, Mondegodon eutrigonus gen. et sp. nov., is described from the earliest Eocene locality of Silveirinha, Portugal. This species shows dental adaptations indicative of a carnivorous diet. M. eutrigonus is referred to the order Acreodi and considered, along with the early Paleocene North American species Oxyclaenus cuspidatus, as a morphological intermediate between two groups of ungulate-like mammals, namely, the triisodontids and mesonychians. Considering that triisodontids are early to early-late Paleocene North American taxa, Mondegodon probably belongs to a group that migrated from North America towards Europe during the first part of the Paleocene. Mondegodon could represent thus a relict genus, belonging to the ante-Eocene European mammalian fauna. The occurrence of such a taxon in Southern Europe may reflect a period of isolation of this continental area during the Paleocene/Eocene transition. In this context, the non-occurrence of closely allied forms of Mondegodon in the Eocene North European mammalian faunas is significant. This strengthens the hypothesis that the mammalian fauna from Southern Europe is characterized by a certain degree of endemism during the earliest Eocene. Mondegodon also presents some striking similarities with an unnamed genus from the early Eocene of India which could represent the first Asian known transitional form between the triisodontids and mesonychians.     

Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania

Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Haţeg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.     

A new bolosaurid parareptile, Belebey chengi sp. nov., from the Middle Permian of China and its paleogeographic significance

A new bolosaurid parareptile, Belebey chengi sp. nov., is described from Dashankou, Gansu Province, China, a Middle Permian locality which is known mostly for its therapsid fauna. The material consists of well-preserved mandibular and anterior skull remains and currently represents the largest and latest surviving member of Bolosauridae. Phylogenetic analysis of bolosaurid interrelationships, the first analysis of any clade of Early Permian parareptiles, indicates that the new taxon groups consistently with the other (Russian) members of the genus Belebey, and forms the sister clade to the genus Bolosaurus from North America. The Early Permian Eudibamus cursoris from Germany falls into the basal most position within Bolosauridae. Our analysis also shows that the split between the main bolosaurid lineages must have occurred near or before the Permo-Carboniferous boundary and that the paleo-equatorial region of Laurasia probably served as the center of origination for these parareptiles. A similar pattern can be found in other clades of Paleozoic amniotes, suggesting that this may be the general trend in early amniote evolution.     

Artiodactyls from the Pondaung Formation (Myanmar): new data and reevaluation of the South Asian Faunal Province during the Middle Eocene

Although Asia is thought to have played a critical role in the radiation of artiodactyls, the fossil record of stem selenodonts (“dichobunoids”) remains dramatically poor in tropical Asian regions. In this study, we report a new dichobunid genus and species Cadutherium kyaukmagyii and a new basal ruminant genus and species Irrawadymeryx pondaungi, from the late Middle Eocene Pondaung Formation, Central Myanmar. Although the scarcity of the present material prevents any attempts to investigate the phylogenetic relationships of Cadutherium with contemporaneous forms from other Holarctic landmasses, this new form shed new light on the diversity of these small rabbit-like ungulates during a key period of their evolutionary history. Reexamination of the small-bodied artiodactyls from Pondaung leads us to propose new identifications of certain published specimens and, in turn, to investigate the temporal and geographic distribution of taxa recognized in the Pondaung Formation. Although fragmentary, these potential new taxa reveal an unsuspected diversity of small forms among artiodactyls of Pondaung. This addition to the Eocene record of dichobunoids and early ruminants provides further insight in the diversity of dental patterns among small artiodactyls from the Pondaung Formation and attests to the antiquity of these groups in Southeast Asia.     

Aggressions and size-related fecundity of queenless workers in the ant <Emphasis Type="Italic">Cataglyphis cursor</Emphasis>

In social hymenoptera, the reproductive division of labor is often linked to differences in individual body size with the reproductive caste (the queen) being larger than the workers. Likewise, the reproductive potential may vary with size within the worker caste and could affect the evolution of worker size in social insects. Here, we tested the relationship between worker size and reproductive potential in the facultative parthenogenetic ant Cataglyphis cursor. Colonies are headed by a multiply mated queen, but workers can produce gynes (virgin queens) and workers by thelytokous parthenogenesis after the queen's death. We observed the behaviour of workers (n = 357) until the production of gynes (212 h over 3 months) in an orphaned colony (mated queen not present). The size of workers was measured, and their paternal lineage determined using six microsatellite markers, to control for an effect of patriline. Larger workers were more likely to reproduce and lay more eggs indicating that individual level selection could take place. However, paternal lineage had no effect on the reproductive potential and worker size. From the behavioural and genetic data, we also show for the first time in this species, evidence of aggressive interactions among workers and a potential for nepotism to occur in orphaned colonies, as the five gynes produced belonged to a single paternal lineage.     

Gamergates in the Australian ant subfamily Myrmeciinae

Ant workers can mate and reproduce in a few hundreds of species belonging to the phylogenetically basal poneromorph subfamilies (sensu Bolton 2003). We report the first occurrence of gamergates (i.e. mated reproductive workers) in a myrmeciomorph subfamily. In a colony of Myrmecia pyriformis that was collected without a queen, workers continued to be produced over a period of 3 years in the laboratory. Behavioural observations and ovarian dissections indicated that three workers were mated and produced the diploid offspring. The Myrmeciinae are thus another taxon in which the selective benefits of sexual reproduction by workers can be investigated.     

Living within fallen palm leaves: the discovery of an unknown Blommersia (Mantellidae: Anura) reveals a new reproductive strategy in the amphibians of Madagascar

We describe a new mantelline frog of the genus Blommersia found in rainforest in North East Madagascar, from the protected areas of Ambatovaky, Betampona, Masoala, and Zahamena. Blommersia angolafa n.sp. is a small frog, with a body size of 17–21 mm, expanded finger and toe tips, and colouration ranging from yellow to dark brown, with pale-bluish spots on the flanks and light tips of fingers and toes. A peculiar aspect characterising this new species is its novel life history and reproductive mode. Both sexes live and breed in a phytotelmic habitat of water accumulated within fallen prophylls and fallen leaf sheaths of at least three species of Dypsis palms. Within these phytotelmata, egg laying and complete larval development occur. Thus, B. angolafa n.sp. represents a new evolutionary lineage of Malagasy frogs in which phytotelmy is known. Up to now, reproduction in phytotelmata in Malagasy frogs has been reported for many cophyline microhylids, most species of Guibemantis, Mantella laevigata, and possibly in a still-undescribed species belonging to the genus Spinomantis. We consider the reproductive mode of B. angolafa as a derived character, having evolved from the more typical reproduction in lentic water bodies. The general scarcity of lentic habitats in Malagasy rainforests may have provided the conditions that favoured the evolution of this phytotelmic breeding strategy. The new species, being specialised to a habitat represented by a few selected Dypsis species, potentially suffers the selective exploitation of these palms.     

Spectacular Batesian mimicry in ants

The mechanism by which palatable species take advantage of their similarity in appearance to those that are unpalatable, in order to avoid predation, is called Batesian mimicry. Several arthropods are thought to be Batesian mimics of social insects; however, social insects that are Batesian mimics among themselves are rare. In Malaysia we found a possible Batesian mimic in an arboreal ant species, Camponotus sp., which was exclusively observed on foraging trails of the myrmicine ant Crematogaster inflata. The bright yellow and black colouring pattern, as well as the walking behaviour, were very similar in both species. We observed general interactions between the two species, and tested their palatability and the significance of the remarkably similar visual colour patterns for predator avoidance. Prey offered to C. inflata was also eaten by Camponotus workers in spite of their being attacked by C. inflata, indicating that Camponotus sp. is a commensal of C. inflata. An experiment with chicks as potential predators suggests that Camponotus sp. is palatable whereas C. inflata is unpalatable. After tasting C. inflata, the chicks no longer attacked Camponotus sp., indicating that Camponotus sp. is a Batesian mimic of Crematogaster inflata.     

The ant genomes have been invaded by several types of mariner transposable elements

To date, only three types of full-length mariner elements have been described in ants, each one in a different genus of the Myrmicinae subfamily: Sinvmar was isolated from various Solenopsis species, Myrmar from Myrmica ruginodis, and Mboumar from Messor bouvieri. In this study, we report the coexistence of three mariner elements (Tnigmar-Si, Tnigmar-Mr, and Tnigmar-Mb) in the genome of a single species, Tapinoma nigerrimum (subfamily Dolichoderinae). Molecular evolutionary analyses of the nucleotide sequence data revealed a general agreement between the evolutionary history of most the elements and the ant species that harbour them, and suggest that they are at the vertical inactivation stage of the so-called Mariner Life Cycle. In contrast, significantly reduced levels of synonymous divergence between Mboumar and Tnigmar-Mb and between Myrmar and Botmar (a mariner element isolated from Bombus terrestris), relative to those observed between their hosts, suggest that these elements arrived to the species that host them by horizontal transfer, long after the species?? split. The horizontal transfer events for the two pairs of elements could be roughly dated within the last 2 million years and about 14 million years, respectively. As would be expected under this scenario, the coding sequences of the youngest elements, Tnigmar-Mb and Mboumar, are intact and, thus, potentially functional. Each mariner element has a different chromosomal distribution pattern according to their stage within the Mariner Life Cycle. Finally, a new defective transposable element (Azteca) has also been found inserted into the Tnigmar-Mr sequences showing that the ant genomes have been invaded by at least four different types of mariner elements.     

Eocene bunoselenodont Artiodactyla from southern Thailand and the early evolution of Ruminantia in South Asia

Although Asia is thought to have played a critical role in the basal radiation of Ruminantia, the fossil record of early selenodont artiodactyls remains poorly documented in this region. Dental remains of a new bunoselenodont artiodactyl are described from the late Eocene of Krabi, southern Thailand. This new form, Krabitherium waileki gen. et sp. nov, is tentatively referred to the Tragulidae (Ruminantia) on the basis of several dental features, including a weak Tragulus fold and the presence of a deep groove on the anterior face of the entoconid. Although this new form is suggestive of the enigmatic ?Gelocus gajensis Pilgrim 1912 from the “base of the Gaj” (lower Chitarwata Formation) of the Bugti Hills (Central Pakistan), K. waileki most likely represents an early representative of a relatively bunodont group of tragulids that includes the genus Dorcabune, known from the Miocene of south Asia. This addition to the Eocene record of early ruminants attests to the antiquity of the group in Southeast Asia and lends support to the hypothesis that the Tragulidae represents one of the first offshoots in the evolutionary history of Ruminantia.