Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes)

Synodontis (Mochokidae, Siluriformes) is a freshwater catfish endemic to Africa. The 118 extant species are present in almost all hydrographic basins. Some species are restricted to a single stream, whereas others have a vast distribution. Synodontis is known in the fossil record since the Miocene, and its history depends on the connections among African basins through time. The identification of species in the fossil record is essential to reconstruct this historical pattern. Catfish pectoral and dorsal spines are robust, they preserve well and they form most of the fossil remains for the genus Synodontis. Unfortunately, the criteria for the identification of extant Synodontis species are not applicable to fossil specimens. Here, we define 11 original morphological characters that permit to discriminate four extant species from the Chad-Chari hydrographic system. Six of these characters are defined on pectoral spines and five on dorsal spines. We then show that these characters can be used successfully for identifying fossil specimens. In particular, we present a case study in which we identify Synodontis cf. schall and Brachysynodontis cf. batensoda in the hominid-bearing sector Toros-Menalla (Late Miocene, northern Chad). We show that spine anatomy can be a powerful tool to recognise catfish species through time and thus to identify historical diversity pattern.     

The emergence of collective foraging in the arboreal<Emphasis Type="Italic"> Gnamptogenys menadensis</Emphasis> (Hymenoptera: Formicidae)

Gnamptogenys menadensis is an arboreal nester that forages opportunistically almost exclusively on vegetation, sometimes recruiting others to participate in prey retrieval. The three-dimensional characteristics of vegetation suggest that functions describing recruitment decision thresholds or the pattern of recruitment in arboreal species may differ from those predicted by optimal foraging theory. To examine the effects of prey abundance and distance on the recruitment dynamics of G. menadensis, we baited nests with one termite, five termites or a number of termites between 20 and 40 either near to or far from the entrance and observed the ensuing behaviors. G. menadensis recruited others when encountering multiple termites regardless of the termite pile's distance from the nest, although a few individuals remained at the site and defended the resource. The pattern of arrivals at the site indicates that the majority and sometimes all arrivals were recruited from the branch trails. In combination, these results suggest that the architecture of the foraging habitat, which limits available return routes to the nest and thus increases encounter probabilities with potential recruits, shaped the process of information transfer and generated a collective pattern of foraging and prey retrieval.     

Termite assemblages in a West-African semi-deciduous forest and teak plantations

Responses of termite assemblages to the conversion of semi-deciduous forest into teak plantations were studied in the Lama Forest Reserve in Benin, West Africa. Four belt transect surveys were run in each of the two forest types, adopting a modified termite diversity assessment protocol. Termite assemblages were remarkably species-poor in both forest types, with only 19 species encountered altogether. The low species richness was due to the rarity of soil-feeders of the soil/humus interface and the absence of true soil-feeders in the compact vertisol soil. Species richness was significantly higher in semi-deciduous forest than in teak plantations, but termite encounters were significantly lower. Termite assemblage and feeding group structure differed significantly among forest types. Wood-feeders were recorded only in semi-deciduous forest. In contrast, fungus-growers were more species-rich and about four times more abundant in teak plantations, mainly due to one Ancistrotermes species. The humification score, which depicts the position of termite assemblages along a gradient of increasing humification of their food substrate, was significantly higher in teak plantations, due to the absence of wood-feeders and the preponderance of fungus-growers. Combined principal components and multiple regression analysis identified two significant predictors of termite assemblages, soil water content and leaf litter biomass. The abundance of fungus-growers in teak plantations seemed to be mainly related to the high leaf litter biomass. Indirect evidence also suggests that lower predation pressure by ants on termites in teak plantations may have contributed to the abundance of termites.     

Dietary characterization of the hominoid Khoratpithecus (Miocene of Thailand): evidence from dental topographic and microwear texture analyses

The genus Khoratpithecus, a hominoid thought to be related to the orangutan lineage, is represented by two known fossil species K. chiangmuanensis and K. piriyai. Both were discovered in Southeast Asia (Thailand) and are dated to the Middle and Late Miocene, respectively. In this study, dental topographic and microwear texture analyses were used to examine molars from both of these species, with the goal of understanding their dietary preferences. Although sample sizes are small for Khoratpithecus, available data are compared to that collected for extant apes. Environmental evidence, such as botanical remains and sedimentological data, is also considered for comparisons with dietary reconstruction. Results from dental topographic analysis suggest that the two fossil species were better adapted to a diet of fruits than to one of leaves, much like the living orangutan or chimpanzee. Results from microwear texture analysis further support this, suggesting that Khoratpithecus preferred soft fruits to hard fruits or seeds. And finally, the botanical and sedimentological evidence point to environments for Khoratpithecus that would have been compatible with a fruit-eating species. Given the small sample sizes available for analysis, however, definitive judgments are not yet possible at this time.     

松属植物地质历史及生物地理演变

松属（Pinus L.）约113种,是松科现代属中最原始的类群。松属植物种类丰富且研究领域广泛,对其已经积累的资料数据进行系统梳理总结十分必要。本文通过总结国内外松属大化石资料,结合分子系统发育、地质背景和地理隔离事件讨论了其地史分布及植物地理学意义;该属化石在早白垩世至全新世地层中均有记录。化石证据表明松属很可能在早白垩世（之前）起源于西欧地区,从这一起源地通过北大西洋陆桥扩散到北美洲东部,而东亚的类群可能是从北美洲西部经过白令陆桥散布的。在晚白垩世分化出双维管束松亚属Subgenus Pinus L.及单维管束松亚属Subgenus Strobus (D. Don) Lemmon,前者更接近祖先类群。古新世由于全球显著增温以及白垩纪末期大灭绝等地质事件的影响使松属数量急剧减少,在晚始新世至中新世时期随着气温转凉转冷再次分化扩散,中新世达到其发展高峰且分布面貌与现代类群近似。松属多样性时空历史可能和新生代气候变迁及晚新生代构造运动塑造的山地隆升等环境变化紧密相关。     

Phenoloxidase activity in the infraorder Isoptera: unraveling life-history correlates of immune investment

A termite colony is usually founded by a pair of alates, the primary reproductives, which produce all the nestmates. In some species, secondary reproductives appear to either replace the primaries or supplement colony reproduction. In termites, secondary reproductives are generally ergatoids derived from workers or nymphoids derived from nymphs. Silvestritermes euamignathus is a termite species that forms multiple nymphoid reproductives, and to date it was hypothesized that these secondary reproductives were the progeny of the primary founding reproductives. We developed markers for 12 microsatellite loci and used COI mitochondrial DNA (mtDNA) to genotype 59 nymphoid neotenics found in a colony of S. euamignathus to test this hypothesis. Our results showed that nymphoids of S. euamignathus are not all siblings. The microsatellite analysis suggests that the secondary reproductives derived from a minimum of four different pairs of reproductives belonging to at least two different matrilines. This is the first record of non-sibling secondary reproductives occupying the same nest in a higher termite. These unrelated reproductives might be the result of either pleometrotic colony foundation or colony fusion.

     

New data on Mustelidae (Carnivora) from Southeast Asia: Siamogale thailandica, a peculiar otter-like mustelid from the late middle Miocene Mae Moh Basin, northern Thailand

We report new dental remains of Mustelidae from the late middle Miocene of Mae Moh Basin, northern Thailand, improving the poor fossil record of the family in Southeast Asia. Siamogale thailandica is a poorly known mustelid, previously recorded from just a single tooth. Here we present over a hundred new specimens attributable to this species. S. thailandica shows a combination of primitive and convergent features of the dentition that makes its original subfamilial assignment to Lutrinae doubtful. Evidence from the dental morphology suggests that it belongs to a bunodont otter-like mustelid that evolved in convergence with “true” otters (Lutrinae) toward a semi-aquatic way of life. Autapomorphic features such as the height and the position of the m1 metaconid and the shape of the P4 lingual shelf make S. thailandica unique among Mustelidae. The morphology of this species is mostly similar to Mionictis species and Lartetictis dubia, reported in the Miocene of North America and Europe, respectively. These similarities could imply immigration events to Thailand in the early or middle Miocene. Alternately, the lineage leading to Siamogale might have deeper origins from an endemic early Miocene Southeast Asian mustelid.     

Agricultural management affects earthworm and termite diversity across humid to semi-arid tropical zones

Earthworm and termite diversity were studied in 12 long-term agricultural field trials across the sub-humid to semi-arid tropical zones of Eastern and Western Africa. In each trial, treatments with high and low soil organic C were chosen to represent contrasts in long-term soil management effects, including tillage intensity, organic matter and nutrient management and crop rotations. For each trial, a fallow representing a relatively undisturbed reference was also sampled. Earthworm taxonomic richness decreased in the direction fallow > high-C soil > low-C soil and earthworm abundance was higher in fallow than under continuous crop production. Termite abundance was not significantly different between fallow and high and low-C treatments and termite taxonomic richness was higher in fallow soil than in the two cropping systems. We concluded that fewer species of earthworms and termites were favored under agricultural management that led to lower soil C. Results indicated that the soil disturbance induced by continuous crop production was more detrimental to earthworms than to termites, when compared to the fallow.     

Resource competition between two fungal parasites in subterranean termites

Subterranean termites live in large groups in underground nests where the pathogenic pressure of the soil environment has led to the evolution of a complex interaction among individual and social immune mechanisms in the colonies. However, groups of termites under stress can show increased susceptibility to opportunistic parasites. In this study, an isolate of Aspergillus nomius Kurtzman, Horn & Hessltine was obtained from a collapsed termite laboratory colony. We determined that it was primarily a saprophyte and, secondarily, a facultative parasite if the termite immunity is undergoing a form of stress. This was determined by stressing individuals of the Formosan subterranean termite Coptotermes formosanus Shiraki via a co-exposure to the virulent fungal parasite Metarhizium anisopliae (Metch.) Sorokin. We also examined the dynamics of a mixed infection of A. nomius and M. anisopliae in a single termite host. The virulent parasite M. anisopliae debilitated the termite immune system, but the facultative, fast growing parasite A. nomius dominated the mixed infection process. The resource utilization strategy of A. nomius during the infection resulted in successful conidia production, while the chance for M. anisopliae to complete its life cycle was reduced. Our results also suggest that the occurrence of opportunistic parasites such as A. nomius in collapsing termite laboratory colonies is the consequence of a previous stress, not the cause of the stress.     

Extended mutualism between termites and gut microbes: nutritional symbionts contribute to nest hygiene

All higher eukaryotes have established symbiotic relationships with diverse microorganisms. One of the most well-characterized symbiotic systems is that of termites and their intestinal microorganisms, which digest cellulose. Recently, diverse types of symbioses between gut microbes and host organisms including humans have received growing attention for various features of their complex interactions beyond nutrition. In termites, researchers are beginning to explore such function of gut symbionts, but only the contribution to internal immunity against entomopathogen is known in a few species. Here, we report that gut symbionts of the dampwood termite Zootermopsis nevadensis protect nests from the spread of the commensal bacterium Serratia marcescens, which has pathogenic potential. Defaunated termites dispersed S. marcescens in the surrounding environment by feeding on the bacteria, which then survived passage through their alimentary tracts, while non-defaunated termites did not. Loss of gut symbionts caused a significant reduction in intestinal acetate, which is an important carbon source for termites. Culture experiments showed that acetate had significant inhibitory effects on S. marcescens at a concentration as low as 12 mM, which indicated that the intestinal acetate of non-defaunated termites (40–130 mM) was capable of suppressing this bacterium. These results suggest that digestive derivatives produced by intestinal symbionts play an essential role in nest hygiene in addition to their nutritional function for termites. Our study provides a better understanding of the multifunctionality of symbiotic relationships in diverse organisms beyond nutrition.     

Cretaceous choristoderan reptiles gave birth to live young

Viviparity (giving birth to live young) in fossil reptiles has been known only in a few marine groups: ichthyosaurs, pachypleurosaurs, and mosasaurs. Here, we report a pregnant specimen of the Early Cretaceous Hyphalosaurus baitaigouensis, a species of Choristodera, a diapsid group known from unequivocal fossil remains from the Middle Jurassic to the early Miocene (about 165 to 20 million years ago). This specimen provides the first evidence of viviparity in choristoderan reptiles and is also the sole record of viviparity in fossil reptiles which lived in freshwater ecosystems. This exquisitely preserved specimen contains up to 18 embryos arranged in pairs. Size comparison with small free-living individuals and the straight posture of the posterior-most pair suggest that those embryos were at term and had probably reached parturition. The posterior-most embryo on the left side has the head positioned toward the rear, contrary to normal position, suggesting a complication that may have contributed to the mother’s death. Viviparity would certainly have freed species of Hyphalosaurus from the need to return to land to deposit eggs; taking this advantage, they would have avoided intense competition with contemporaneous terrestrial carnivores such as dinosaurs.     

The first fossil fungus gardens of Isoptera: oldest evidence of symbiotic termite fungiculture (Miocene, Chad basin)

Higher termites of the subfamily Macrotermitinae (fungus-growing termites) are known to build fungus gardens where a symbiotic fungus (Termitomyces sp.) is cultivated. The fungus grows on a substrate called fungus comb, a structure built with the termites’ own faeces. Here we present the first fossil fungus combs ever found in the world. They were extracted from 7-million-year-old continental sandstone (Chad basin). Fossilized fungus combs have an ovoid morphology with a more or less flattened concave base and a characteristic general alveolar aspect. Under lens, they display a typical millimetre-scale pelletal structure. The latter, as well as the general shape and alveolar aspect, are similar to the morphology of fungus combs from extant fungus-growing termites.     

A chicken-sized crane precursor from the early Oligocene of France

A skeleton of a chicken-sized crane precursor is described from the Lower Oligocene of the Lubéron in Southern France. Parvigrus pohli gen. et sp. nov. is the most substantial Paleogene fossil record of the Grues (Aramidae [limpkin] + Gruidae [cranes]), and among its oldest representatives. The fossil species is classified in the new taxon Parvigruidae, which is shown to be the sister group of extant Grues. It is the first fossil record of a stem lineage representative of the Grues and, among others, differs from modern Grues in its smaller size, shorter beak, and rail-like limb proportions. Size increase in the stem lineage of the Gruidae may be related to the spread of grasslands during the Oligocene and Miocene. Occurrence of stem lineage Grues in the Lower Oligocene of Europe is in concordance with the fact that there is no evidence for the presence of crown group members of modern avian “families” in pre-Oligocene fossil deposits. Electronic Supplementary Material  Supplementary material is available for this article at     

Scent of a queen—cuticular hydrocarbons specific for female reproductives in lower termites

In social insects, it is assumed that signals of the queen inform nestmates about her reproductive status. Thus, workers forego their own reproduction if the queen signals high fertility. In hemimetabolous termites, little is known about reproductive inhibition, but evidence exists for a royal-pair control. Workers of lower termites exhibit a high developmental flexibility and are potentially able to become reproductives, but the presence of a fertile reproductive restrains them from reaching sexual maturity. The nature of this control, however, remains unknown. Here, we report on qualitative differences in cuticular hydrocarbon profiles between queens and workers of the basal drywood termite Cryptotermes secundus. Queens were characterized by a shift to long-chained and branched hydrocarbons. Most remarkably, similar chemical patterns are regarded as fertility cues of reproductives in social Hymenoptera. This might suggest that both groups of social insects convergently evolved similar chemical signatures. The present study provides deeper insights into how termites might have socially exploited these signatures from sexual communication in their cockroach-like ancestor. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A new fossil thryonomyid from the Late Miocene of the United Arab Emirates and the origin of African cane rats

Cane rats (Thryonomyidae) are represented today by two species inhabiting sub-Saharan Africa. Their fossil record is predominately African, but includes several Miocene species from Arabia and continental Asia that represent dispersal events from Africa. For example, Paraulacodus indicus, known from the Miocene of Pakistan, is closely related to living Thryonomys. Here we describe a new thryonomyid, Protohummus dango, gen. et sp. nov., from the late Miocene Baynunah Formation of the United Arab Emirates. The new thryonomyid is less derived than “Thryonomys” asakomae from the latest Miocene of Ethiopia and clarifies the origin of crown Thryonomys and the evolutionary transition from Paraulacodus. A phylogenetic analysis shows Protohummus dango to be morphologically intermediate between Paraulacodus spp. and extinct and living Thryonomys spp. The morphological grade and phylogenetic position of Protohummus dango further supports previous biochronological estimates of the age of the Baynunah Formation (ca. 6–8 Ma).     

A gravid lizard from the Cretaceous of China and the early history of squamate viviparity

Although viviparity is most often associated with mammals, roughly one fifth of extant squamate reptiles give birth to live young. Phylogenetic analyses indicate that the trait evolved more than 100 times within Squamata, a frequency greater than that of all other vertebrate clades combined. However, there is debate as to the antiquity of the trait and, until now, the only direct fossil evidence of squamate viviparity was in Late Cretaceous mosasauroids, specialised marine lizards without modern equivalents. Here, we document viviparity in a specimen of a more generalised lizard, Yabeinosaurus, from the Early Cretaceous of China. The gravid female contains more than 15 young at a level of skeletal development corresponding to that of late embryos of living viviparous lizards. This specimen documents the first occurrence of viviparity in a fossil reptile that was largely terrestrial in life, and extends the temporal distribution of the trait in squamates by at least 30 Ma. As Yabeinosaurus occupies a relatively basal position within crown-group squamates, it suggests that the anatomical and physiological preconditions for viviparity arose early within Squamata.     

Phylogeography of Australia’s king brown snake (Pseudechis australis) reveals Pliocene divergence and Pleistocene dispersal of a top predator

King brown snakes or mulga snakes (Pseudechis australis) are the largest and among the most dangerous and wide-ranging venomous snakes in Australia and New Guinea. They occur in diverse habitats, are important predators, and exhibit considerable morphological variation. We infer the relationships and historical biogeography of P. australis based on phylogenetic analysis of 1,249 base pairs from the mitochondrial cytochrome b, NADH dehydrogenase subunit 4 and three adjacent tRNA genes using Bayesian, maximum-likelihood, and maximum-parsimony methods. All methods reveal deep phylogenetic structure with four strongly supported clades comprising snakes from New Guinea (I), localities all over Australia (II), the Kimberleys of Western Australia (III), and north-central Australia (IV), suggesting a much more ancient radiation than previously believed. This conclusion is robust to different molecular clock estimations indicating divergence in Pliocene or Late Miocene, after landbridge dispersal to New Guinea had occurred. While members of clades I, III and IV are medium-sized, slender snakes, those of clade II attain large sizes and a robust build, rendering them top predators in their ecosystems. Genetic differentiation within clade II is low and haplotype distribution largely incongruent with geography or colour morphs, suggesting Pleistocene dispersal and recent ecomorph evolution. Significant haplotype diversity exists in clades III and IV, implying that clade IV comprises two species. Members of clade II are broadly sympatric with members of both northern Australian clades. Thus, our data support the recognition of at least five species from within P. australis (auct.) under various criteria. We discuss biogeographical, ecological and medical implications of our findings.Electronic Supplementary Material  Supplementary material is available in the online version of this article at     

Grooves to tubes: evolution of the venom delivery system in a Late Triassic “reptile”

Venom delivery systems occur in a wide range of extant and fossil vertebrates and are primarily based on oral adaptations. Teeth range from unmodified (Komodo dragons) to highly specialized fangs similar to hypodermic needles (protero- and solenoglyphous snakes). Developmental biologists have documented evidence for an infolding pathway of fang evolution, where the groove folds over to create the more derived condition. However, the oldest known members of venomous clades retain the same condition as their extant relatives, resulting in no fossil evidence for the transition. Based on a comparison of previously known specimens with newly discovered teeth from North Carolina, we describe a new species of the Late Triassic archosauriform Uatchitodon and provide detailed analyses that provide evidence for both venom conduction and document a complete structural series from shallow grooves to fully enclosed tubular canals. While known only from teeth, Uatchitodon is highly diagnostic in possessing compound serrations and for having two venom canals on each tooth in the dentition. Further, although not a snake, Uatchitodon sheds light on the evolutionary trajectory of venom delivery systems in amniotes and provide solid evidence for venom conduction in archosaur-line diapsids.     

The earliest modern mongoose (Carnivora, Herpestidae) from Africa (late Miocene of Chad)

We report on the earliest modern mongooses of Africa, from the late Miocene (ca. 7 Ma) of the hominid locality TM 266, Toros-Menalla, Chad. The material is based on fragmentary dentitions of three individuals. The main diagnostic feature of the Chadian species is the great development of the shear in the carnassials, which distinguishes the Chadian specimens from all extant herpestids except Herpestes and Galerella. In comparison with most extinct and extant Herpestes, the species from Toros-Menalla differs by a markedly smaller size and, depending on the species, relatively more elongated carnassials, more transversely elongated M1 and more reduced p4. On the basis of a great morphological similarity and the absence of significant differences, we assign our material to Galerella sanguinea; the Chadian finding therefore represents the earliest appearance of an extant species of Herpestidae. This record ties the first appearance of the genus to a minimum age of ca. 7 Ma, which is consistent with the estimated divergence date of 11.4 Ma known from the literature for the species of Galerella.