Biogeography and evolution of social parasitism in Australian Myrmecia bulldog ants revealed by phylogenomics.

Studying the historical biogeography and life history transitions from eusocial colony life to social parasitism contributes to our understanding of the evolutionary mechanisms generating biodiversity in eusocial insects. The ants in the genus Myrmecia are a well-suited system for testing evolutionary hypotheses about how their species diversity was assembled through time because the genus is endemic to Australia with the single exception of the species M. apicalis inhabiting the Pacific Island of New Caledonia, and because at least one social parasite species exists in the genus. However, the evolutionary mechanisms underlying the disjunct biogeographic distribution of M. apicalis and the life history transition(s) to social parasitism remain unexplored. To study the biogeographic origin of the isolated, oceanic species M. apicalis and to reveal the origin and evolution of social parasitism in the genus, we reconstructed a comprehensive phylogeny of the ant subfamily Myrmeciinae. We utilized Ultra Conserved Elements (UCEs) as molecular markers to generate a molecular genetic dataset consisting of 2,287 loci per taxon on average for 66 out of the 93 known Myrmecia species as well as for the sister lineage Nothomyrmecia macrops and selected outgroups. Our time-calibrated phylogeny inferred that: (i) stem Myrmeciinae originated during the Paleocene âˆ¼ 58 Ma ago; (ii) the current disjunct biogeographic distribution of M. apicalis was driven by long-distance dispersal from Australia to New Caledonia during the Miocene âˆ¼ 14 Ma ago; (iii) the single social parasite species, M. inquilina, evolved directly from one of the two known host species, M. nigriceps, in sympatry via the intraspecific route of social parasite evolution; and (iv) 5 of the 9 previously established taxonomic species groups are non-monophyletic. We suggest minor changes to reconcile the molecular phylogenetic results with the taxonomic classification. Our study enhances our understanding of the evolution and biogeography of Australian bulldog ants, contributes to our knowledge about the evolution of social parasitism in ants, and provides a solid phylogenetic foundation for future inquiries into the biology, taxonomy, and classification of Myrmeciinae.

Cretaceous environmental changes led to high extinction rates in a hyperdiverse beetle family.

BACKGROUND: As attested by the fossil record, Cretaceous environmental changes have significantly impacted the diversification dynamics of several groups of organisms. A major biome turnover that occurred during this period was the rise of angiosperms starting ca. 125 million years ago. Though there is evidence that the latter promoted the diversification of phytophagous insects, the response of other insect groups to Cretaceous environmental changes is still largely unknown. To gain novel insights on this issue, we assess the diversification dynamics of a hyperdiverse family of detritivorous beetles (Tenebrionidae) using molecular dating and diversification analyses. RESULTS: Age estimates reveal an origin after the Triassic-Jurassic mass extinction (older than previously thought), followed by the diversification of major lineages during Pangaean and Gondwanan breakups. Dating analyses indicate that arid-adapted species diversified early, while most of the lineages that are adapted to more humid conditions diversified much later. Contrary to other insect groups, we found no support for a positive shift in diversification rates during the Cretaceous; instead there is evidence for an 8.5-fold increase in extinction rates that was not compensated by a joint increase in speciation rates. CONCLUSIONS: We hypothesize that this pattern is better explained by the concomitant reduction of arid environments starting in the mid-Cretaceous, which likely negatively impacted the diversification of arid-adapted species that were predominant at that time.

Leptospirosis risk increases with changes in species composition of rat populations.

Rats are major reservoirs of leptospirosis and considered as a main threat to biodiversity. A recent introduction of Rattus rattus to the island of Futuna (Western Polynesia) provided the opportunity to test if a possible change in species composition of rat populations would increase the risk of leptospirosis to humans. We trapped rodents on Wallis and Futuna and assessed Leptospira carriage in 357 rodents (Rattus norvegicus, R. rattus, Rattus exulans, and Mus domesticus) from 2008 to 2012. While Leptospira prevalence in rodents and the composition of rat populations on Futuna fluctuated with rainfall, the biomass of Leptospira-carrying rodents has been continuously rising from 2008 to 2012. Our results suggest that the introduction of R. rattus increases the risk to humans being infected with leptospirosis by rats.

Checklist of the bees (Hymenoptera, Apoidea) of New Caledonia.

Background: In a world where insects and notably bees are declining, assessing their distribution over time and space is crucial to evaluate species status and highlight conservation priorities. However, this can be a daunting task, especially in areas such as tropical oceanic islands where exhaustive samplings over time have been lacking. This is the case in New Caledonia, an archipelago located in the southwest Pacific. Historical records of bee species are piecemeal and, although contemporary samplings have significantly advanced our knowledge of the bee fauna of New Caledonia, the status of several species remains to be elucidated. New information: Here, we provide an updated checklist of the 51 bee species recorded for New Caledonia using previous publications and personal samplings. We documented their distribution, origin (i.e. endemic, native or alien) and the year and location of their occurrences. Based on the year of their first capture and the year of their last capture, we determined an occurrence status for each species. Thus, 10 years after the last checklist of the New Caledonian bee fauna, the literature review and recent samplings allowed us to add six new species to the list. Half of them are recently introduced species including one firstly mentioned in this paper (i.e. Hylaeusalbonitens). We consider here that 30 species are effectively present on the territory and the presence of 21 species could not be determined due to a lack of data, which highlights the need to increase sampling efforts across New Caledonia. Given the difficulty of exhaustively sampling the entire archipelago, we would recommend taking, as a starting point, altitude environments and areas where data-deficient species were captured. In a broader perspective, biomolecular analyses are crucial to confirm species identifications. This is also needed to make comparisons between archipelagoes and thus clarify the distribution and status of species at the scale of the southwest Pacific.

Sequential invasions by fruit flies (Diptera: Tephritidae) in Pacific and Indian Ocean islands: A systematic review.

The aim of our review was to examine the cases of Tephritidae invasions across island systems in order to determine whether they follow a hierarchical mode of invasion. We reviewed the literature on factors and mechanisms driving invasion sequences in Pacific and Southwest Indian Ocean islands and gathered every record of invasion by a polyphagous tephritid in island groups. From invasion date or period, we defined an invasion link when a new fruit fly established on an island where another polyphagous tephritid is already resident (that was indigenous or a previous invader). Across surveyed islands, we documented 67 invasion links, involving 24 tephritid species. All invasion links were directional, i.e., they involved a series of invasions by invaders that were closely related to a resident species but were increasingly more competitive. These sequential establishments of species are driven by interspecific competition between resident and exotic species but are also influenced by history, routes, and flows of commercial exchanges and the bridgehead effect. This information should be used to improve biosecurity measures. Interactions between trade flow, invasive routes, and the presence of invasive and resident species should be integrated into large-scale studies.

Clonal reproduction by males and females in the little fire ant.

Sexual reproduction can lead to major conflicts between sexes and within genomes. Here we report an extreme case of such conflicts in the little fire ant Wasmannia auropunctata. We found that sterile workers are produced by normal sexual reproduction, whereas daughter queens are invariably clonally produced. Because males usually develop from unfertilized maternal eggs in ants and other haplodiploid species, they normally achieve direct fitness only through diploid female offspring. Hence, although the clonal production of queens increases the queen's relatedness to reproductive daughters, it potentially reduces male reproductive success to zero. In an apparent response to this conflict between sexes, genetic analyses reveal that males reproduce clonally, most likely by eliminating the maternal half of the genome in diploid eggs. As a result, all sons have nuclear genomes identical to those of their father. The obligate clonal production of males and queens from individuals of the same sex effectively results in a complete separation of the male and female gene pools. These findings show that the haplodiploid sex-determination system provides grounds for the evolution of extraordinary genetic systems and new types of sexual conflict.

New fossil discoveries illustrate the diversity of past terrestrial ecosystems in New Caledonia.

New Caledonia was, until recently, considered an old continental island harbouring a rich biota with outstanding Gondwanan relicts. However, deep marine sedimentation and tectonic evidence suggest complete submergence of the island during the latest Cretaceous to the Paleocene. Molecular phylogenies provide evidence for some deeply-diverging clades that may predate the Eocene and abundant post-Oligocene colonisation events. Extinction and colonization biases, as well as survival of some groups in refuges on neighbouring paleo-islands, may have obscured biogeographic trends over long time scales. Fossil data are therefore crucial for understanding the history of the New Caledonian biota, but occurrences are sparse and have received only limited attention. Here we describe five exceptional fossil assemblages that provide important new insights into New Caledonia's terrestrial paleobiota from three key time intervals: prior to the submersion of the island, following re-emergence, and prior to Pleistocene climatic shifts. These will be of major importance for elucidating changes in New Caledonia's floristic composition over time.

Effects of Urbanization on Plant-Pollinator Interactions in the Tropics: An Experimental Approach Using Exotic Plants.

Land-use changes through urbanization and biological invasions both threaten plant-pollinator networks. Urban areas host modified bee communities and are characterized by high proportions of exotic plants. Exotic species, either animals or plants, may compete with native species and disrupt plant-pollinator interactions. These threats are heightened in insular systems of the Southwest Pacific, where the bee fauna is generally poor and ecological networks are simplified. However, the impacts of these factors have seldom been studied in tropical contexts. To explore those questions, we installed experimental exotic plant communities in urban and natural contexts in New Caledonia, a plant diversity hotspot. For four weeks, we observed plant-pollinator interactions between local pollinators and our experimental exotic plant communities. We found a significantly higher foraging activity of exotic wild bees within the city, together with a strong plant-pollinator association between two exotic species. However, contrary to our expectations, the landscape context (urban vs. natural) had no effect on the activity of native bees. These results raise issues concerning how species introduced in plant-pollinator networks will impact the reproductive success of both native and exotic plants. Furthermore, the urban system could act as a springboard for alien species to disperse in natural systems and even invade them, leading to conservation concerns.

New data on the aphid (Hemiptera, Aphididae) fauna of New Caledonia: some new biosecurity threats in a biodiversity hotspot.

Thirty-three species of aphids are now established in New Caledonia. All species appear to have been introduced accidentally by human activity in the last century. Here, 17 aphid species are recorded for the first time: Aphis eugeniae, Aphis glycines, Aphis odinae, Aulacorthum solani, Brachycaudus helichrysi, Cerataphis orchidearum, Greenidea psidii, Hyperomyzus carduellinus, Hysteroneura setariae, Lipaphis pseudobrassicae, Micromyzus katoi, Myzus ornatus, Pentalonia caladii, Rhopalosiphum nymphaeae, Rhopalosiphum rufiabdominale, Schizaphis rotundiventris, and Tetraneura fusiformis. Thirteen more species are also more or less regularly intercepted at the borders through biosecurity surveys, without further establishment. This demonstrates that aphids represent a major biosecurity threat, including a threat as potential plant virus vectors. The reinforcement of biosecurity is a priority for such biodiversity hotspots, from the perspectives of both agriculture and the native environment. Prioritisation and promotion of local development of vegetable and fruit production, rather than their risky importation from abroad, is desirable. Such an approach also should be promoted and extended to other Pacific islands, which all share the lack of native aphid fauna and their associated plant disease vector risks.

RésuméTrente trois (33) espèces de pucerons sont aujourd'hui recensées de Nouvelle-Calédonie. Toutes ces espèces sont exotiques et ont été introduites accidentellement par les activités humaines. Dix-sept (17) espèces y sont ainsi recensées pour la première fois : Aphis eugeniae, Aphis glycines, Aphis odinae, Aulacorthum solani, Brachycaudus helichrysi, Cerataphis orchidearum, Greenidea psidii, Hyperomyzus carduellinus, Hysteroneura setariae, Lipaphis pseudobrassicae, Micromyzus katoi, Myzus ornatus, Pentalonia caladii, Rhopalosiphum nymphaeae, Rhopalosiphum rufiabdominale, Schizaphis rotundiventris et Tetraneura fusiformis. Par ailleurs, au moins 13 autres espèces sont régulièrement interceptées par la biosécurité sans établissement actuel de populations. Les pucerons apparaissent donc comme une menace croissante pour la biosécurité de l'archipel. Aussi, le renforcement des mesures de biosécurité aux frontières apparaît prioritaire en association à une promotion du développement local de productions maraîchères et fruitières. Ainsi, la limitation de ces importations à risque, devrait contribuer à une meilleure protection des productions agricoles et de la biodiversité. Une telle approche devrait également être promue dans les pays insulaires du Pacifique, qui se caractérisent par la même disharmonie de peuplements, l'absence de communautés natives de pucerons et du risque associé de vectorisation de maladies phytopathogènes.

Latrodectism in New Caledonia: first report of presumed redback spider (Latrodectus hasselti) envenomation.

We describe a recent case of presumed redback spider (Latrodectus hasselti) envenomation observed near Noumea in New Caledonia. This is the first local reported case in this archipelago, where L. hasselti is currently considered a native species. We highlight the challenge of providing appropriate analgesia in this situation and believe that the use of specific redback spider antivenom should be considered in New Caledonia.

Distribution, habitats, phenology and conservation of New Caledonian Odonata.

Compared to other archipelagos of the Pacific, the New Caledonian Odonata fauna is rich and diverse with 56 valid species or subspecies (23 endemics, 41%) from eight families (four Zygoptera: Argiolestidae, Coenagrionidae, Isostictidae, Lestidae, and four Anisoptera: Aeshnidae, Corduliidae, Synthemistidae, Libellulidae) and 31 genera (including four endemics, 13%). In Zygoptera, we record 19 species including 12 endemics (63%), and among Anisoptera, we record 37 species or subspecies, including 11 endemics (30%). We removed five species from the list that had been erroneously recorded as occurring in New Caledonia: Tramea carolina (Linnaeus, 1763), Austroargiolestes icteromelas (Selys-Longchamps, 1862), Ischnura torresiana Tillyard, 1913, Xiphiagrion cyanomelas Selys-Longchamps, 1876 and Hemicordulia oceanica Selys-Longchamps, 1871. The occurrence of Tramea limbata (Desjardins, 1835) appears also doubtful, but we were unable to clarify to which taxon this record referred hence we excluded it from our update. From a biogeographic perspective, the New Caledonian fauna has mostly Australian affinities with some connections with southeast Asia and the Pacific region. We provide for each species, whenever information was available, a distribution map with a brief review of its known ecology, behaviour and phenology. We also evaluated each species' conservation status, in light of known threats (range restriction, scarcity and human activity including altered water flow). We consider seventeen species (30%) endangered. The most immediate threats concern water pollution including alteration to the flow of water courses caused by mining, deforestation and fires. Invasive species, such as alien fish, may be predators of concern for odonata larva, although this has not yet been proven in New Caledonia.

Globally important islands where eradicating invasive mammals will benefit highly threatened vertebrates.

Invasive alien species are a major threat to native insular species. Eradicating invasive mammals from islands is a feasible and proven approach to prevent biodiversity loss. We developed a conceptual framework to identify globally important islands for invasive mammal eradications to prevent imminent extinctions of highly threatened species using biogeographic and technical factors, plus a novel approach to consider socio-political feasibility. We applied this framework using a comprehensive dataset describing the distribution of 1,184 highly threatened native vertebrate species (i.e. those listed as Critically Endangered or Endangered on the IUCN Red List) and 184 non-native mammals on 1,279 islands worldwide. Based on extinction risk, irreplaceability, severity of impact from invasive species, and technical feasibility of eradication, we identified and ranked 292 of the most important islands where eradicating invasive mammals would benefit highly threatened vertebrates. When socio-political feasibility was considered, we identified 169 of these islands where eradication planning or operation could be initiated by 2020 or 2030 and would improve the survival prospects of 9.4% of the Earth's most highly threatened terrestrial insular vertebrates (111 of 1,184 species). Of these, 107 islands were in 34 countries and territories and could have eradication projects initiated by 2020. Concentrating efforts to eradicate invasive mammals on these 107 islands would benefit 151 populations of 80 highly threatened vertebrates and make a major contribution towards achieving global conservation targets adopted by the world's nations.

Updating the Phylogenetic Dating of New Caledonian Biodiversity with a Meta-analysis of the Available Evidence.

For a long time, New Caledonia was considered a continental island, a fragment of Gondwana harbouring old clades that originated by vicariance and so were thought to be locally ancient. Recent molecular phylogenetic studies dating diversification and geological data indicating important events of submergence during the Paleocene and Eocene (until 37 Ma) brought evidence to dismiss this old hypothesis. In spite of this, some authors still insist on the idea of a local permanence of a Gondwanan biota, justifying this assumption through a complex scenario of survival by hopping to and from nearby and now-vanished islands. Based on a comprehensive review of the literature, we found 40 studies dating regional clades of diverse organisms and we used them to test the hypothesis that New Caledonian and inclusive Pacific island clades are older than 37 Ma. The results of this meta-analysis provide strong evidence for refuting the hypothesis of a Gondwanan refuge with a biota that originated by vicariance. Only a few inclusive Pacific clades (6 out of 40) were older than the oldest existing island. We suggest that these clades could have extinct members either on vanished islands or nearby continents, emphasizing the role of dispersal and extinction in shaping the present-day biota.

Seabirds supply nitrogen to reef-building corals on remote Pacific islets.

Seabirds concentrate nutrients from large marine areas on their nesting islands playing an important ecological role in nutrient transfer between marine and terrestrial ecosystems. Here we investigate the role of guano on corals reefs across scales by analyzing the stable nitrogen isotopic (δ15N) values of the scleractinian coral Pocillopora damicornis on fringing reefs around two Pacific remote islets with large seabird colonies. Marine stations closest to the seabird colonies had higher nitrate + nitrite concentrations compared to more distant stations. Coral and zooxanthellae δ15N values were also higher at these sites, suggesting that guano-derived nitrogen is assimilated into corals and contributes to their nitrogen requirements. The spatial extent of guano influence was however restricted to a local scale. Our results demonstrate that seabird-derived nutrients not only spread across the terrestrial ecosystem, but also affect components of the adjacent marine ecosystem. Further studies are now needed to assess if this nutrient input has a positive or negative effect for corals. Such studies on remote islets also open fresh perspectives to understand how nutrients affect coral reefs isolated from other anthropogenic stressors.

Rare sexual reproduction events in the clonal reproduction system of introduced populations of the little fire ant.

A unique reproductive system has previously been described in Wasmannia auropunctata, a widespread invasive ant species, where males are produced clonally, female queens are parthenogens, and female workers are produced sexually. However, these findings were mostly based on samples originating from only a limited part of the native range of the species in South America. We used microsatellite markers to uncover the reproductive modes displayed by a large number of nests collected in various invasive W. auropunctata populations introduced 40 years ago into New Caledonia, where the species now forms a single 450-km-long supercolony. Although the main reproduction system in New Caledonia remained clonality for both male and female reproductives, we found evidence of rare sexual reproduction events that led to the production of both new queen and male clonal lineages. All clonal lineages observed in New Caledonia potentially derived from sexual reproduction, recombination, and mutation events from a single female and a single male genotype. Hence, the male and female gene pools are not strictly separated in New Caledonia and the two sexes do not follow independent evolutionary trajectories. Our results also suggest genetic determination for both parthenogenesis and caste. We discuss the evolutionary implications of the emergence of sex in the clonal reproduction system of introduced populations of W. auropunctata.

A review of the genus Kanakia Distant, 1892 (Insecta: Hemiptera, Cicadoidea, Cicadidae) from New Caledonia.

Among New Caledonian cicada taxa, the endemic genus Kanakia Distant, 1892 appears as the most spectacular one, especially because of the large size of its species (length up to 50 mm for males). Recent new specimen collections helped to clarify the taxonomy of this spectacular genus. According to morphological characteristics, we were able to redefine and to split the present genus into three distinct genera: Kanakia Distant, 1892, Pseudokanakia Delorme gen. nov. and Panialna Delorme gen. nov. These two new genera are monotypic and have been respectively established from revision of Kanakia flavoannulata (Distant 1920) and Kanakia parva Boulard 1991. Also, the type species, Kanakia typica Distant, 1892, appears to be a complex of cryptic species. New acoustic and morphological observations allowed us to redefine K. typica and to describe four new species in this complex: Kanakia paniensis Delorme sp. nov., Kanakia rana Delorme sp. nov., Kanakia salesnii Delorme sp. nov. and Kanakia fuscocosta Delorme sp. nov. Kanakia gigas Boulard 1988 is also briefly discussed and the female is described. An identification key of the Kanakia species and allied genera is also provided.

Description of a new genus and two new species of high frequency cicada from New Caledonia (Insecta: Hemiptera, Cicadoidea, Cicadidae).

The new genus Murmurillana Delorme gen. nov., is described within the tribe Cicadettini Buckton, 1889, designating Murmurillana inaudibilis Delorme sp. nov., as the type species. Murmurillana inaudibilis Delorme sp. nov. and Murmurillana paenetacita Delorme sp. nov. are described from New Caledonia. They are respectively found in mid altitude dense Niaouli shrub (Melaleuca quinquenervia, Myrtaceae) vegetation, mixed with dense fern cover (Pteridium sp., Dennstaedtiaceae) on the Massif of Aoupinié (800 m) and on foothills of the Mont Panié (570 m). Male calling songs of the two new species are described from field recordings. These calling songs exhibit unusually high dominant frequencies. A key to the species of Murmurillana Delorme gen. nov., is also provided.

Crickets (Insecta, Orthoptera, Grylloidea) from Southern New Caledonia, with descriptions of new taxa.

Intensive sampling of cricket communities has been undertaken in southern New Caledonia in selected plots of vegetation, i.e. rain forest, preforest and maquis shrubland. This leads to the discovery of many new taxa, which are described in the present paper, together with closely related species from nearby areas. Descriptions are based on general morphology and characters of genitalia. Calling songs are described for all acoustic taxa but two, and observations about species habitats are given. In total, 35 species belonging to 13 genera are studied, including 21 new species and two new genera. The pattern of assemblages of cricket species in New Caledonia is discussed.

Knocking on Heaven's Door: Are Novel Invaders Necessarily Facing Naïve Native Species on Islands?

The impact of alien predator species on insular native biota has often been attributed to island prey naïveté (i.e. lack of, or inefficient, anti-predator behavior). Only rarely, however, has the concept of island prey naïveté been tested, and then only a posteriori (i.e. hundreds or thousands of years after alien species introduction). The presence of native or anciently introduced predators or competitors may be crucial for the recognition and development of adaptive behavior toward unknown predators or competitors of the same archetype (i.e. a set of species that occupy a similar ecological niche and show similar morphological and behavioral traits when interacting with other species). Here, we tested whether two squamates endemic to New Caledonia, a skink, Caledoniscincus austrocaledonicus, and a gecko, Bavayia septuiclavis, recognized and responded to the odor of two major invaders introduced into the Pacific islands, but not yet into New Caledonia. We chose one predator, the small Indian mongoose Herpestes javanicus and one competitor, the cane toad Rhinella marina, which belong respectively to the same archetype as the following two species already introduced into New Caledonia in the nineteenth century: the feral cat Felis catus and the golden bell frog Litoria aurea. Our experiment reveals that geckos are naïve with respect to the odors of both an unknown predator and an unknown competitor, as well as to the odors of a predator and a competitor they have lived with for centuries. In contrast, skinks seem to have lost some naïveté regarding the odor of a predator they have lived with for centuries and seem "predisposed" to avoid the odor of an unknown potential competitor. These results indicate that insular species living in contact with invasive alien species for centuries may be, although not systematically, predisposed toward developing adaptive behavior with respect to species belonging to the same archetype and introduced into their native range.

The gasteruptiid wasp fauna of New Caledonia, with description of three new species of Gasteruption (Hymenoptera: Evanioidea: Gasteruptiidae).

The rarely collected gasteruptiid wasp fauna (Evanioidea: Gasteruptiidae) of New Caledonia is reviewed. Previously only two species of Pseudofoenus (Hyptiogastrinae) were known. Here, we record the subfamily Gasteruptiinae from New Caledonia for the first time and describe three new species of Gasteruption: G. lacoulee Jennings, Krogmann & Parslow, sp. nov., G. maquis Jennings, Krogmann & Parslow, sp. nov., and G. sarramea Jennings, Krogmann & Parslow, sp. nov. An identification key to the Gasteruptiidae of New Caledonia is provided.