Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura)

<p>Abstract</p> <p>Background</p> <p>Sucking lice (Phthiraptera: Anoplura) are obligate, permanent ectoparasites of eutherian mammals, parasitizing members of 12 of the 29 recognized mammalian orders and approximately 20% of all mammalian species. These host specific, blood-sucking insects are morphologically adapted for life on mammals: they are wingless, dorso-ventrally flattened, possess tibio-tarsal claws for clinging to host hair, and have piercing mouthparts for feeding. Although there are more than 540 described species of Anoplura and despite the potential economical and medical implications of sucking louse infestations, this study represents the first attempt to examine higher-level anopluran relationships using molecular data. In this study, we use molecular data to reconstruct the evolutionary history of 65 sucking louse taxa with phylogenetic analyses and compare the results to findings based on morphological data. We also estimate divergence times among anopluran taxa and compare our results to host (mammal) relationships.</p> <p>Results</p> <p>This study represents the first phylogenetic hypothesis of sucking louse relationships using molecular data and we find significant conflict between phylogenies constructed using molecular and morphological data. We also find that multiple families and genera of sucking lice are not monophyletic and that extensive taxonomic revision will be necessary for this group. Based on our divergence dating analyses, sucking lice diversified in the late Cretaceous, approximately 77 Ma, and soon after the Cretaceous-Paleogene boundary (ca. 65 Ma) these lice proliferated rapidly to parasitize multiple mammalian orders and families.</p> <p>Conclusions</p> <p>The diversification time of sucking lice approximately 77 Ma is in agreement with mammalian evolutionary history: all modern mammal orders are hypothesized to have diverged by 75 Ma thus providing suitable habitat for the colonization and radiation of sucking lice. Despite the concordant timing of diversification events early in the association between anoplurans and mammals, there is substantial conflict between the host and parasite phylogenies. This conflict is likely the result of a complex history of host switching and extinction events that occurred throughout the evolutionary association between sucking lice and their mammalian hosts. It is unlikely that there are any ectoparasite groups (including lice) that tracked the early and rapid radiation of eutherian mammals.</p

Review of endemic Sulawesi squirrels.

260 p. : ill. (some col.), maps ; 26 cm. "Issued June 11, 2010." Includes bibliographical references (p. 246-255).Analyses of fur color patterns, morphometric data derived from external, cranial, and dental dimensions, and distributions of collection sites for voucher specimens form the basis for a taxonomic revision of Sulawesi’s endemic squirrel fauna. Eight species of tree squirrels in Rubrisciurus and Prosciurillus and two species of ground squirrels in Hyosciurus are recognized. All are diurnal and inhabit primary forest formations. Diet consists of fruit, nuts, seeds, and arthropods. Rubrisciurus rubriventer, the largest in body size, forages on the ground and in the lower canopy layer, is found throughout Sulawesi where primary forest persists, and occurs through an altitudinal range embracing tropical lowland evergreen and lower montane rain forests; it is absent from upper montane rain forest. Five species of arboreal squirrels comprise the Prosciurillus leucomus group, a cluster of species occupying the upper forest canopy: P. leucomus, known only from lowland and montane habitats in the northern peninsula and one offshore island; P. alstoni, recorded from lowland tropical evergreen rain forest in the eastern section of Sulawesi’s central core, the east-central and southeastern arms, and two southeastern islands; P. weberi, represented by a few specimens from the coastal lowlands of the southern core of Sulawesi; P. topapuensis, endemic to the western mountain block in Sulawesi’s central core and occurring along an altitudinal gradient from lowland evergreen rain forest to upper montane rain forest; and P. rosenbergii, the only species of squirrel collected on islands in the Sangihe Archipelago north of the northeastern tip of the northern peninsula. The Prosciurillus murinus group contains two species of small body size: P. murinus, found throughout Sulawesi and in all forest formations, from the coastal lowlands to mountaintops, and a forager in the lower canopy layers; and P. abstrusus, known only from montane forest habitats on Pegunungan Mekongga in the southeastern peninsula. Of two species of ground squirrels, Hyosciurus heinrichi occupies montane forest habitats in the western mountain block of Sulawesi’s central core. It is altitudinally parapatric to H. ileile, which inhabits lowland evergreen and lower montane rain forests in the western mountain block and northeastern lowlands of central Sulawesi, and montane forest on the northern peninsula. A slightly revised classification of Sciuridae is provided in which a new tribe, Exilisciurini, is proposed for the Bornean and Philippine Exilisciurus. Previously published results of morphological and molecular analyses point to Rubrisciurus, Prosciurillus, and Hyosciurus as a monophyletic cluster, the Hyosciurina, nested within a larger clade, the Nannosciurini, which along with Exilisciurini n. tribe and Funambulini, comprise the Nannosciurinae, one of the three subfamilies constituting Sciuridae, and one that contains most of the Indomalayan genera. The present diversity of species endemic to Sulawesi was derived from an ancient lineage that crossed a sea barrier from the Sunda Shelf to Sulawesi during the late Miocene. Eight new species of hoplopleurid sucking lice ‪(‬Insecta, Anoplura‪)‬ are described as parasitizing 8 of the 10 species of squirrels endemic to Sulawesi: Hoplopleura rubrisciuri from Rubrisciurus rubriventer, Hoplopleura leucomus from Prosciurillus leucomus, Hoplopleura alstoni from Prosciurillus alstoni, Hoplopleura topapuensis from Prosciurillus topapuensis, Hoplopleura murinus from Prosciurillus murinus, Hoplopleura abstrusus from Prosciurillus abstrusus, Hoplopleura heinrichi from Hyosciurus heinrichi, and Hoplopleura ileile from Hyosciurus ileile. Examples of Prosciurillus weberi and P. rosenbergii were surveyed but no lice were recovered. A phylogenetic analysis based on cladistic principles for six species of Sulawesian squirrel lice for which both sexes were available is presented and the results discussed with respect to host relationships. These new data are incorporated into a discussion covering zoogeography of global sciurid-sucking louse associations, emphasizing the Indomalayan squirrel fauna. Globally, members of 11 genera of Anoplura parasitize sciurids, a figure far exceeding the number of anopluran genera associated with any other mammalian family. Nine of these ‪(‬the enderleinellids, Atopophthirus, Enderleinellus, Microphthirus, Phthirunculus, and Werneckia; the hoplopleurid, Paradoxophthirus; and the polyplacids, Johnsonpthirus, Linognathoides, and Neohaematopinus‪)‬ are primary parasites of sciurids. The remaining two ‪(‬the hoplopleurid Hoplopleura, and the polyplacid Polyplax‪)‬ include representatives that are acquired ‪(‬secondary‪)‬ parasites of sciurids--the majority of species in these two louse genera parasitize other groups of mammals but a small number of species have transferred to squirrel hosts. Sciurid hosts and geographic distributions of these 11 anopluran genera are discussed. Historically, representatives of Hoplopleura colonized different sciurid hosts on several separate occasions with one known species on a North American tree squirrel ‪(‬Sciurus‪)‬, two described species on North American flying squirrels ‪(‬Glaucomys‪)‬, two species parasitizing North American species of chipmunks ‪(‬Tamias and Eutamias‪)‬, 14 described species from Indomalayan nannosciurine squirrels ‪(‬Callosciurus, Tamiops, Rubrisciurus, Prosciurillus, Hyosciurus, and Funambulus‪)‬, and one species parasitizing a Chinese xerine ground squirrel ‪(‬Sciurotamias‪)‬. The zoogeography of the seven sciurid-infesting louse genera known from Southeast Asia is discussed using data from nine different countries or regions ‪(‬China, Taiwan, Thailand, peninsular Malaysia, Sumatra, Java, Borneo, the Philippines, and Sulawesi‪)‬. A reduction in the number of sciuirid-infesting anopluran genera and species is apparent from mainland northern/western regions to insular southern/eastern regions with members of seven genera and 23 species described from China but only one genus and eight species from Sulawesi. The absence of known species of Hoplopleura from Bornean and Javanese squirrels suggests that such a fauna may await discovery on one or both of these islands. Six of the eight species of Hoplopleura found parasitizing species of endemic Sulawesi squirrels were recovered as a monophyletic clade from a phylogenetic analysis employing anatomical structures associated with male and female lice. ‪(‬Two species of Sulawesi Hoplopleura are based on females and nymphs only and were not incorporated into the analysis.‪)‬ The monophyletic cluster formed by the Sulawesian squirrel lice joined with the monophyletic assemblage containing the three Sulawesi squirrel genera--Rubrisciurus, Prosciurillus, and Hyosciurus--suggest that the ancestral squirrel lineage that arrived in Sulawesi during the late Miocene may have been carrying its unique Hoplopleura parasite

Integrating “Big Data” into Aquatic Ecology: Challenges and Opportunities

Got “Big Data”? Not sure how best to use it? Big Data is becoming an important facet of aquatic ecology, and researchers must learn to harness it to reap the rewards of using it. The benefits of using Big Data are many, and include advancements in scientific understanding at larger scales and higher resolution, applications to improving environmental management and policy, and public engagement. We aim to demystify the use of Big Data for individual scientists, and provide some food for thought for the aquatic ecology community on how to develop this sphere. To achieve this, we highlight six key challenges: (1) how to recognize if you have Big Data, (2) handling Big Data, (3) issues with classical analytical techniques, (4) verification of Big Data, (5) considerations for data sharing, and (6) community development of knowledge infrastructures. We then present approaches and tools which have been successfully applied to these challenges in aquatic ecology and other scientific fields

Report of the Workshop Evaluating the Nature of Midwater Mining Plumes and Their Potential Effects on Midwater Ecosystems

The International Seabed Authority (ISA) is developing regulations to control the future exploitation of deep-sea mineral resources including sulphide deposits near hydrothermal vents, polymetallic nodules on the abyssal seafloor, and cobalt crusts on seamounts. Under the UN Convention on the Law of the Sea the ISA is required to adopt are taking measures to ensure the effective protection of the marine environment from harmful effects arising from mining-related activities. Contractors are required to generate environmental baselines and assess the potential environmental consequences of deep seafloor mining. Understandably, nearly all environmental research has focused on the seafloor where the most direct mining effects will occur. However, sediment plumes and other impacts (e.g., noise) from seafloor mining are likely to be extensive in the water column. Sediment plumes created on the seafloor will affect the benthic boundary layer which extends 10s to 100s of meters above the seafloor. Separation or dewatering of ore from sediment and seawater aboard ships will require discharge of a dewatering plume at some depth in the water column. It is important to consider the potential impacts of mining on the ocean’s midwaters (depths from ~200 m to the seafloor) because they provide vital ecosystem services and harbor substantial biodiversity. The better known epipelagic or sunlit surface ocean provisions the rest of the water column through primary production and export flux (This was not the focus at this workshop as the subject was considered too large and surface discharges are unlikely). It is also home to a diverse community of organisms including commercially important fishes such as tunas, billfish, and cephalopods that contribute to the economies of many countries. The mesopelagic or twilight zone (200-1000 m) is dimly lit and home to very diverse and abundant communities of organisms. Mesopelagic plankton and small nekton form the forage base for many deep-diving marine mammals and commercially harvested epipelagic species. Furthermore, detritus from the epipelagic zone falls through the mesopelagic where it is either recycled, providing the vital process of nutrient regeneration, or sinks to greater depths sequestering carbon from short-term atmospheric cycles. The waters below the mesopelagic down to the seafloor (both the bathypelagic and abyssopelagic) are very poorly characterized but are likely large reservoirs of novel biodiversity and link the surface and benthic ecosystems. Great strides have been made in understanding the biodiversity and ecosystem function of the ocean’s midwaters, but large regions, including those containing many exploration license areas and the greater depths where mining plumes will occur, remain very poorly studied. It is clear that pelagic communities are distinct from those on the seafloor and in the benthic boundary layer. They are often sampled with different instrumentation. The fauna have relatively large biogeographic ranges and they are more apt to mix freely across stakeholder boundaries, reference areas and other spatial management zones. Pelagic organisms live in a three-dimensional habitat and their food webs and populations are vertically connected by daily or lifetime migrations and the sinking flux of detritus from the epipelagic. The fauna do not normally encounter hard surfaces, making them fragile, and difficult to capture and maintain for sensitivity or toxicity studies. Despite some existing general knowledge, ecological baselines for midwater communities and ecosystems that likely will be impacted by mining have not been documented. There is an urgent need to conduct more research and evaluate the midwater biota (microbes to fishes) in regions where mining is likely to occur. Deep-sea mining activities may affect midwater organisms in a number of ways, but it is still unclear at what scale perturbations may occur. The sediment plumes both from collectors on the seafloor and from midwater discharge will have a host of negative consequences. They may cause respiratory distress from clogged gills or respiratory surfaces. Suspension feeders, such as copepods, polychaetes, salps, and appendicularians, that filter small particles from the water and form an important basal group of the food web, may suffer from dilution of their food by inorganic sediments and/or clogging of their fragile mucous filter nets. Small particles may settle on gelatinous plankton causing buoyancy issues. Metals, including toxic elements that will enter the food web, will be released from pore waters and crushed ore materials. Sediment plumes will also absorb light and change backscatter properties, reducing visual communication and bioluminescent signaling that are very important for prey capture and reproduction in midwater animals. Noise from mining activities may alter the behaviors of marine mammals and other animals. Small particles have high surface area to volume ratios, high pelagic persistence and dispersal and as a result greater potential to result in pelagic impacts. All of these potential effects will result in mortality, migration (both horizontal and vertical), decreased fitness, and shifts in community composition. Depending on the scale and duration of these effects, there could be reduction in provisioning to commercial fish species, delivery of toxic metals to pelagic food webs and hence human seafood supply, and alterations to carbon transport and nutrient regeneration services. After four days of presentations and discussions, the workshop participants came to several conclusions and synthesized recommendations. 1. Assuming no discharge in the epipelagic zone, it is essential to minimize mining effects in the mesopelagic zone because of links to our human seafood supply as well as other ecosystem services provided by the mesopelagic fauna. This minimization could be accomplished by delivering dewatering discharge well below the mesopelagic/bathypelagic transition (below ~1000 m depth). 2. Research should be promoted by the ISA and other bodies to study the bathypelagic and abyssopelagic zones (from ~1000 m depths to just above the seafloor). It is likely that both collector plumes and dewatering plumes will be created in the bathypelagic, yet this zone is extremely understudied and contains major unknowns for evaluating mining impacts. 3. Management objectives, regulations and management actions need to prevent the creation of a persistent regional scale “haze” (enhanced suspended particle concentrations) in pelagic midwaters. Such a haze would very likely cause chronic harm to deep midwater ecosystem biodiversity, structure and function. 4. Effort is needed to craft suitable standards, thresholds, and indicators of harmful environmental effects that are appropriate to pelagic ecosystems. In particular, suspension feeders are very important ecologically and are likely to be very sensitive to sediment plumes. They are a high priority for study. 5. Particularly noisy mining activities such as ore grinding at seamounts and hydrothermal vents is of concern to deep diving marine mammals and other species. One way to minimize sound impacts would be to minimize activities in the sound-fixing-and-ranging (SOFAR) channel (typically at depths of ~1000 m) which transmits sounds over very long distances. 6. A Lagrangian (drifting) perspective is needed in monitoring and management because the pelagic ecosystem is not a fixed habitat and mining effects are likely to cross spatial management boundaries. For example, potential broad-scale impacts to pelagic ecosystems should be considered in the deliberations over preservation reference zones, the choice of stations for environmental baseline and monitoring studies and other area-based management and conservation measures. 7. Much more modeling and empirical study of realistic mining sediment plumes is needed. Plume models will help evaluate the spatial and temporal extent of pelagic (as well as benthic) ecosystem effects and help to assess risks from different technologies and mining scenarios. Plume modeling should include realistic mining scenarios (including duration) and assess the spatial-temporal scales over which particle concentrations exceed baseline levels and interfere with light transmission to elucidate potential stresses on communities and ecosystem services. Models should include both near and far field-phases, incorporating realistic near field parameters of plume generation, flocculation, particle sinking, and other processes. It is important to note that some inputs to these models such as physical oceanographic parameters are lacking and should be acquired in the near-term. Plume models need to be complemented by studies to understand effects on biological components by certain particle sizes and concentrations

Too Big to Fail — U.S. Banks’ Regulatory Alchemy: Converting an Obscure Agency Footnote into an “At Will” Nullification of Dodd-Frank’s Regulation of the Multi-Trillion Dollar Financial Swaps Market

The multi-trillion-dollar market for, what was at that time wholly unregulated, over-the-counter derivatives (“swaps”) is widely viewed as a principal cause of the 2008 worldwide financial meltdown. The Dodd-Frank Act, signed into law on July 21, 2010, was expressly considered by Congress to be a remedy for this troublesome deregulatory problem. The legislation required the swaps market to comply with a host of business conduct and anti-competitive protections, including that the swaps market be fully transparent to U.S. financial regulators, collateralized, and capitalized. The statute also expressly provides that it would cover foreign subsidiaries of big U.S. financial institutions if their swaps trading could adversely impact the U.S. economy or represent the use of extraterritorial trades as an attempt to “evade” Dodd-Frank. In July 2013, the CFTC promulgated an 80-page, triple-columned, and single-spaced “guidance” implementing Dodd-Frank’s extraterritorial reach, i.e., that manner in which Dodd-Frank would apply to swaps transactions executed outside the United States. The key point of that guidance was that swaps trading within the “guaranteed” foreign subsidiaries of U.S. bank holding company swaps dealers were subject to all of Dodd-Frank’s swaps regulations wherever in the world those subsidiaries’ swaps were executed. At that time, the standardized industry swaps agreement contemplated that, inter alia, U.S. bank holding company swaps dealers’ foreign subsidiaries would be “guaranteed” by their corporate parent, as was true since 1992. In August 2013, without notifying the CFTC, the principal U.S. bank holding company swaps dealer trade association privately circulated to its members standard contractual language that would, for the first time, “deguarantee” their foreign subsidiaries. By relying only on the obscure footnote 563 of the CFTC guidance’s 662 footnotes, the trade association assured its swaps dealer members that the newly deguaranteed foreign subsidiaries could (if they so chose) no longer be subject to Dodd-Frank. As a result, it has been reported (and it also has been understood by many experts within the swaps industry) that a substantial portion of the U.S. swaps market has shifted from the large U.S. bank holding companies swaps dealers and their U.S. affiliates to their newly deguaranteed “foreign” subsidiaries, with the attendant claim by these huge big U.S. bank swaps dealers that Dodd-Frank swaps regulation would not apply to these transactions. The CFTC also soon discovered that these huge U.S. bank holding company swaps dealers were “arranging, negotiating, and executing” (“ANE”) these swaps in the United States with U.S. bank personnel and, only after execution in the U.S., were these swaps formally “assigned” to the U.S. banks’ newly “deguaranteed” foreign subsidiaries with the accompanying claim that these swaps, even though executed in the U.S., were not covered by Dodd-Frank. In October 2016, the CFTC proposed a rule that would have closed the “deguarantee” and “ANE” loopholes completely. However, because it usually takes at least a year to finalize a “proposed” rule, this proposed rule closing the loopholes in question was not finalized prior to the inauguration of President Trump. All indications are that it will never be finalized during a Trump Administration. Thus, in the shadow of the recent tenth anniversary of the Lehman failure, there is an understanding among many market regulators and swaps trading experts that large portions of the swaps market have moved from U.S. bank holding company swaps dealers and their U.S. affiliates to their newly deguaranteed foreign affiliates where Dodd- Frank swaps regulation is not being followed. However, what has not moved abroad is the very real obligation of the lender of last resort to rescue these U.S. swaps dealer bank holding companies if they fail because of poorly regulated swaps in their deguaranteed foreign subsidiaries, i.e., the U.S. taxpayer. While relief is unlikely to be forthcoming from the Trump Administration or the Republican-controlled Senate, some other means will have to be found to avert another multi-trillion-dollar bank bailout and/or a financial calamity caused by poorly regulated swaps on the books of big U.S. banks. This paper notes that the relevant statutory framework affords state attorneys general and state financial regulators the right to bring so-called “parens patriae” actions in federal district court to enforce, inter alia, Dodd- Frank on behalf of a state’s citizens. That kind of litigation to enforce the statute’s extraterritorial provisions is now badly needed