Harold Heatwole

Harold Heatwole
  • North Carolina State University

About

215
Publications
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Introduction
Harold Heatwole currently works at the Department of Zoology, University of New England, Armidale NSW 2351, Australia, and in the Department of Biology, North Carolina State University, Raleigh, NC 27695-7617, USA. His research is in herpetology, especially sea snakes; the ecology of ants; ecology of tardigrades; and island biogeography.
Current institution
North Carolina State University

Publications

Publications (215)
Article
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Assemblages of terrestrial biotas in Antarctica have low species-diversity, taxonomic breadth, and number of trophic links and may provide insights not only into adaptation to extreme environments, but also into an understanding of community structure and dynamics not readily achieved by studying more complex, less tractable, systems. To this end,...
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Marine snakes represent the most speciose group of marine reptiles and are a significant component of reef and coastal ecosystems in tropical oceans. Research on this group has historically been challenging due to the difficulty in capturing, handling, and keeping these animals for field- and lab-based research. Inexplicable declines in marine snak...
Article
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Roughly 70 species of sea snakes inhabit the Indo-Pacific but are absent from the Atlantic Ocean. Paleoclimatic conditions in the Coral Triangle were favorable for evolutionary transitions to the sea, while those in the Caribbean region and coastlines bordering the Atlantic Ocean were less favorable. The dispersal of sea snakes from the Indian to A...
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The geographic range of sea kraits encompasses one of the geologically most-complex regions of the world. At its center lies Wallacea (the transition between the terrestrial biotas of the Asian and Australian tectonic plates) and the Indonesian Throughflow (nexus of the equatorial marine biotas of the Indian and Pacific oceans). The aim of this stu...
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Biology, including herpetology, has made greater strides in recent decades than in any time in history. It has progressed from a largely inductive science performed on an expeditionary basis to a laboratory-based discipline in which preformed hypotheses are tested empirically. My research has spanned that change in paradigm and an example of expedi...
Article
Water is an essential resource affecting behavior and the acquisition of energy, especially in environments where water is spatially or temporally restricted or unavailable. Recent investigations have shown that several species of marine snakes dehydrate at sea and are dependent on environmental sources of fresh water to maintain water balance. How...
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Dehydration and drinking behaviors were investigated in the little file snake (Acrochordus granulatus) collected from marine populations in the Philippines and in Australia. File snakes dehydrate in seawater and do not drink seawater when dehydrated in air and offered seawater to drink. Dehydrated file snakes drink freshwater, and the threshold of...
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Point diversity provides essential information about structuring of assemblages and interactions of component species, not achieved using other scales of analysis. A baited grid was used (1) to ascertain number of species of scavenging ants at point resources of food in an Australian eucalypt woodland at three times in the diel cycle and (2) to ass...
Chapter
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Leaves undergo ontogenetic changes in their toughness, chemical composition, and texture, all of which affect their palatability to herbivores and pathogens and their resistance to physical agents (Lowman and Box 1983; Lowman 1995; Coley and Kursar 1996; Coley et al. 2006). Consequently, rates of herbivory and other damage would be expected to chan...
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Abstract.—An assessment of marine elapid snakes found 9% of marine elapids are threatened with extinction, and an additional 6% are Near Threatened. A large portion (34%) is Data Deficient. An analysis of distributions revealed the greatest species diversity is found in Southeast Asia and northern Australia. Three of the seven threatened species oc...
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Previous studies have treated the status of decline and conservation in amphibians, generally and on a country by country basis, for the Western Hemisphere. The present volume presents a series of papers for the Eastern Hemisphere, and this issue covers North Africa. An annotated checklist to current nomenclature of regional taxa is also provided.
Article
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Erroneous locality records and those based solely on vagrants have provided an unrealistic assessment of the true distribution (area of occupancy) of breeding populations of sea kraits (genus Laticauda). This distortion over-estimates the extent of their geographic ranges and seriously under-estimates their conservation status.
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Spot checks and periodic day-long surveys of the wild mammals, birds, reptiles, and amphibians sold for food were carried out in the Lak Xao market and other markets in Laos. Birds were the predominant offering in terms of number of species, but amphibians, although represented by few species, were predominant in numbers of individuals. Mammals wer...
Article
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Temperature probably had no direct effect on the evolution of sea kraits within their center of origin, a geologically stable thermal zone straddling the equator, but may have indirectly affected expansions and contractions in distributions beyond that zone through global fluctuations that caused alternation of higher and lower sea levels. The nort...
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Why should we conserve biodiversity?: Peter Sale, in his important book Our Dying Planet (2011), presented and critically examined economic and ethical/esthetic arguments for conserving biodiversity. We refer the reader to that book for an overview. An ethical responsibility of humans towards nature is usually denied: man has responsibility towards...
Article
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It is clear that nature is undergoing rapid changes as a result of human activities such as industry, agriculture, travel, fisheries and urbanisation. What effects do these activities have? Are they disturbing equilibria in ecological populations and communities, thus upsetting the balance of nature, or are they enhancing naturally occurring disequ...
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ZusammenfassungIn freier Natur wurden bei Megarhyssa macrurus lunator, M. atrata lineata und M. greenei, die sich alle als Parasiten der in Holz bohrenden Larven von Tremex columba entwickeln, Verhaltensbeobachtungen gemacht, beschrieben und funktionell gedeutet. Alle drei Arten zeigten nahezu gleiche Verhaltensweisen, außer wenn Bauunterschiede, z...
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A sample of 1834 leaves from 83 plants in 26 families was collected in tropical rain forest in Madagascar from three vertical strata: top of emergent trees (up to 37 m), top of trees in upper canopy (about 22 m), and shrubs and saplings at ground level. These leaves were examined for damage by seven different agents: fungi, epiphyllae, mechanical i...
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Six species of tardigrades, Pseudechiniscus suillus, Macrobiotus sp., Hypsibius antarcticus, Ramajendas frigidus, Diphascon chilenense and Diphascon pingue were extracted from mosses and lichens from the ice-free regions of the Windmill Islands near Casey Base, East Antarctica. Significant positive associations were found between the three common s...
Article
Cold Pseudechis porphyriacus aid heating by basking, flattening and tilting. When body temperatures are higher, thermoregulation is achieved by shuttling between sun and shade. A warm snake in a cooling environment frequently coils. The major factors associated with rate of temperature change in the body core were (1) horizontal gradient between bo...
Article
The direction of coiling was periodically recorded for two species of viperid snakes--copperheads (Agkistrodon contortrix) and cottonmouths (Agkistrodon piscivorus). Overall, neither species showed a significant preference for coiling in a particular direction. Only 1 of 22 snakes exhibited an individual preference, a result within expectation for...
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Aipysurus laevis and Emydocephalus annulatus typically occur in spatially discrete populations, characteristic of metapopulations; however, little is known about the factors influencing the spatial and temporal stability of populations or whether specific conservation strategies, such as networks of marine protected areas, will ensure the persisten...
Data
Table 1: Status of Aipysurus laevis and Emydocephalus annulatus on 90 reefs in the Pompey and Swain reefs complexes, southern Great Barrier Reef surveyed between 1967 and 2002, including current and prior zoning status.
Article
The effect of spraying insecticide on eucalypt saplings was experimentally tested in a New England woodland. Replicate branches of five species were sprayed fortnightly with Thiodan over a period of 4 months. Their leaf damage, leaf flushes, stem growth, and bud production were compared with those of control branches. Loss of leaf area to insect gr...
Article
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The sea krait Laticauda colubrina is the most widespread member of its genus, extending from the Bay of Bengal through much of Asia and the Indo-Malayan Archipelago to New Guinea and many islands of the western Pacific Ocean. Unconfirmed records of the species may extend the range to the western coast of Central America. The species is subject to m...
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Seabirds were counted during four 24-hour periods, two in summer and two in winter, on each of two cays (Price Cay and Frigate Cay) in the Swain Reefs, Great Barrier Reef, Australia. There were six breeding species of seabirds and 13 nonbreeding ones. Each species showed its own pattern of diel change in number of individuals on the islands, some b...
Article
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The Laticauda colubrina complex previously consisted of three species, Laticauda saintgironsi from New Caledonia and the Loyalty Islands, Laticauda frontalis from Vanuatu, and Laticauda colubrina, a widespread species ranging from the Andaman and Nicobar Islands and the Myanmar-Thai-Malaysian peninsula, through the Indonesian archipelago to New Gui...
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A new Philautus is described from Phou Dendin National Biodiversity Conservation Area in northern Laos. Philautus petilus sp. nov. is most remarkable by having a very slender, elongate habitus. Other distinguishing characteristics include having a tympanum diameter 80% of the eye diameter, white asperities on the dorsum, and distinctive coloration...
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The callipodidan milliped, Sinocallipus simplipodicus Zhang, 1993, previously known only from a cave in Yunnan Province, China, is redescribed based on specimens from an epigean habitat in southern Laos, some 600 mi (960 km) south of the type locality. SEM photos of the gonopods, ovi- positor, gnathochilarium, midbody exoskeleton, an ozopore, and l...
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A 34-year study (1966-2000) of the lizards and snakes of the New England region of New South Wales yielded 14 species of geckos, 4 of pygopodids, 8 of dragons, 3 of goannas, 47 of skinks, and 30 of snakes, for a total of 106 species. Four additional species reported by Swan (1990), and an additional one from the Atlas of NSW Wildlife brings the tot...
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Sub-Antarctic Macquarie Island lies in the Southern Ocean between Tasmania and Antarctica and just above the Antarctic Convergence. Extensive flora and fauna samples were collected during the 1977—78 Australian Museum Expedition. Thirteen genera and 25 species of tardigrades are reported from among the 8725 specimens recovered from the 72 samples....
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The ultrastructure of two kinds of mechanoreceptive organs, pit organs and neuromasts, in the skin of adult giant salamanders (Andrias davidianus) was studied by transmission electron microscopy. Neuromasts and pit organs differ in their types of synapses, the spatial distribution of kinocilia on sensory cells, and in the degree to which sensory ce...
Article
The olive sea snake, Aipysurus laevis (Lacépède) grows at a rate of 0.22-0.95 cm/month, with young animals growing faster than older ones. Males reach sexual maturity in their third year and females in their fourth or fifth year. There is sexual dimorphism in size, with females larger than males; at snout-vent lengths greater than 80 cm, females ar...
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Ultrastructure of cutaneous glands is described in the Australian hylid Litoria caerulea. Three main types of glands could be distinguished in both ventral and dorsal skin: mucous, serous or granular, and lipid glands. Both mucous, and to some extent, serous glands show a PAS-positive reaction. Some of the granular-serous glands react to lipid stai...
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An ecological survey of the ant fauna of the southern part of the Junggar Basin and adjacent mountains, Xinjiang, China, revealed 46 species of which 27 (59%) were new records for China. Most of the species are widespread and no endemics were found. A largely boreal fauna occupies the spruce forest zone at high elevations of the Tienshan Mountains,...
Article
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: Resistance of bullfrogs (Rana catesbeiana) to the venom of two viperid snakes, the cottonmouth (Agkistrodon piscivorus piscivorus) and the copperhead (Agkistrodon contortrix contortrix), was measured by two criteria: (1) the LD<sub>50</sub> of the venom in various stages of frogs; and (2) the survival time of animals after envenomation. Tadpoles...
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Short-term movements of 12 olive sea snakes, Aipysurus laevis, were tracked using ultrasonic transmitters fed to snakes. This species restricts itself to a home range of about 1500-1800 m2. Three individuals displaced from their home ranges did not home to them. The home range often is linear along a reef face where the reef abuts open sand. Snakes...
Article
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Eels of the genus Gymnothorax from the Pacific are selectively preyed upon by banded sea kraits (Laticauda colubrina) and have been reported to sustain massive doses of sea krait venom without ill effect. By contrast, the present study found that Gymnothorax moringa from the Caribbean, where no sea snakes occur, are sensitive to sea krait venom, wi...
Article
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1. Harvesting the existing by-catch from the prawn industry, rather than returning captured snakes to the sea, would more than double the current impact of prawning upon snake populations. 2. Recovery of depleted populations from either reproduction or immigration would be slow for many species, and may not keep pace with losses. 3. Harvesting rate...
Article
Ants do not occur in the rocky desert of the Nazca Plain, Peru, except in the vicinity of shrubs in outwash gulleys, where there are two species that have similar activity periods and microhabitats and exhibit interference competition. Only one species of ant was found on the Chott El Djerid, Tunisia, a large, dry salt lake, devoid of vegetation.
Article
During the austral summer of 1990-91, a survey of Tardigrada inhabiting terrestrial mosses and lichens was conducted in the northern Prince Charles Mountains, East Antarctica. Five genera and six species were recovered including Echiniscus jenningsi, Diphascon sanae, Hypsibius antarcticus, Macrobiotus blocki, Macrobiotus stuckenbergi, and Milnesium...
Chapter
In previous chapters, the acquisition and expenditure of energy were discussed at the individual or species level. Such budgets are merely individual links in the overall exchanges within a biotic community. In the present chapter, the flow of energy among species within invertebrate assemblages and between them and other components of desert commu...
Chapter
Because resources in deserts are linked to the infrequent and erratic occurrence of rain and are unpredictable and ephemeral, animals that live there must be adapted to deal with episodes of food shortage and to exploit temporary abundances. One way is to reduce energy expenditure during lean times by becoming dormant (see Chap. 6). Another is to s...
Chapter
Energy taken in as food is apportioned in various ways. Some builds new tissue as growth or is deposited in energy storage products such as fat or glycogen. Some is used in production of young. Much of it Fuels metabolism and is expended in bodily maintenance. Finally, what is not assimilated is lost in the faeces. An animal’s individual energy bud...
Chapter
Between rains, water may be scarce, and high temperatures, even if not lethal, can contribute to metabolic excesses and wastage of energy by active animals. Production is reduced and concentration of resources may be too low for an organism to maintain vital activities. All of this means that periodically energy is scarce, the time available for it...
Chapter
Invertebrates are ectotherms and therefore depend on their external environment for body heat. Consequently, they expend little energy in metabolic thermogenesis and their total energy demands are reduced accordingly. Their metabolic rates are lower and a greater proportion of their food can go directly into such vital functions as activity, growth...
Chapter
Animals are unable to produce their own food and must rely either on (1) attracting, ambushing or trapping it, or (2) actively ferreting it out. Sedentary techniques variously have been termed ambush or sit-and-wait strategies; more mobile ones have been called cruise searching, widely ranging searching, active foraging or simply foraging. Some spe...
Chapter
A mobile animal has the advantage of being able to escape harsh conditions and seek out momentarily favourable sites (Riechert 1979; Farrow 1990). Nomadism is a strategy traditionally employed by desert tribes and their grazing domestic animals, and among wild animals is more characteristic of vertebrates, especially birds and large mammals, than i...
Book
Desert invertebrates live in an environment where resources alternate unpredictably between brief periods of plenty and prolonged scarcity. This text describes the adaptive strategies of desert invertebrates in acquiring energy and sustaining life under such vicissitudes. Some co-operate in foraging; others compete for resources. Some are nomadic a...
Article
The lateral-line sense organs in the skin of larval, juvenile and adult salamanders (Andrias davidianus) were examined by light and scanning electron microscopy. In addition to mechanoreceptive neuromasts, there are electroreceptive ampullary organs. Anatomically, the latter are similar to the ampullary organs of some other urodeles. In the giant s...
Article
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Abstract: Responses of four species of eels to the venoms of two species of sea snakes, a dietary generalist (Aipysurus laevis) and an eel specialist (Laticauda colubrina), were compared. All eels tested were more resistant to the specialist venom than to that of the generalist. Two muraenid eels (Gymnothorax hepaticus and G. undulatus), both symp...
Article
A survey of the terrestrial tardigrades inhabiting soil, algae, lichens, and mosses was conducted during the austral summer of 1990 at the Australian Research Station at Mawson and other localities on the Mawson Coast. Five genera and six species of tardigrades were recovered: Echiniscus jenningsi, Diphascon sanae, Hypsibius antarcticus, Macrobiotu...
Article
Volume two consists of seven individually authored chapters (separately abstracted). The first chapter reviews intrasexual selection and alternative mating behaviours. Next female choice and mating system structure is examined. The third chapter covers parental care. Chapter four looks at aggregation and kin recognition; next interspecific interact...
Article
A survey of the terrestrial tardigrades inhabiting algae, lichens, and mosses in the Larsemann Hills, East Antarctica was conducted at 61 sites during the austral summer of 1987. Five genera and six species of Tardigrada were recovered. Statistical analysis of biotic association was conducted and for most species-pairs, tardigrades occur randomly w...
Article
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Sea-snakes were collected from research trawlers and commercial prawn trawlers in the Gulf of Carpentaria during the period from April 1976 to December 1991. The data were analysed on the basis of CPUE (catch per unit effort) for depth, latitude and season. The research trawlers, operating in the eastern Gulf of Carpentaria, and the commercial praw...
Chapter
The dieback syndrome in the New England district of Australia is considered to be one of the most severe tree declines worldwide (Heatwole and Lowman 1986; Mueller-Dombois 1990/91; Fig. 1). Over the last 100 years, dramatic alterations in land use and agricultural practices have resulted in a landscape devoid of living trees, and also devoid of see...
Article
Aipysurus laevis venom has been shown to have a direct nephrotoxic effect in mice. A single subcutaneous injection (0.075 mg/kg body wt.) of the whole venom caused acute renal tubular degeneration and proliferative glomerulonephritis. The tubular changes appeared within 1 hour and remained for at least 14 days. Mesangial proliferative glomeruloneph...
Article
1. Three prey species of fish (Chromis nitida, Dascyllus aruanus and Istiblennius meleagris) were envenomated with Aipysurus laevis venom at three dose levels.2. Controls and low venom dosage groups maintained relatively constant ventilation rates with minimal differences among these three groups.3. Medium and high venom dosage groups exhibited ven...
Article
The LD50 values and survival times of three pomacentrid species and two blennies were measured after being subjected to the venom of one of their predators, the olive sea snake, Aipysurus laevis. The species differed significantly in the speed of their responses to the venom. At high venom doses, blennies had higher survival times than pomacentrids...
Article
Insect defoliation is commonly associated with and assumed to be a cause of mortality in Australian eucalypts, particularly in rural regions where trees suffer from the eucalypt dieback syndrome. To test this, leaf growth and defoliation were measured in the canopies of Eucalpyptus trees from June 1982 to June 1986, and related to tree health and e...
Article
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A single species accounted for about half the total ant biomass and the number of individuals. Less common species rank differently by biomass than by abundance because of body size differences. Among the scavengers, large-bodied species live in larger colonies than small-bodied species; the reverse applies to two seed-gathering species. There were...
Article
Development, survival, reproduction and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum (Hausmann) (Hemiptera: Aphididae) at constant temperatures of 10, 13, 15, 20, 25, 30 and 32C are reported. The developmental times of all life stages were inversely related to temperature ranging from 10 to 30C. Sp...
Article
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An assemblage of 8 species of ants contained 4 diurnal species, 2 nocturnal ones and 2 active both by day and night. More species and individuals came to baits at night and in the morning than during the heat of the day. Monomorium sp., the commonest species, found 65% of baits and had the highest visitation rates at all times of day except midday....
Article
A survey of eighty-two cays and ten continental islands of the Great Barrier Reef and Torres Straits, Australia, yielded 603 species of ferns and flowering plants. The numbers of species on islands were multiply regressed, separately for cays and continental islands, againts seven independent variables: latitude, longitude, insular area, insular he...
Article
The venom apparatus of Ethmostigmus rubripes, a generalized predator, consists of the telopodites of the postcephalic segment, the basal article of w which contains the venom gland. Within the gland, venom granules are concentrated in intracellular secretory granules, from which they are discharged into vacuoles in the cytoplasm of the secretory ce...
Article
Venom from the olive sea snake, Aipysurus laevis, was injected into three species of prey fish, Chromis nitida, Dascyllus aruanus and Istiblennius meleagris. Their behaviour and ventilatory patterns were observed for 48 hr. Six progressive stages of envenomation, involving impairment of the locomotory and ventilatory systems, were identified. There...
Article
Drift disseminules were collected on cays of the Swain Reefs (Great Barrier Reef, Australia) during twelve expeditions over 7 years. At least thirty-four species not belonging to the small established cay floras are represented in the collections. Assemblages varied between visits and between cays, in both relative proportions of species and in tot...

Questions

Questions (2)
Question
I am the editor of the series "Amphibian Biology" and am contemplating an issue on the "Status of Amphibians of the Atlantic Islands". Would you be interested in contributing a chapter on Bermuda? So far the series has a volume each on the following regions: South America, Europe,
Asia, North Africa and volumes are in progress on: The Caribbean (ready for press), Central America, SubSaharan Africa, and North America. I will be happy to supply any additional information you may require.
Best regards,
Harold Heatwole
Dept. or Zoology,
University or New England,
Armidale, NSW 2351, Australia
Question
I would like to email you about your project.
I am the editor of the series Amphibian Biology, now in its 12th volume. I am contemplating an issue on the Status of the Amphibians of the Atlantic Islands and wonder whether you would be interested in contributing a chapter on Bermuda.
Thank you.
Harold Heatwole
Dept. Zoology, UnIversity oF New England, Armidale, NSW 2351, Australia

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