Professor Frank N Carrick AM | The University of Queensland, Australia (original) (raw)
Papers by Professor Frank N Carrick AM
The Proceedings of the Royal Society of Queensland
South East Queensland (SEQ) is experiencing the fastest human population growth in Australia, wit... more South East Queensland (SEQ) is experiencing the fastest human population growth in Australia, with attendant challenges for wildlife conservation due to expanding urbanisation. The documented dramatic decline of koalas Phascolarctos cinereus on mainland SEQ has provoked popular suggestions that North Stradbroke Island (NSI) should become an "island ark" for koalas. A multidisciplinary study was undertaken to determine the status of koalas on NSI. Aboriginal and European references to koalas on NSI were collected and analysed. To study koala distribution, direct and indirect visual surveys were conducted, whilst habitat use and home ranges were determined by fitting 33 koalas with VHF collars and radio-tracking them. Population characteristics, including health status, proximate causes of mortality and genetic profile, were gathered from radio-tracked koalas and from hospital databases of the Department of Environment and Resource Management.; ;
Ecological Society of …, 2011
Carrick, Frank, Ellis, Bill, Fitzgibbon, Sean and Melzer, Alistair (2007). Planned Long-term Ecol... more Carrick, Frank, Ellis, Bill, Fitzgibbon, Sean and Melzer, Alistair (2007). Planned Long-term Ecological Research Sites Preview Likely Effects of Climate Change on Tropical Koalas. In:, Proceedings of the Ecological Society of Australia Conference 2007: Adapting to Change. ...
Australian Journal of Zoology, 2012
ABSTRACT Island populations of animals are expected to show reduced genetic variation and increas... more ABSTRACT Island populations of animals are expected to show reduced genetic variation and increased incidence of inbreeding because of founder effects and the susceptibility of small populations to the effects of genetic drift. Koalas (Phascolarctos cinereus) occur naturally in a patchy distribution across much of the eastern Australian mainland and on a small number of islands near the Australian coast. We compared the genetic diversity of the naturally occurring population of koalas on North Stradbroke Island in south-east Queensland with other island populations including the introduced group on St Bees Island in central Queensland. The population on St Bees Island shows higher diversity (allelic richness 4.1, He = 0.67) than the North Stradbroke Island population (allelic richness 3.2, He = 0.55). Koalas on Brampton, Newry and Rabbit Islands possessed microsatellite alleles that were not identified from St Bees Island koalas, indicating that it is most unlikely that these populations were established by a sole secondary introduction from St Bees Island. Mitochondrial haplotypes on the central Queensland islands were more similar to a haplotype found at Springsure in central Queensland and the inland clades in southeast Queensland, rather than the coastal clade in south-east Queensland.
Australian Journal of Chemistry, 1993
7-Hydroxy-1, 8-cineole (3a) and the corresponding carboxylic acid (3b) have been isolated from th... more 7-Hydroxy-1, 8-cineole (3a) and the corresponding carboxylic acid (3b) have been isolated from the urine of brushtail possums (Trichosurus vulpecula: Marsupialia) fed a diet enhanced with 1,8-cineole. Chemical syntheses of these two novel metabolites are described.
Australian Mammalogy, 2002
The coefficients of digestion and volume classify both O. anatinus and T. aculeatus as Snipes RL,... more The coefficients of digestion and volume classify both O. anatinus and T. aculeatus as Snipes RL, Snipes H and Carrick FN, 2002. Morphometric data on the intestines of the platypus Ornithorhynchus anatinus and the short-beaked echidna Tachyglossus aculeatus. Australian Mammalogy 24: 229-231.
Acta Zoologica, 2004
Abstract Johnston, S.D., Daddow, L., Carrick, F.N. and Jamieson, B. 2004. Observations of sper... more Abstract Johnston, S.D., Daddow, L., Carrick, F.N. and Jamieson, B. 2004. Observations of spermiogenesis and epididymal sperm maturation in the rufous hare wallaby, Lagorchestes hirsutus (Metatheria, Mammalia). -Acta Zoologica (Stockholm) 85 : 53 -58 Acrosomal development in the early spermatid of the rufous hare wallaby shows evidence of formation of an acrosomal granule, similar to that found in eutherian mammals, the Phascolarctidae and Vombatidae. Unlike the other members of the Macropodidae so far examined, the acrosome of this species appears to be fully compacted at spermiation and extends evenly over 90% of the dorsal aspect of the nucleus. During spermiogenesis, the nucleus of the rufous hare wallaby spermatid showed evidence of uneven condensation of chromatin; this may also be related to the appearance of unusual nucleoplasm evaginations from the surface of the fully condensed spermatid. This study was unable to find evidence of the presence of Sertoli cell spurs or nuclear rotation during spermiogenesis in the rufous hare wallaby. The majority of spermatozoa immediately before spermiation had a nucleus that was essentially perpendicular to the long axis of the sperm tail. Nuclei of spermatozoa found in the process of being released or isolated in the lumen of the seminiferous tubule were rotated almost parallel to the long axis of the flagellum; complete parallel alignment occurred during epididymal maturation. At spermiation spermatozoa have characteristically small cytoplasmic remnants compared to those of other macropods. Unlike the majority of macropodid spermatozoa so far described, the spermatozoa of the rufous hare wallaby showed little evidence of morphological change during epididymal transit. There was no formation of a fibre network around the midpiece or of plasma membrane specializations in this region; the only notable change was a distinctive flattening of midpiece mitochondria and scalloping of the anterior mitochondrial sheath to accommodate the sperm head. Preliminary evidence from spermiogenesis and epididymal sperm maturation supports the classification of the rufous hare wallaby as a separate genus but also indicates that its higher taxonomic position may need to be re-evaluated.
Acta Zoologica, 2003
Appearance of cauda epididymal spermatozoa of the rufous hare wallaby, Lagorchestes hirsutus (Met... more Appearance of cauda epididymal spermatozoa of the rufous hare wallaby, Lagorchestes hirsutus (Metatheria, Mammalia) imaged by electron and confocal microscopy. -Acta Zoologica (Stockholm) 84 : 139-143
Acta Zoologica, 2002
Lloyd, S., Carrick, F. and Hall, L. 2002. Ultrastructure of the mature spermatozoon of the musky ... more Lloyd, S., Carrick, F. and Hall, L. 2002. Ultrastructure of the mature spermatozoon of the musky rat-kangaroo, Hypsiprymnodon moschatus (Potoroidae: Marsupialia). -Acta Zoologica (Stockholm) 83 : 167-174
Reproduction, Fertility and Development, 2010
The effects of breeding season (late spring to early autumn) on south-east Queensland male koala ... more The effects of breeding season (late spring to early autumn) on south-east Queensland male koala fertility were examined to improve the efficacy of the AI procedure and to determine the practicality of using free-range animals as semen donors for a genome resource bank. Seasonal changes in male koala reproductive function were assessed in a wild free-range population (n = 14; obtained every 6 weeks from January to November 2005), a necropsied healthy wild population (n = 84; obtained monthly from September 2004 to August 2005) and a captive population (n = 7; obtained monthly from October 2005 to October 2006). Reproductive parameters investigated included bodyweight, coat score, sternal gland area and activity, testosterone secretion, reproductive anatomy volume and semen quality (before and after cryopreservation). Collectively, these findings show that reproduction in male koalas from south-east Queensland changes seasonally and that winter appears to be the optimal season in which to collect semen samples by electroejaculation. While it was possible to repeatedly collect semen from free-range koalas for future genetic management via potential storage in a genome resource bank, the survival of these spermatozoa after cryopreservation was poor and will require further improvement.
Wildlife Research, 2008
ABSTRACT North Stradbroke Island lies 4 km off the coast of Queensland, Australia. It is home to ... more ABSTRACT North Stradbroke Island lies 4 km off the coast of Queensland, Australia. It is home to one of the only naturally occurring island populations of koalas and it is mined for mineral sands. We analysed the diet and day use tree selections of koalas and recorded the tree species composition of revegetated and undisturbed landscapes at this location. We used faecal cuticle examination to compare the diet of koalas that used reconstructed landscapes with that of koalas that used undisturbed areas. Reconstructed landscapes that were composed of more than 95% diet and/or roost tree species had evidence of use by koalas. Eucalyptus robusta was the most commonly eaten and utilised species and there was no difference in general diet composition between koalas that used the revegetated landscape and those inhabiting undisturbed areas. Other species that were used for roosting and forage included E. racemosa, E. pilularis, Lophostemon confertus and Melaleuca quinquinerva. We observed individual differences and seasonal variation in the diet composition of radio-tracked koalas. These results suggest either flexibility in the diet choices of koalas, or individual preferences within groups of koalas. Our results also indicate that post-mining landscapes can provide habitat that will be used by koalas, which should encourage further efforts into habitat re-creation for native species.
Wildlife Research, 2003
Koala dispersal was investigated as part of a detailed ecological study of a nationally significa... more Koala dispersal was investigated as part of a detailed ecological study of a nationally significant koala population located 20 km south-east of Brisbane, Queensland. From 1996 to 2000, 195 koalas from three sites were captured and fitted with radio-collars. A total of 40 koalas (23 ...
Reproduction, Fertility and Development, 2008
The aim of the present study was to determine whether analogues of gonadotrophin-releasing hormon... more The aim of the present study was to determine whether analogues of gonadotrophin-releasing hormone (GnRH) could be used to both induce an acute testosterone response and suppress anterior pituitary function in male koalas, and induce a luteal phase in female koalas. Experiment 1 characterised the steroidogenic response of male koalas to administration of 30 microg (4.3 microg kg(-1)) natural-sequence GnRH. Intra-muscular injection of natural-sequence GnRH induced the release of LH and testosterone with peak concentrations at 30 min (3.7 +/- 1.9 ng mL(-1)) and 2 h (5.4 +/- 0.5 ng mL(-1)), respectively. In Experiment 2, a single injection of the GnRH antagonist acyline (100 microg (14.3 microg kg(-1)) or 500 microg (71.4 microg kg(-1))) did not influence the testosterone response to subsequent injections of natural-sequence GnRH. In Experiment 3, 4 microg (~0.67 microg kg(-1)) of the GnRH agonist buserelin induced a luteal phase in five female koalas based on a LH surge, secretion of progestogen, and a normal-length oestrous cycle. The findings have shown that (1) natural-sequence GnRH can be used to test gonadotroph cell function and determine the testosterone-secreting capacity of male koalas, (2) the GnRH antagonist, acyline, at the dose rates used, does not suppress the pituitary-testis axis in male koalas, and (3) the GnRH agonist, buserelin, induces a normal luteal phase in female koalas.
Fish Physiology and Biochemistry, 2000
The cDNAs coding for the brain GnRHs (AY373449-51), pituitary GH, SL and PRL, and liver IGFs (AY4... more The cDNAs coding for the brain GnRHs (AY373449-51), pituitary GH, SL and PRL, and liver IGFs (AY427954-5) were isolated. Partial cDNA sequences of the brain (Cyp19b) and gonadal (Cyp19a) aromatases have also been obtained. These tools would be utilized to study the endocrine regulation of puberty in the grey mullet.
Annual Meeting of …, 2011
... microsatellite data. Furthermore, we present results from two programs that incorporate spati... more ... microsatellite data. Furthermore, we present results from two programs that incorporate spatial data, TESS and Geneland, demonstrating the effect of the distribution of samples on the number of inferred clusters and their distribution. ...
Conservation Biology, 2000
1800s and then declined dramatically in the early 1900s. The decline was associated with habitat ... more 1800s and then declined dramatically in the early 1900s. The decline was associated with habitat loss, hunting, disease, fire, and drought. Declines are continuing in Queensland and New South Wales. In contrast, dense koala populations in habitat isolates in Victoria and South Australia are managed to reduce population size and browse damage. Current understanding of koala distribution and abundance suggests that the species does not meet Australian criteria as endangered or vulnerable fauna. Its conservation status needs to be reviewed, however, in light of the extensive land clearing in New South Wales and Queensland since the last (1980s) broadscale surveys. Consequently, we recommend that broadacre clearing be curtailed in New South Wales and Queensland and that regular, comprehensive, standardized, national koala surveys be undertaken. Given the fragmentation of koala habitat and regional differences in the status of the koala, we recommended that studies on regional variation in the koala be intensified and that koala ecology in fragmented and naturally restricted habitats be developed. More generally, the National Koala Conservation Strategy should be implemented.
… Meeting and Annual …, 2011
The characteristics of areas of natural vegetation that appeal to a species may be absent from re... more The characteristics of areas of natural vegetation that appeal to a species may be absent from reconstructed landscapes, either by design or due to elements beyond the control of the land manager. In such cases a novel environment may be created; does this represent ...
The Proceedings of the Royal Society of Queensland
South East Queensland (SEQ) is experiencing the fastest human population growth in Australia, wit... more South East Queensland (SEQ) is experiencing the fastest human population growth in Australia, with attendant challenges for wildlife conservation due to expanding urbanisation. The documented dramatic decline of koalas Phascolarctos cinereus on mainland SEQ has provoked popular suggestions that North Stradbroke Island (NSI) should become an "island ark" for koalas. A multidisciplinary study was undertaken to determine the status of koalas on NSI. Aboriginal and European references to koalas on NSI were collected and analysed. To study koala distribution, direct and indirect visual surveys were conducted, whilst habitat use and home ranges were determined by fitting 33 koalas with VHF collars and radio-tracking them. Population characteristics, including health status, proximate causes of mortality and genetic profile, were gathered from radio-tracked koalas and from hospital databases of the Department of Environment and Resource Management.; ;
Ecological Society of …, 2011
Carrick, Frank, Ellis, Bill, Fitzgibbon, Sean and Melzer, Alistair (2007). Planned Long-term Ecol... more Carrick, Frank, Ellis, Bill, Fitzgibbon, Sean and Melzer, Alistair (2007). Planned Long-term Ecological Research Sites Preview Likely Effects of Climate Change on Tropical Koalas. In:, Proceedings of the Ecological Society of Australia Conference 2007: Adapting to Change. ...
Australian Journal of Zoology, 2012
ABSTRACT Island populations of animals are expected to show reduced genetic variation and increas... more ABSTRACT Island populations of animals are expected to show reduced genetic variation and increased incidence of inbreeding because of founder effects and the susceptibility of small populations to the effects of genetic drift. Koalas (Phascolarctos cinereus) occur naturally in a patchy distribution across much of the eastern Australian mainland and on a small number of islands near the Australian coast. We compared the genetic diversity of the naturally occurring population of koalas on North Stradbroke Island in south-east Queensland with other island populations including the introduced group on St Bees Island in central Queensland. The population on St Bees Island shows higher diversity (allelic richness 4.1, He = 0.67) than the North Stradbroke Island population (allelic richness 3.2, He = 0.55). Koalas on Brampton, Newry and Rabbit Islands possessed microsatellite alleles that were not identified from St Bees Island koalas, indicating that it is most unlikely that these populations were established by a sole secondary introduction from St Bees Island. Mitochondrial haplotypes on the central Queensland islands were more similar to a haplotype found at Springsure in central Queensland and the inland clades in southeast Queensland, rather than the coastal clade in south-east Queensland.
Australian Journal of Chemistry, 1993
7-Hydroxy-1, 8-cineole (3a) and the corresponding carboxylic acid (3b) have been isolated from th... more 7-Hydroxy-1, 8-cineole (3a) and the corresponding carboxylic acid (3b) have been isolated from the urine of brushtail possums (Trichosurus vulpecula: Marsupialia) fed a diet enhanced with 1,8-cineole. Chemical syntheses of these two novel metabolites are described.
Australian Mammalogy, 2002
The coefficients of digestion and volume classify both O. anatinus and T. aculeatus as Snipes RL,... more The coefficients of digestion and volume classify both O. anatinus and T. aculeatus as Snipes RL, Snipes H and Carrick FN, 2002. Morphometric data on the intestines of the platypus Ornithorhynchus anatinus and the short-beaked echidna Tachyglossus aculeatus. Australian Mammalogy 24: 229-231.
Acta Zoologica, 2004
Abstract Johnston, S.D., Daddow, L., Carrick, F.N. and Jamieson, B. 2004. Observations of sper... more Abstract Johnston, S.D., Daddow, L., Carrick, F.N. and Jamieson, B. 2004. Observations of spermiogenesis and epididymal sperm maturation in the rufous hare wallaby, Lagorchestes hirsutus (Metatheria, Mammalia). -Acta Zoologica (Stockholm) 85 : 53 -58 Acrosomal development in the early spermatid of the rufous hare wallaby shows evidence of formation of an acrosomal granule, similar to that found in eutherian mammals, the Phascolarctidae and Vombatidae. Unlike the other members of the Macropodidae so far examined, the acrosome of this species appears to be fully compacted at spermiation and extends evenly over 90% of the dorsal aspect of the nucleus. During spermiogenesis, the nucleus of the rufous hare wallaby spermatid showed evidence of uneven condensation of chromatin; this may also be related to the appearance of unusual nucleoplasm evaginations from the surface of the fully condensed spermatid. This study was unable to find evidence of the presence of Sertoli cell spurs or nuclear rotation during spermiogenesis in the rufous hare wallaby. The majority of spermatozoa immediately before spermiation had a nucleus that was essentially perpendicular to the long axis of the sperm tail. Nuclei of spermatozoa found in the process of being released or isolated in the lumen of the seminiferous tubule were rotated almost parallel to the long axis of the flagellum; complete parallel alignment occurred during epididymal maturation. At spermiation spermatozoa have characteristically small cytoplasmic remnants compared to those of other macropods. Unlike the majority of macropodid spermatozoa so far described, the spermatozoa of the rufous hare wallaby showed little evidence of morphological change during epididymal transit. There was no formation of a fibre network around the midpiece or of plasma membrane specializations in this region; the only notable change was a distinctive flattening of midpiece mitochondria and scalloping of the anterior mitochondrial sheath to accommodate the sperm head. Preliminary evidence from spermiogenesis and epididymal sperm maturation supports the classification of the rufous hare wallaby as a separate genus but also indicates that its higher taxonomic position may need to be re-evaluated.
Acta Zoologica, 2003
Appearance of cauda epididymal spermatozoa of the rufous hare wallaby, Lagorchestes hirsutus (Met... more Appearance of cauda epididymal spermatozoa of the rufous hare wallaby, Lagorchestes hirsutus (Metatheria, Mammalia) imaged by electron and confocal microscopy. -Acta Zoologica (Stockholm) 84 : 139-143
Acta Zoologica, 2002
Lloyd, S., Carrick, F. and Hall, L. 2002. Ultrastructure of the mature spermatozoon of the musky ... more Lloyd, S., Carrick, F. and Hall, L. 2002. Ultrastructure of the mature spermatozoon of the musky rat-kangaroo, Hypsiprymnodon moschatus (Potoroidae: Marsupialia). -Acta Zoologica (Stockholm) 83 : 167-174
Reproduction, Fertility and Development, 2010
The effects of breeding season (late spring to early autumn) on south-east Queensland male koala ... more The effects of breeding season (late spring to early autumn) on south-east Queensland male koala fertility were examined to improve the efficacy of the AI procedure and to determine the practicality of using free-range animals as semen donors for a genome resource bank. Seasonal changes in male koala reproductive function were assessed in a wild free-range population (n = 14; obtained every 6 weeks from January to November 2005), a necropsied healthy wild population (n = 84; obtained monthly from September 2004 to August 2005) and a captive population (n = 7; obtained monthly from October 2005 to October 2006). Reproductive parameters investigated included bodyweight, coat score, sternal gland area and activity, testosterone secretion, reproductive anatomy volume and semen quality (before and after cryopreservation). Collectively, these findings show that reproduction in male koalas from south-east Queensland changes seasonally and that winter appears to be the optimal season in which to collect semen samples by electroejaculation. While it was possible to repeatedly collect semen from free-range koalas for future genetic management via potential storage in a genome resource bank, the survival of these spermatozoa after cryopreservation was poor and will require further improvement.
Wildlife Research, 2008
ABSTRACT North Stradbroke Island lies 4 km off the coast of Queensland, Australia. It is home to ... more ABSTRACT North Stradbroke Island lies 4 km off the coast of Queensland, Australia. It is home to one of the only naturally occurring island populations of koalas and it is mined for mineral sands. We analysed the diet and day use tree selections of koalas and recorded the tree species composition of revegetated and undisturbed landscapes at this location. We used faecal cuticle examination to compare the diet of koalas that used reconstructed landscapes with that of koalas that used undisturbed areas. Reconstructed landscapes that were composed of more than 95% diet and/or roost tree species had evidence of use by koalas. Eucalyptus robusta was the most commonly eaten and utilised species and there was no difference in general diet composition between koalas that used the revegetated landscape and those inhabiting undisturbed areas. Other species that were used for roosting and forage included E. racemosa, E. pilularis, Lophostemon confertus and Melaleuca quinquinerva. We observed individual differences and seasonal variation in the diet composition of radio-tracked koalas. These results suggest either flexibility in the diet choices of koalas, or individual preferences within groups of koalas. Our results also indicate that post-mining landscapes can provide habitat that will be used by koalas, which should encourage further efforts into habitat re-creation for native species.
Wildlife Research, 2003
Koala dispersal was investigated as part of a detailed ecological study of a nationally significa... more Koala dispersal was investigated as part of a detailed ecological study of a nationally significant koala population located 20 km south-east of Brisbane, Queensland. From 1996 to 2000, 195 koalas from three sites were captured and fitted with radio-collars. A total of 40 koalas (23 ...
Reproduction, Fertility and Development, 2008
The aim of the present study was to determine whether analogues of gonadotrophin-releasing hormon... more The aim of the present study was to determine whether analogues of gonadotrophin-releasing hormone (GnRH) could be used to both induce an acute testosterone response and suppress anterior pituitary function in male koalas, and induce a luteal phase in female koalas. Experiment 1 characterised the steroidogenic response of male koalas to administration of 30 microg (4.3 microg kg(-1)) natural-sequence GnRH. Intra-muscular injection of natural-sequence GnRH induced the release of LH and testosterone with peak concentrations at 30 min (3.7 +/- 1.9 ng mL(-1)) and 2 h (5.4 +/- 0.5 ng mL(-1)), respectively. In Experiment 2, a single injection of the GnRH antagonist acyline (100 microg (14.3 microg kg(-1)) or 500 microg (71.4 microg kg(-1))) did not influence the testosterone response to subsequent injections of natural-sequence GnRH. In Experiment 3, 4 microg (~0.67 microg kg(-1)) of the GnRH agonist buserelin induced a luteal phase in five female koalas based on a LH surge, secretion of progestogen, and a normal-length oestrous cycle. The findings have shown that (1) natural-sequence GnRH can be used to test gonadotroph cell function and determine the testosterone-secreting capacity of male koalas, (2) the GnRH antagonist, acyline, at the dose rates used, does not suppress the pituitary-testis axis in male koalas, and (3) the GnRH agonist, buserelin, induces a normal luteal phase in female koalas.
Fish Physiology and Biochemistry, 2000
The cDNAs coding for the brain GnRHs (AY373449-51), pituitary GH, SL and PRL, and liver IGFs (AY4... more The cDNAs coding for the brain GnRHs (AY373449-51), pituitary GH, SL and PRL, and liver IGFs (AY427954-5) were isolated. Partial cDNA sequences of the brain (Cyp19b) and gonadal (Cyp19a) aromatases have also been obtained. These tools would be utilized to study the endocrine regulation of puberty in the grey mullet.
Annual Meeting of …, 2011
... microsatellite data. Furthermore, we present results from two programs that incorporate spati... more ... microsatellite data. Furthermore, we present results from two programs that incorporate spatial data, TESS and Geneland, demonstrating the effect of the distribution of samples on the number of inferred clusters and their distribution. ...
Conservation Biology, 2000
1800s and then declined dramatically in the early 1900s. The decline was associated with habitat ... more 1800s and then declined dramatically in the early 1900s. The decline was associated with habitat loss, hunting, disease, fire, and drought. Declines are continuing in Queensland and New South Wales. In contrast, dense koala populations in habitat isolates in Victoria and South Australia are managed to reduce population size and browse damage. Current understanding of koala distribution and abundance suggests that the species does not meet Australian criteria as endangered or vulnerable fauna. Its conservation status needs to be reviewed, however, in light of the extensive land clearing in New South Wales and Queensland since the last (1980s) broadscale surveys. Consequently, we recommend that broadacre clearing be curtailed in New South Wales and Queensland and that regular, comprehensive, standardized, national koala surveys be undertaken. Given the fragmentation of koala habitat and regional differences in the status of the koala, we recommended that studies on regional variation in the koala be intensified and that koala ecology in fragmented and naturally restricted habitats be developed. More generally, the National Koala Conservation Strategy should be implemented.
… Meeting and Annual …, 2011
The characteristics of areas of natural vegetation that appeal to a species may be absent from re... more The characteristics of areas of natural vegetation that appeal to a species may be absent from reconstructed landscapes, either by design or due to elements beyond the control of the land manager. In such cases a novel environment may be created; does this represent ...