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{{Short description|Extinct species of lizard}} |
{{Short description|Extinct species of lizard}} |
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{{Speciesbox |
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{{speciesbox |
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| name = ''Gigarcanum'' |
| name = ''Gigarcanum'' |
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| image = Gecko de Delcourt Hoplodactylus delcourti GLAM MHNL 2016 3741.jpg |
| image = Gecko de Delcourt Hoplodactylus delcourti GLAM MHNL 2016 3741.jpg |
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| status_system = IUCN3.1 |
| status_system = IUCN3.1 |
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| status_ref = <ref name=Hitchmough2019>{{cite iucn |last1=Hitchmough |first1=R. |author1-link=species:Rodney Arthur Hitchmough |last2=van Winkel |first2=D. |author2-link=species:Dylan van Winkel |last3=Lettink |first3=M. |author3-link=species:Marieke Lettink |last4=Chapple |first4=D. |author4-link=species:David G. Chapple |date=2019 |title=''Hoplodactylus delcourti '' |doi=10.2305/IUCN.UK.2019-2.RLTS.T10254A120158840.en |volume=2019 |page=e.T10254A120158840}}</ref> |
| status_ref = <ref name=Hitchmough2019>{{cite iucn |last1=Hitchmough |first1=R. |author1-link=species:Rodney Arthur Hitchmough |last2=van Winkel |first2=D. |author2-link=species:Dylan van Winkel |last3=Lettink |first3=M. |author3-link=species:Marieke Lettink |last4=Chapple |first4=D. |author4-link=species:David G. Chapple |date=2019 |title=''Hoplodactylus delcourti '' |doi=10.2305/IUCN.UK.2019-2.RLTS.T10254A120158840.en |volume=2019 |page=e.T10254A120158840}}</ref> |
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| extinct = |
| extinct = 19th century |
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| parent_authority = Heinicke, et al, 2023 |
| parent_authority = Heinicke, et al, 2023 |
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| genus = Gigarcanum |
| genus = Gigarcanum |
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| species = delcourti |
| species = delcourti |
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| authority = ([[:fr:Aaron Matthew Bauer|Bauer]] & [[species:Anthony Patrick Russell|Russell]], 1986) |
| authority = ([[:fr:Aaron Matthew Bauer|Bauer]] & [[species:Anthony Patrick Russell|Russell]], 1986) |
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| synonyms = *''Hoplodactylus delcourti'' |
| synonyms = *''Hoplodactylus delcourti'' <small>Bauer & Russell, 1986</small> |
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}} |
}} |
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'''''Gigarcanum delcourti''''', formerly '''''Hoplodactylus delcourti''''', is an extinct [[species]] of gecko in the [[Family (biology)|family]] [[Diplodactylidae]]. It is [[List of largest extant lizards|the largest known of all geckos]], with a [[snout-to-vent length]] (SVL) of {{convert|37|cm|in|abbr=on|sigfig=3}} and an overall length (including tail) of at least {{convert|60|cm|in|abbr=on|sigfig=3}}.<ref name=Wilson2004>{{cite book |last=Wilson |first=Kerry-Jayne |year=2004 |title=Flight of the Huia: Ecology and Conservation of New Zealand's Frogs, Reptiles, Birds and Mammals |publisher=Canterbury University Press |isbn=0-908812-52-3 |oclc=937349394 |url=https://books.google.com/books?id=BXbwAAAAMAAJ}}</ref> It is only known a single taxidermied specimen collected in the 19th century that was rediscovered unlabelled in a museum in France. The origin of the specimen was undocumented.<ref name=Hitchmough2019/> While originally suggested to have been from [[New Zealand]] and the ''kawekaweau'' of [[Māori people|Māori]] oral tradition,<ref>{{cite journal |last=Worthy |first=T.H. |author-link=Trevor Worthy |date=March 1997 |title=Quaternary fossil fauna of South Canterbury, South Island, New Zealand |journal=Journal of the Royal Society of New Zealand |doi=10.1080/03014223.1997.9517528 |doi-access=free |volume=27 |issue=1 |pages=67–162}}</ref><ref>{{cite journal |last=Tennyson |first=Alan J.D. |author-link=species:Alan James Drummond Tennyson |date=2010 |title=The origin and history of New Zealand's terrestrial vertebrates |journal=New Zealand Ecological Society |volume=34 |issue=1 |pages=6–27 |url=http://newzealandecology.org/nzje/2914.pdf}}</ref><ref name=Worthy2016>{{cite book |last=Worthy |first=Trevor H. |date=2016 | |
'''''Gigarcanum delcourti''''', formerly '''''Hoplodactylus delcourti''''', is an extinct [[species]] of gecko in the [[Family (biology)|family]] [[Diplodactylidae]]. It is [[List of largest extant lizards|the largest known of all geckos]], with a [[snout-to-vent length]] (SVL) of {{convert|37|cm|in|abbr=on|sigfig=3}} and an overall length (including tail) of at least {{convert|60|cm|in|abbr=on|sigfig=3}}.<ref name=Wilson2004>{{cite book |last=Wilson |first=Kerry-Jayne |year=2004 |title=Flight of the Huia: Ecology and Conservation of New Zealand's Frogs, Reptiles, Birds and Mammals |publisher=Canterbury University Press |isbn=0-908812-52-3 |oclc=937349394 |url=https://books.google.com/books?id=BXbwAAAAMAAJ}}</ref> It is only known from a single taxidermied specimen collected in the 19th century that was rediscovered unlabelled in a museum in France. The origin of the specimen was undocumented.<ref name=Hitchmough2019/> While originally suggested to have been from [[New Zealand]] and the ''kawekaweau'' of [[Māori people|Māori]] oral tradition,<ref>{{cite journal |last=Worthy |first=T.H. |author-link=Trevor Worthy |date=March 1997 |title=Quaternary fossil fauna of South Canterbury, South Island, New Zealand |journal=Journal of the Royal Society of New Zealand |doi=10.1080/03014223.1997.9517528 |doi-access=free |volume=27 |issue=1 |pages=67–162|bibcode=1997JRSNZ..27...67W }}</ref><ref>{{cite journal |last=Tennyson |first=Alan J.D. |author-link=species:Alan James Drummond Tennyson |date=2010 |title=The origin and history of New Zealand's terrestrial vertebrates |journal=New Zealand Ecological Society |volume=34 |issue=1 |pages=6–27 |url=http://newzealandecology.org/nzje/2914.pdf}}</ref><ref name=Worthy2016>{{cite book |last=Worthy |first=Trevor H. |date=2016 |chapter=A Review of the Fossil Record of New Zealand Lizards |title=New Zealand Lizards |publisher=Springer International Publishing |doi=10.1007/978-3-319-41674-8_3 |isbn=978-3-319-41672-4 |pages=65–86}}</ref> DNA evidence from the specimen suggests that it originates from [[New Caledonia]].<ref name=Heinicke2023>{{cite journal |last1=Heinicke |first1=Matthew P. |last2=Nielsen |first2=Stuart V. |last3=Bauer |first3=Aaron M. |last4=Kelly |first4=Ryan |last5=Geneva |first5=Anthony J. |last6=Daza |first6=Juan D. |last7=Keating |first7=Shannon E. |last8=Gamble |first8=Tony |date=2023-06-19 |title=Reappraising the evolutionary history of the largest known gecko, the presumably extinct ''Hoplodactylus delcourti'', via high-throughput sequencing of archival DNA |journal=Scientific Reports |issn=2045-2322 |doi=10.1038/s41598-023-35210-8 |volume=13 |issue=1 |page=9141 |pmid=37336900 |pmc=10279644 |bibcode=2023NatSR..13.9141H }}</ref> |
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==History== |
==History== |
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According to the report of Major W. G. Mair in 1873, in 1870, a [[Māori people|Māori]] chief said that he had killed a ''kawekaweau'' he found under the bark of a dead [[Metrosideros robusta| |
According to the report of Major W. G. Mair in 1873, in 1870, a [[Māori people|Māori]] chief said that he had killed a ''kawekaweau'' he found under the bark of a dead [[Metrosideros robusta|rātā tree]] in the [[Waimana|Waimana Valley]] in [[Te Urewera]] on the North Island of New Zealand. This is the only documented report of anyone ever seeing a ''kawekaweau'' alive. Mair reported the chiefs description of the animal as being "two feet long and as thick as a man’s wrist; colour brown, striped longitudinally with dull red".<ref name=Bauer1986>{{cite journal |last1=Bauer |first1=Aaron M. |last2=Russell |first2=Anthony P. |date=1986 |title=''Hoplodactylus delcourti'' n. sp. (Reptilia: Gekkonidae), the largest known gecko |journal=New Zealand Journal of Zoology |volume=13 |pages=141–148 |doi=10.1080/03014223.1986.10422655 |url=http://www.bio.ucalgary.ca/contact/faculty/pdf/russell/38.pdf |archive-url=https://web.archive.org/web/20130420190127/http://www.bio.ucalgary.ca/contact/faculty/pdf/russell/38.pdf |archive-date=2013-04-20}}</ref> |
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A single stuffed specimen was "discovered" in the basement of the [[Muséum d'histoire naturelle de Marseille|Natural History Museum of Marseille]] in 1986; the origins and date of collection of the specimen remain a mystery, as it was unlabelled when it was found.<ref name=Bauer1986/> It has been present in the collection of the museum since at least the 1870s, and likely since the 1830s based on its unusual preservation style of being eviscerated, dried and mounted, rather than being kept in spirits as is more common for preserved specimens. The specimen is missing the internal organs and most of the [[axial skeleton]], but retains the skull and [[appendicular skeleton]].<ref name=Heinicke2023/> It was described as the new species ''[[Hoplodactylus]] delcourti''. Initially, scientists examining the specimen suggested that it was from New Zealand and was in fact the lost ''kawekaweau'', a giant and mysterious forest lizard of |
A single stuffed specimen was "discovered" in the basement of the [[Muséum d'histoire naturelle de Marseille|Natural History Museum of Marseille]] in 1986; the origins and date of collection of the specimen remain a mystery, as it was unlabelled when it was found.<ref name=Bauer1986/> It has been present in the collection of the museum since at least the 1870s, and likely since the 1830s based on its unusual preservation style of being eviscerated, dried and mounted, rather than being kept in spirits as is more common for preserved specimens. The specimen is missing the internal organs and most of the [[axial skeleton]], but retains the skull and [[appendicular skeleton]].<ref name=Heinicke2023/> It was described as the new species ''[[Hoplodactylus]] delcourti''. Initially, scientists examining the specimen suggested that it was from New Zealand and was in fact the lost ''kawekaweau'', a giant and mysterious forest lizard of Māori oral tradition.<ref name=Bauer1986/> |
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Attempts to extract DNA from the sole specimen in 1994 were unsuccessful.<ref>{{cite book |last=Cooper |first=Alan |title=Ancient DNA : Recovery and Analysis of Genetic Material from Paleontological, Archaeological, Museum, Medical, and Forensic Specimens |publisher=Springer New York |year=1994 |isbn=978-1-4612-4318-2 |editor1-last=Herrmann |editor1-first=Bernd |location=New York |pages=149–165 |chapter=DNA from Museum Specimens |doi=10.1007/978-1-4612-4318-2_10 |oclc=852788348 |editor2-last=Hummel |editor2-first=Susanne}}</ref> [[Trevor Worthy]] suggested in 2016 that the specimen originated on an island of [[New Caledonia]] rather than New Zealand, due to a lack of fossil evidence for the lizard in New Zealand caves despite abundant remains of all other known species of New Zealand gecko.<ref name="Worthy2016" /> It was omitted from the ''Conservation Status of New Zealand Reptiles, 2021'' on the basis that it was likely to be from New Caledonia.<ref>{{cite book |last1=Hitchmough |first1=Rod |last2=Barr |first2=Ben |last3=Knox |first3=Carey |last4=Lettink |first4=Marieke |last5=Monks |first5=Joanne M. |last6=Patterson |first6=Geoff B. |author6-link=species:Geoff B. Patterson |last7=Reardon |first7=James T. |last8=van Winkel |first8=Dylan |last9=Rolfe |first9=Jeremy |last10=Michel |first10=Pascale |date=2021 |title=Conservation Status of New Zealand Reptiles, 2021 |series=New Zealand Threat Classification Series |publisher=[[Department of Conservation (New Zealand)|Department of Conservation]] |volume=35 |pages=3–6 |url=https://www.doc.govt.nz/globalassets/documents/science-and-technical/nztcs35entire.pdf |access-date=13 January 2022}}</ref> This was confirmed by the successful sequencing of the specimen's [[mitochondrial DNA]] in 2023, which found that it was nested within the New Caledonian species of [[Diplodactylidae]] rather than the New Zealand species, and distinctive enough to warrant placement in the new genus '''''Gigarcanum'''''.<ref name="Heinicke2023" /> In the DNA analysis, the relationships of New Caledonian geckos were poorly resolved, but ''Gigarcanum'' was usually found to be most closely related to the New Caledonia genera ''[[Eurydactylodes]]'', ''[[Mniarogekko]]'' and/or ''[[Rhacodactylus]]''.<ref name="Heinicke2023" /> |
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| ⚫ | The [[specific name (zoology)|specific epithet]] ''delcourti'' is taken from the surname of French museum worker Alain Delcourt, who discovered the forgotten specimen in the Marseille museum.<ref>{{cite book |last1=Beolens |first1=Bo |last2=Watkins |first2=Michael |last3=Grayson |first3=Michael |date=2011 |title=The Eponym Dictionary of Reptiles |publisher=Johns Hopkins University Press |location=Baltimore |isbn=978-1-4214-0135-5 |page=69}}</ref> According to the authors, the genus name ''Gigarcanum'' derives from "a combination of two words: the Latin adjective ''gigas'', meaning giant and taken from the Ancient Greek Γίγᾱς, and the Latin noun ''arcanum'', meaning secret or mystery. The combination refers to the size of the type species and the unknown provenance of the only known specimen".<ref name=Heinicke2023/> |
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== Description == |
== Description == |
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[[File:Gigarcanum size comparison.jpg|thumb|Size comparison of the holotype (top) with ''[[Rhacodactylus leachianus]]'']] |
[[File:Gigarcanum size comparison.jpg|thumb|Size comparison of the holotype (top) with ''[[Rhacodactylus leachianus]]'']] |
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''Gigarcanum delcourti'' is 50% longer and was likely several times heavier than the largest living gecko, the also New Caledonian ''[[Rhacodactylus leachianus]]<ref name=Heinicke2023/> |
''Gigarcanum delcourti'' is 50% longer and was likely several times heavier than the largest living gecko, the also New Caledonian ''[[Rhacodactylus leachianus]]'',<ref name=Heinicke2023/> with a [[snout-to-vent length]] (SVL) of {{convert|37|cm|in|abbr=on|sigfig=3}} and an overall length (including tail) of at least {{convert|60|cm|in|abbr=on|sigfig=3}}.<ref name=Wilson2004/> The body is robust, and the tail is tapering, cylindrical and weakly annulated. The skull is large, and makes up about 20% of the SVL. The digits bear claws, and are weakly-moderately webbed. The digit pads are rectangular and broad.<ref name=Heinicke2023/> The body colour is yellowish-brown, with dark reddish-brown stripes running along the length of the upper body.<ref name=Bauer1986/> |
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== Ecology == |
== Ecology == |
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Based on comparison with its living relatives, it was probably a [[Nocturnality|nocturnal]] arboreal animal that climbed trees. It probably had diet mainly of arthropods, but possibly also seasonally consumed fruit. It likely had a clutch size of two, as all other known New Caledonian geckos do, though whether it was [[oviparous]] or [[viviparous]] is uncertain.<ref name=Heinicke2023/> |
Based on comparison with its living relatives, it was probably a [[Nocturnality|nocturnal]] arboreal animal that climbed trees. It probably had a diet mainly of arthropods, but possibly also seasonally consumed fruit. It likely had a clutch size of two, as all other known New Caledonian geckos do, though whether it was [[oviparous]] or [[viviparous]] is uncertain.<ref name=Heinicke2023/> |
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== Extinction == |
== Extinction == |
||
''Gigarcanum delcourti'' was likely extinct or extremely rare by the time of the colonisation of New Caledonia in the mid 19th century, due to the absence of any other records of the species.<ref name=Heinicke2023/> |
''Gigarcanum delcourti'' was likely extinct or extremely rare by the time of the colonisation of New Caledonia in the mid 19th century, due to the absence of any other records of the species.<ref name=Heinicke2023/> |
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==Taxonomy== |
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In the DNA analysis, the relationships of New Caledonian geckos were poorly resolved, but ''Gigarcanum'' was usually found to most closely related to the New Caledonia genera ''[[Eurydactylodes]]'', ''[[Mniarogekko]]'' and/or ''[[Rhacodactylus]]''.<ref name=Heinicke2023/> |
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==References== |
==References== |
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{{Reflist}} |
{{Reflist}} |
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{{Wikispecies|Hoplodactylus delcourti}} |
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{{Taxonbar|from=Q1737353}} |
{{Taxonbar|from=Q1737353}} |
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Latest revision as of 11:05, 21 June 2024
| Gigarcanum | |
|---|---|
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| Model in Lille Natural History Museum | |
Scientific classification 
| |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Order: | Squamata |
| Family: | Diplodactylidae |
| Genus: | †Gigarcanum Heinicke, et al, 2023 |
| Species: | †G. delcourti
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| Binomial name | |
| †Gigarcanum delcourti | |
| Synonyms | |
| |
Gigarcanum delcourti, formerly Hoplodactylus delcourti, is an extinct species of gecko in the family Diplodactylidae. It is the largest known of all geckos, with a snout-to-vent length (SVL) of 37 cm (14.6 in) and an overall length (including tail) of at least 60 cm (23.6 in).[2] It is only known from a single taxidermied specimen collected in the 19th century that was rediscovered unlabelled in a museum in France. The origin of the specimen was undocumented.[1] While originally suggested to have been from New Zealand and the kawekaweau of Māori oral tradition,[3][4][5] DNA evidence from the specimen suggests that it originates from New Caledonia.[6]
History
[edit]According to the report of Major W. G. Mair in 1873, in 1870, a Māori chief said that he had killed a kawekaweau he found under the bark of a dead rātā tree in the Waimana Valley in Te Urewera on the North Island of New Zealand. This is the only documented report of anyone ever seeing a kawekaweau alive. Mair reported the chiefs description of the animal as being "two feet long and as thick as a man’s wrist; colour brown, striped longitudinally with dull red".[7]
A single stuffed specimen was "discovered" in the basement of the Natural History Museum of Marseille in 1986; the origins and date of collection of the specimen remain a mystery, as it was unlabelled when it was found.[7] It has been present in the collection of the museum since at least the 1870s, and likely since the 1830s based on its unusual preservation style of being eviscerated, dried and mounted, rather than being kept in spirits as is more common for preserved specimens. The specimen is missing the internal organs and most of the axial skeleton, but retains the skull and appendicular skeleton.[6] It was described as the new species Hoplodactylus delcourti. Initially, scientists examining the specimen suggested that it was from New Zealand and was in fact the lost kawekaweau, a giant and mysterious forest lizard of Māori oral tradition.[7]
Attempts to extract DNA from the sole specimen in 1994 were unsuccessful.[8] Trevor Worthy suggested in 2016 that the specimen originated on an island of New Caledonia rather than New Zealand, due to a lack of fossil evidence for the lizard in New Zealand caves despite abundant remains of all other known species of New Zealand gecko.[5] It was omitted from the Conservation Status of New Zealand Reptiles, 2021 on the basis that it was likely to be from New Caledonia.[9] This was confirmed by the successful sequencing of the specimen's mitochondrial DNA in 2023, which found that it was nested within the New Caledonian species of Diplodactylidae rather than the New Zealand species, and distinctive enough to warrant placement in the new genus Gigarcanum.[6] In the DNA analysis, the relationships of New Caledonian geckos were poorly resolved, but Gigarcanum was usually found to be most closely related to the New Caledonia genera Eurydactylodes, Mniarogekko and/or Rhacodactylus.[6]
The specific epithet delcourti is taken from the surname of French museum worker Alain Delcourt, who discovered the forgotten specimen in the Marseille museum.[10] According to the authors, the genus name Gigarcanum derives from "a combination of two words: the Latin adjective gigas, meaning giant and taken from the Ancient Greek Γίγᾱς, and the Latin noun arcanum, meaning secret or mystery. The combination refers to the size of the type species and the unknown provenance of the only known specimen".[6]
Description
[edit]
Gigarcanum delcourti is 50% longer and was likely several times heavier than the largest living gecko, the also New Caledonian Rhacodactylus leachianus,[6] with a snout-to-vent length (SVL) of 37 cm (14.6 in) and an overall length (including tail) of at least 60 cm (23.6 in).[2] The body is robust, and the tail is tapering, cylindrical and weakly annulated. The skull is large, and makes up about 20% of the SVL. The digits bear claws, and are weakly-moderately webbed. The digit pads are rectangular and broad.[6] The body colour is yellowish-brown, with dark reddish-brown stripes running along the length of the upper body.[7]
Ecology
[edit]Based on comparison with its living relatives, it was probably a nocturnal arboreal animal that climbed trees. It probably had a diet mainly of arthropods, but possibly also seasonally consumed fruit. It likely had a clutch size of two, as all other known New Caledonian geckos do, though whether it was oviparous or viviparous is uncertain.[6]
Extinction
[edit]Gigarcanum delcourti was likely extinct or extremely rare by the time of the colonisation of New Caledonia in the mid 19th century, due to the absence of any other records of the species.[6]
References
[edit]- ^ a b Hitchmough, R.; van Winkel, D.; Lettink, M.; Chapple, D. (2019). "Hoplodactylus delcourti ". IUCN Red List of Threatened Species. 2019: e.T10254A120158840. doi:10.2305/IUCN.UK.2019-2.RLTS.T10254A120158840.en.
- ^ a b Wilson, Kerry-Jayne (2004). Flight of the Huia: Ecology and Conservation of New Zealand's Frogs, Reptiles, Birds and Mammals. Canterbury University Press. ISBN 0-908812-52-3. OCLC 937349394.
- ^ Worthy, T.H. (March 1997). "Quaternary fossil fauna of South Canterbury, South Island, New Zealand". Journal of the Royal Society of New Zealand. 27 (1): 67–162. Bibcode:1997JRSNZ..27...67W. doi:10.1080/03014223.1997.9517528.
- ^ Tennyson, Alan J.D. (2010). "The origin and history of New Zealand's terrestrial vertebrates" (PDF). New Zealand Ecological Society. 34 (1): 6–27.
- ^ a b Worthy, Trevor H. (2016). "A Review of the Fossil Record of New Zealand Lizards". New Zealand Lizards. Springer International Publishing. pp. 65–86. doi:10.1007/978-3-319-41674-8_3. ISBN 978-3-319-41672-4.
- ^ a b c d e f g h i Heinicke, Matthew P.; Nielsen, Stuart V.; Bauer, Aaron M.; Kelly, Ryan; Geneva, Anthony J.; Daza, Juan D.; Keating, Shannon E.; Gamble, Tony (2023-06-19). "Reappraising the evolutionary history of the largest known gecko, the presumably extinct Hoplodactylus delcourti, via high-throughput sequencing of archival DNA". Scientific Reports. 13 (1): 9141. Bibcode:2023NatSR..13.9141H. doi:10.1038/s41598-023-35210-8. ISSN 2045-2322. PMC 10279644. PMID 37336900.
- ^ a b c d Bauer, Aaron M.; Russell, Anthony P. (1986). "Hoplodactylus delcourti n. sp. (Reptilia: Gekkonidae), the largest known gecko" (PDF). New Zealand Journal of Zoology. 13: 141–148. doi:10.1080/03014223.1986.10422655. Archived from the original (PDF) on 2013-04-20.
- ^ Cooper, Alan (1994). "DNA from Museum Specimens". In Herrmann, Bernd; Hummel, Susanne (eds.). Ancient DNA : Recovery and Analysis of Genetic Material from Paleontological, Archaeological, Museum, Medical, and Forensic Specimens. New York: Springer New York. pp. 149–165. doi:10.1007/978-1-4612-4318-2_10. ISBN 978-1-4612-4318-2. OCLC 852788348.
- ^ Hitchmough, Rod; Barr, Ben; Knox, Carey; Lettink, Marieke; Monks, Joanne M.; Patterson, Geoff B.; Reardon, James T.; van Winkel, Dylan; Rolfe, Jeremy; Michel, Pascale (2021). Conservation Status of New Zealand Reptiles, 2021 (PDF). New Zealand Threat Classification Series. Vol. 35. Department of Conservation. pp. 3–6. Retrieved 13 January 2022.
- ^ Beolens, Bo; Watkins, Michael; Grayson, Michael (2011). The Eponym Dictionary of Reptiles. Baltimore: Johns Hopkins University Press. p. 69. ISBN 978-1-4214-0135-5.