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<mods:title id="6A2A26B9BA70FF6CB7266BBEAD43200D">The freshwater crabs of Danum Valley Conservation Area in Sabah, East Malaysia, with a description of a new species of Thelphusula Bott, 1969 (Crustacea, Brachyura, Gecarcinucidae, Potamidae, Sesarmidae)</mods:title>
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<mods:namePart id="C01509BBC5B61EB9822EBC7F0012EFA3">Ng, Peter K. L.</mods:namePart>
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<mods:namePart id="9D5B85AE5F3ECD1B662B2A388FE42E6F">Ng, Paul Y. C.</mods:namePart>
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<treatment id="1F52DF65FAC270F55F190926B99313CB" ID-GBIF-Taxon="144148301" LSID="urn:lsid:plazi:treatment:1F52DF65FAC270F55F190926B99313CB" httpUri="http://treatment.plazi.org/id/1F52DF65FAC270F55F190926B99313CB" lastPageNumber="89" pageId="0" pageNumber="89">
<subSubSection id="2933745CD1DC8F9BE223EB7F315C60DC" pageId="0" pageNumber="89" type="nomenclature">
<paragraph id="98CE07AB3088B30B4448A547A9580AB5" pageId="0" pageNumber="89">
<taxonomicName id="A8BBCC84BCBAFEDA4545B9DA66F1A7EA" ID-CoL="8W6D7" authority="Ng, 1991" authorityName="Ng" authorityYear="1991" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa terrapes" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="terrapes">Arachnothelphusa terrapes Ng, 1991</taxonomicName>
Figure 5
</paragraph>
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<subSubSection id="76FE8EDCB3A328D40BE7B9AC7DE9C760" pageId="0" pageNumber="89" type="reference_group">
<paragraph id="FADBC3ED7B1CD7C9FE6465E980E22984" pageId="0" pageNumber="89">
<taxonomicName id="A9F73707F44AE0EA973B1E3481CF6FE5" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa terrapes" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="terrapes">Arachnothelphusa terrapes</taxonomicName>
Ng, 1991: 8, figs 3-6;
<bibRefCitation id="8F9874BCFFB308EDAA611C8BF80539E3" author="Ng, PKL" journalOrPublisher="Raffles Bulletin of Zoology Supplement" pageId="0" pageNumber="89" pagination="1 - 286" title="Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world." volume="17" year="2008">Ng et al. 2008</bibRefCitation>
: 69.
</paragraph>
</subSubSection>
<subSubSection id="C112D72A7FB244EDE57F0E45B0B28EF0" pageId="0" pageNumber="89" type="materials_examined">
<paragraph id="EA390F8C1879EF53073CAC7ACE7DA5C0" pageId="0" pageNumber="89">Material examined.</paragraph>
<paragraph id="E62368529DE739D6ED9F46CE9434AE75" pageId="0" pageNumber="89">
Holotype: male (17.6
<normalizedToken id="B87FCBF91FFF20CB63B7CF3101E2464A" originalValue="×">x</normalizedToken>
13.3 mm) (ZRC 1992.7918), Danum Valley Field Centre, station 507, in dry stump on ridge, Lahad Datu, Sabah, Borneo, leg. H.K. Voris, 23 October 1990. Paratype: female (25.7
<normalizedToken id="DE577C1893BDC359FC2C2339E16EFC06" originalValue="×">x</normalizedToken>
18.6 mm) (ZRC 1992.7919), Danum Valley, Lahad Datu, Sabah, Borneo, leg. S.C. Choy, 21 July 1989. Others: 1 male (30.8
<normalizedToken id="E7C31F47B5E8689C4858CFCC16A897C5" originalValue="×">x</normalizedToken>
20.5 mm), 1 female (30.1
<normalizedToken id="EEA8EA0E03D4DED2E8ECAF32DF92E673" originalValue="×">x</normalizedToken>
20.5 mm, with 26 juvenile crabs) (ZRC 2017.1205), from water-filled tree buttress, ca. 35 cm above ground Danum Valley, Lahad Datu, Sabah, Borneo, Malaysia, 20 July 2017.
</paragraph>
</subSubSection>
<subSubSection id="1A354E69E0FD36A3B0869946953DA62F" pageId="0" pageNumber="89" type="materials_examined">
<paragraph id="9B9C60127AC02C91580EC268EE8F84AA" pageId="0" pageNumber="89">Comparative material.</paragraph>
<paragraph id="E2E7308D638CA86341A442A0F4934358" pageId="0" pageNumber="89">
<taxonomicName id="DEC6797A37C693FF24B3DDE856E82703" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa kadamaiana" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="kadamaiana">Arachnothelphusa kadamaiana</taxonomicName>
(Borradaile, 1900): 1 female (23.2
<normalizedToken id="E481A6FC0911163D51F756B26A45FDBE" originalValue="×">x</normalizedToken>
17.1 mm) (ZRC 2009.0094), Poring, Basin 1A, Sabah, Malaysia, Borneo, coll. R.F. Inger et al., 12 August 1992; 3 males (21.1
<normalizedToken id="D6990BF1F15F3E0434963B9D317522D8" originalValue="×">x</normalizedToken>
15.8 mm, 22.8
<normalizedToken id="F96D251EAFF70F10F0426296B6CD1BB6" originalValue="×">x</normalizedToken>
16.5 mm, 25.3
<normalizedToken id="8C2ECED4DD46CDEE3CCAB21CDC3C93D5" originalValue="×">x</normalizedToken>
18.5 mm) (ZRC 2002.0097), Crocker Range, Sabah,
<geoCoordinate id="8B99356C3FE5C46B0DE58920DFCAE46A" direction="north" orientation="latitude" precision="925" value="5.45">5°27'N</geoCoordinate>
<geoCoordinate id="4246C15884373D5CB7E72F5B1857FEAE" direction="east" orientation="longitude" precision="925" value="116.05">116°03'E</geoCoordinate>
, coll. I. Das, 24 April 2001.
<taxonomicName id="327BDC2103002A2CB2E178D5E5413B52" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa kadamaiana" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="kadamaiana">Arachnothelphusa aff. kadamaiana</taxonomicName>
: 1 female (19.0
<normalizedToken id="E56D7711AC03CCBB7DEE415B5D1C7D85" originalValue="×">x</normalizedToken>
14.2 mm) (ZRC 2002.0098), Bako National Park, Sarawak, coll. I. Das and L. Grismer, 27 March 2001.
<taxonomicName id="B5E2B04CA1280AFA972632414FABE8BA" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa merarapensis" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="merarapensis">Arachnothelphusa merarapensis</taxonomicName>
Grinang, Pui &amp; Ng, 2015: Holotype male (22.5
<normalizedToken id="39E19FD196AE760A71973C42284698B7" originalValue="×">x</normalizedToken>
16.8 mm) (ZRC 2016.0297), water-filled tree-hole, ca. 100 cm above ground, steep dipterocarp forest, Merarap Hot Spring Resort, Lawas, northern Sarawak, Malaysia, Borneo,
<geoCoordinate id="D803908872905790AE7C7ECAA0E0BA55" direction="north" orientation="latitude" precision="1" value="4.3737226">4°22'25.4&quot;N</geoCoordinate>
,
<geoCoordinate id="326EFECBF4117E9F00F21CFC5E179276" direction="east" orientation="longitude" precision="1" value="115.43614">115°26'10.1&quot;E</geoCoordinate>
, 485 m asl, coll. J. Grinang and Y.M. Pui, 31 October 2014.
</paragraph>
</subSubSection>
<subSubSection id="2E9CDCC70E73C8120B05646C5B3BECE5" pageId="0" pageNumber="89" type="colour">
<paragraph id="F2344D2B81506CB038EF8D325A6A7C22" pageId="0" pageNumber="89">Colour.</paragraph>
<paragraph id="6B175D3B6C91814FB87926F36F598793" pageId="0" pageNumber="89">
The live coloration of this species observed in the recent pair of specimens is a uniform dark purple colour on the dorsal surface of the carapace, ambulatory legs and chelipeds, with a pale purple to dull white on the thoracic sternum, pleon and distal portions of the ambulatory legs and cheliped fingers (Fig. 5
<normalizedToken id="EE5CDB29A66E29D929BF35DD9C00884D" originalValue="B–F">B-F</normalizedToken>
). The dark purple colouration appears considerably darker when the animal is dry, explaining the original paler colour observation by
<bibRefCitation id="29A712FE4195D9967A2B8A9F7EC50494" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng (1991)</bibRefCitation>
.
</paragraph>
<caption id="1B87B166ADF5FC6C12CB5362D5EFD415" pageId="0" pageNumber="89">
<paragraph id="18C204BD1CD3827EA2F737C8FB8777FC" pageId="0" pageNumber="89">
Figure 5. Habitat and life colour of
<taxonomicName id="4213DECC77985CC3105DC733C8722D11" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa terrapes" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="terrapes">Arachnothelphusa terrapes</taxonomicName>
. A water-filled tree hole at base of tree in Danum Valley where crab was found B water filled tree hole where crabs were hiding C, D male (30.8
<normalizedToken id="85DD3A1B745B757F82F601ECE4D71EF0" originalValue="×">x</normalizedToken>
20.5 mm) (ZRC 2017.1205) E, F female (30.1
<normalizedToken id="7CC9673D46206FAA88886EC90057CC6A" originalValue="×">x</normalizedToken>
20.5 mm, with juvenile crabs) (ZRC 2017.1205).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="9000C73635992871D0820B7DAF0128F8" pageId="0" pageNumber="89" type="remarks">
<paragraph id="966E7D9C46ADFC84D3D6DAA6703117F5" pageId="0" pageNumber="89">Remarks.</paragraph>
<paragraph id="2097BE5E256989B2009D7528F08F703D" pageId="0" pageNumber="89">
<bibRefCitation id="5D202D0E1BE74AB80691C67B36D49AE7" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng (1991)</bibRefCitation>
established
<taxonomicName id="4B4245777A81D4C56D036C6FFA7263BE" genus="Arachnothelphsua" lsidName="Arachnothelphsua" pageId="0" pageNumber="89" rank="genus">Arachnothelphsua</taxonomicName>
for several Bornean species previously classified as
<taxonomicName id="FACD025DF85215CF376143D64364CE60" class="Malacostraca" family="Gecarcinucidae" genus="Thelphusula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Thelphusula" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Thelphusula</taxonomicName>
Bott, 1969, with
<taxonomicName id="4BEB82B9D31A35520D836BD9A82C73DB" class="Malacostraca" family="Potamidae" genus="Potamon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Potamon (Potamon) melanippe" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="melanippe" subGenus="Potamon">Potamon (Potamon) melanippe</taxonomicName>
De Man, 1899, as the type species. Currently, four other species are recognised:
<taxonomicName id="F9AB86CFFBE5C5CC50E0825AAB7B39FE" lsidName="A. kadamaiana" pageId="0" pageNumber="89" rank="species" species="kadamaiana">A. kadamaiana</taxonomicName>
(Borradaile, 1900),
<taxonomicName id="99E4B075F2354CFB6C04967E5005D214" lsidName="A. rhadamanthysi" pageId="0" pageNumber="89" rank="species" species="rhadamanthysi">A. rhadamanthysi</taxonomicName>
(Ng &amp; Goh, 1987),
<taxonomicName id="2E85265B19056802D787ED83350B659B" lsidName="A. terrapes" pageId="0" pageNumber="89" rank="species" species="terrapes">A. terrapes</taxonomicName>
Ng, 1991, and
<taxonomicName id="CC9A4E41DBDCE059092B0461934CBD0C" lsidName="A. merarapensis" pageId="0" pageNumber="89" rank="species" species="merarapensis">A. merarapensis</taxonomicName>
Grinang, Pui &amp; Ng, 2015, all from northern Borneo. One species originally included by
<bibRefCitation id="5A386E5EFFF8CEFD59B15C6857D1CFD0" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng (1991)</bibRefCitation>
in
<taxonomicName id="D789F861994460425B5C3EF412C782DF" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Arachnothelphusa</taxonomicName>
,
<taxonomicName id="A22EC1C0B2075A08C9C85AC2003B195D" class="Malacostraca" family="Gecarcinucidae" genus="Parathelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parathelphusa (Liotelphusa) nobilii" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="nobilii" subGenus="Liotelphusa">Parathelphusa (Liotelphusa) nobilii</taxonomicName>
Colosi, 1920, was transferred to
<taxonomicName id="4AC2B413400150D423438852FFC91649" class="Malacostraca" family="Gecarcinucidae" genus="Stygothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Stygothelphusa" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Stygothelphusa</taxonomicName>
Ng, 1989, by
<bibRefCitation id="E6D4D4F8E473F8ADD48436E979CCF4EF" pageId="0" pageNumber="89">Ng and Álvarez (2000)</bibRefCitation>
(see also
<bibRefCitation id="C4CEA38FE8FD267D683D5162FE926B67" author="Ng, PKL" journalOrPublisher="Raffles Bulletin of Zoology Supplement" pageId="0" pageNumber="89" pagination="91 - 97" title="Stygothelphusacranbrooki, a new species of cave crab from Gua Sireh, Sarawak, Malaysia (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="29" year="2013">Ng 2013</bibRefCitation>
;
<bibRefCitation id="FF962C9E40AEEAE0832665F98BC7D194" author="Ng, PKL" journalOrPublisher="Zootaxa" pageId="0" pageNumber="89" pagination="90 - 96" title="On a new species of troglobitic crab of the genus Stygothelphusa Ng, 1989, from Sarawak, Malaysia (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." url="https://doi.org/10.11646/zootaxa.3774.1.7" volume="3774" year="2014">Ng and Grinang 2014</bibRefCitation>
).
</paragraph>
<paragraph id="A74EDE685F6DDFF4188608ED5787A2BC" pageId="0" pageNumber="89">
<taxonomicName id="8F592BCDB8A242212B1D02BB68313052" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa terrapes" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="terrapes">Arachnothelphusa terrapes</taxonomicName>
is easily distinguished from congeners by the deep U-shaped sinus separating the truncate external orbital tooth from the epibranchial tooth (
<bibRefCitation id="DE3767A463CA94655F29264D7CEB36B7" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng 1991</bibRefCitation>
: fig. 3).
<taxonomicName id="DB42513E30EBA0290EBA3E0E9A9289F7" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa merarapensis" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="merarapensis">Arachnothelphusa merarapensis</taxonomicName>
has a superficially similar anterolateral margin except that the two teeth are separated by an obtusely triangular broad cleft instead (
<bibRefCitation id="40C0A9A9CC79375F3FC37734D37A2D54" author="Grinang, J" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="454 - 460" title="A new species of tree-hole dwelling crab of the genus Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Brachyura: Gecarcinucidae) from northern Sarawak, Malaysia, Borneo." volume="63" year="2015">Grinang et al. 2015</bibRefCitation>
: fig. 1A, B). Other congeners have the epibranchial tooth separated by a V-shaped notch or the margin is almost entire (De
<bibRefCitation id="090E83E11409F54C896C487446F38AB2" author="Man, JG De" journalOrPublisher="Notes Leyden Museum" pageId="0" pageNumber="89" pagination="53 - 144" title="Zoological Results of the Dutch Expedition to Central Borneo. The Crustacea. Part II, Brachyura." volume="21" year="1899">Man 1899</bibRefCitation>
: pl. 9;
<bibRefCitation id="7708F8B064F07E4D9DF7C8636218FCD6" author="Ng, PKL" journalOrPublisher="Stygologia" pageId="0" pageNumber="89" pagination="313 - 330" title="Cavernicolous freshwater crabs (CrustaceaDecapoda, Brachyura) from Sabah, Borneo." volume="3" year="1987">Ng and Goh 1987</bibRefCitation>
: pl. 3A;
<bibRefCitation id="4F73BEC949A6A1A30F6F0EC88FCAA30E" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng 1991</bibRefCitation>
: fig. 1A;
<bibRefCitation id="92986AE189C02A929CFC13E31E8E36C9" author="Grinang, J" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="454 - 460" title="A new species of tree-hole dwelling crab of the genus Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Brachyura: Gecarcinucidae) from northern Sarawak, Malaysia, Borneo." volume="63" year="2015">Grinang et al. 2015</bibRefCitation>
: fig. 6A).
</paragraph>
<paragraph id="57DC71AFECF839BD436ADDAD55A8C24E" pageId="0" pageNumber="89">
The biology of species of
<taxonomicName id="FF79E6AC1297E37FC457BE5C471BB900" genus="Arachnothelphsua" lsidName="Arachnothelphsua" pageId="0" pageNumber="89" rank="genus">Arachnothelphsua</taxonomicName>
is not well known. All known species are represented by only very few specimens (
<bibRefCitation id="30412CD33B593C0F65D28279F4F986F0" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng 1991</bibRefCitation>
) and there is often no accompanying ecological data.
<taxonomicName id="6FA97A4FB737A19EB3515FEB558C75C9" genus="Arachnothelphsua" lsidName="Arachnothelphsua melanippe" pageId="0" pageNumber="89" rank="species" species="melanippe">Arachnothelphsua melanippe</taxonomicName>
and
<taxonomicName id="27469E598CACFC8075D931356549B388" lsidName="A. kadaimana" pageId="0" pageNumber="89" rank="species" species="kadaimana">A. kadaimana</taxonomicName>
were both described without any indication of their biology (
<bibRefCitation id="5F2D48F8C88BD796147554C68854AE7E" author="Man, JG De" journalOrPublisher="Notes Leyden Museum" pageId="0" pageNumber="89" pagination="53 - 144" title="Zoological Results of the Dutch Expedition to Central Borneo. The Crustacea. Part II, Brachyura." volume="21" year="1899">De Man 1899</bibRefCitation>
;
<bibRefCitation id="8E854DE8D118556BB57A30B8DA05D275" author="Borradaile, LA" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="0" pageNumber="89" pagination="93 - 95" title="On a small collection of decapod crustaceans from freshwater in North Borneo." url="https://doi.org/10.1111/j.1096-3642.1890.tb01706.x" volume="1900" year="1900">Borradaile 1900</bibRefCitation>
).
<bibRefCitation id="D80C205A09AD7D4678CE71EF1E82C3D5" author="Grinang, J" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="454 - 460" title="A new species of tree-hole dwelling crab of the genus Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Brachyura: Gecarcinucidae) from northern Sarawak, Malaysia, Borneo." volume="63" year="2015">Grinang et al. (2015)</bibRefCitation>
reported on a female specimen from Poring in Sabah but there was no information on where it was found. In the ZRC there are two lots of
<taxonomicName id="CD91F7E942D42EBF002863A28D988A8E" lsidName="A. kadaimana" pageId="0" pageNumber="89" rank="species" species="kadaimana">A. kadaimana</taxonomicName>
(ZRC 2009.0094, ZRC 2002.0097) from Sabah, also without specific habitat data. A female specimen of
<taxonomicName id="AF33C99749AC5D2CF5DE22895C3F12A7" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Arachnothelphusa</taxonomicName>
, close to but not conspecific with
<taxonomicName id="0657F09ADEDB6E5B81817C6FC3DE7A7E" lsidName="A. kadaimana" pageId="0" pageNumber="89" rank="species" species="kadaimana">A. kadaimana</taxonomicName>
(ZRC 2002.0098) from Bako National Park in Sarawak was collected from a tree trunk (I. Das, per. comm.).
<taxonomicName id="46BBC9102106551EF51E216C48AF5B2B" genus="Arachnothelphsua" lsidName="Arachnothelphsua terrapes" pageId="0" pageNumber="89" rank="species" species="terrapes">Arachnothelphsua terrapes</taxonomicName>
was found low on shrubs (
<bibRefCitation id="2178FA9A3B9C92EB726EC96C401477AF" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng 1991</bibRefCitation>
) while
<taxonomicName id="51CCFDE53F817CEF33D370BAECF61E9B" lsidName="A. rhadamanthysi" pageId="0" pageNumber="89" rank="species" species="rhadamanthysi">A. rhadamanthysi</taxonomicName>
was collected on a stalagmite wall inside a cave (
<bibRefCitation id="F71DF4E2B323052BC68EDBEA44D5221F" author="Ng, PKL" journalOrPublisher="Stygologia" pageId="0" pageNumber="89" pagination="313 - 330" title="Cavernicolous freshwater crabs (CrustaceaDecapoda, Brachyura) from Sabah, Borneo." volume="3" year="1987">Ng and Goh 1987</bibRefCitation>
). The most detailed account so far was that by
<bibRefCitation id="E2CE45A0C97F41D5D50F605BB4B81533" author="Grinang, J" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="454 - 460" title="A new species of tree-hole dwelling crab of the genus Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Brachyura: Gecarcinucidae) from northern Sarawak, Malaysia, Borneo." volume="63" year="2015">Grinang et al. (2015)</bibRefCitation>
for
<taxonomicName id="4AEF7D58E52475A7957A32FFF9F077D5" lsidName="A. merarapensis" pageId="0" pageNumber="89" rank="species" species="merarapensis">A. merarapensis</taxonomicName>
from Merarap Hot Springs in Sarawak, who obtained the species from low tree holes approx. 150 cm from the ground. It is not known if the crabs live in phytotelms higher up on the forest trees.
<taxonomicName id="B946DAB383097FCAB44F6B3780551E55" genus="Arachnothelphsua" lsidName="Arachnothelphsua rhadamanthysi" pageId="0" pageNumber="89" rank="species" species="rhadamanthysi">Arachnothelphsua rhadamanthysi</taxonomicName>
has since been photographed by naturalists in the forested area outside Gomantong caves where it was first found, suggesting it is only a facultative cave dweller (
<bibRefCitation id="E4FF5276C9D8DF9AB6CA10928A727F30" author="Christensen, K" journalOrPublisher="Abhandlungen der Senckenbergischen naturforschenden Gesellschaft" pageId="0" pageNumber="89" url="http://www.tropicalbats.com/tropicalbats-blog/borneo-caving1" year="2015">Christensen 2015</bibRefCitation>
).
</paragraph>
<paragraph id="B3A3F5C8A7BDD446E3860138AFC97628" pageId="0" pageNumber="89">
<taxonomicName id="8E1BCE72B2956B4EF306B7326D771298" class="Malacostraca" family="Gecarcinucidae" genus="Arachnothelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arachnothelphusa terrapes" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="terrapes">Arachnothelphusa terrapes</taxonomicName>
was described from a pair of specimens, the first, a female collected in 1989 which moulted shortly after capture and died, leaving both the animal and exuvium in poor condition. The male holotype was collected a year later from a dry tree stump, with the live coloration being a deep reddish brown on dorsal surfaces, chelipeds and legs (
<bibRefCitation id="078460016706147BAB5D2189E6E7B3D7" author="Ng, PKL" journalOrPublisher="Zoologische Mededelingen" pageId="0" pageNumber="89" pagination="1 - 12" title="Bornean freshwater crabs of the genus Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura: Gecarcinucidae)." volume="65" year="1991">Ng 1991</bibRefCitation>
: 11). In view of the present observations of this species as a tree hole specialist, it is likely the holotype male was only taking temporary refuge in the tree stump when it was found.
</paragraph>
<paragraph id="7FCEAECBDAD34C1A50AC343C011A968B" pageId="0" pageNumber="89">
Two individuals of
<taxonomicName id="4C65FA941BA56762A82ED9DEC551E504" lsidName="A. terrapes" pageId="0" pageNumber="89" rank="species" species="terrapes">A. terrapes</taxonomicName>
were observed at 0030 hours in Danum Valley, less than 50 m apart. The first, a large adult male was observed at the edge of a water-filled hole on a tree buttress, roughly 35 cm above the ground (Fig. 5A). A second, a female carrying newly hatched young under its pleon, was found inside a water filled tree hole approx. 150 cm above ground (Fig. 5F). This species is nocturnal and highly sensitive to light, swiftly retreating into their holes when disturbed. Additional observations by other naturalists who have photographed this species in Danum Valley suggest it is always found on trees and never on the forest floor itself (unpublished data). It is clear that
<taxonomicName id="38AEEFC136F13D3E066FE613D26D833C" lsidName="A. terrapes" pageId="0" pageNumber="89" rank="species" species="terrapes">A. terrapes</taxonomicName>
is a true phytotelm species and predominantly arboreal in nature, living exclusively in tree-holes; although they will move in search of other tree holes if the one they are residing begins to dry up or when searching for a mate. All specimens have been observed on the lower parts of trees and it is not known if they climb much higher up. All specimens recorded so far have been solitary. The present observation of an adult female carrying young (Fig. 5F) is notable and confirms the species breeds in the phytotelm.
</paragraph>
<paragraph id="57B34643DFAE490CFA9C2782A837BA0D" pageId="0" pageNumber="89">
The biology of obligate arboreal crabs has been discussed at length by
<bibRefCitation id="97AB01C168C0DD2E787C3DA91AFAE9B9" pageId="0" pageNumber="89">Sivasothi et al. (1993)</bibRefCitation>
,
<bibRefCitation id="BA2AA063995DB0E976141B6C09AB9CEE" author="Sivasothi, N" journalOrPublisher="Crustaceana" pageId="0" pageNumber="89" pagination="25 - 38" title="Niche preferences of tree-climbing crabs in Singapore mangroves." url="https://doi.org/10.1163/156854000504093" volume="73" year="2000">Sivasothi (2000)</bibRefCitation>
,
<bibRefCitation id="93A6B4DA3B888FAAD1D5A98D740509B7" author="Cumberlidge, N" journalOrPublisher="Journal of Crustacean Biology" pageId="0" pageNumber="89" pagination="302 - 308" title="Tree-climbing crabs (Potamonautidae and Sesarmidae) from phytotelmic microhabitats in rainforest canopy in Madagascar." url="https://doi.org/10.1651/C-2532" volume="25" year="2005">Cumberlidge et al. (2005)</bibRefCitation>
,
<bibRefCitation id="ED66CB2B9943A6BBA727C3FF3395E83F" author="Fratini, S" journalOrPublisher="Evolutionary Ecology Research" pageId="0" pageNumber="89" pagination="219 - 233" title="Tree-climbing mangrove crabs: a case of convergent evolution." volume="7" year="2005">Fratini et al. (2005)</bibRefCitation>
,
<bibRefCitation id="D3B55AFD43B47C6B8C860A857FF5AA66" author="Grinang, J" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="454 - 460" title="A new species of tree-hole dwelling crab of the genus Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Brachyura: Gecarcinucidae) from northern Sarawak, Malaysia, Borneo." volume="63" year="2015">Grinang et al. (2015)</bibRefCitation>
,
<bibRefCitation id="10343CBC24A65249DE51488625DCFAD3" author="Ng, PKL" journalOrPublisher="Raffles Bulletin of Zoology Supplement" pageId="0" pageNumber="89" pagination="216 - 225" title="Observations on the taxonomy and ecology of Labuaniumpolitum (De Man, 1887) (Crustacea: Decapoda: Sesarmidae), an obligate arboreal crab on the nipah palm, Nypafructicans (Arecales: Arecaceae)." volume="31" year="2015">Ng et al. (2015)</bibRefCitation>
,
<bibRefCitation id="2151DA743B75BB58872DCD37F2196C90" author="Wehrtmann, IS" journalOrPublisher="Journal of Crustacean Biology" pageId="0" pageNumber="89" pagination="303 - 309" title="First confirmed report of a primary freshwater crab (Brachyura: Pseudothelphusidae) associated with bromeliads in the Neotropics." url="https://doi.org/10.1163/1937240X-00002429" volume="36" year="2016">Wehrtmann et al. (2016)</bibRefCitation>
and
<bibRefCitation id="3FE69573A65C6F4AF7134CF611778424" author="Kumar, AB" journalOrPublisher="Journal of Crustacean Biology" pageId="0" pageNumber="89" pagination="157 - 167" title="Description of a new genus and new species of a completely arboreal crab (Decapoda: Brachyura: Gecarcinucidae) from the Western Ghats in India, with notes on the ecology of arboreal crabs." url="https://doi.org/10.1093/jcbiol/rux012" volume="37" year="2017">Kumar et al. (2017)</bibRefCitation>
. While most are primary freshwater crabs (sensu
<bibRefCitation id="43F919B0D921A3C23A0724FA667C8339" author="Yeo, DCJ" editor="Yeo, DCJ" journalOrPublisher="Crustaceana Monographs" pageId="0" pageNumber="89" pagination="1 - 6" title="Preface-Freshwater Decapod Biology in the 21 st Century." url="https://doi.org/10.1163/9789004207615_002" volume="19" volumeTitle="Advances in freshwater decapod systematics and biology." year="2014">Yeo et al. 2014</bibRefCitation>
) of the families
<taxonomicName id="799B97F9223074965F6A1A3C16139FBF" family="Potamidae" lsidName="" pageId="0" pageNumber="89" rank="family">Potamidae</taxonomicName>
and
<taxonomicName id="447A925701413C5A085A579278B391BC" family="Gecarcinucidae" lsidName="" pageId="0" pageNumber="89" rank="family">Gecarcinucidae</taxonomicName>
, members of two South East and East Asian sesarmid genera,
<taxonomicName id="A4F7B809B765AD47BF69D4D0D34FA42C" class="Malacostraca" family="Sesarmidae" genus="Geosesarma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Geosesarma" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Geosesarma</taxonomicName>
De Man, 1892, and
<taxonomicName id="D915ED0CC18DE4C77129E5BEBD7BA58D" class="Malacostraca" family="Sesarmidae" genus="Scandarma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scandarma" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Scandarma</taxonomicName>
Schubart, Liu &amp; Cuesta, 2003, are primarily arboreal in habits (see
<bibRefCitation id="5DA981F47D24622DD4B1CE666E06883A" author="Schubart, CD" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="49 - 59" title="A new genus and species of tree-climbing crab (Crustacea: Brachyura: Sesarmidae) from Taiwan with notes on its ecology and larval morphology." volume="51" year="2003">Schubart et al. 2003</bibRefCitation>
;
<bibRefCitation id="932239B38A50E09292AF2C63BA1DF397" author="Naruse, T" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="337 - 341" title="Scandarmasplendidum, a new species of tree-climbing crab (Decapoda: Brachyura: Sesarmidae) from Sarawak, Malaysia, Borneo." volume="55" year="2007">Naruse and Ng 2007</bibRefCitation>
;
<bibRefCitation id="BC9A60FD586FB2EA0026C066D59DA2DF" author="Ng, PKL" journalOrPublisher="Raffles Bulletin of Zoology" pageId="0" pageNumber="89" pagination="226 - 242" title="On the identities of the highland vampire crabs, Geosesarmafoxi (Kemp, 1918) and G. serenei Ng, 1986, with description of a new phytotelmic species from Penang, Peninsular Malaysia (Crustacea: Decapoda: Brachyura: Sesarmidae)." volume="65" year="2017">Ng 2017</bibRefCitation>
). There are of course some species of freshwater crabs that occasionally climb trees and use phytotelms but can also be found on the forest floor or nearby streams, and thus are not obligate arboreal species. In Asia,
<taxonomicName id="86CB0CA94718F22165EE34BC3E7E3130" class="Malacostraca" family="Gecarcinucidae" genus="Sundathelphusa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sundathelphusa celer" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="celer">Sundathelphusa celer</taxonomicName>
(Ng, 1991) (
<taxonomicName id="C719CC0F0CE6DAD8BCAC11B61CBDB9FF" family="Gecarcinucidae" lsidName="" pageId="0" pageNumber="89" rank="family">Gecarcinucidae</taxonomicName>
), from the Philippines was collected in a tree hollow above ground, but it is not certain if it is a wholly arboreal species (
<bibRefCitation id="0627F076A8652F354BD04E090CA76B65" author="Ng, PKL" editor="Fransen, CHJM" journalOrPublisher="Crustaceana Monographs" pageId="0" pageNumber="89" pagination="561 - 571" title="On the identity of Para-Bary-thelphusa grapsoides longipes Balss, 1937, with description of a new species from the Philippines (Brachyura, Gecarcinucidae)." volume="14" volumeTitle="Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume." year="2010">Ng 2010</bibRefCitation>
).
<taxonomicName id="CC2407C905EA116DC423E9AD2DC8C218" class="Malacostraca" family="Gecarcinucidae" genus="Perbrinckia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Perbrinckia scansor" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="scansor">Perbrinckia scansor</taxonomicName>
(Ng, 1995) from Sri Lanka has also been noted to have arboreal tendencies but is clearly a terrestrial species that occasionally climbs trees (
<bibRefCitation id="3700FED6FF1C6D09CE5965FEAD228136" pageId="0" pageNumber="89">Ng 1995</bibRefCitation>
: 183;
<bibRefCitation id="6BFB50910EBC523900F57A14D63393A3" author="Ng, PKL" journalOrPublisher="Zeylanica" pageId="0" pageNumber="89" pagination="113 - 199" title="The freshwater crabs of Sri Lanka (Decapoda: Brachyura: Paratelphusidae)." volume="6" year="2001">Ng and Tay 2001</bibRefCitation>
: 148-149). In Hainan, China, some species of
<taxonomicName id="40B2D900C12D1E97B8CC04C0FD5A2D73" class="Malacostraca" family="Potamidae" genus="Neotiwaripotamon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Neotiwaripotamon" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="genus">Neotiwaripotamon</taxonomicName>
Dai &amp; Naiyanetr, 1994, are known to be primarily arboreal (unpublished data;
<bibRefCitation id="E7CCAF2461435EF848101545ECBCA08C" author="Shih, H-T" journalOrPublisher="Science Development, Taiwan" pageId="0" pageNumber="89" pagination="40 - 46" title="East Asian Freshwater Crabs." volume="428" year="2008">Shih 2008</bibRefCitation>
). In India, the only known true arboreal phytotelm species is the recently described
<taxonomicName id="DBE33201EAA1C9BA88D99477475FE2BD" class="Malacostraca" family="Gecarcinucidae" genus="Kani" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kani maranjandu" order="Decapoda" pageId="0" pageNumber="89" phylum="Arthropoda" rank="species" species="maranjandu">Kani maranjandu</taxonomicName>
Kumar, Raj &amp; Ng, 2017.
</paragraph>
</subSubSection>
</treatment>
</document>
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