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<mods:title id="A37FADF83FC8FE1911F5DCB8531D05E0">Revision of the freshwater crabs of the Johora tahanensis (Bott, 1966) species group (Crustacea, Brachyura, Potamidae), with a key to the genus</mods:title>
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<mods:namePart id="04403955275B9DC602C08A2A205E9FB1">Ng, Peter K. L.</mods:namePart>
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<taxonomicName id="B0C7ACA5F0834D9BCE8514792E13243F" ID-CoL="8YXLN" LSID="EF6BEDB2-6B88-5D0D-9F4C-B693F42CC32C" authority="Ng, 1990" authorityName="Ng" authorityYear="1990" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora thoi" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="thoi">Johora thoi Ng, 1990</taxonomicName>
<figureCitation id="325F0BEBEE3CF94A5D92B0A18546BC99" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Overall dorsal view A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure1" httpUri="https://binary.pensoft.net/fig/476312" pageId="0" pageNumber="1">Figures 1C</figureCitation>
<figureCitation id="0D9E5A190152E64ED9D3003DDDA510FE" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">, 2C</figureCitation>
<figureCitation id="3DDBF51FCA57D2B7C23308FA00B0351B" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Frontal view of cephalothorax A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure3" httpUri="https://binary.pensoft.net/fig/476314" pageId="0" pageNumber="1">, 3C</figureCitation>
<figureCitation id="D2B35E01C22C6A0C40024C3F63DCBA72" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Left third maxilliped A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure4" httpUri="https://binary.pensoft.net/fig/476315" pageId="0" pageNumber="1">, 4C</figureCitation>
<figureCitation id="6BC3B67AB99454424570193A511F61E2" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Anterior thoracic sternum and pleon A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure5" httpUri="https://binary.pensoft.net/fig/476316" pageId="0" pageNumber="1">, 5C</figureCitation>
<figureCitation id="243FE13FB50CE5276271AE9593A5FE2D" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">, 6</figureCitation>
<figureCitation id="6D3AD1C8D09747EBDB6BC9AB120BA477" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. A-D Johora thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu E-H J. thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu I-L J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2020.0361), Terengganu A, E, I left G 1 (ventral view) B, F, J distal part of left G 1 (ventral view) C, G, K distal part of left G 1 (dorsal view) D, H, L left G 2 (ventral view) A-D after Ng (1990: fig. 2 A, C-E). Scale bars: 1.0 mm (A, D, E, H, I, L); 0.5 mm (B, C, F, G, J, K)." figureDoi="10.3897/zookeys.994.56810.figure9" httpUri="https://binary.pensoft.net/fig/476320" pageId="0" pageNumber="1">, 9A-H</figureCitation>
<figureCitation id="7923D58493726F1511CDDA35F4C37536" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Overall dorsal view A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure12" httpUri="https://binary.pensoft.net/fig/476323" pageId="0" pageNumber="1">, 12C</figureCitation>
<figureCitation id="F0272CBB5304176877E321CF62967215" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Female pleon A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure13" httpUri="https://binary.pensoft.net/fig/476324" pageId="0" pageNumber="1">, 13C</figureCitation>
<figureCitation id="227E1D802398BA6C5A81DEE8706207C5" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Sternopleonal cavity and vulvae A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure14" httpUri="https://binary.pensoft.net/fig/476325" pageId="0" pageNumber="1">, 14C</figureCitation>
</paragraph>
</subSubSection>
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<paragraph id="00456306E592F9D6A0745B8C11A438FA" pageId="0" pageNumber="1">
<taxonomicName id="CCB1444257AAB37308E94823F7A7E24B" authorityName="Ng" authorityYear="1990" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora thoi" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="thoi">Johora thoi</taxonomicName>
Ng, 1990: 305, figs 1, 2; 
<bibRefCitation id="845FF152420F93F08A46C1FE748D533B" author="Ng, PKL" editor="Yule, C" journalOrPublisher="Malaysian Academy of Sciences, city" pageId="0" pageNumber="1" pagination="311 - 336" refId="B14" refString="Ng, PKL, 2004. Crustacea: Decapoda, Brachyura. In: Yule, C, Yong, HS, Eds., Freshwater Invertebrates of the Malaysian Region. Malaysian Academy of Sciences, city: 311 - 336" title="Crustacea: Decapoda, Brachyura." volumeTitle="Freshwater Invertebrates of the Malaysian Region." year="2004">Ng 2004</bibRefCitation>
: 321; 
<bibRefCitation id="F1390CB09DCD779DACA7E027FAB4D1DC" author="Ng, PKL" editor="Chua, L" journalOrPublisher="28 - 30 June 2005, Forest Research Institute Malaysia, Kepong" pageId="0" pageNumber="1" pagination="95 - 120" refId="B21" refString="Ng, PKL, Yeo, DCJ, 2007. Malaysian freshwater crabs: conservation prospects and challenges. In: Chua, L, Ed., Proceedings of the Seminar on the Status of Biological Diversity in Malaysia &amp; Threat Assessment of Plant Species in Malaysia. 28-30 June 2005, Forest Research Institute Malaysia, Kepong: 95 - 120" title="Malaysian freshwater crabs: conservation prospects and challenges." volumeTitle="Proceedings of the Seminar on the Status of Biological Diversity in Malaysia &amp; Threat Assessment of Plant Species in Malaysia." year="2007">Ng and Yeo 2007</bibRefCitation>
: 102; 
<bibRefCitation id="CDD4528B4DC111A14E903F133AC73B5A" author="Ng, PKL" journalOrPublisher="Raffles Bulletin of Zoology, Supplement" pageId="0" pageNumber="1" pagination="1 - 286" refId="B18" refString="Ng, PKL, Guinot, D, Davie, PJF, 2008. Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology, Supplement 17: 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>
: 163; 
<bibRefCitation id="6202FA278BA0F24D24C9329C399A12DF" DOI="https://doi.org/10.1016/j.biocon.2009.02.038" author="Cumberlidge, N" journalOrPublisher="Biological Conservation" pageId="0" pageNumber="1" pagination="1665 - 1673" refId="B3" refString="Cumberlidge, N, Ng, PKL, Yeo, DCJ, Magalhaes, C, Campos, MR, Alvarez, F, Naruse, T, Daniels, SR, Esser, LJ, Attipoe, FYK, Clotilde-Ba, F-L, Darwall, W, Mcivor, A, Collen, B, Ram, M, 2009. Freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges. Biological Conservation 142: 1665 - 1673, DOI: https://doi.org/10.1016/j.biocon.2009.02.038" title="Freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges." url="https://doi.org/10.1016/j.biocon.2009.02.038" volume="142" year="2009">Cumberlidge et al. 2009</bibRefCitation>
: table.
</paragraph>
</subSubSection>
<subSubSection id="4924918342C6F1C02B3458096E9AA203" pageId="0" pageNumber="1" type="materials_examined">
<paragraph id="4B38E826096C0C634803018B55AFAF3E" pageId="0" pageNumber="1">Material examined.</paragraph>
<paragraph id="39017CD37BA93FAB65866B794FE26A56" pageId="0" pageNumber="1">
<emphasis id="548BF7E384F4D51EA55BEE2322249AEC" bold="true" italics="true" pageId="0" pageNumber="1">Holotype</emphasis>
: male (41.2 
<normalizedToken id="9EBF060017D60C8231F9B6B9DA9DDEEA" originalValue="×">x</normalizedToken>
33.3 mm) (ZRC 1989.2249), Telok Kalong Besar, Pulau Redang, Terengganu, ca. 
<geoCoordinate id="0D07DEE5AC80EDC3C385C23D730DB8D6" degrees="5" direction="north" minutes="46" orientation="latitude" precision="15" seconds="7" value="5.768611">5°46'7&quot;N</geoCoordinate>
, 
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, coll. Tho YP, 8 March 1989. Others: 1 male (ZRC 1989.3758), in freshwater stream, Telok Kalong Besar, Pulau Redang, Terengganu, coll. Saw LG, 1 August 1989; 1 male (ZRC 1989.3740), Telok Kalong Besar, Pulau Redang, Terengganu, coll. Saw LG, 1 August 1989; 1 male, 1 female (ZRC 1996.2085), site 5, stream behind Pasir Panjang, Pulau Redang, Terengganu, Lim KKP et al., 25 June 1992; 1 male (ZRC 1996.2087), Pulau Redang, Terengganu, Lim KKP et al., 23 June 1992; 3 males (ZRC 1996.2086), site 7, stream behind Telok Kalong Besar, Pulau Redang, Terengganu, coll. Lim KKP et al., 25 June 1992; 6 males, 2 females, 1 juvenile (ZRC 2001.1167), from freshwater rocky stream, East Coast forest trail, ca. 1.2-1.6 km from Pasir Panjang to Telok Dalam, Pulau Redang, Terengganu, coll. Tan HH and Koh LL, 20 June 2001. All locations in Peninsular Malaysia.
</paragraph>
</subSubSection>
<subSubSection id="2D71E9952D90E758AA92FF9EF47436FB" pageId="0" pageNumber="1" type="diagnosis">
<paragraph id="EF2F22B913DE65D67348496430002C87" pageId="0" pageNumber="1">Diagnosis.</paragraph>
<paragraph id="DE11AB99A4D609A7A567EF9E8A53A6E6" pageId="0" pageNumber="1">
Adult carapace width to length ratio 1.19-1.24 (Figs 
<figureCitation id="8EABF6DCFBD2279C23837A673704852E" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Overall dorsal view A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure1" httpUri="https://binary.pensoft.net/fig/476312" pageId="0" pageNumber="1">1C</figureCitation>
, 
<figureCitation id="C14E6592B1A906A08BB25BBD2A2AB34C" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">2C</figureCitation>
, 
<figureCitation id="FF61918F5CCBD33D02C8195E092FBC5D" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6A, B</figureCitation>
, 
<figureCitation id="4C5DBAEC614750B38AE02FD896B6518D" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Overall dorsal view A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure12" httpUri="https://binary.pensoft.net/fig/476323" pageId="0" pageNumber="1">12C</figureCitation>
); dorsal surface gently convex in frontal view, not inflated (Figs 
<figureCitation id="802234E3BDF88C2FDC9BDC27918D902F" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Frontal view of cephalothorax A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure3" httpUri="https://binary.pensoft.net/fig/476314" pageId="0" pageNumber="1">3C</figureCitation>
, 
<figureCitation id="1D9E4266236424E18BA0915A51D4C3EF" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6C</figureCitation>
); frontal margin almost straight or slightly sinuous (Figs 
<figureCitation id="1E05AF36E81F91E6F84C4032EAB85C33" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">2C</figureCitation>
, 
<figureCitation id="361396635DB62D60318EBEB13A7FCE17" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6B</figureCitation>
); suborbital, pterygostomial and sub-branchial regions rugose, pterygostomial region covered with dense setae (Figs 
<figureCitation id="900D3362A065B225736C8DF81DB826AD" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Frontal view of cephalothorax A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure3" httpUri="https://binary.pensoft.net/fig/476314" pageId="0" pageNumber="1">3C</figureCitation>
, 
<figureCitation id="D27E4F383B3847DA5FAFF509EDCB2351" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6C</figureCitation>
); epigastric cristae distinct, distinctly anterior to sharp postorbital cristae, postorbital cristae with lateral edges low, not joining lateral margin (Figs 
<figureCitation id="90EBD767E1432F71679BE7348E4EC041" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">2C</figureCitation>
, 
<figureCitation id="F0FF70800C1BA0531ECF75478FC64FD2" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6B</figureCitation>
); external orbital tooth separated from epibranchial tooth by prominent cleft, epibranchial tooth sharp, distinct (Figs 
<figureCitation id="63CF62A986B86CAC58A398DE59BC56F2" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">2C</figureCitation>
, 
<figureCitation id="D0581271CB6F91B302A00149E15D9DC8" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6B</figureCitation>
); anterolateral margin distinctly convex (Figs 
<figureCitation id="714A54A7A2AAEECEA7EC29F72CFA1D9A" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">2C</figureCitation>
, 
<figureCitation id="F5B8503B9F82FCF66ABF6A6BC829B764" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6B</figureCitation>
); posterolateral margin gently sinuous to almost straight, distinctly converging towards gently convex, entire posterior carapace margin (Figs 
<figureCitation id="294E4D37EE7C85C40A46E0075760E8B3" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsal view of carapace A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249) Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure2" httpUri="https://binary.pensoft.net/fig/476313" pageId="0" pageNumber="1">2C</figureCitation>
, 
<figureCitation id="8621C46769ED07FB3C800A4FAD3706B4" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6B</figureCitation>
); posterior margin of epistome with triangular median triangle, lateral margin obliquely sloping (Figs 
<figureCitation id="97373F470EDBAD7B3CFEB2691EC7E18E" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Frontal view of cephalothorax A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure3" httpUri="https://binary.pensoft.net/fig/476314" pageId="0" pageNumber="1">3C</figureCitation>
, 
<figureCitation id="005E1C38BC5B74FDF2933EED57BE146F" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6C</figureCitation>
); outer surfaces of third maxillipeds with dense, long stiff setae; ischium subrectangular, with shallow median oblique groove (Figs 
<figureCitation id="54EF17259EBA77FE7753E55E9A1EB0EF" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Frontal view of cephalothorax A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure3" httpUri="https://binary.pensoft.net/fig/476314" pageId="0" pageNumber="1">3C</figureCitation>
, 
<figureCitation id="A4918B15C6FD03AB1F7F42DD28CF66A1" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Left third maxilliped A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure4" httpUri="https://binary.pensoft.net/fig/476315" pageId="0" pageNumber="1">4C</figureCitation>
); ambulatory legs relatively longer, length to width ratio of merus of fourth ambulatory leg 3.0-3.2 (Figs 
<figureCitation id="CA22BD4A368191CB5533C5DB976DC207" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Overall dorsal view A Johora intermedia Ng, 1986, holotype male (19.8 x 15.1 mm) (ZRC 1984.6529), Pahang B J. tahanensis (Bott, 1966), male (30.7 x 25.5 mm) (ZRC 1984.6795), Pahang C J. thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, holotype male (25.1 x 21.1 mm) (ZRC 1984.6673), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype male (22.2 x 18.6 mm) (ZRC 2006.0052), Thailand F J. booliati sp. nov., holotype male (34.6 x 30.0 mm) (ZRC 2020.0072), Pahang G J. erici sp. nov., holotype male (38.3 x 32.0 mm) (ZRC 2020.0360), Perak H J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure1" httpUri="https://binary.pensoft.net/fig/476312" pageId="0" pageNumber="1">1C</figureCitation>
, 
<figureCitation id="165026F515BD8CE8BFEEFAF2DCF6A034" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Johora thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu A overall dorsal view B dorsal view of carapace C frontal view of cephalothorax D anterior thoracic sternum and pleon." figureDoi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1">6A</figureCitation>
, 
<figureCitation id="E48B9EA1E47D313E38573F2252537ED5" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Overall dorsal view A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure12" httpUri="https://binary.pensoft.net/fig/476323" pageId="0" pageNumber="1">12C</figureCitation>
); G1 subterminal segment with broad proximal part, tapering suddenly to slender distal part, with distinct shelf-like structure along outer margin; terminal segment straight, long, slender, subequal in length to subterminal segment, surfaces with scattered short setae (Fig. 
<figureCitation id="065D76CB3118CA2BF9CD921038CEE26B" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. A-D Johora thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu E-H J. thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu I-L J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2020.0361), Terengganu A, E, I left G 1 (ventral view) B, F, J distal part of left G 1 (ventral view) C, G, K distal part of left G 1 (dorsal view) D, H, L left G 2 (ventral view) A-D after Ng (1990: fig. 2 A, C-E). Scale bars: 1.0 mm (A, D, E, H, I, L); 0.5 mm (B, C, F, G, J, K)." figureDoi="10.3897/zookeys.994.56810.figure9" httpUri="https://binary.pensoft.net/fig/476320" pageId="0" pageNumber="1">9A-C, E-G</figureCitation>
); G2 shorter than G1, distal segment long, about two-thirds length of basal segment (Fig. 
<figureCitation id="B64F235FCB6F02265CAA431AFFF6A6EF" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. A-D Johora thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu E-H J. thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu I-L J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2020.0361), Terengganu A, E, I left G 1 (ventral view) B, F, J distal part of left G 1 (ventral view) C, G, K distal part of left G 1 (dorsal view) D, H, L left G 2 (ventral view) A-D after Ng (1990: fig. 2 A, C-E). Scale bars: 1.0 mm (A, D, E, H, I, L); 0.5 mm (B, C, F, G, J, K)." figureDoi="10.3897/zookeys.994.56810.figure9" httpUri="https://binary.pensoft.net/fig/476320" pageId="0" pageNumber="1">9D, H</figureCitation>
). Female pleon ovate; somites 3-6 progressively narrower; telson semi-circular (Fig. 
<figureCitation id="F0E8BBFF77B6F9852542FC48458CB133" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Female pleon A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure13" httpUri="https://binary.pensoft.net/fig/476324" pageId="0" pageNumber="1">13C</figureCitation>
). Vulvae large, on anterior half of sternite 6, adjacent to suture with sternite 5, lateral sternal vulvar cover triangular, partially overlapping sternite 5 (Fig. 
<figureCitation id="A76EBD56DC625E7F1D0A3E72756521A6" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Sternopleonal cavity and vulvae A Johora intermedia Ng, 1986, female (17.9 x 14.7 mm) (ZRC 2001.1002), Selangor B J. tahanensis (Bott, 1966), female (24.4 x 21.0 mm) (ZRC 1984.6765), Pahang C J. thoi Ng, 1990, female (31.7 x 25.5 mm) (ZRC 2001.1167), Terengganu D J. hoiseni Ng &amp; Takeda, 1992, paratype female (22.1 x 19.0 mm) (ZRC 1984.6675), Kelantan E J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005, paratype female (21.0 x 17.4 mm) (ZRC 2006.0053), Thailand F J. booliati sp. nov., paratype female (40.4 x 34.4 mm) (ZRC 1995.270), Pahang G J. erici sp. nov., paratype female (32.8 x 26.9 mm) (ZRC 1995.268), Perak H J. michaeli sp. nov., paratype female (29.9 x 25.0 mm) (ZRC 2010.0047), Terengganu." figureDoi="10.3897/zookeys.994.56810.figure14" httpUri="https://binary.pensoft.net/fig/476325" pageId="0" pageNumber="1">14C</figureCitation>
).
</paragraph>
<caption id="274E76D0E0AB654B255A117E6D6FEC94" doi="10.3897/zookeys.994.56810.figure6" httpUri="https://binary.pensoft.net/fig/476317" pageId="0" pageNumber="1" start="Figure 6" startId="F6">
<paragraph id="3D71BFDFC5B1CF4D3C73BA370279DB5F" pageId="0" pageNumber="1">
<emphasis id="4D332E22BE92666F0F2575B0B06927CC" bold="true" pageId="0" pageNumber="1">Figure 6.</emphasis>
<taxonomicName id="452BC560CEFF2058D72B509F1C1BC85C" authorityName="Ng" authorityYear="1990" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora thoi" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="thoi">
<emphasis id="0B4E2AE8491EC0948EE846E332EE0790" italics="true" pageId="0" pageNumber="1">Johora thoi</emphasis>
</taxonomicName>
Ng, 1990, male (21.1 
<normalizedToken id="7F5B2B6C56E4FF0765D7CBC2EA474686" originalValue="×">x</normalizedToken>
17.7 mm) (ZRC 2001.1167), Terengganu 
<emphasis id="EE45805F770709E948CA570F46E6F59A" bold="true" pageId="0" pageNumber="1">A</emphasis>
overall dorsal view 
<emphasis id="D9954CC8CEC47803893C066A3A7147F4" bold="true" pageId="0" pageNumber="1">B</emphasis>
dorsal view of carapace 
<emphasis id="A46B20A05969A128430C1B399A4E4E6F" bold="true" pageId="0" pageNumber="1">C</emphasis>
frontal view of cephalothorax 
<emphasis id="B2FDF5500411C7B780C7B78C751D5387" bold="true" pageId="0" pageNumber="1">D</emphasis>
anterior thoracic sternum and pleon.
</paragraph>
</caption>
<caption id="F39C82ACB61E7B79A755C25DDB12EB93" doi="10.3897/zookeys.994.56810.figure7" httpUri="https://binary.pensoft.net/fig/476318" pageId="0" pageNumber="1" start="Figure 7" startId="F7">
<paragraph id="F5CEDC9BACA3C547E382422AA8CE587F" pageId="0" pageNumber="1">
<emphasis id="B87BD41CBF560962C819659CA1C29C96" bold="true" pageId="0" pageNumber="1">Figure 7.</emphasis>
<taxonomicName id="EABAA9C8A82DC594B7751A9217D9AE93" authorityName="Ng" authorityYear="2020" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora erici" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="erici">
<emphasis id="9CA7A1F4B41E9DECA699764049FDEA85" italics="true" pageId="0" pageNumber="1">Johora erici</emphasis>
</taxonomicName>
sp. nov., male (41.2 
<normalizedToken id="3A430B8ED8ACB91788A0913A82FBF6E5" originalValue="×">x</normalizedToken>
33.8 mm) (ZRC 1995.0269), Kelantan 
<emphasis id="AACF7E4815742424A2D38F2E060AD31C" bold="true" pageId="0" pageNumber="1">A</emphasis>
overall dorsal view 
<emphasis id="9A3003CBAA219E9693324D6104168AA2" bold="true" pageId="0" pageNumber="1">B</emphasis>
dorsal view of carapace 
<emphasis id="B914FB01B66B3BC73B7145FE6F547487" bold="true" pageId="0" pageNumber="1">C</emphasis>
frontal view of cephalothorax 
<emphasis id="EAD78B07AC7E3515E7662E9DF4B412EB" bold="true" pageId="0" pageNumber="1">D</emphasis>
anterior thoracic sternum and pleon.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="A7EF794539EC54A3680CA6BED1E08792" pageId="0" pageNumber="1" type="remarks">
<paragraph id="BBA03FBB8CC7A6B5C70FEF005499AFD7" pageId="0" pageNumber="1">Remarks.</paragraph>
<paragraph id="EAE62E714A643A4EA07B3DD21E237D15" pageId="0" pageNumber="1">
This is one of the largest species of 
<taxonomicName id="0990734F805AE48FC41F173FD3C484DD" authorityName="Bott" authorityYear="1966" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="B8C5F0BCE76A460F2078A373CD27ECA6" italics="true" pageId="0" pageNumber="1">Johora</emphasis>
</taxonomicName>
and is rivalled in size only by 
<taxonomicName id="D30BEF33F6CC1A2F5E8ECFFFC78731F6" lsidName="J. tiomanensis" pageId="0" pageNumber="1" rank="species" species="tiomanensis">
<emphasis id="AA8C3ABF669001030E4E86C3056D955B" italics="true" pageId="0" pageNumber="1">J. tiomanensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="90D0CE9A10B15C351DFF767CA0C0EFF8" lsidName="J. counsilmani" pageId="0" pageNumber="1" rank="species" species="counsilmani">
<emphasis id="77D7BD809497F7765B5F2A1E22453EF4" italics="true" pageId="0" pageNumber="1">J. counsilmani</emphasis>
</taxonomicName>
from Pulau Tioman. The G1 structure of 
<taxonomicName id="17D4A63795B74E865AE3A65586B7229C" lsidName="J. thoi" pageId="0" pageNumber="1" rank="species" species="thoi">
<emphasis id="01D59490CD333AB72FA426C1FA15DE1B" italics="true" pageId="0" pageNumber="1">J. thoi</emphasis>
</taxonomicName>
is distinctive and consistent, the long and straight terminal segment being evident even in small subadult specimens (Fig. 
<figureCitation id="A61425C8EAFAF957FD7341EE300693D1" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. A-D Johora thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu E-H J. thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu I-L J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2020.0361), Terengganu A, E, I left G 1 (ventral view) B, F, J distal part of left G 1 (ventral view) C, G, K distal part of left G 1 (dorsal view) D, H, L left G 2 (ventral view) A-D after Ng (1990: fig. 2 A, C-E). Scale bars: 1.0 mm (A, D, E, H, I, L); 0.5 mm (B, C, F, G, J, K)." figureDoi="10.3897/zookeys.994.56810.figure9" httpUri="https://binary.pensoft.net/fig/476320" pageId="0" pageNumber="1">9E-G</figureCitation>
). The only other species with superficially similar G1s are 
<taxonomicName id="33D49A93C8B64197E790FCD42373BAA9" lsidName="J. singaporensis" pageId="0" pageNumber="1" rank="species" species="singaporensis">
<emphasis id="C8E928655311E5730B2C153330F6F107" italics="true" pageId="0" pageNumber="1">J. singaporensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="0928BB35E5DD11318836652C008E9A81" lsidName="J. michaeli" pageId="0" pageNumber="1" rank="species" species="michaeli">
<emphasis id="716C4F7C91972A7BADC4690F9652C2F5" italics="true" pageId="0" pageNumber="1">J. michaeli</emphasis>
</taxonomicName>
sp. nov. from Singapore and mainland Terengganu, respectively. In these species, however, the G1 terminal segment is prominently shorter, being only two-thirds or less the length of the subterminal segment (Fig. 
<figureCitation id="C1C06BCA71CA688B6EA1F348BF9AAF1C" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. A-D Johora thoi Ng, 1990, holotype male (41.2 x 33.3 mm) (ZRC 1989.2249), Terengganu E-H J. thoi Ng, 1990, male (21.1 x 17.7 mm) (ZRC 2001.1167), Terengganu I-L J. michaeli sp. nov., holotype male (22.7 x 19.2 mm) (ZRC 2020.0361), Terengganu A, E, I left G 1 (ventral view) B, F, J distal part of left G 1 (ventral view) C, G, K distal part of left G 1 (dorsal view) D, H, L left G 2 (ventral view) A-D after Ng (1990: fig. 2 A, C-E). Scale bars: 1.0 mm (A, D, E, H, I, L); 0.5 mm (B, C, F, G, J, K)." figureDoi="10.3897/zookeys.994.56810.figure9" httpUri="https://binary.pensoft.net/fig/476320" pageId="0" pageNumber="1">9I-K</figureCitation>
; 
<bibRefCitation id="C33FD4087CB0BD7700E1B7EA49D50A9B" author="Ng, PKL" journalOrPublisher="Malayan Nature Journal" pageId="0" pageNumber="1" pagination="13 - 44" refId="B11" refString="Ng, PKL, 1987. A revision of the Malayan freshwater crabs of the genus Johora Bott, 1966 stat. nov. (Decapoda: Brachyura: Potamidae). Malayan Nature Journal 41: 13 - 44" title="A revision of the Malayan freshwater crabs of the genus Johora Bott, 1966 stat. nov. (Decapoda: Brachyura: Potamidae)." volume="41" year="1987">Ng 1987</bibRefCitation>
: fig. 8A, B).
</paragraph>
<caption id="8CBF2616BFCDF9FEC8AD9C421244B6A0" doi="10.3897/zookeys.994.56810.figure8" httpUri="https://binary.pensoft.net/fig/476319" pageId="0" pageNumber="1" start="Figure 8" startId="F8">
<paragraph id="96D5F7DE81BCB1D06248756941D2F783" pageId="0" pageNumber="1">
<emphasis id="88CA127F14E3A1405FE45CB2373AC28C" bold="true" pageId="0" pageNumber="1">Figure 8.</emphasis>
<emphasis id="A96EE8DD5EF329C3300DF191D6E79863" bold="true" pageId="0" pageNumber="1">A-D</emphasis>
<taxonomicName id="60033F482B19624D60F1AA9538EC1A9A" authorityName="Bott" authorityYear="1966" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora tahanensis" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="tahanensis">
<emphasis id="D474B8D7C301B17264A0254B2896CC47" italics="true" pageId="0" pageNumber="1">Johora tahanensis</emphasis>
</taxonomicName>
(Bott, 1966), male (30.7 
<normalizedToken id="A0920429D2F2F5527169AEDC970F625E" originalValue="×">x</normalizedToken>
25.5 mm) (ZRC 1984.6795), Pahang 
<emphasis id="8BB4021A691BA95AB731FD940B8A1683" bold="true" pageId="0" pageNumber="1">E-H</emphasis>
<taxonomicName id="16E16D13BD7259168488C51CAC63FE9D" lsidName="J. hoiseni" pageId="0" pageNumber="1" rank="species" species="hoiseni">
<emphasis id="DD699ADFFC1FFFC10D9D043C4751B252" italics="true" pageId="0" pageNumber="1">J. hoiseni</emphasis>
</taxonomicName>
Ng &amp; Takeda, 1992, holotype male (25.1 
<normalizedToken id="435EE3DA6503DED93ADF75F98A7E2F35" originalValue="×">x</normalizedToken>
21.1 mm) (ZRC 1984.6673), Kelantan 
<emphasis id="0123E82F9B38EE260112EE052CD4682D" bold="true" pageId="0" pageNumber="1">A, E</emphasis>
left G1 (ventral view) 
<emphasis id="F17F5851B0408A3982E005B1C9E87199" bold="true" pageId="0" pageNumber="1">B, F</emphasis>
distal part of left G1 (ventral view) 
<emphasis id="F4511E6BF02C162B4B9C6EBFFA73A8F8" bold="true" pageId="0" pageNumber="1">C, G</emphasis>
distal part of left G1 (dorsal view) 
<emphasis id="C9C7D41535ECDF93E90C6E52EC3861BD" bold="true" pageId="0" pageNumber="1">D, H</emphasis>
left G2 (ventral view) 
<emphasis id="FC75706262225AD7D20504DDB87B1645" bold="true" pageId="0" pageNumber="1">E-H</emphasis>
after 
<bibRefCitation id="43E4419D2DB9038A4CC397C4A0151E58" author="Ng, PKL" journalOrPublisher="Bulletin of the National Science Museum, Tokyo, Series A" pageId="0" pageNumber="1" pagination="103 - 116" refId="B19" refString="Ng, PKL, Takeda, M, 1992. On some freshwater crabs (Crustacea, Brachyura, Potamidae, Parathelphusidae and Grapsidae) from Peninsular Malaysia. Bulletin of the National Science Museum, Tokyo, Series A 18 (3): 103 - 116" title="On some freshwater crabs (Crustacea, Brachyura, Potamidae, Parathelphusidae and Grapsidae) from Peninsular Malaysia." volume="18" year="1992">Ng and Takeda (1992</bibRefCitation>
: fig. 3B, D-F). Scale bars: 1.0 mm (
<emphasis id="249350F7EC0E7F1BE65346F59E726B06" bold="true" pageId="0" pageNumber="1">A, D, E, H</emphasis>
); 0.5 mm (
<emphasis id="DB052441220E1E606A6CB3C6034B6A76" bold="true" pageId="0" pageNumber="1">B, C, F, G</emphasis>
).
</paragraph>
</caption>
<caption id="E098D9D1C93831FAAAA4E1824374BB6F" doi="10.3897/zookeys.994.56810.figure9" httpUri="https://binary.pensoft.net/fig/476320" pageId="0" pageNumber="1" start="Figure 9" startId="F9">
<paragraph id="DCEC662F76C928C11C5496A7165D2888" pageId="0" pageNumber="1">
<emphasis id="683E223A265F19C8F90A54D1F07F0DA6" bold="true" pageId="0" pageNumber="1">Figure 9.</emphasis>
<emphasis id="43B05380F1EE3AEE646B5DC5EA9420E2" bold="true" pageId="0" pageNumber="1">A-D</emphasis>
<taxonomicName id="46812558DE0DF5B75ABC6DF9C92C6F1C" authorityName="Ng" authorityYear="1990" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora thoi" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="thoi">
<emphasis id="589CF688D30EBD6696E9CC094A0EF7DA" italics="true" pageId="0" pageNumber="1">Johora thoi</emphasis>
</taxonomicName>
Ng, 1990, holotype male (41.2 
<normalizedToken id="156B79224B60C0FB96BBD0358BCB2DD7" originalValue="×">x</normalizedToken>
33.3 mm) (ZRC 1989.2249), Terengganu 
<emphasis id="3F589A4FF99676E01E8A234126055E39" bold="true" pageId="0" pageNumber="1">E-H</emphasis>
<taxonomicName id="E7FDF49AB1C0CD47565A12F6ED59B698" lsidName="J. thoi" pageId="0" pageNumber="1" rank="species" species="thoi">
<emphasis id="3F569222537F166DDD23302C01AD05A6" italics="true" pageId="0" pageNumber="1">J. thoi</emphasis>
</taxonomicName>
Ng, 1990, male (21.1 
<normalizedToken id="DDDCF345647A62BB9E1787D18A5E8C8D" originalValue="×">x</normalizedToken>
17.7 mm) (ZRC 2001.1167), Terengganu 
<emphasis id="4323EE932DE1008954EAB722D05700BF" bold="true" pageId="0" pageNumber="1">I-L</emphasis>
<taxonomicName id="5BCFD9218BB4A427BA09D8C0B7C40630" lsidName="J. michaeli" pageId="0" pageNumber="1" rank="species" species="michaeli">
<emphasis id="D87727A716E2C17A492D77C2D60C2CE2" italics="true" pageId="0" pageNumber="1">J. michaeli</emphasis>
</taxonomicName>
sp. nov., holotype male (22.7 
<normalizedToken id="9BE64E82E80F120D04270813FA70868A" originalValue="×">x</normalizedToken>
19.2 mm) (ZRC 2020.0361), Terengganu 
<emphasis id="5B9D24B0A02A4E328E2CD4DA6B6D8FF1" bold="true" pageId="0" pageNumber="1">A, E, I</emphasis>
left G1 (ventral view) 
<emphasis id="FC67653547CED1CC94D0E53F250C20CD" bold="true" pageId="0" pageNumber="1">B, F, J</emphasis>
distal part of left G1 (ventral view) 
<emphasis id="6C0CF59B0911E82E8B92D564FB4D92BA" bold="true" pageId="0" pageNumber="1">C, G, K</emphasis>
distal part of left G1 (dorsal view) 
<emphasis id="6B3582F0B50988F8B9562AFE755601C4" bold="true" pageId="0" pageNumber="1">D, H, L</emphasis>
left G2 (ventral view) 
<emphasis id="E4382F25874AABD7CC547FB2D4265E15" bold="true" pageId="0" pageNumber="1">A-D</emphasis>
after 
<bibRefCitation id="789AA945B7B800EBFBDC1C88E7452E72" DOI="https://doi.org/10.1080/00222939000770221" author="Ng, PKL" journalOrPublisher="Journal of Natural History" pageId="0" pageNumber="1" pagination="305 - 310" refId="B13" refString="Ng, PKL, 1990. A new species of Johora Bott, 1966 (Crustacea: Decapoda: Brachyura: Potamidae) from Pulau Redang, Trengganu, Peninsular Malaysia. Journal of Natural History 24: 305 - 310, DOI: https://doi.org/10.1080/00222939000770221" title="A new species of Johora Bott, 1966 (Crustacea: Decapoda: Brachyura: Potamidae) from Pulau Redang, Trengganu, Peninsular Malaysia." url="https://doi.org/10.1080/00222939000770221" volume="24" year="1990">Ng (1990</bibRefCitation>
: fig. 2A, C-E). Scale bars: 1.0 mm (
<emphasis id="B318F1E5B1C867EFF08DD4DC065EA7BE" bold="true" pageId="0" pageNumber="1">A, D, E, H, I, L</emphasis>
); 0.5 mm (
<emphasis id="2C6636724AE8DD3B2AE444B01C1CFE9E" bold="true" pageId="0" pageNumber="1">B, C, F, G, J, K)</emphasis>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="BE351263011EB1CE013A7E6612039AAE" pageId="0" pageNumber="1" type="distribution">
<paragraph id="151BB173A23589EB9782E43478289CDC" pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph id="8F72C257BAEBCC13647161EDF6EEFCD7" pageId="0" pageNumber="1">
<taxonomicName id="3B6148EBD9FFDDFD7F61B00DD8DD8BD0" authorityName="Ng" authorityYear="1990" class="Malacostraca" family="Potamidae" genus="Johora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Johora thoi" order="Decapoda" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="thoi">
<emphasis id="5C5CE5A7D4F8E04AF0D97BE4F4405BA4" italics="true" pageId="0" pageNumber="1">Johora thoi</emphasis>
</taxonomicName>
is endemic to Pulau Redang, the largest island in the Redang Archipelago, a group of nine islands about 25 km off the northeastern coast of Peninsular Malaysia. The species has been recorded from most parts of the island, including its highest point (359 m), occurring in all clean waters there (Fig. 
<figureCitation id="9DD57C6E6F3E48346CAA6BCAD9CCB75E" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Distribution of species of the Johora tahanensis species group 1 J. tahanensis (Bott, 1966) 2 J. thoi Ng, 1990 3 J. hoiseni Ng &amp; Takeda, 1992 4 J. thaiana Leelawathanagoon, Lheknim &amp; Ng, 2005 5 J. booliati sp. nov. 6 J. erici sp. nov. 7 J. michaeli sp. nov. Map data 2020 Google." figureDoi="10.3897/zookeys.994.56810.figure15" httpUri="https://binary.pensoft.net/fig/476326" pageId="0" pageNumber="1">15</figureCitation>
). It lives under rocks and can be found along the stream banks at night.
</paragraph>
</subSubSection>
<subSubSection id="C520A491FF054551219786072280F0E6" pageId="0" pageNumber="1" type="conservation">
<paragraph id="7BEE5B882F6230E54596B43D81755BE6" pageId="0" pageNumber="1">Conservation.</paragraph>
<paragraph id="76AF33D1E3B34F4CA684DE3405682EB9" pageId="0" pageNumber="1">
Although the entire Redang Archipelago is a marine park, the forests are not fully protected, and development of the land for tourism and excessive freshwater use are concerns. As the species is only known from one island only 7 km long and 6 km wide, it is treated as endangered by 
<bibRefCitation id="BFBC85C389FFDC55A2604905AABC8F0A" author="Ng, PKL" editor="Chua, L" journalOrPublisher="28 - 30 June 2005, Forest Research Institute Malaysia, Kepong" pageId="0" pageNumber="1" pagination="95 - 120" refId="B21" refString="Ng, PKL, Yeo, DCJ, 2007. Malaysian freshwater crabs: conservation prospects and challenges. In: Chua, L, Ed., Proceedings of the Seminar on the Status of Biological Diversity in Malaysia &amp; Threat Assessment of Plant Species in Malaysia. 28-30 June 2005, Forest Research Institute Malaysia, Kepong: 95 - 120" title="Malaysian freshwater crabs: conservation prospects and challenges." volumeTitle="Proceedings of the Seminar on the Status of Biological Diversity in Malaysia &amp; Threat Assessment of Plant Species in Malaysia." year="2007">Ng and Yeo (2007)</bibRefCitation>
, but 
<bibRefCitation id="5328F7B07456ED38334DCE55FF9B3252" DOI="https://doi.org/10.1016/j.biocon.2009.02.038" author="Cumberlidge, N" journalOrPublisher="Biological Conservation" pageId="0" pageNumber="1" pagination="1665 - 1673" refId="B3" refString="Cumberlidge, N, Ng, PKL, Yeo, DCJ, Magalhaes, C, Campos, MR, Alvarez, F, Naruse, T, Daniels, SR, Esser, LJ, Attipoe, FYK, Clotilde-Ba, F-L, Darwall, W, Mcivor, A, Collen, B, Ram, M, 2009. Freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges. Biological Conservation 142: 1665 - 1673, DOI: https://doi.org/10.1016/j.biocon.2009.02.038" title="Freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges." url="https://doi.org/10.1016/j.biocon.2009.02.038" volume="142" year="2009">Cumberlidge et al. (2009)</bibRefCitation>
noted it was vulnerable at best as the area is technically protected.
</paragraph>
</subSubSection>
</treatment>
</document>