Pan et al.: Aiyunamon, a new freshwater crab genus
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 70: 320–328

Date of publication: 27 May 2022
DOI: 10.26107/RBZ-2022-0016
http://zoobank.org/urn:lsid:zoobank.org:pub:1E7D8362-E5CF-49F2-B4EA-778E67B5CC94

Aiyunamon, a new genus for southern Chinese potamid species
previously placed in Eosamon Yeo & Ng, 2007 (Decapoda: Brachyura:
Potamidae)
Da Pan1, Peter K.L. Ng2 & Hongying Sun1*
Abstract. Four species of Chinese potamid freshwater crabs currently placed in Eosamon Yeo & Ng, 2007 areassigned to a new genus, Aiyunamon. While the new genus is morphologically close to Eosamon, Indochinamon
Yeo & Ng, 2007, and Doimon Yeo & Ng, 2007, it possesses a distinctive combination of carapace, male pleon,
male first gonopod, and vulva characters that easily distinguishes it. This distinction is supported by genetic data.
The new genus is described and compared with allied genera. Eosamon species are now restricted to Indochina.
Key words. Potamoidea, Potamiscinae, taxonomy, Yunnan, freshwater crab

INTRODUCTION

MATERIAL AND METHODS

The freshwater crab genus Eosamon Yeo & Ng, 2007,
was established for a group of species earlier attributed toPotamon sensu lato (Yeo & Ng, 2007). It currently contains
12 recognised species with a disjunct distribution (Yeo &Ng, 2007; Yeo, 2010; Zhang et al., 2020). Eight species, E.
boonyaratae (Naiyanetr, 1987), E. brousmichei (Rathbun,
1904), E. hafniense (Bott, 1966), E. nominathuis Yeo, 2010,
E. paludosum (Rathbun, 1904), E. phuphanense (Naiyanetr,
1992), E. smithianum (Kemp, 1923) (type species), and E.
yotdomense (Naiyanetr, 1984), are distributed in easternnortheastern Thailand and southern Vietnam, while the otherfour species, E. daiae Zhang & Sun, 2020, E. lushuiense(Dai & Chen, 1985), E. tengchongense (Dai & Chen, 1985),
and E. tumidum (Wood-Mason, 1871), are found primarily
in southwestern China.

The specimens that were examined are deposited in
the Nanjing Normal University, Jiangsu, China (NNU);
National Zoological Museum of China, Institute of Zoology,
Chinese Academy of Sciences, Beijing, China (IZCAS);
and Zoological Reference Collection, Lee Kong Chian
Natural History Museum, National University of Singapore,
Singapore (ZRC). The terminology used follows Ng (1988)
and Davie et al. (2015). The following abbreviations areused: G1, male first gonopod; G2, male second gonopod.
Measurements provided are of the maximum carapace widthand length, respectively.
Genomic DNA of Aiyunamon tumidum was extracted fromgill tissue using the Cell and Tissue DNA Extraction Kit(Generay Biotech). Partial sequence of 16S rRNA gene wasamplified following Ji et al. (2016) and deposited under the
GenBank accession number: OL853518 (voucher number:
NNU 319301), OL853819 (voucher number: NNU 324401),
and OL853520 (voucher number: NNU 314101). After
combining with publicly available sequences downloaded
from GenBank, sequences were aligned using MAFFT 7.310(Katoh & Standley, 2013). Maximum Likelihood (ML)analysis was performed using IQ-TREE 1.6.12 (Nguyen
et al., 2015). Bootstrap values (BS) were obtained through1,000 ultra-fast bootstrap replicates (Minh et al., 2013).
Best substitution model was estimated with the ‘-m TEST’
option (Kalyaanamoorthy et al., 2017). Phylogenetic tree wasvisualised using iTOL 5 (Letunic & Bork, 2021).

Even as they established Eosamon, Yeo & Ng (2007)highlighted that the type specimens of three species from
China were not well studied and their inclusion in the genus
was provisional. Studies by Zhang et al. (2020: fig. 12)
and Pan et al. (2022: fig. 4) have confirmed their concerns,
showing that Eosamon species from China are geneticallydistinct. In this study, the four Chinese Eosamon species areformally transferred to their own genus, Aiyunamon (Fig. 1).

Accepted by: Tohru Naruse
Jiangsu Key Laboratory for Biodiversity and Biotechnology, College of Life
Sciences, Nanjing Normal University, 1 Wenyuan Rd, Nanjing 210023, China; Email:sunhongying@njnu.edu.cn (*corresponding author)
1

Lee Kong Chian Natural History Museum, National University of Singapore, 2Conservatory Drive, Singapore 117377, Republic of Singapore
2© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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Fig. 1. Distribution of Aiyunamon species. Species occurrence records were obtained through long-term field surveys by the Chinese authors.

SYSTEMATICS

Material examined. Aiyunamon daiae (Zhang & Sun,
2020): 1 male (26.6 × 22.2 mm, NNU 190503) (holotype),
Bangyang Village, Longba Town, Longchuan County,
Dehong Prefecture, Yunnan Province, China, coll. X. Hao& Z. Zhang, 5 May 2019; 1 female (20.1 × 16.5 mm, NNU
190505) (paratype), same data as holotype. Aiyunamon
lushuiense (Dai & Chen, 1985): 1 male (26.5 × 22.1 mm,
NNU 148401), Wagu Village, Liuku Town, Lushui City,
Yunnan Province, China, coll. K. Chu, P. Wang & H. Sun,
4 May 2016; 1 female (21.4 × 18.3 mm, NNU 282103),
Shuishan Village, Lushui City, Yunnan Province, China,

Family Potamidae Ortmann, 1896
Subfamily Potamiscinae Bott, 1970
Aiyunamon, new genus
Type species. Eosamon daiae Zhang & Sun, 2020, bypresent designation.

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Pan et al.: Aiyunamon, a new freshwater crab genus

Fig. 2. Overall habitus of species of Aiyunamon, new genus, and Eosamon smithianum (Kemp, 1923). A, A. daiae (Zhang & Sun, 2020),
NNU 190503; B, A. lushuiense (Dai & Chen, 1985), NNU 148401; C, A. tengchongense (Dai & Chen, 1985), NNU 193261; D, A. tumidum(Wood-Mason, 1871), IZCAS CB11382; E, E. smithianum, ZRC 1990.498.

coll. K. Chu, P. Wang & H. Sun, 4 May 2016. Aiyunamon
tengchongense (Dai & Chen, 1985): 1 male (37.9 × 30.1 mm,
NNU 193261), Lianghe County, Yunnan Province, China,coll. X. Hao & Z. Zhang, 9 May 2019; 1 female (25.5 ×
20.5 mm, NNU 282202), Hongdoushu Village, MenglianCounty, Tengchong City, Yunnan Province, China, coll. K.
Chu, P. Wang & H. Sun, 5 May 2016. Aiyunamon tumidum(Wood-Mason, 1871): 1 male (23.2 × 18.7 mm, IZCASCB11382), Sipai Mountain, Yunnan Province, China, coll.
1964; 1 female (23.6 × 18.4 mm, IZCAS CB11383), Sipai
Mountain, Yunnan Province, China, coll. 1964. Eosamon
smithianum (Kemp, 1923): 1 male (49.6 × 39.1 mm, ZRC
1990.498–514), 1 female (39.5 × 33.4 mm, ZRC 1990.498–514), Krating Waterfall area, near Kitchagood National Park,

eastern Thailand, fast flowing water with smooth rocks, sandsubstrate, coll. P. K. L. Ng, December 1988.
Diagnosis. Carapace slightly broader than long, dorsal surface
slightly convex, regions distinctly defined; epigastric cristae
distinct, rugose, separated from postorbital cristae by shallow
groove; postorbital cristae distinct, rugose; H-shaped groove
distinct (Figs. 2A–D, 3A–D). Frontal margin slightly sinuous,
gently concave medially, placed anterior to level of external
orbital angles; external orbital angle low, separated fromrest of margin by small or shallow cleft; epibranchial tooth
low, not prominent; anterolateral margins slightly convex,
cristate, lined with distinct granules; posterolateral margins
gently converging posteriorly (Fig. 2A–D). Third maxilliped
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Fig. 3. Frontal view of carapace of species of Aiyunamon, new genus, and Eosamon smithianum (Kemp, 1923). A, A. daiae (Zhang &Sun, 2020), NNU 190503; B, A. lushuiense (Dai & Chen, 1985), NNU 148401; C, A. tengchongense (Dai & Chen, 1985), NNU 193261;
D, A. tumidum (Wood-Mason, 1871), IZCAS CB11382; E, E. smithianum, ZRC 1990.498.

exopod with long flagellum. Male pleon triangular, lateral
margins almost straight (Fig. 4A–D). G1 curved outwards;
distal part of outer margin of subterminal segment at junction
with terminal segment deeply concave; terminal segment
conical, straight to slightly curved, without dorsal flap (Fig.
5A–D, F–I); G2 slender. Female vulvae with anterior margin
appressed against suture of thoracic sternites 5/6, ovate,
without operculum (Fig. 6A–D).

fig. 5); epibranchial tooth indistinct (Fig. 2A–D) (versus
epibranchial tooth distinct, sharp in Eosamon; Fig. 2E; cf.
Yeo & Ng, 2007: fig. 5); male pleon lateral margins straight(Fig. 4A–D) (versus male pleon lateral margins slightly
concave in Eosamon; Fig. 4E; cf. Yeo & Ng, 2007: fig. 5);G1 subterminal segment with distal part of outer margin,
at junction with terminal segment, deeply concave (Fig.
5A–D, F–I) (versus G1 outer margin of subterminal segment
gradually tapering to junction with terminal segment, marginappearing gently concave in Eosamon; Fig. 5E, J; cf. Yeo& Ng, 2007: fig. 5); G1 terminal segment without any
trace of dorsal flap (Fig. 5A–D, F–I) (versus G1 terminal
segment with low, broad dorsal flap in Eosamon; Fig. 5E, J;
cf. Yeo & Ng, 2007: fig. 5); and anterior margin of female
vulvae appressed on suture of thoracic sternites 5/6, without
operculum (Fig. 6A–D) (versus only anterior edge of female
vulvae touching suture of thoracic sternites 5/6, with distinct
lateral operculum present in Eosamon; Fig. 6E).

Etymology. The genus is named after the late Prof. Aiyun
Dai, in honour of her immense contributions to Chinese
freshwater crab taxonomy. Gender of genus neuter.
Remarks. This new genus currently contains four species,
Aiyunamon daiae (Zhang & Sun, 2020) (type species), A.
lushuiense (Dai & Chen, 1985), A. tengchongense (Dai &Chen, 1985), and A. tumidum (Wood-Mason, 1871).
Morphologically, Aiyunamon is most similar to Eosamon
but can be separated by the following characters: postorbitalcristae rugose (Figs. 2A–D, 3A–D) (versus postorbital cristae
sharp in Eosamon; Figs. 2E, 3E; cf. Yeo & Ng, 2007: fig.
5); external orbital angle low, separated from the lateral
margin by shallow cleft (Figs. 2A–D, 3A–D) (versus external
orbital angle prominent, separated from lateral margin bydeep cleft in Eosamon; Figs. 2E, 3E; cf. Yeo & Ng, 2007:

Aiyunamon is also superficially similar to Indochinamon Yeo& Ng, 2007 which is also present in southwestern China.
Aiyunamon can easily be distinguished by its proportionately
less transverse carapace (Fig. 2A–D) (versus more transverse
carapace in Indochinamon; cf. Yeo & Ng, 1998: fig. 7D);
the epigastric cristae being separated from postorbitalcristae by shallow groove (Fig. 2A–D) (versus epigastric323

Pan et al.: Aiyunamon, a new freshwater crab genus

Fig. 4. Male pleon of species of Aiyunamon, new genus, and Eosamon smithianum (Kemp, 1923). A, A. daiae (Zhang & Sun, 2020), NNU
190503; B, A. lushuiense (Dai & Chen, 1985), NNU 148401; C, A. tengchongense (Dai & Chen, 1985), NNU 193261; D, A. tumidum(Wood-Mason, 1871), IZCAS CB11382; E, E. smithianum, ZRC 1990.498.

suite of features as in Indochinamon (cf. Naiyanetr & Ng,
1990: fig. 4); and the G1 terminal segment is gently curved tostraight and without any dorsal flap (Fig. 5A–D, F–I) (versus
more curved terminal segment with a clear dorsal flap in
Doimon; cf. Naiyanetr & Ng, 1990: fig. 5A−C). Aiyunamon
differs from Iomon Yeo & Ng, 2007 in having the external
orbital angle low and separated from the rest of the margin
only by a shallow cleft (Fig. 2A–D) (versus external orbital
angle relatively large, broadly triangular in Iomon; cf. Ng &Naiyanetr, 1993: fig. 13), and the G1 slightly bent outwards(Fig. 5A–D, F–I) (versus G1 distinctly bent outwards in
Iomon; cf. Ng & Naiyanetr, 1993: fig. 48).

cristae being separated from postorbital cristae by distinct
groove in Indochinamon; cf. Yeo & Ng, 1998: fig. 7D; theexternal orbital angle is low and separated from the rest of
the margin only by a shallow cleft (Figs. 2A–D, 3A–D)(versus well-developed external orbital angle separated
from rest of margin by a deep cleft in Indochinamon; cf.
Yeo & Ng, 1998: fig. 7D); the low epibranchial teeth (Fig.
2A–D) (versus more distinct, sharp epibranchial teeth inIndochinamon; cf. Yeo & Ng, 1998: fig. 7D) and relatively
more slender G1 terminal segment (Fig. 5A–D, F–I) (versus
relatively stouter terminal segment in Indochinamon; cf. Yeo& Ng, 1998: fig. 5I, J). The carapace of Aiyunamon can be
distinguished from Doimon Yeo & Ng, 2007 by the same
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Fig. 5. Male first gonopod (G1) of species of Aiyunamon, new genus, and Eosamon smithianum (Kemp, 1923). A, F, A. daiae (Zhang &Sun, 2020), NNU 190503; B, G, A. lushuiense (Dai & Chen, 1985), NNU 148401; C, H, A. tengchongense (Dai & Chen, 1985), NNU
193261; D, I, A. tumidum (Wood-Mason, 1871), IZCAS CB11382; E, J, E. smithianum, ZRC 1990.498. A–D, right G1, dorsal view; F–I,
right G1, ventral view; E, J, left G1, dorsal and ventral views, respectively (laterally transposed for comparisons).

In its external features, Aiyunamon is also superficiallysimilar to Potamiscus montosus Dai, Song, He, Cao, Xu& Zhong, 1975, and P. yiwuensis Dai & Cai, 1998. The
new genus, however, is easily separated by possessing a
distinct flagellum on the third maxilliped exopod (cf. Dai,
1999: figs. 91, 92, 93; Zhang et al., 2020: fig. 4) (versus
absent or vestigial flagellum in Potamiscus; cf. Dai & Cai,
1998: fig. 2; Dai, 1999: fig. 99) and the relatively slenderG1 terminal segment (Fig. 5A–D, F–I) (versus relatively
stouter G1 terminal segment in Potamiscus; cf. Dai & Cai,
1998: figs. 5, 6; Dai, 1999: fig. 99).

hilly streams, and they have also been found in holes on
the ridges of rice fields. The burrows in the rice fields areabout 30–50 cm deep.
The dispersal abilities and fecundity of freshwater crabs are
relatively low. As such, genera usually do not have disjunctdistributions, especially for highland taxa. Therefore, as
exemplified by this study, genera with disjunct distributions
should be restudied to see if the taxa are monophyletic.
Another case is Tenuipotamon Dai, 1990, which also has a
disjunct distribution, but studies have since shown that thegenus is not monophyletic (cf. Pan et al., 2022: figs. 2, 3).

The confirmed distribution of Aiyunamon is southwestern
Yunnan (Fig. 1), about 1,300–1,700 m above sea level.
Aiyunamon tumidum was described from Yunnan and upper
Myanmar (Wood-Mason, 1871: 454) but we are not certain
if all his specimens belong to the same species; they willneed to be re-examined. Yeo & Ng (2007: 304) recorded
one syntype male and seven non-type specimens of A.
tumidum in their revision of the Potamidae, but all werefrom Yunnan. Aiyunamon species are usually found hiding
under stones and/or burrowing on the banks of shallow

MOLECULAR RESULTS & DISCUSSION
All four Aiyunamon species as well as three Eosamon species,
including the type species of the genus, E. smithianum(Kemp, 1923), were included in our phylogenetic analysis.
While all Aiyunamon species were clustered together withmoderate support value (bootstrap value = 70), Aiyunamon
and Eosamon species segregated to different clades with the325Pan et al.: Aiyunamon, a new freshwater crab genus

Fig. 6. Vulvae of species of Aiyunamon, new genus, and Eosamon smithianum (Kemp, 1923). A, A. daiae (Zhang & Sun, 2020), NNU
190505; B, A. lushuiense (Dai & Chen, 1985), NNU 282103; C, A. tengchongense (Dai & Chen, 1985), NNU 282202; D, A. tumidum(Wood-Mason, 1871), IZCAS CB11383; E, E. smithianum, ZRC 1990.498.

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Tree scale: 0.1

Paratelphusula gibbosa AB428512
Socotrapotamon nojidense AB428493
Eosamon yotdomense AB428485
Kukrimon cucphuongensis AB428483
99
Eosamon boonyaratae AB428487
98
100
Eosamon smithianum AB428486
Parapotamon spinescens AB428467
Yuebeipotamon calciatile LC176064
Megacephalomon kittikooni AB428462
Minpotamon nasicum AB428450
71
57
99
Nanhaipotamon nanriense AB212868
54
Apotamonautes hainanensis AB428459
69
Candidiopotamon rathbunae AB208598
Demanietta
renongensis AB428475
100
Tomaculamon pygmaeus AB428473
100
Ovitamon artifrons AB428466
Arquatopotamon jizushanense KY963596
75
87
Tenuipotamon huaningense AB428491
100
Pararanguna semilunata AB428490
75
82
Potamiscus loshingense AB428488
Shanphusa curtobates AB428478
Indochinamon malipoense MT887278
Indochinamon ahkense MZ350923
99
85
Iomon luangprabangense MZ350929
65
Beccumon jarujini AB428479
58
Pupamon nayung AB428477
94
100
Potamiscus montosus MN737133
Aiyunamon tumidum OL853519
70
Aiyunamon tumidum OL853520
100
99 Aiyunamon tumidum OL853518
Aiyunamon tengchongense MT887285
70
95
Aiyunamon lushuiense MT887284
Aiyunamon daiae MT887281
88
100
Aiyunamon daiae MT887280
100
Aiyunamon daiae MT887283
97
Aiyunamon daiae MT887282
Fig. 7. Maximum Likelihood phylogenetic tree based on 16S rRNA gene sequences of potamid crabs including Aiyunamon, new genus, andEosamon Yeo & Ng, 2007. Bootstrap values over 50 are shown on branches. The accession numbers are provided along with species names.
86

divergence being quite major (Fig. 7). The result confirmed
the genetic distinctiveness between Aiyunamon and Eosamon(cf. Pan et al., 2022: fig. 4). Our phylogenetic analysis, basedon 16S rRNA gene sequences, supports the establishmentof the new genus, Aiyunamon.

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ACKNOWLEDGEMENTS
Thanks are due to Jun Chen and Kaibayier Meng (National
Zoological Museum of China, Institute of Zoology, Chinese
Academy of Sciences, Beijing, China) for permission to
examine the specimens deposited in the IZCAS. We would
also like to thank Tohru Naruse for his helpful comments
to the manuscript. This study was funded by the National
Natural Science Foundation of China (No. 32170454) to SHY.
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