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Uca crassipes (White, 1847). Live coloration of adults (A–D, F) and juvenile (E). Photos taken from Dongsha Island, Taiwan (A–E) and Cocos-Keeling, Australia (F).
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The fiddler crab, Uca splendida (Stimpson, 1858) has been synonymized under Uca crassipes (White, 1847) since Crane (1975). Studies of specimens from the Hong Kong type locality and adjacent areas of China, Taiwan and Vietnam show that U. splendida is a valid species, with a characteristic suite of carapace and gonopod features as well as a distinc...
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... (Fig. 4H), although their chelae are identical (Figs. 3B, 4B, 5B). The carapace of most adult U. splendida has similar transverse black bands on the bluish carapace and in some females, the anterior part of the carapace is orange-red ( Fig. 4A-E). The carapace coloration of U. crassipes varies in color, including being entirely scarlet red (Fig. 5A); with different degrees of red on a black, blue or green background; blue or green bands on a dark background (Fig. 5B-E); and mottled dark spots on a white background (Fig. 5F). Whereas the eyestalks of U. splendida are invariably red (Fig. 4A-F, H), U. crassipes tends to have green or white eyestalks (Figs. 4H, 5A-F), although a few ...
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... transverse black bands on the bluish carapace and in some females, the anterior part of the carapace is orange-red ( Fig. 4A-E). The carapace coloration of U. crassipes varies in color, including being entirely scarlet red (Fig. 5A); with different degrees of red on a black, blue or green background; blue or green bands on a dark background (Fig. 5B-E); and mottled dark spots on a white background (Fig. 5F). Whereas the eyestalks of U. splendida are invariably red (Fig. 4A-F, H), U. crassipes tends to have green or white eyestalks (Figs. 4H, 5A-F), although a few specimens sometimes possess red eyestalks. Some specimens of U. crassipes may have a carapace that has a mixture of blue ...
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... some females, the anterior part of the carapace is orange-red ( Fig. 4A-E). The carapace coloration of U. crassipes varies in color, including being entirely scarlet red (Fig. 5A); with different degrees of red on a black, blue or green background; blue or green bands on a dark background (Fig. 5B-E); and mottled dark spots on a white background (Fig. 5F). Whereas the eyestalks of U. splendida are invariably red (Fig. 4A-F, H), U. crassipes tends to have green or white eyestalks (Figs. 4H, 5A-F), although a few specimens sometimes possess red eyestalks. Some specimens of U. crassipes may have a carapace that has a mixture of blue and black patterns (Fig. 5C, E), similar to that of U. ...
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... dark spots on a white background (Fig. 5F). Whereas the eyestalks of U. splendida are invariably red (Fig. 4A-F, H), U. crassipes tends to have green or white eyestalks (Figs. 4H, 5A-F), although a few specimens sometimes possess red eyestalks. Some specimens of U. crassipes may have a carapace that has a mixture of blue and black patterns (Fig. 5C, E), similar to that of U. splendida, but the dark part is invariably more irregular and the transverse bands not as prominent as the latter. In addition, the blue coloration is always relatively darker in U. crassipes (Fig. ...
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... possess red eyestalks. Some specimens of U. crassipes may have a carapace that has a mixture of blue and black patterns (Fig. 5C, E), similar to that of U. splendida, but the dark part is invariably more irregular and the transverse bands not as prominent as the latter. In addition, the blue coloration is always relatively darker in U. crassipes (Fig. ...
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Citations
... These values are comparable with threshold values previously reported for thoracotreme crabs (1.49%-6.25%), from studies of groups in which taxonomy is relatively well resolved, and the examined taxa show discrete morphologies (see Davie et al., 2010;Shih et al., 2012Shih et al., , 2019KJH Wong et al., 2012). However, cryptochirid taxonomy is nowhere well resolved in two sense: numerous previously unrecognized species-level taxa (Wei et al., 2016;Bähr et al., 2021;Xu et al., 2021; see below) and previous morphology-based generic placement insufficient in encapsulate true species richness, thus rendered composite (van der Meij and Nieman, 2016; see Remarks under L. doughnut above). ...
Highly specialized cryptochirid crabs are obligate symbionts of scleractinian corals in tropical and subtropical seas. General morphologies of cryptochirid crabs remain poorly described due to their small size and difficulties in collection; thus, the current inventory is probably an underestimation. In the present study, we sampled cryptochirid crabs from coral communities in Hong Kong. In the literature, only Cryptochirus hongkongensis (now Neotroglocarcinus hongkongensis) with unknown hosts had been recorded in Hong Kong since 1936. In addition to morphological examination, identification in the present study is further supported by sequence divergence of mitochondrial cytochrome c oxidase I (COI) and 16S ribosomal DNA markers. Six operative taxonomic units (OTUs), representing four species and one species complex with two species, were revealed among our material: Cryptochirus coralliodytes, Lithoscaptus paradoxus, Lithoscaptus doughnut sp. nov., Lithoscaptus scottae sp. nov., and Xynomaia sheni species complex. Morphological description of these species is provided, including description of the two new pseudocryptic species. The hosts of the genus Lithoscaptus belong largely to the Merulinidae, while L. doughnut sp. nov. inhabits the Plesiastreidae.
... Therefore, they may provide us with a way to test whether the predation risk of crabs is associated with body colouration. Here, we conducted a laboratory study to compare the predation risk from a fish predator in two sympatric fiddler crab species with different body colour patterns: Austruca perplexa with brownish-white colourations and Paraleptuca crassipes with red-and-black colour patterns (Shih et al., 2012(Shih et al., , 2016Shih and Poupin, 2020). ...
... A. perplexa and P. crassipes are distributed sympatrically in the Indo-West Pacific (Crane, 1975;Shih et al., 2012Shih et al., , 2016Shih and Poupin, 2020). In the Ryukyu Archipelago in subtropical Japan, they are found in mangrove forests with muddy sediment, and A. perplexa also occurs on bare muddy flats in mangrove estuaries (Kawaida et al., 2017;K. ...
Many animals show conspicuous colour patterns that inform potential predators about their unprofitability due to their toxicity or unpalatability (aposematism). However, no systematic work has evaluated the colouration of crabs in terms of aposematism, even though many species exhibit conspicuous colouration. We conducted a laboratory study to compare predation risk from a fish predator, the barred mudskipper (Periophthalmus argentilineatus), in two sympatric fiddler crab species with different body colour patterns: Austruca perplexa with brownish-white colourations and Paraleptuca crassipes with red-and-black colour patterns. Fiddler crabs and mudskippers are active on tidal flats during low tides. A predation trial by stocking one crab and one mudskipper in a test arena was repeated five times for each species–sex combination. The animal behaviour was video recorded for 24 h, and crabs were confirmed to be alive or dead after 72 h. A. perplexa females were most vulnerable to predation from mudskippers. The attack intensity from mudskippers was stronger during the nighttime than daytime against P. crassipes females and males and A. perplexa males, suggesting that mudskippers may avoid them by visually detecting the body colouration in P. crassipes and the body structure in males of both species (i.e. a large weaponry claw) during the daytime. Mudskippers eventually preyed on and consumed some individuals in both species–sexes. Mudskippers are known to shift their main food components from small crustaceans through polychaete worms to crabs, such as fiddler crabs, and unpalatable polychaete worms tend to be brightly coloured. We suggest that the red colouration of P. crassipes may be incidental mimicry of the unpalatable red/orange-coloured polychaete worms that mudskippers might have learned to avoid.
... However, this species complex has one terminal seta on the antennal exopod, which is different from G. tetragonon which has two terminal setae. No distinct character could be found that distinguished Paraleptuca crassipes from Pa. splendida, reflecting their close phylogeny (Shih et al. 2012(Shih et al. 2016b. Undistinguished morphology of zoea I among closely related species are also reported in other taxa, such as Ocypode ceratophthalmus (Pallas, 1772) and O. stimpsoni Ortmann, 1897 (Jiang et al. 2014). ...
Fiddler crabs (Brachyura: Ocypodidae: Gelasiminae) in Taiwan consist of 5 genera and 15 species, but knowledge of their larval development is limited to just 3 species, namely Austruca lactea, Tubuca arcuata, and Xeruca formosensis. In our study, the morphology of the first zoeal stage (zoea I) of the 15 species is described and compared to previous studies. The results show that the characters of zoea I can be used to distinguish the five studied genera and most species (except three groups, "Gelasimus borealis, G. jocelynae and G. vocans", "Paraleptuca crassipes and P. splendida" and "Tubuca arcuata, T. coarctata and T. paradussumieri "). The lateral spines on the carapace in zoea I are suggested to be a unique character in the Ocypodinae because they are absent in the Gelasiminae and Ucidinae, which supports the current systematics of the Ocypodidae.
... Milne Edwards, 1852) (≥ 1.29%, Shih and Poupin, 2020); Gelasimus excisus (Nobili, 1906) (= G. neocultrimanus (Bott, 1973) and G. jocelynae Shih, Naruse & Ng, 2010(≥ 4.77%, recalculated from Shih et al. 2010; Minuca rapax (Smith, 1870) and M. virens (Salmon & Atsaides, 1968) (≥ 3.29%, Thurman et al. 2018); the Minuca burgersi complex (3.77%-4.78% (mean p-distance), Thurman et al. 2021); Paraleptuca splendida (Stimpson, 1858) and P. crassipes (White, 1847) (≥ 2.49%, Shih et al. 2012); Paraleptuca boninensis (Shih, Komai & Liu, 2013) and P. splendida (≥ 2.33%, Shih et al. 2013); and Tubuca alcocki T. urvillei (H. Milne Edwards, 1852) (3.78%, Shih et al. 2018), although the distance between Austruca bengali (Crane, 1975) and its sister species, A. variegata (Heller, 1862) (Shih et al. 2019), is larger (≥ 13.7%). ...
Nine species of fiddler crabs (Crustacea: Ocypodidae: Gelasiminae) are known from the Arabian Sea and adjacent waters (Red Sea, Gulf of Aden, Gulf of Oman and Arabian/Persian Gulf): five species of Austruca, one Cranuca, two Gelasimus and one Tubuca. COI sequence data match morphological species boundaries and shows high connectivity within each. The fauna is highly endemic, with three species of Austruca (A. albimana, A. iranica and A. sindensis) confined to this region, and four others restricted to the Indian Ocean. restricted to the Indian Ocean. Austruca albimana and A. iranica speciated locally and now have narrowly overlapping ranges in Oman. Our results confirm the westernmost distributions of Austruca annulipes and Tubuca alcocki are Pakistan and the Red Sea, respectively. A key for the nine species is also provided to help identification.
... Milne-Edwards 1873) and A. perplexa (H. Milne Edwards 1852) (Shih et al. 2019;Shih and Poupin 2020), Paraleptuca crassipes (White 1847) (Shih et al. 2012(Shih et al. 2013, Leptuca uruguayensis (Nobili 1901) (Laurenzano et al. 2012) and Uca maracoani (Latreille, 1803) (Wieman et al. 2014). Several species exhibit low genetic diversity across their ranges implying extensive connectivity via larval transport. ...
For this study, in addition to museum vouchers, 1437 specimens of Minuca burgersi (Holthuis, 1967) were collected from crab colonies at 105 locations in the western Atlantic Ocean to examine diversity in a species with a large geographic range. Both allometric and geometric morphometry were coupled with the molecular analysis of DNA to give a broader perspective of intraspecific variability in this species. A total of 1153 specimens from the Caribbean Sea and the Atlantic coast of South America demonstrated that M. burgersi from both regions are very similar in their pattern of growth. The average carapace width (CW) for Caribbean is larger than the average for South American males and females. However, size distribution based on CW is unimodal in Caribbean and bimodal in South American populations. The carapace length-width ratio is about 0.68 in females and 0.66 in males. South American males express asymmetric elongation of the cheliped in smaller CW intervals than Caribbean males. In a sample of 259 females, carapace shape is distinct between South American and Caribbean populations. Caribbean populations have less swelling in the branchial regions than South American populations. The swelling correlates primarily with geographic region and to a lesser degree with substrate and salinity. Molecular data from the 16S rDNA and cytochrome c oxidase subunit I (COI) reveal three clades within Minuca burgersi. Two clades are distributed in the Caribbean and the third in eastern South America. The timing of divergence between Caribbean and South American clades is coincident with an increased rate of water and sediment outflow from the Amazon as inferred from the geologic record. Current patterns and associated gene flow within the Caribbean were subsequently influenced by the closing of the Isthmus of Panama. We speculate that various populations may employ different larval dispersion mechanisms resulting in genetic heterogeneity. Consequently, there is considerable biological divergence among populations of M. burgersi in the Caribbean and South America.
... Although fiddler crabs have been studied in many of their natural distribution areas, the study of fiddler crabs in mainland China is almost nonexistent. Most of the fiddler crab studies have been conducted in Hong Kong and Taiwan and were related to crab distribution, diversity, and burrow morphology (Li et al., 2018;Shih et al., 2010;Shih et al., 2012). The study of fiddler crabs in mainland China may help advance the protection and restoration of mangroves in China. ...
... As with many other groups of organisms, the taxonomy of fiddler crabs has gradually shifted over time as new studies, technologies, and scientific attitudes have led to new and different insights into the evolutionary history of these species. The last truly comprehensive revision and study of the taxonomy of fiddler crabs (Crane, 1975) has served as a benchmark for all studies since and has allowed subsequent researchers to use a more piecemeal approach to updating the taxonomy and systematics as necessary (Thurman, 1981(Thurman, , 1982George & Jones, 1982;Barnwell & Thurman, 1984;von Prahl & Toro, 1985;von Hagen, 1987;von Hagen & Jones, 1989;Levinton et al., 1996;Sturmbauer et al., 1996;Rosenberg, 2001Rosenberg, , 2013Beinlich & von Hagen, 2006;Shih et al., 2009Shih et al., , 2010Shih et al., , 2012Shih et al., , 2013aShih et al., , 2015Shih et al., , 2016bShih et al., , 2018Shih et al., , 2019Spivak & Cuesta, 2009;Landstorfer & Schubart, 2010;Naderloo et al., 2010Naderloo et al., , 2016Thurman et al., 2018). Today we recognize 105 extant species (Rosenberg, 2014), with five additional named fossil taxa, a major shift from the 62 extant species (92 taxa with subspecies included) and two fossils recognized by Crane (1975). ...
... As with many other groups, named species of fiddler crab have gone through waves of consolidation and expansion. Over the last few decades, the relationships among most of the historical names have largely stabilized with taxonomic advances mostly revolving around the recognition/discovery of cryptic species within formerly recognized single species (Novak & Salmon, 1974;Thurman, 1981;Naderloo et al., 2010Naderloo et al., , 2016Shih et al., 2009Shih et al., , 2010Shih et al., , 2012Shih et al., , 2013aShih et al., , 2018Shih et al., , 2019Thurman et al., 2018), although a few purely novel species have been described as well (George & Jones, 1982;von Prahl & Toro, 1985;von Hagen, 1987;Landstorfer & Schubart, 2010). ...
... 3) Where will the next cryptic species be found? As molecular systematics has been more broadly applied to fiddler crabs, a number of geographically widespread species have recently been split into sets of more regional similar/cryptic species (Shih et al., 2009(Shih et al., , 2010(Shih et al., , 2012(Shih et al., , 2019Naderloo et al., 2010Naderloo et al., , 2016Thurman et al., 2018), and it is likely more are waiting to be found. Species such as Uca (Uca) princeps (Crane, 1975;MSR, unpublished data), Tubuca (Tubuca) forcipata (MSR, unpublished data; H.T. Shih, personal communication), and Minuca ecuadoriensis (Barnwell, 1988) have all been observed to encompass enough variation to raise questions as to whether they represent multi-species complexes currently hidden under a single name. ...
Fiddler crabs (Ocypodidae) have gone through a gradual series of taxonomic revisions and refinements over the last 40 years, culminating most recently with an expansion from a single genus into eleven different genera. I examine the opportunities presented by these revisions with respect to establishing formal names for previously established clades at a variety of taxonomic levels that were otherwise previously impossible to name due to historical compression of these crabs into a single genus, including the establishment or reestablishment of three tribes (Ucini, Gelasimini, and Minucini) and ten subgenera: Uca (Uca), Uca (Acanthoplax), Gelasimus (Gelasimus), Gelasimus (Mesuca), Austruca (Austruca), Austruca (Cuneatuca), Austruca (Sinduca), Tubuca (Tubuca), Tubuca (Australuca), and Tubuca (Angustuca). A previously overlooked synonymy between Gelasimus excisa (Nobili, 1906) and G. neocultrimana (Bott, 1973) is discussed, and the former name is adopted as valid.
... The value is higher than some species within the Ocypodoidea, e.g., the minimum genetic distance of K2P between two species is 2.79 % between Paraleptuca crassipes (White, 1847) and P. splendida (Stimpson, 1858); 3.62 % between Gelasimus hesperiae (Crane, 1975) and "Clade U"; and 3.62 % between Mictyris brevidactylus Stimpson, 1858 and M. guinotae Davie, Shih & Chan, 2010, but still smaller than 6.25 % between Ocypode stimpsoni Ortmann, 1897 andO. mortoni George, 1982;and 4.43 % between Scopimera globosa (De Haan, 1835) and S. ryukyuensis Wong, Chan & Shih, 2010(see Davie et al. 2010Shih et al. 2010Shih et al. , 2012Wong et al. 2010Wong et al. , 2012Chu et al. 2015). ...
A new pseudocryptic species of fiddler crab, Tubuca alcocki sp. n., is described from the northern Indian Ocean. The new species was previously identified with T. urvillei (H. Milne Edwards, 1852), but can be distinguished by the structures of the anterolateral angle of the carapace and male first gonopod. The molecular data of the mitochondrial cytochrome oxidase subunit I gene shows that both are sister taxa and the divergence time is estimated at 2.2 million years ago, around the beginning of the Pleistocene. While the new species is widely distributed in the northern part of Indian Ocean, occurring from the Red Sea to India and the Anda-man Sea; T. urvillei sensu stricto has a more restricted range, and is known only from southeastern Africa.
... For example, ≤ 0.61% for Mictyris thailandensis Davie, Wisespongpand & Shih, 2013 (Mictyridae, Davie et al., 2013), ≤ 0.61% for Scopimera spp. (Dotillidae, Wong et al., 2011), ≤ 0.61% for Helice latimera complex (Varunidae, Shih & Suzuki, 2008) and ≤ 1.86% for Paraleptuca splendida (Stimpson, 1858) (Ocypodidae,Shih et al., 2012Shih et al., , 2016. It shows that all Guam haplotypes (including holotype of E. villosus) as well as other E. notatus/E. ...
... villosus should be regarded as the same species.In summary, considering the entire suite of morphological and molecular evidence, E. villosus Ng, 2003, is here treated as a junior subjective synonym of E. notatus (Heller, 1865).Adult Epigrapsus notatus are known to inhabit the supralittoral zone, from vegetated sandy coasts to coastal forests, sometimes foraging during the day. They are usually not encountered in open habitats and are often observed under large objects (e.g., rocks, wood, etc.) on soft substrates and/ or in and around their burrows under forest cover(Ng et al., 1998; Liu & Jeng, 2005; Naruse, 2005;Shih, 2012 Shih, , 2013 Fujita, 2017a).Liu & Jeng (2005) noted that E. notatus is rarely observed outside the forest even on rainy nights.Small individuals of E. notatus had been discovered in rather unusual environments.Ng (2003) noted that the small male holotype of E. villosus(ZRC 2002.0011, 8.9 × 9.4 mm) was collected from about 50 metres from the entrance of a cave, which itself is situated about 100-200 metres from the shoreline in Guam. ...
... 50 m from the coast (JCEM & TN, pers. obs.).Shih (2012) noted that young individuals could be found under driftwood in the supralittoral zone of Dongsha Island, northern South China Sea, while Shih (2013) mentioned that adults were collected under rocks in the tropical coastal forest of southern Taiwan. Fujita (2017b), however, collected young and adult forms of E. notatus from the lower supralittoral zone, under boulder/rubble substratum in the southern Ryukyus, which was occasionally splashed with seawater by wave action. ...
The identities of what have been referred to as Epigrapsus notatus (Heller, 1865) and E. villosus Ng, 2003, are evaluated using morphological and molecular methods. Morphological comparisons showed that the main characters used to diagnose E. villosus (viz. the setation of the carapace, the condition of the epigastric cristae, the granulation on the branchial region, and proportions of the ambulatory legs) can be attributed to the small size of the type specimen; and the extreme condition of these structures can be connected to adult forms of E. notatus through transitional phases. One morphological feature (the laterally expanded rst epibranchial tooth) is seen only in the holotype of E. villosus, but it can also be regarded as a character state speci c to small individuals of E. notatus, and signifying the extreme end of possible variation in that species. The analysis of mitochondrial cytochrome c oxidase subunit I (COI) reveals three haplotypes in the E. notatus/E. villosus complex, but with only as much as 0.61% of nucleotide divergence between them. In conclusion, it is clear that E. notatus undergoes dramatic morphological change during its growth, and that E. villosus should be considered a junior synonym of E. notatus.
... To establish a criterion, Lefébure et al. (2006) and Costa et al. (2007) proposed that an interspecific-intrageneric sequence threshold could be set at 16 to 17.2% divergence of K2P to separate cryptic crustacean taxa. For fiddler crabs from the Indo-West Pacific, Shih et al. (2009Shih et al. ( , 2010Shih et al. ( , 2012Shih et al. ( , 2018 reported interspecific COI divergences of 2.79 to 11.9% among morphological species. Using new molecular tools, several wide-ranging species were shown recently to be a mosaic of closely related pseudocryptic species (e.g. ...
... Shih et al. 2010Shih et al. , 2018. In addition, COI data have been useful in resurrecting synonymised species (Shih et al. 2012). Thus, geographic studies of specimens using DNA sequence data have produced a more accurate assessment of fiddler crab biodiversity. ...
A classic dilemma in taxonomy is distinguishing intraspecific from interspecific variation. In order to better comprehend the process of divergence and speciation, we examine morphological, genetic, developmental and behavioural variation among related fiddler crab populations from eastern North America, the Caribbean and South America. We chose geographically remote populations that appear related to Minuca rapax (Smith, 1870). First, using females from across the range of the species, we use geometric morphometric techniques to identify regional differences in carapace shape. Second, in the northern portion of the range, the Caribbean into the Gulf of Mexico, we report variation in the relationship between corporal size and cheliped length in males. Third, we examine the major components of the courtship waves produced by males from several locations in the western Gulf of Mexico. Fourth, we compare the structure of the gastric mill between different populations in the Gulf of Mexico, the Caribbean and the Atlantic Ocean. And, fifth, we use mitochondrial 16S rDNA and cytochrome oxidase subunit I as genetic markers to define the phylogeographic relationship among specimens from more than 20 populations. From these studies, we find discrete, distinct populations across the original range of the species. In particular, populations in the northern Gulf of Mexico appear to represent a lineage that has resulted from limited gene flow and sustained selection pressures. On the basis of the observed degree of divergence, it is apparent that some separated populations in M. rapax should be recognised as evolutionary significant units. The geographic range of these populations is consistent with the historical range for Minuca virens (Salmon & Atsaides, 1968), a putative species that otherwise cannot be consistently distinguished from M. rapax based on discrete external morphological characters. This study provides evidence for M. virens as an emergent but possibly not completely isolated subclade of the M. rapax species complex.
