INTRODUCTION
Grapsoid crabs are one of the most important groups of intertidal brachyurans in the Gulf of Oman and the Persian Gulf, and are one of the most highly investigated groups (Naderloo, Reference Naderloo2011). Naderloo (Reference Naderloo2011) reported 11 grapsoid species from the gulfs: Grapsus albolineatus Latreille in Milbert, 1812, Grapsus granulosus (H. Milne Edwards, 1853), Metopograpsus messor (Forskål, 1775), Metopograpsus thukuhar (Owen, 1839), Nanosesarma sari (Naderloo & Türkay, 2009), Nanosesarma jousseaumei (Nobili, 1905), Parasesarma persicum (Naderloo & Schubart, 2010), Perisesarma guttatum (A. Milne-Edwards, 1869), Metaplax indica (H. Milne Edwards, 1852), Thalassograpsus harpax (Hilgendorf, 1892) and Plagusia squamosa (Herbst, 1790). Of which, Plagusia guttatum and Plagusia squamosa occur exclusively in the Gulf of Oman, while all remaining species are widely distributed in both the Persian Gulf and Gulf of Oman (Apel & Türkay, Reference Apel and Türkay1999; Naderloo, Reference Naderloo2011; Naderloo & Türkay, Reference Naderloo and Türkay2012). The genus Percnon currently contains eight species, of which four species have been recorded from the Western Indian Ocean. These are Percnon abbreviatum (Dana, 1851), Percnon affine (H. Milne Edwards, 1853), Percnon guinotae Crosnier, Reference Crosnier1965 and Percnon planissimum (Herbst, 1804). The two latter species are distributed northwardly in the Indian Ocean and were recorded from Socotra Island by Simões et al. (Reference Simões, Jones and Apel2001). Percnon affine was first described by H. Milne Edwards (1853) from Madagascar in the western Indian Ocean, but has now been recorded from the Indo-West Pacific, as far-reaching as South Africa to New Caledonia, the Hawaiian Islands, Tahiti, Fiji, the Niue Islands and the Cocos-Keeling Islands (Crosnier, Reference Crosnier1965; Mclay, Reference McLay1989; Woodroffe, Reference Woodroffe1994). The present record of Percnon affine from the Gulf of Oman further widens its distribution range to the northern Indian Ocean.
SYSTEMATICS
Order DECAPODA Latreille, 1803
Infraorder BRACHYURA Linnaeus, 1758
Family PLAGUSIIDAE Dana, 1851
Genus Percnon Gistel, 1848
Percnon affine (H. Milne Edwards, 1853)
(Figures 1–5)
Map showing the distribution of Percnon affine (H. Milne Edwards, 1853) in the Indo-West Pacific. Solid circles indicate the already recorded area; hollow circles indicate the sampling point of the species in the present study in the northern Indian Ocean.
Typical habitat of Percnon affine (H. Milne Edwards, 1853) at Large Beach, Chabahar, Iran, Gulf of Oman. Photo credit: Yaser Fatemi.
Living specimen of Percnon affine (H. Milne Edwards, 1853) in the sampling site at Large Beach, Chabahar, Iran, Gulf of Oman. Frontal view. Photo credit: Yaser Fatemi.
Living specimen of Percnon affine (H. Milne Edwards, 1853) in sampling site at Large Beach, Chabahar, Iran, Gulf of Oman. Dorsal view, same specimen as Figure 2. Photo credit: Yaser Fatemi.
Apical part of the first male gonopod of Percnon affine (H. Milne Edwards, 1853), lateral view. Scale bar: 40 μm.
MATERIAL EXAMINED
Six males and two females were collected by hand in mid-intertidal zone of rocky habitat in Large Beach, Chabahar (Iran, Gulf of Oman, 25°16′N 60°40′E, Figure 1). The specimens were living under stones, in crevices and on the algal bed (Figure 2). The materials collected for taxonomic examination were deposited in the Zoological Museum, University of Tehran, under voucher number ‘ZUTC 5314’.
RESULTS
The morphology of the collected specimens fits well with the available taxonomic descriptions (Crosnier, Reference Crosnier1965; Mclay, Reference McLay1989). Percnon affine has distinctive features that can be easily distinguished from the other closely related species, such as Percnon guinotae (Crosnier, Reference Crosnier1965), Percnon abbreviatum (Dana, 1851) and Percnon planissimum (Herbst, 1804). The anterolateral teeth are strongly developed. Six small spines are present along the anterolateral margins terminating in a large spine (Figures 3 & 4). Rostral area equipped with two large, upwardly curved sharp spines. Five small spines present along the lateral margins of the rostral spines. Pre-orbital margins bear a single large spine and two smaller ones. Epistome has three equal spines, walking legs with six or seven large spines on the anterior margin of merus. Male gonopods have an apical process bending outward with 45° (Figure 5).
One to two specimens per m2 were found along the outthrust rock veneer. The specimens were found under stones, in crevices and on the algal bed.
DISCUSSION
Despite intensive sampling along the Iranian coasts of the Gulf of Oman and the Persian Gulf during the last decade, we did not encounter this species in these intertidal rocky habitats. Whereas, the two most common grapsoid species, Grapsus albolineatus and Plagusia squamosal, were prominent inhabitants of the rocky intertidal zones on the coasts of the Gulf of Oman. It seems that the occurrence of Percnon affine (H. Milne Edwards, 1853) has been overlooked during these years and that it was established in this region long before 2013. Surprisingly, the newly arrived species now has a well-established population in sympatricity with the two native species. The alien species even appears to be more common than Percnon squamosa in the mid-intertidal zone, where the two species have denser populations. Whereas, G. albolineatus has a wider spatial distribution upward to the higher intertidal zone. Most grapsoid crabs live in shallow rocky shores up to a depth of 2 m, with or without macroalgal cover (Thessalou-Legaki et al., Reference Thessalou-Legaki, Zenetos, Kambouroglou, Corsini-Foka, Kouraklis, Dounas and Nicolaidou2006). The Iranian coasts of the Gulf of Oman have rocky structures, which are likely to act suitable habitats for such grapsoid crabs. Percnon affine is easily distinguishable from Percnon guinotae Crosnier, Reference Crosnier1965 and Percnon planissimum (Herbst, 1804), as it has a large patch of dense setae on the proximal inner surface of its cheliped pal, whereas, the setal patch is small in Percnon guinotae and absent in Percnon planissimum (see Crosnier, Reference Crosnier1965: Figures 136–138). Regarding the carapace morphology, Percnon affine is close to Percnon guinotae, but distinct from the other species by the number of features it has; in particular, by having distinct spines and an anterior margin of epistome (Crosnier, Reference Crosnier1965: Figures 131, 133).
The Gulf of Oman is located in a heavy traffic line of oil/gas transportation. Most of the oil and gas of the Persian Gulf is transported using tankers through the Strait of Hormuz and the Gulf of Oman to all over the world (Haapkylai et al., Reference Haapkylai, Ramade and Salvat2007). Shipping plays a very important role in introducing alien species to Iranian waters (Naderloo, Reference Naderloo2014). Percnon affine can be transported by surface currents (Pipitone et al., Reference Pipitone, Badalamenti and Sparrow2001) and ballast water (Galil et al., Reference Galil, Froglia, Noel and Briand2002; Azzurro et al., Reference Azzurro, Milazzo, Maynou, Abelló and Temraz2011). Although, it can also be easily transported on ship hulls, similar to other species of rafting crabs of the family Plagusiidae. Primarily, ballast water is a potential vector for the larval dispersal of most marine organisms, and particularly in regions such as those of the Persian Gulf and the Gulf of Oman, which experience a large amount of exchanged ballast water (Naderloo, in press). The Hepu mitten crab Eriochier hepuensis (Clark et al., Reference Clark, Abdul-Sahib and Al-Asadi2006; Naser et al., Reference Naser, Timothy, Ng, Apel, Yasser, Bishop, Ng and Clark2012; Naderloo, Reference Naderloo2014) was most likely introduced to the northern Persian Gulf estuarine system via ballast water. Likewise, ballast water can play the same role in the dispersal of Percnon affine. Adults of this species can also reach the northern coast of the Indian Ocean via drifting debris, buoys and ship hulls. The wide distribution of Percnon squamosa has been explained by such anthropogenic activities. Due to ever-increasing regional anthropogenic activities and recent global warming, more species are expected to disperse to higher latitudes. Therefore, necessary risk assessments should be carried out in this region. Furthermore, molecular studies may be needed to find the source population of already introduced species, such as Percnon affine.
ACKNOWLEDGEMENT
We are very grateful to T. Rashidi and Rouholah Fatemi for their valuable help during this study.