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Intertidal habitats and decapod (Crustacea) diversity of Qeshm Island, a biodiversity hotspot within the Persian Gulf Reza Naderloo, Michael Türkay & Alireza Sari Marine Biodiversity ISSN 1867-1616 Volume 43 Number 4 Mar Biodiv (2013) 43:445-462 DOI 10.1007/s12526-013-0174-3 1 23 Your article is protected by copyright and all rights are held exclusively by Senckenberg Gesellschaft für Naturforschung and SpringerVerlag Berlin Heidelberg. This e-offprint is for personal use only and shall not be selfarchived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Mar Biodiv (2013) 43:445–462 DOI 10.1007/s12526-013-0174-3 ORIGINAL PAPER Intertidal habitats and decapod (Crustacea) diversity of Qeshm Island, a biodiversity hotspot within the Persian Gulf Reza Naderloo & Michael Türkay & Alireza Sari Received: 21 February 2013 / Revised: 10 July 2013 / Accepted: 12 July 2013 / Published online: 7 August 2013 # Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2013 Abstract Qualitative rapid assessments and taxonomic surveys of decapod crustaceans were carried out along the entire coastline of Qeshm Island in the Persian Gulf. Shore morphology and habitat distribution were examined. Simultaneously, decapod samples were collected from 40 selected sites. In total, 131 species from five different infraorders were identified, of which 61 were recorded for the first time from Qeshm Island and 18 species were new records for the Persian Gulf. The Brachyura possessed the highest species richness (73 species, 56 %) within the different infraorders. Among the surveyed habitats, the combined habitat “rocky/cobble”, occurring mainly along the south coast, had the highest species diversity. Rocky shores were dominated by Grapsus albolineatus, Metopograpsus messor, and Eriphia smithii. Cobble beaches were dominated by Leptodius exaratus, Epixanthus frontalis, Clibanarius signatus, Nanosesarma sarii, Petrolisthes spp. and Alpheus lobidens. Mudflats and mangrove forests, typical habitats of the north coast of Qeshm Island, were bordered along their landward fringe with Nasima dotilliformis and Uca sindensis, which were accompanied by Uca iranica in some places with coarser sediments. On muddy substrate and among mangroves, Metopograpsus messor, Parasesarma persicum, Eurycarcinus orientalis, Macrophthalmus depressus, Metaplax indica, Ilyoplax stevensi, Manningis arabicum, Opusia indica and Alpheus lobidens were the most common species. Exposed sandy beaches, mainly R. Naderloo (*) : A. Sari School of Biology and Center of Excellence in Phylogeny of Living Organisms, College of Science, University of Tehran, 14155-6455 Tehran, Iran e-mail: rnaderloo@ut.ac.ir A. Sari e-mail: sari@ut.ac.ir M. Türkay Research Institute and Natural Museum of Senckenberg, Frankfurt am Main, Germany e-mail: mtuerkay@senckenberg.de found on the south coast, were inhabited by Ocypode rotundata and Coenobita scaevola in high-intertidal and supralittoral zones, respectively. Emerita holthuisi occurred slightly lower in the mid-intertidal zone of relatively steep beaches where Diogenes avarus and Ryphila cancellus were found whenever the sandy beach was relatively flat. Intertidal habitats of the island are generally in relatively good condition compared with the other Persian Gulf states. Keywords Biodiversity . Crustacea . Decapoda . Intertidal habitat . Iran . Persian Gulf . Qeshm Island Introduction Intertidal habitats of the Persian Gulf and the associated species assemblages had been considerably understudied until late 1970s, when Basson et al. (1977) provided a basic description of the Saudi Arabian part of the Persian Gulf, together with their associated fauna. Later, Jones (1986b) and Clayton (1986) conducted ecological studies on the shores of Kuwait (the Northern Persian Gulf) that mainly dealt with the intertidal habitat structure and its vertical faunal zonation. A huge oil spill following the Gulf War of 1991 was the main trigger for some ecological and faunistic works in the coastal zone of the Arabian coast, mainly on the muddy shore along Saudi Arabia (e.g. Apel and Türkay 1992, 1999; Apel 1994a, b, 1996). The mangal fauna of the Persian Gulf is even more poorly studied; indeed, no comprehensive data are available on the decapod crustacean fauna and their zonation in this important ecosystem. Al-Ghais and Cooper (1996) and Cooper (1997) published on the mangal associated Brachyura of Abu Dhabi (UAE) and Al-Khayat and Jones (1999) compared macrofauna in natural and planted mangroves in Qatar. A few ecological and biotopeoriented studies, however, focused on the western and southern coasts of the Persian Gulf. Their results are not fully applicable to the Eastern Persian Gulf, because of considerably different Author's personal copy 446 biological and physical characteristics (Sheppard et al. 1992). The knowledge of the intertidal habitat and associated decapod assemblages in the eastern Persian Gulf is still scarce and there is almost no information on zonation and community structure. Some taxonomic/faunistic studies on the decapod fauna in the intertidal zone of the Iranian coasts have been performed (e.g. Pretzmann 1971; Bahmani 1997; Hosseini 2009; Naderloo and Schubart 2009, 2010; Naderloo and Türkay 2009, 2011; Anker et al. 2010; Naderloo et al. 2011). These mostly deal with a single species/taxon or focus on a small patchy habitat. Since October 2007, a research program is being undertaken along the Iranian coastline of the Persian Gulf, including Qeshm Island, focusing mainly on the intertidal habitat and their associated decapod crustacean assemblages. The results regarding decapod crustacean communities relating to different types of habitats of Qeshm Island are presented herewith. Currently, no detailed data are available on the physical and biological characteristics of the shoreline of this island. Höpner et al. (2000) presented a very short account on the intertidal area of the Persian Gulf, with brief reference to the intertidal habitats of Qeshm Island. Kavousi et al. (2011) provided basic information on the coral reefs at two locations along the south coast of Qeshm Island. The decapod crustacean fauna of the island, like in most parts of the mainland coast of Iran, was completely unknown. Naderloo and Türkay (2009) were the first to describe a decapod crustacean from Qeshm Island, Nanosesarma sarii Naderloo and Türkay 2009, and to redescribe N. jousseaumei Nobili, 1905, as a new record for the Persian Gulf. Naderloo and Schubart (2010) described a further sesarmid crab, Parasesarma persicum, from the Persian Gulf with records from Qeshm Island, as did Anker et al. (2010) by describing a new caridean shrimp (Athanas iranicus) from the island. Macrophthalmus sinuspersici was recently described from the Persian Gulf and recorded from the island (Naderloo and Türkay 2011). However, these are all isolated taxonomic studies not dealing with the whole decapod crustacean communities and providing no information on their occurrence and distribution around the island. The present study provides a basic description of habitat characteristics of intertidal regions together with their associated decapod crustacean assemblages. It is intended to inform future ecological surveys and underpin management decisions regarding the sustainable use of marine resources and regional conservation programs. Materials and methods The research area is located in the south-western part of the Persian Gulf, a few kilometres (about 1.8 km at the closest position between Laft Port of Qeshm Island and Pohl Village on the mainland) off the southern coast of Iran, at the entrance to the Strait of Hormuz. It is the largest island in the Persian Mar Biodiv (2013) 43:445–462 Gulf, with a length of about 110 km, a maximum breadth of 34 km (between Laft and Shibderaz) and a minimum breadth of 10 km (between Salakh and Tabl); with an area of 1,491 km2, it is twice as large as one of the western coastal states, Bahrain. Climatic conditions are arid to semi-arid, with a mean annual rainfall of 160 mm (http://www.irimet.net). The largest mangal ecosystem of the Persian Gulf is located along the north coast of the island (26° 40′-27° 00′N, 55° 21′55° 52′E). It has been assigned in “the Ramsar Convention list” under the name of “Khuran straits” with a total protected area of 100,000 ha. This ecosystem is now regarded as a Biosphere Reserve of IUCN Management Category IV and IX (Höpner et al. 2000). Sampling surveys were primarily conducted to investigate the biophysical structure of the intertidal habitats of Qeshm Island. In a rapid scanning survey, 120 coastal sites around the island were visited, covering almost the entire coastline of the island. Habitat structure was defined mainly using four general substrate categories, comprising: muddy, sandy, rocky/ cobble and mangrove. In most cases, the width of the intertidal region together with the relative slope was measured and their general profile sketched. The salinity of shore water was measured using a hand refractometer. Air and water temperatures were recorded. Accurate site locations were recorded using a hand-held GPS (Garmin, Olathe, KS, USA), and all sites were photographed. Surveys were usually performed during low spring tides, when the major extent of the shore was exposed. A tidal time-table was extracted from http:// www.iranhydrography.org. The tidal system of Qeshm Island completely follows the usual tidal regime of the Persian Gulf, which is generally semidiurnal. Tidal range is fairly regular and considerable, varying from about 4.5 m in the north-eastern end (e.g. Dargahan) to about 3.8 m in northeastern end (e.g. Basaeedu). Most of the sampling surveys were performed in the summer months, when two low tides occur during daylight, one early in the morning and a successive one in the late afternoon. Mapping of the coastal structure was performed by Geographic Information Service (GIS) using software ArcGIS 10. Since 2007, samples have been collected around the Qeshm Island, irregularly from October to April. Taxonomic surveys, assessing the crustacean faunal assemblages, were performed at 40 selected sites, covering all representative habitats (Fig. 1). Our sampling transects ran from land to the sea, i.e. we collected samples along a vertical line. Hence, we refer to the vertical distribution of species along the transects throughout the manuscript. Here vertical does not mean vertically in the sediment, but vertical to the coast line. Samples were collected by a group of at least four biologists, who spent 2–3 h at each site, depending on the shore type. Samples were collected at least twice from all the selected sites in order to ensure a high probability that all existing species were collected. It must be noted that caridean shrimps received less Author's personal copy Mar Biodiv (2013) 43:445–462 447 Fig. 1 The location of Qeshm Island in the Persian Gulf, indicating 40 taxonomic sampling sites. Major cities and villages along the island are indicated attention than other decapods and that the real species numbers for this group will therefore be higher than the ones presented here. Intertidal regions were divided based upon tidal inundation into three main categories: high, mid, and low intertidal zones. In most cases, the supralittoral zone, and in a few cases the sublittoral zone, were also included in this survey. Specimens from the different zones were collected in separate jars or plastic bags. Material was mainly collected by hand, digging in muddy substrate and sieving the fine sediments. At some sites, a hand-made pushnet (metal frame, 75×27 cm; length of net, 130 cm; mesh size, 1 cm) (Manning 1975) was used to Author's personal copy 448 Mar Biodiv (2013) 43:445–462 Table 1 List of families and species collected around Qeshm Island Familya Species No. sitesb Alpheidae (16) Alpheus lobidens De Haan, 1850 Alpheus edamensis De Man, 1888 Alpheus edwardsi Audouin, 1827 Alpheus macrodactylus Ortmann, 1890 Alpheus paracrinitus Miers, 1881 Alpheus lutosus Anker & De Grave, 2009 Alpheus n. sp. 1 Alpheus n. sp. 2 Athanas dimorphus Ortmann 1894 Athanas djiboutensis Coutiere, 1897 Athanas iranicus Anker, Naderloo & Marin, 2010 Athanas n. sp. Salmoneus gracilipes Miya, 1972 Synalpheus coutierei A. H. Banner, 1953 Synalpheus gracilirostris De Man, 1910 Synalpheus hastilicrassus Coutiere, 1905 Latreutes anoplonyx Kemp, 1914 27 5 5 1 1 2 5 1 6 1 1 1 1 2 2 1 1 Lysmata vittata (Stimpson, 1860) Lysmata sp. 1 Lysmata sp. 2 Thor amboinensis (De Man, 1888) Cuapetes elegans (Paulson, 1895) Cuapetes grandis Stimpson, 1860 Palaemon cf. serrifer (Stimpson, 1860) Palaemon cf. debilis Dana, 1852 Palaemonetes sp. 1 Philocheras Parvirostris Kemp, 1916 Microprosthema validum Stimpson, 1860 Upogebia carinicauda (Stimpson, 1860) Upogebia pseudochelata Tattersal, 1921 Upogebia darwinii (Miers, 1884) Upogebia cf. savignyi (Strahl, 1862) Callianidea typa H. Milne Edwards, 1837 Michaelcallianassa indica Sakai, 2002 Neocallichirus jousseaumei (Nobili, 1904) Emerita holthuisi Sankoli, 1965 2 1 1 2 1 1 2 1 1 1 1 3 1 3 1 2 2 5 4 Coenobita scaevola (Forskål, 1775) Calcinus latens (Randall, 1840) Clibanarius signatus Heller, 1861 Diogenes avarus Heller, 1865 Diogenes karwarensis Nayak & Neelakantan, 1989 Diogenes planimanus Henderson, 1893 Diogenes tirmiziae Siddiqui & McLaughlin, 2003 25 1 16 13 7 2 Sandy (sup, high) Rocky, cobble (mid, low) Rocky, cobble (high, mid) Sandy, muddy-sand, cobble (high, mid) Muddy-sand (high, mid) Sandy, cobble (high, mid) 3 2 1 1 3 2 5 Sandy, cobble (high, mid) Sandy, cobble (mid) Cobble (mid) Cobble (high, mid) Rocky, cobble (high, mid) Rocky, cobble (high, mid) Rocky, cobble (high, mid) Hippolytidae (5) Palaemonidae (5) Crangonidae (1) Spongicolidae (1) Upogebiidae (4) Callianassidae (3) Hippidae (1) Coenobitidae (1) Diogenidae (8) Paguridae (1) Porcellanidae (11) Areopaguristes perspicax (Nobili, 1906) Areopaguristes sp. Pagurus kulkarnii Sankolli, 1961 Pachycheles natalensis (Krauss, 1843) Pachycheles tomentosus Henderson, 1893 Petrolisthes boscii (Audouin, 1826) PGc + + + + + + + QId Habitat and zonee + + + + + + Muddy, cobble (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Muddy (mid) Sandy, cobble (mid) Muddy (mid, low) Rocky, cobble (mid) Rocky bed (mid) Sandy, rocky (mid, low) Sandy, cobble (mid, low) Muddy-sand with shells (mid) Muddy-sand with shells (mid) Cobble (mid, low) Cobble (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Jellyfish (low) + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + Rocky (mid, low) Cobble (mid, low) Rocky (sponge) (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Muddy-sand (mid, low) Muddy-sand (mid, low) Rocky, cobble (mid, low) Cobble (mid, low) Muddy-sand (high, mid) Rocky bed (high, mid) Rocky (sponge) (mid, low) Rocky (high, mid) Rocky, cobble (mid, low) Muddy-sand (mid, low) Rocky (mid, low) Exposed sandy (mid) Author's personal copy Mar Biodiv (2013) 43:445–462 449 Table 1 (continued) Familya Dromiidae (1) Matutidae (1) Leucosiidae (4) Hymenosomatidae (1) Majidae (7) Portunidae (10) Eriphidae (1) Menippidae (1) Oziidae (2) Pseudoziidae (1) Xanthidae (7) Pilumnidae (10) Species No. sitesb Petrolisthes leptocheles (Heller, 1861) Petrolisthes rufescens (Heller, 1861) Pisidia dehaani (Krauss, 1843) 12 18 8 Pisidia gordoni (Johnson, 1970) Pisidia inaequalis (Heller, 1861) Polyonyx loimicola Sankolli, 1965 Polyonyx obesulus Miers, 1884 Raphidopus indicus Henderson, 1893 Cryptodromia fallax (Lamarck, 1818) Matuta victor (Fabricius, 1781) Ebalia abdominalis Nobili, 1906 Nursia rubifera Müller, 1887 Hiplyra variegata (Rüppel, 1830) Ryphila cancellus (Herbst, 1783) Elamena sindensis Alcock, 1900 Achaeus lacertosus Stimpson, 1907 Menaethiops nodulosa (Nobili, 1905) Menaethiops n. sp. 1 Menaethiops n. sp. 2 Menaethius monocerus (Latreille, 1825) Micippa platipes (Rüppel, 1830) 3 3 2 2 1 1 3 1 6 2 5 2 1 1 1 2 1 2 Schizophrys aspera (H. Milne Edwards, 1834) Charybdis helleri (A. Milne-Edwards, 1867) Gonioinfradens paucidentata (A. Milne-Edwards 1861) Portunus segnis (Forskål, 1775) Scylla serrata (Forskål, 1775) Thalamita admete (Herbst, 1803) Thalamita crenata Rüppell, 1830 Thalamita poissonii (Savigny, 1817) Thalamita prymna (Herbst, 1803) Thalamita rubridens Apel & Spiridonov, 1998 Thalamita savignyi A. Milne-Edwards, 1861 Eriphia smithi MacLeay, 1838 Menippe rumphii (Fabricius, 1798) Epixanthus frontalis (H. Milne Edwards, 1834) Lydia tenax (Rüppell, 1830) Pseudozius caystrus (Adams & White, 1849) Actaea jacquelinae Guinot, 1976 Atergatis laevigatus A. Milne-Edwards, 1865 Leptodius exaratus (H. Milne Edwards, 1834) 2 2 1 14 2 1 2 1 4 1 1 10 1 6 1 2 1 2 19 + Medeaops neglectus (Balss, 1920) Phymodius drachi Guinot, 1964 Xanthias sinensis (A. Milne-Edwards, 1867) Zozymodes xanthoides (Krauss, 1843) Eurycarcinus integrifrons De Man, 1879 2 3 2 2 + + Eurycarcinus orientalis A. Milne-Edwards, 1867 Heteropanope glabra (Stimpson, 1858) Heteropilumnus trichophoroides (De Man, 1895) Pilumnopeus convexus (Maccagno, 1936) 6 15 2 2 26 PGc + QId Habitat and zonee + Rocky, cobble (high, mid) Rocky, cobble (high, mid) Rocky, cobble (high, mid) + + + + + + + + + + + + + + + + + + + + + + + + + + + + Rocky, cobble (high, mid) Rocky, cobble (high, mid) Muddy (high, mid) Rocky, (sponge) (mid, low) Rocky, muddy (mid, low) Rocky bed (mid) Exposed sandy beach (low) Sandy, cobble (mid, low) Rocky, cobble (mid) Sandy (mid, low) Sandy (mid, low) Rocky, cobble (mid, low) Rocky (mid, low) Rocky bed, cobble (mid, low) Rocky bed, cobble (mid, low) Rocky bed, cobble (mid, low) Rocky bed, cobble (mid, low) Rocky (mid, low) Rocky (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Sandy, rocky (mid, low) Mangroves (mid, low) Rocky, cobble (mid, low) Sandy, cobble (mid, low) Rocky (mid, low) Rocky, cobble (mid, low) Rocky, cobble (mid, low) Cobble (mid, low) Rocky, cobble (high, mid) Rocky, cobble (high, mid) Rocky, cobble (high, mid) Rocky, cobble (high, mid) Rocky, cobble (high, mid) Oyster bank (low) Sandy (low) Rocky, cobble (high, mid, low) Rocky, cobble (mid) Rocky, cobble (mid) Rocky, cobble (mid) Rocky, cobble (mid) Muddy (mid, low) Muddy (mid, low) Cobble (mid) Cobble (mid) Rocky, cobble, muddy (mid, low) Author's personal copy 450 Mar Biodiv (2013) 43:445–462 Table 1 (continued) Familya Dotillidae (4) Macrophthalmidae (7) Camptandriidae (3) Ocypodidae (4) Grapsidae (4) Sesarmidae (3) Varunidae (2) Pinnotheridae (1) a Species No. sitesb Pilumnopeus vauquelini (Audoun, 1826) Pilumnus longicornis Hilgendorf, 1879 Pilumnus savignyi Heller, 1861 1 2 6 Pilumnus sp. Pilumnidae sp. Dotilla blanfordi Alcock, 1900 Scopimera crabricauda Alcock, 1900 Ilyoplax frater (Kemp, 1919) Ilyoplax stevensi (Kemp, 1919) Ilyograpsus rhizophorae (Barnard, 1950) Macrophthalmus sinuspersici Naderloo & Türkay, 2011 Macrophthalmus grandidieri A. Milne-Edwards, 1867 Macrophthalmus depressus Rüppell, 1830 Macrophthalmus dentipes Lucas, 1836 Macrophthalmus laevis A. Milne-Edwards, 1867 Macrophthalmus sulcatus H. Milne Edwards,1852 Manningis arabicum (Jones & Clayton, 1983) Nasima dotilliformis (Alcock, 1900) Opusia indica (Alcock, 1900) Ocypode rotundata Miers, 1882 Uca inversa (Hoffman, 1874) 1 2 2 11 9 13 15 4 1 10 5 5 3 12 13 11 19 1 Uca iranica Pretzmann, 1971 Uca sindensis Alcock, 1900 Grapsus albolineatus Lamarck, 1818 Grapsus granulosus H. Milne Edward, 1853 Metopograpsus messor (Forskål, 1775) Metopograpsus thukuhar (Owen, 1839) Nanosesarma sarii Naderloo & Türkay, 2009 Nanosesarma jousseaumei (Nobili, 1906) Parasesarma persicum Naderloo & Schubart, 2010 Metaplax indica H. Milne Edwards, 1852 Thalassograpsus harpax (Hilgendorf, 1892) Arcotheres placunae (Hornell & Southwell, 1909) 13 12 15 5 28 4 27 5 14 10 7 4 PGc QId Habitat and zonee + + + Rocky, cobble, (mid) Rocky, cobble (mid, low) Rocky, cobble (mid, low) + Rocky, cobble (mid) Rocky (mid) Sandy flat (high) Sandy flat (high) Muddy, mangroves (mid, low) Muddy, mangroves (high, mid) Muddy, mangroves (mid, low) Rocky, cobble (mid) Sandy-mud (high, mid) Muddy, mangroves (mid, low) Muddy, muddy-sand (mid, low) Muddy-sand (high, mid) Muddy-sand, sandy (high, mid) Muddy, mangroves (mid, low) Muddy, mangroves (high) Muddy, mangroves (mid, low) Sandy (high) Sandy-mud (high) + Muddy-sand, mangroves (high) Sandy-mud, mangroves (high) Rocky (high) Rocky, cobble (high, mid) Rocky, muddy, mangroves Muddy, mangroves (high, mid) Cobble, muddy, mangroves (high, mid) Cobble (high, mid) Muddy, mangroves (high, mid) Muddy, mangroves (mid, low) Rocky, cobble (high, mid) Muddy-sandy/sandy flat (high) + + Total number of species in each family is given in parentheses b Number of sites where species have been found c Indicating first records from the Persian Gulf d Indicating first records from Qeshm Island e High, mid and low included in parentheses refer to high-intertidal, mid-intertidal and low-intertidal, respectively sample in shallow subtidal zones of sandy beaches. In a few cases, shallow subtidal zones of rocky shores were investigated by snorkelling. Samples were preserved in 70 % ethanol and some specimens were preserved in 96 % ethanol for further genetic analyses. The material was first shipped to the Zoology Museum of the University of Tehran (ZUTC), where it was sorted and identified preliminarily. Taxonomic examinations of the species were undertaken in the Senckenberg Museum in Frankfurt am Main, Germany (SMF), where they were compared with specimens deposited in the collection or with loaned material from other museums. The material of the present survey is deposited at the ZUTC and SMF. A multivariate cluster analysis of all 40 taxonomic sites was conducted for the decapod species composition using the software package Brodgar 2.6.6. Hierarchical clustering based on Euclidean distance was applied using Ward’s method in order to group the sites with similar species compositions. Taxa which could not be identified to species level were Author's personal copy Mar Biodiv (2013) 43:445–462 ignored. Species which were found on more than four sites and also species with ten occurrences at any site (if occurring at less than four sites) were included in the statistical analyses. Results Faunistic survey A total of 131 decapod species were collected along the intertidal coast of Qeshm Island (Table 1), of which 11 species could not be exactly assigned reliably to hitherto known species. Of these, three alpheid shrimps, including two from the genus Alpheus Fabricius, 1798 and one from the genus Athanas Leach, 1814, are currently being described as new to science. Three caridean species from the families Hippolytidae and Palaemonidae are subject of an ongoing study. The remaining five species in different suborders were unidentifiable, mostly due to being small-sized juveniles. Taxonomic composition The 131 species collected belonged to five different infraorders, i.e. Caridea, Stenopodidea, Thalassinidea, Anomura and Brachyura. The highest number of species (74) or 56 % of all species belonged to the Brachyura (Fig. 2). The species within Brachyura fell into 20 different families, of which Portunidae and Pilumnidae, each with ten species (14 %), were the best represented along the coast of Qeshm Island. Two families Macrophthalmidae (with species characteristics of muddy/ sandy habitats) and Xanthidae (mostly occurring on stony substrate), each with seven species (11 %), had a considerable diversity (Fig. 3). Seven families each were represented by one species only, including: Dromiidae, Matutidae, Hymenosomatidae, Eriphidae, Menippidae, Pseudoziidae and Pinnotheridae. Of seven rare brachyuran families, two (Dromiidae and Menippidae) occurred at a single site (Table 1). The infraorder Stenopodidea was represented by a single species (Microprosthema validum). Caridea and Anomura comprised 27 species (20 %) and 22 species (17 %), respectively, which were relatively well represented in the intertidal region of Qeshm Island. Four families were recorded among caridean shrimps. Alpheidae had the highest species richness, comprising 62 % (16 species) of all collected shrimps, with eight species belonged to the well represented genus Alpheus. The families Hippolytidae and Palaemonidae each with five species (19 %), and Crangonidae with only one species (2 %) were further caridean families found in the survey area (Fig. 2). The infraorder Thalassinidea with seven species (5 %) belonging to the two families, Upogebiidae and Callianassidae, had a relatively low species richness. Within the thalassinidian shrimps, the genus Upogebia with four species, namely U. carinicauda, U. darwinii, U. pseudochelata and 451 Upogebia cf. savignyi, was well represented along the coast of Qeshm Island. Anomuran crabs belonged to five different families, including: Hippidae, Coenobitidae, Diogenidae, Paguridae and Porcellanidae. The families Hippidae (Emerita holthuisi), Coenobitidae (Coenobita scaevola), and Paguridae (Pagurus kulkarnii) were represented by a single species each. The majority of the anomuran species belonged to the two families Diogenidae and Porcellanidae, with eight species (36 %) and 11 species (50 %), respectively. These families had the highest species richness amongst Anomura. Five species of the hermit crabs of the family Diogenidae belonged to the genus Diogenes Dana, 1851, while the genera Calcinus Dana, 1851, Clibanarius Dana, 1852, and Areopaguristes Thomson, 1899, were represented with one, one and two species, respectively. The crab-like Anomura of the family Porcellanidae were particularly diverse at Qeshm Island, and were represented by five genera. Of these, Petrolisthes Stimpson, 1858 and Pisidia Leach, 1820 each comprised three species, and Pachycheles Stimpson, 1858 and Polyonyx Stimpson, 1858 each comprised two species, while Raphidopus indicus was the only species within the genus, occurring in the survey area (Table 1). Evaluating the family-level occurrence (i.e. number of sites at which a taxon was found) and distribution, it is apparent that the Pilumnidae (33 of 40 sites), Sesarmidae (32 sites), Grapsidae (31 sites), Alpheidae (29 sites) and Ocypodidae (28 sites) were the most widely distributed families around Qeshm Island. Diogenidae (26 sites), Coenobitidae (25 sites), Xanthidae (24 sites), Macrophthalmidae (23 sites) Porcellanidae (20 sites), and Portunidae (18 sites) were also common. The family Coenobitidae was represented by as single species (Coenobita scaevola), but occurred at more than half of the surveyed sites (71 %). Several abundant families, which were not as diverse in terms of species richness, were represented by low numbers of species (e.g. Sesarmidae with three species, Grapsidae and Ocypodidae with four species each). The most frequently collected decapod species were Metopograpsus messor (28 sites), Nanosesarma sarii (27 sites), Alpheus lobidens (27 sites), Pilumnopeus convexus (26 sites), and Coenobita scaevola (25 sites) (Table 1). Description of coastal environments Four main types of substrate (muddy, sandy, rocky/cobble and mangrove) occurred along the intertidal zone of Qeshm Island. The majority of the study sites were a combination of these four substrates. Generally, the north coast was sedimentary, mostly sheltered sandy-mud (covering 38 % of the coastline) or muddy (covering 25 % of the coastline), together covering more than half of the northern coastal line (Fig. 4). Mangroves (Tabl mangal system together with the patchily distributed mangrove stands) were the second most common habitat along the north coast covering about 33 % of the northern coastline. Rocky/cobble Author's personal copy 452 coasts were less common and showed a more patchy distribution, their coverage hardly reached 5 %. Pure sandy beaches were totally lacking on the north coast, while sandy/rocky was the least represented substrate (3 %) (Fig. 4). Over than half of the more exposed south coast was sandy in combination with rocky and cobbles, which covered about 53 % of the southern coastline. In contrast, pure sandy beaches were a less common habitat, covering 18 % of the southern coastline. Rocky shores were the second most common coastal habitat (22 %), generally found in association with cobble. Sedimentary flats occurred only in the central part of the south coast (between Shib Deraz and Nagasheh), where a small stand of planted mangrove was present (Fig. 4). Multivariate cluster analyses of the 40 sites with reference to their species composition (Fig. 5), separated the northern sites from the southern ones. Group A includes mostly northern coastal sites plus one site (station 11) from the south coast comprised sedimentary habitats with muddy, sandy-mud or muddy-sand substrates. Group A is further divided into two distinct subgroups: subgroup I includes sedimentary sites with much coarser substrate (usually sandy-mud or muddy-sand). Station 11 (habitat with artificial mangroves at Naghasheh), which had a muddy-sand substrate, is included in this subgroup. Sites included in subgroup II had muddy substrate, containing a similar species composition. Group B comprises more than 70 % of all study sites, including all southern sites and five sites from the north coast. This group is also clearly subdivided into two subgroups. Subgroup III (Fig. 5) contains the seven most diverse sites, which were generally composed of rocky/cobble substrate. The species composition of station 24 (Basaeedu) was closely related to the diverse southern sites and was included with them in the same group. Amongst these seven diverse sites, four sites were considerably decapod crustaceans diverse [station 1 (43 species), station 5 (36 species), station 21 (39 species), and station 24 (34 species)]. Subgroup IV (Fig. 5) of group B contains sites with rocky/sandy substrate. Within this subgroup two smaller groups are distinguishable, a small group (with 6 sites) comprising sandy substrate, and a larger group (with 13 sites) comprising sandy/rocky substrate. Main habitats and decapod communities Mud flat The north coast of Qeshm Island lays in an extremely sheltered location without wave action and with very limited tidal energy, resulting in a wide, gently sloping, muddy-sandy/ sandy-mud flat. Sedimentary flats of the northern coastline were generally soft with a high proportion of coarse particles (sand and shell fragments) in the high-intertidal zone, usually fringed with narrow bands of sandy beach towards the supralittoral zone. Different intertidal zones of extensive Mar Biodiv (2013) 43:445–462 sedimentary flats are very difficult to determine according to spring and neep tide levels. But the zonation of the muddy flat was clearly defined by the vertical distribution of decapod species belonging to the superfamily Ocypodoidea (Table 2). Burrows of Nasima dotilliformis were characteristic of the highintertidal zone, extending normally landwards into the supralittoral zone, where they burrowed in relatively consolidated substrate among salt-marsh plants. In the high intertidal zone, Uca sindensis was sympatric with N. dotilliformis, which was also sympatric with (or sometimes replaced by) Uca iranica in regions with high proportions of coarse particles. Two smallsized species, Ilyoplax stevensi and Opusia indica overlapped in their occurrences with N. dotilliformis in its lower limit in the high-intertidal zone. High-intertidal zones were characterised by muddy-sand mixed with shell fragments patchily fringing with a thin veneer of limestones in their higher limit. Such zones were dominated by a newly recorded thalassinid shrimp, Upogebia carinicauda. Chimney-shaped structures of burrows of this mud shrimp with an average height of 4 cm were characteristic in the summer months (April-June). Macrophthalmus laevis and M. sulcatus were two common macrophthalmid crabs of this zone, occurring slightly below in the high-intertidal zone of this habitat. The mid-intertidal zones were generally marked by burrows of a large macrophthalmid crab (Macrophthalmus dentipes) which tended to burrow in loose sandy-mud substrates. The most distinct associate of M. dentipes was the mudskipper (Periophthlamus waltoni). A wide variety of decapod crustaceans was found sympatric with M. dentipes in the mid-intertidal zone (see Table 2). The low-intertidal zones of mud flats could be determined by the occurrence of Macrophthalmus depressus. This mediumsized macrophthalmid crab was usually found in a narrow band of low-intertidal zones, together with some common decapod crustaceans of the mid-intertidal zone, mainly Metaplax indica, Eurycarcinus orientalis, Eurycarcinus integrifrons and Opusia indica, Alpheus lobidens, Michaelcallianassa indica (Table 2). Mangroves The Qeshm mangal ecosystem “Harra of Tabl” is the largest mangrove system in the Persian Gulf covering over 6,800 ha. The mangrove forest was monospecific with Avicennia marina (Forskål, 1775), containing well-developed trees reaching up to 4–6 m in height and located in a fully sheltered embayment in the northwestern coast of Qeshm. The muddy banks of the water channels in most areas of the Qeshm mangal system were occupied by oyster. Presence of the dead shells promote the establishment of mangroves via the trapping of sediments and hosts oyster-specific brachyurans (e.g. Actaea jacquelinae, Pilumnopeus convexus and Nanosesarma sarii). Sometimes, they were replaced by the barnacle Microeuraphia permitini, which was found growing only on the trunks of the trees. On the muddy banks of the mangroves (low-intertidal zone), the most Author's personal copy Mar Biodiv (2013) 43:445–462 453 Fig. 2 Percentages of the different suborders of Decapoda of Qeshm Island (above); percentages of four families of Caridea of Qeshm Island (below, left); percentages of five families of Anomura of Qeshm Island (below, right) common decapod species were Macrophthalmus depressus, Metaplax indica, Eurycarcinus orientalis, Ilyograpsus rhizophorae, Opusia indica, Alpheus lobidens and Manningis arabicum (Fig. 6). Among the pneumatophores and trunk zones, usually impenetrable, was a habitat for the most common grapsoid crabs, Parasesarma persicum and M. messor, with juveniles distributed on the sediment surface, while adults were generally found in the canopy zone of trees. The rare species Metopograpsus thukuhar was found at only two sites in syntopy with M. messor. Other decapod species found in this zone are presented in Table 1. The landward edge of the Qeshm mangal stand was usually a wide band of muddy flats with sparse halophytic plants, mainly Salicornia sinus-persica, Arthrocnemum macrostachyum and Halocnemum strobilaceum. This zone was dominated by two ocypodoid crabs, Nasima dotilliformis and Uca sindensis. Remarkable burrow structures of N. dotilliformis were one of the prominent features of this zone, which extended to the high-intertidal zone. There were also patchy mangrove swamps which were artificially planted along most parts of the north coast. Most of these fringing artificial mangroves are exposed to the open sea; the majority of planted mangrove bushes had not established or had Author's personal copy 454 Mar Biodiv (2013) 43:445–462 Fig. 3 Percentages of different families of the Brachyura collected from Qeshm Island. Seven families, each with one species, are included in Others developed very poorly, most likely due to the loose nature of muddy-sand/sandy substrate, and partly perhaps because of their immediate exposure to the sea. The single best developed mangrove stand occurred at Holor coast. The substrate was dominantly muddy-sand, trees reaching up to 2 m in height, but not densely populated. This was inhabited by some mangalcharacteristic decapod crustaceans, common fiddler crab, Uca iranica in the N. dotilliformis zone, where it replaced U. sindensis in relatively coarser sediments. On the southern coast of the island, a small stand of A. marina occurred between Band-Chapi and Naghasheh (Fig. 4), where mangrove shrubs were planted along banks of artificial tidal channels running parallel to the sea coast in the sandy or muddy-sand flat. Sea grasses, dominantly Halodule uninervis, were abundant in the sheltered unconsolidated sandy or muddysand substrata. South coast mangroves had some of the brachyuran crabs that are also characteristic for the well-developed mangroves of the north coast. A very small population of the mud crab, Scylla serrata, with medium-sized specimens was found in this mangal ecosystem. Apparently this species has established itself in this narrow habitat quite recently. A further rare species was the fiddler crab Uca inversa, which occurred in a small patch on consolidated substrate along the landward edge. Sandy beaches Most of the sandy beaches were found, usually together with rocky patches, along the south coast of Qeshm Island. Pure sandy beaches were a less common habitat covering about 18 % of the southern coastline (Fig. 4). To the south-east of the island, most beaches had a relatively steep profile, with the narrow intertidal zone sloping down, in most parts, to a rocky subtidal. Rocky structures in the shallow subtidal or lower intertidal zone of the sandy beaches are exposed only during low spring tides in some regions. Supralittoral zones of the sandy beaches were usually characterised by sandy dune belts, vegetated mainly with halophytes (e.g. Suaeda maritima, and Seidlitzia rosmarinus), and bordered mostly along their landward fringe with Acacia spp. The typical decapod crustacean inhabitant of this zone was a semi-terrestrial anomuran known as the land hermit crab, Coenobita scaevola (Table 3). Footprints of this nocturnally active hermit were a main feature of this supralittoral zone, while all collected specimens were found buried in the shaded area of coastal vegetation during day-time, emerging usually at night to scavenge in the intertidal zone. This species was often seen crossing the coastal road, and its footprints were found several hundred metres further landward. The highintertidal zone of such a steep sandy beach was dominated by the only ghost crab of the Persian Gulf, Ocypode rotundata. Conical small pyramids of male animals, in particular, were seen during the breeding season (e.g. with a density of two burrows per 1.0 m2 counted in May in Namakdan). Further down on a steeper slope, the suspension feeding mole crab Emerita holthuisi inhabited a swash active zone. Similar exposed sandy beaches, favourably populated by E. holthuisi, were fully lacking on the north coast. This species was Author's personal copy Mar Biodiv (2013) 43:445–462 455 Fig. 4 Distribution and extent of different habitats along the coastline of Qeshm Island collected only at four sites along the eastern part of the south coast in a dense population of 25 specimens per 0.25 m2. Midintertidal continued to low intertidal and shallow subtidal zones with a gentle profile were favourite habitats of a small leucosiid crab Ryphila cancellus. Matuta victor is a large-sized brachyuran crab usually found sympatric with R. cancellus, but distributed slightly deeper than the latter. Two small-sized brachyuran species of the family Dotillidae, namely Dotilla blanfordi and Scopimera crabicauda were common inhabitants of sandy flats of seaward fringe of mangroves along the south coast. Only the pinnotherid species Arcotheres placunae living in bivalve molluscs Amiantis umbonella also occurred in such sheltered sandy flats. The small-sized hermit crab Diogenes avarus usually occurred on all sandy beaches, being particularly abundant on sandy flats. Rocky shore Rocky shores were merely found along the south coast of Qeshm Island but not as long stretches, rather interrupting sandy beaches in the south-eastern half and cobble-shores mainly in the southwestern half. Nevertheless, rocky shores covered about 18 % of the southern coastline (Fig. 4). Very small patches of rocky substrate occurred in the northeast of the island. These consisted mainly of a thin veneer of sedimentary limestone rocks, and often flourished with typical rocky fauna (e.g. in Kuweii). The coastal rocks around Qeshm Island were mainly sedimentary (limestone), composed of marine organisms’ fragments, mainly those of Foraminifera, echinoderms and mollusc shells. Rocky substrates were often combined with dead corals on the south coast, while no signs of dead corals were found along the north coast. An interesting limestone rocky substrate with coquina rocks occurred in the northwestern part of the island on the Basaeedu headland. Rocky shores of the south coast mainly included rocky cliffs in high-intertidal zones with large boulders seawards (e.g. Namakdan, Kani) and rocky platforms (e.g. Shib-Deraz). Common species of the rocky shore and their relative distribution are shown in Table 3. Common species found in the rocky beds of the Shib Deraz were Nanosesarma sarii, Leptodius exaratus, Pilumnopeus convexus, Eriphia smithi, Petrolisthes rufescens, and Petrolisthes leptocheles. A narrow band (5-10 m wide) of about a 1-km long rocky bed stretched from about 2 km of Qeshm city westwards along the south coast. This rocky bed was a very poor habitat for the crustacean species. The herbivorous crab Grapsus albolineatus Author's personal copy 456 Mar Biodiv (2013) 43:445–462 Fig. 5 Hierarchical cluster analysis using the Squared Euclidean distance performed with Ward’s method in order to group the sites with similar species compositions. Cluster A includes sites on the north coast with mangrove, muddy, sandymud or muddy-sand substrate. Cluster B includes all south coast sites together with some from the north coast, which were mostly combination structure of sandy/ rocky or exclusively rocky or sandy occurred in the high-intertidal zones spreading towards the mid-intertidal zones of nearly all rocky shores, where this highly active crab was sympatric with Metopograpsus messor. The scavenger eriphid crab, E. smithi was found exclusively in crevices, slightly lower on the shore than G. albolineatus. The most common anomuran crab Clibanarius signatus was generally present in shaded areas. Grapsus granulosus was a less common species (found at five sites), recorded for the first time in the Persian Gulf and coexisting with G. albolineatus in the mid-intertidal zone. Table 2 Vertical distribution patterns of the common species of decapod crustaceans in muddy and sandy/muddy shores on Qeshm Island Author's personal copy Mar Biodiv (2013) 43:445–462 457 Fig. 6 Schematic diagram of the main mangrove system along north-western Qeshm Island in Tabl area. Most common decapod crustacean species are shown together with their relative vertical distribution Rocky shores at the south coast, in most cases, supported growth of the brown algae Sargassum sp. in the sublittoral zone. These algae reached up to 60 cm of height in some locations, particularly during the cold season, and were found washed-up on the shore. On the contrary, rocks of the northern coast supported the growth of a green algal tuft. Sedimentary rocks were densely covered with a filamentous green lawn (Table 4). present as well. Common species of the cobble-beaches and their vertical distribution are presented in Table 3, together with rocky shore species. Cobble beaches The faunistic results of the present survey illustrate the scarcity of the information on the taxonomy and assemblage structure of the decapod crustaceans along the intertidal habitats of the Persian Gulf, particularly along the eastern coast (Iranian coast). Of 131 decapod crustaceans collected, about 47 % (61 species) were new records for Qeshm Island. About 14 % (18 species) of all species collected were recorded for the first time from the Persian Gulf. These new records include caridean shrimps: Alpheus edamensis De Man, 1888, A. macrodactylus Ortmann, 1890, A. paracrinitus Miers, 1881, Athanas djiboutensis Coutiere, 1897, Salmoneus gracilipes Miya, 1972, Synalpheus gracilirostris De Man, 1910 and S. hastilicrassus Coutiere, 1905, Cuapetes elegans (Paulson, 1895), Cuapetes grandis Stimpson, 1860, Palaemon serrifer (Stimpson, 1860), P. debilis Dana, 1852, Philocheras parvirostris Kemp, 1916; Stenopodidea: Microprosthema validum Stimpson, 1860; Thalassinidea: Upogebia darwinii (Miers, 1884) and U. pseudochelata Tattersall, 1921; Anomura: Polyonyx loimicola Sankolli, 1965; Brachyura: Heteropanope glabra (Stimpson, 1858), Arcotheres placunae (Hornell and Southwell, 1909). Nine species (7 %) are new to the science, of which four had Cobble beaches, composed of small limestone rocks, occurred scattered only along the south coast. This kind of habitat was usually found in the proximity of rocky shores and sometimes in combination with sandy substrate. Such a complex habitat, usually trapping sediments between loose rocks and supporting algal cover, provided diverse microhabitats allowing different species to occur together in the same locality. Consequently, this environment was the most diverse among the coastal habitats around the island. High-intertidal zones were characterised either by rocks (e.g. Zeytun Park Beach) or by sandy beaches (e.g. Dustku). Cobbles were mainly present in the mid-intertidal zone of the shore, hosting many of the species also occurring at the rocky shores and rock beds, as well as a few on the sandy flat. The common species of this zone, which were mainly found under stones, are Clibanarius signatus, Leptodius exaratus, Nanosesarma sarii, Petrolisthes leptocheles and P. rufescens. The eriphid species Eriphia smithi and different alpheid shrimps were also characteristic of this habitat. Several different species of the families Xanthidae, Pilumnidae and Upogebiidae were Discussion Species richness Author's personal copy 458 Mar Biodiv (2013) 43:445–462 Table 3 Vertical distribution patterns of the common species of decapod crustaceans in exposed sandy shores of Qeshm Island already been described from among the material of the present survey, including: Nanosesarma sarii Naderloo and Türkay, 2009, Parasesarma persicum Naderloo and Schubart, 2010, Athanas iranicus Anker et al., 2010 and Macrophthalmus sinuspersici Naderloo and Türkay, 2011. The island surprisingly includes the highest number of decapod species known from the Iranian coast, where 150 species were recorded (Naderloo and Türkay 2012). Of these, 87 % were recorded from Qeshm Island. In terms of species numbers, Qeshm Island is comparable with other regions in the Indian Ocean. For instance, Socotra Island, located in the tropical Indian Ocean with a total area of 3.796 km2, has 130 recorded decapod species (Simões et al. 2001), while Qeshm Island (1,491 km2) with slightly larger than half the Socotra area has roughly the same number (131) of the decapod species. This occurs in spite of the generally harsh environmental conditions of the Persian Gulf, i.e. high salinity and fluctuating temperatures (e.g. Jones 1986a; Sheppard et al. 1992; Apel and Türkay 1999; Naderloo et al. 2010). However, the location of Qeshm Island at the entrance of the Persian Gulf results in an inflow of fresh low-salinity oceanic waters through the Gulf of Oman and this makes this island very different from the other parts of the Gulf. The high diversity of the island was mainly a result of the high decapod diversity in stony habitats along the south coast, whereas the muddy substrate of the northern coast was about as diverse as other muddy shores within the Gulf (Apel 1994a, 1996; Clayton 1986; Naderloo and Türkay 2012). This is confirmed by comparing the number of decapod species recorded from a rocky/cobble site in Qeshm Island (e.g. 43 species at station Table 4 Vertical distribution patterns of the common species of decapod crustaceans in rocky and rocky/cobble shores of Qeshm Island Author's personal copy Mar Biodiv (2013) 43:445–462 1), with similar sites along the mainland coast of Iran (37 species from Bushehr, 32 species from Moghdan, Naderloo and Türkay 2012). Intertidal habitat and associated decapod fauna The intertidal area of Qeshm Island includes a considerable ecosystem diversity, varying from extensive mangroves, gently sloping sedimentary flats, exposed sandy beaches to cobble/rocky shore. Different habitats with varying physical characteristics favour different species communities; hence the north coast of the island with a predominantly sedimentary structure hosted completely different assemblages of decapod crustaceans compared with the south coast. This was shown by results from the cluster analysis (Fig. 5). The sites clustered according to the nature of the physical structures, which were favoured by different species assemblages. The high decapod species diversity of Qeshm Island is thus a consequence of its high ecosystem diversity. The highest species richness was present in stony habitats, in particular in the mixed substrates of rocky/cobble, rather than is softer sedimentary habitats. This is consistent with results of other local studies (e.g. Basson et al. 1977; Titgen 1982) and generally supported by findings of similar surveys from other parts of the world (Raffaelli and Hawkins 1996; Simões et al. 2001). The high richness is most likely due to the availability of different microenvironments in stony substrates. Of 131 identified decapod species, 88 (67 %) were found in stony shores (rocky, cobble, rocky/cobble), 76 of which exclusively inhabited rocky/cobble (Table 1) in the mid-intertidal zone. Four more common species of the rocky/cobble habitat found to be also common along muddy shores were: Alpheus lobidens, Metopograpsus messor, Nanosesarma sarii and Pilumnopeus convexus. Four fairly common species in stony habitats included Macrophthalmus sinuspersici, Nanosesarma jousseaumei, N. sarii, Thalassograpsus harpax. These species are only known from the Iranian coast of the Gulf and not recorded from the Arabian side, despite of the quite extensive samplings conducted along the western coast (e.g. Basson et al. 1977; Titgen 1982; Jones 1986a, b; Apel 2001). Mud-flats were less diverse, with only about 23 % (30 species) of all collected decapod crustaceans. This is less than half of the rocky/cobble species richness. The brachyuran community of the mud flat and the vertical distribution of different crab species is comparable with those recorded from the northern Persian Gulf coast of Kuwait (Clayton 1986), Saudi Arabia (Apel 1994a, 1996) and from other Iranian coasts of the Persian Gulf (Naderloo and Türkay 2012). The only brachyuran species which is dominant on the high intertidal zone of mud flats on the northern part of the Gulf (Iran, Iraq and Kuwait) is Leptochryseus kuwaitensis (Al-Khayat and Jones 1996; Ng et al. 2009; Naderloo and Türkay 2012). This species 459 is completely missing along the southern coast of the Persian Gulf and replaced by Nasima dotilliformis. The latter has been recorded from all the Gulf States [Iran (Stephensen 1946; present study), Iraq (Ng et al. 2009), Kuwait (Jones 1986a), Saudi Arabia (Basson et al. 1977; Apel 1994a, 1996), Qatar (Al-Khayat and Jones 1996, 1999)]. Although decapod diversity is relatively low in this habitat, the abundance and biomass is considerably high. Some species (e.g. Uca sindensis, Uca iranica and Macrophthalmus dentipes) had the highest density, which can be attributed to the high content of organic material in the muddy substrate. The mangroves of Qeshm Island were not as diverse regarding the decapod crustaceans as other intertidal habitats, but still many more species have been recorded from here compared with other parts of the Persian Gulf. Of the 23 brachyuran crab species recorded from Qeshm Island, 13 and 14 species have been recorded from mangroves in Abu Dhabi (Cooper 1997) and Qatar (Al-Khayat and Jones 1999), respectively. The mangrove habitat shared many specific decapod assemblages with the mud flat. Mangroves ecologically are important because of their high primary productivity, importance as nursery grounds for shrimps and fish larvae and as providing feeding ground for many resident and migratory birds. Two species, Metopograpsus thukuhar and Scylla serrata were exclusively found in mangroves. Metopograpsus messor and Parasesarma persicum were the two most common crabs amongst the trunks and pneumatophores in the study area, whereas in east African mangroves two species Metopograpsus thukuhar and Perisesarma guttatum are the most common and ecologically important species (Vannini and Valmori 1981; Gillikin and Schubart 2004). An interesting record of the present study is the giant mud crab Scylla serrata, which is commercially important. This species is widely distributed in the Indo-West Pacific and has already been recorded from the south coast of the Gulf of Oman (Apel and Spiridonov 1998), its north coast along the Iranian coast in the mangroves of Jask (Rezaie-Atagholipour et al. 2013) and mangroves of Khorkhalasi (R.N. pers. obs.). Here it is the largest and most common species and is used as a food source by local people. Scylla serrata has recently been recorded from the southeast Persian Gulf in Ras al-Khaimah (UAE) by Hogarth and Beech (2001). This species is thought to have been very common in the past when extensive mangroves were widely distributed along the coast of the Persian Gulf (Hogarth and Beech 2001). Numbers declined following the disappearance of mangroves from most parts of the Persian Gulf, but there are sign of recovery. We found this commercially important crab in the planted mangroves along the south coast (Naghasheh), while, despite extensive sampling survey, no individual was recorded in the largest mangal system along the north coast (Tabl). Sandy beaches are the least diverse habitat within the Persian Gulf (Jones 1986b) and all over the world (Raffaelli and Author's personal copy 460 Hawkins 1996). The Decapod community of the sandy beach of Qeshm Island is fully comparable with those of the coast of Kuwait (Jones 1986b) and Saudi Arabia (Basson et al. 1977). In comparison to the adjacent regions, the only genus which appears to be under-represented in the Persian Gulf is Ocypode, which is represented with only one species Ocypode rotundata, while two species of the genus occur in the sandy beaches of Pakistan (Türkay et al. 1996), four species were recorded from Socotra Islands (Türkay et al. 1996; Simões et al. 2001) and two species from the Red Sea (Türkay et al. 1996). The terrestrial hermit crab Coenobita scaevola is the only hermit crab occurring in the supralittoral zone of sandy beaches in the Persian Gulf and Gulf of Oman. This hermit crab was a common species in the sandy beaches of Qeshm Island. Vertical zonation pattern The division of intertidal habitats into three different zones (high, mid and low intertidal) is generally difficult, especially when comparing sites with different physical and topographical characteristics (English et al. 1997). Many workers believe that the zonation should be based on biological elements with determined species compositions and distribution of the key species (e.g. Raffaelli and Hawkins 1996). As this study was the first to determine the species composition of decapod crustaceans and their vertical distribution along the coast of Qeshm Island, it was only possible to use water movement to define different zones. Therefore, we advocate that in using the present data in comparison with data from other regions, differences in tidal regimes and regional topography as well as the physical nature of the shores must be taken into consideration. The vertical distribution of the different brachyuran species in the mud flats of Qeshm Island is relatively similar with that recorded from Kuwait (Clayton 1986), Saudi Arabia (Apel 1994a, 1996) and Iran mainland (Naderloo and Türkay 2012). Unfortunately, there is no documentation on the intertidal rocky/cobble crustacean assemblage from other parts of the Persian Gulf, except a brief study of Jones (1986b) from Kuwait that deals with the most common rocky fauna and flora and this includes few crustacean decapod species. Intertidal zone and biodiversity conservation Monitoring and documenting the biodiversity along the coasts of the Persian Gulf is very important since the area is under extreme pressure due to several environmental and anthropogenic factors, including oil-related industries, coastal developments, urban sewage, developing tourist industry, immigration of invasive species and, most importantly, global warming (e.g. Facey 2008; Khan 2008; Krupp and Abuzinada 2008). Mar Biodiv (2013) 43:445–462 Two recent reviews by Sheppard et al. (2010) and by Sale et al. (2011) clearly address the environmental problems of the Persian Gulf and the necessity for an integrative conservation program. Mangroves and reefs of the Persian Gulf are under greatest threat (Sheppard et al. 2010). Qeshm Island, possessing the largest mangal ecosystem with about a 6,800ha area within the Gulf, is of major importance for conservation of this environment (Spalding et al. 2010). In the past, mangroves were used as building material, firewood and forage for livestock by traditional users, which is possibly the main destructive reason for this ecosystem in the world (Saenger et al. 1983). Currently, only the leaves and young branches of mangroves are used as livestock fodder by rural people. Camels are sometimes also being seen foraging among the arid zones of the system, which locally seem to have a small destructive effect on the mangroves. Illegal shrimp catching is occasionally seen in the Tabl mangal ecosystem, which seems to have the second rank in destructing the mangroves’ fauna and flora. Further, Tabl mangrove forest is now the most important recreational site around the island, and the ecosystem is being opened up to tourism, which needs an integrative tourism management. Nonetheless, Qeshm mangroves are in relatively good condition compared with the mangroves of the other Persian Gulf states like Saudi Arabia and UAE (Spalding et al. 2010). No published record is available on the oil-related pollution of intertidal habitats of Qeshm Island. Considering the location of the island in the Strait of Hormoz, which is a main oil transporting passage, it has the potential to be subject to a heavy oil spill. There are some pollution impacts along the northern coast due to its vicinity to the Bandar-Abbas port. In general, anthropogenic effects presently cannot be considered as a big threat to the intertidal habitats of the island, mainly because of the small rural population inhabiting the coastal area. Developing the free economic zone with subsequent impacts on the island coastal environments is by far the greatest threat to the region. Clear evidence for such a possible threat is seen in the coastal situation of Dubai (Sheppard et al. 2010). Acknowledgments We would like to thank Dr. Ranjbaran (School of Geology, University of Tehran) for helping with identification of the coastal rocks and boulders. The help of Mr. Abbas Kazemi (University of Tehran) in field works is highly acknowledged. We are grateful to Moritz Sonnewald (Senckenberg Museum, Frankfurt) for his help with statistical analysis and working with Brodgar software. The English of the paper was checked by “GRAGE Language Service” of University of Frankfurt, which is especially acknowledged. This biodiversity project was mainly funded by Iran National Science Foundation (INSF) and University of Tehran. Their financial help for facilitating the biodiversity project (including the present one) in the Iranian coast of the Persian Gulf is highly appreciated. We are indebted to DAAD (German Academic Exchange Service) for financial support in the form of a PhD Scholarship for R. Naderloo facilitated extensive data analysis. Author's personal copy Mar Biodiv (2013) 43:445–462 References Al-Ghais SM, Cooper RT (1996) Brachyura (Grapsidae, Ocypodidae, Portunidae, Xanthidae and Leucosiidae) of Umm Al Quwain mangal, United Arab Emirates. Trop Zool 9:409–430 Al-Khayat JA, Jones DA (1996) Two new genera, Manningis and Leptochryseus (Decapoda: Camptandriinae), and descriptions of the first zoea of six Brachyurans from the Arabian Gulf. J Crustac Biol 16(4):797–813 Al-Khayat JA, Jones DA (1999) A comparison of the macrofauna of natural and replanted mangroves in Qatar. 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