Intertidal habitats and decapod (Crustacea) diversity of Qeshm Island, a biodiversity hotspot within...

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Marine Biodiversity ISSN 1867-1616Volume 43Number 4 Mar Biodiv (2013) 43:445-462DOI 10.1007/s12526-013-0174-3

Intertidal habitats and decapod(Crustacea) diversity of Qeshm Island, abiodiversity hotspot within the Persian Gulf

Reza Naderloo, Michael Türkay &Alireza Sari

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ORIGINAL PAPER

Intertidal habitats and decapod (Crustacea) diversityof 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 surveysof decapod crustaceanswere carried out along the entire coastlineof Qeshm Island in the Persian Gulf. Shore morphology andhabitat distribution were examined. Simultaneously, decapodsamples were collected from 40 selected sites. In total, 131species from five different infraorders were identified, of which61 were recorded for the first time from Qeshm Island and 18species were new records for the Persian Gulf. The Brachyurapossessed the highest species richness (73 species, 56 %) withinthe different infraorders. Among the surveyed habitats, the com-bined habitat “rocky/cobble”, occurring mainly along the southcoast, had the highest species diversity. Rocky shores weredominated by Grapsus albolineatus, Metopograpsus messor,and Eriphia smithii. Cobble beaches were dominated byLeptodius exaratus, Epixanthus frontalis, Clibanarius signatus,Nanosesarma sarii, Petrolisthes spp. and Alpheus lobidens.Mudflats andmangrove forests, typical habitats of the north coastof Qeshm Island, were bordered along their landward fringe withNasima dotilliformis and Uca sindensis, which were accompa-nied by Uca iranica in some places with coarser sediments. Onmuddy substrate and amongmangroves,Metopograpsus messor,Parasesarma persicum, Eurycarcinus orientalis, Macro-phthalmus depressus, Metaplax indica, Ilyoplax stevensi,Manningis arabicum, Opusia indica and Alpheus lobidens werethe most common species. Exposed sandy beaches, mainly

found on the south coast, were inhabited by Ocypode rotundataandCoenobita scaevola in high-intertidal and supralittoral zones,respectively. Emerita holthuisi occurred slightly lower in themid-intertidal zone of relatively steep beaches where Diogenesavarus and Ryphila cancellus were found whenever the sandybeach was relatively flat. Intertidal habitats of the island aregenerally in relatively good condition compared with the otherPersian Gulf states.

Keywords Biodiversity . Crustacea . Decapoda . Intertidalhabitat . Iran . Persian Gulf . Qeshm Island

Introduction

Intertidal habitats of the Persian Gulf and the associated speciesassemblages had been considerably understudied until late1970s, when Basson et al. (1977) provided a basic descriptionof the Saudi Arabian part of the Persian Gulf, together withtheir associated fauna. Later, Jones (1986b) and Clayton (1986)conducted ecological studies on the shores of Kuwait (theNorthern Persian Gulf) that mainly dealt with the intertidalhabitat structure and its vertical faunal zonation. A huge oilspill following the Gulf War of 1991 was the main trigger forsome ecological and faunistic works in the coastal zone of theArabian coast, mainly on the muddy shore along Saudi Arabia(e.g. Apel and Türkay 1992, 1999; Apel 1994a, b, 1996). Themangal fauna of the Persian Gulf is even more poorly studied;indeed, no comprehensive data are available on the decapodcrustacean fauna and their zonation in this important ecosys-tem. Al-Ghais andCooper (1996) andCooper (1997) publishedon the mangal associated Brachyura of Abu Dhabi (UAE) andAl-Khayat and Jones (1999) compared macrofauna in naturaland planted mangroves in Qatar. A few ecological and biotope-oriented studies, however, focused on the western and southerncoasts of the Persian Gulf. Their results are not fully applicableto the Eastern Persian Gulf, because of considerably different

R. Naderloo (*) :A. SariSchool of Biology and Center of Excellence in Phylogeny of LivingOrganisms, College of Science, University of Tehran,14155-6455 Tehran, Irane-mail: [email protected]

A. Sarie-mail: [email protected]

M. TürkayResearch Institute and Natural Museum of Senckenberg,Frankfurt am Main, Germanye-mail: [email protected]

Mar Biodiv (2013) 43:445–462DOI 10.1007/s12526-013-0174-3

Author's personal copy

biological and physical characteristics (Sheppard et al. 1992).The knowledge of the intertidal habitat and associated decapodassemblages in the eastern Persian Gulf is still scarce and thereis almost no information on zonation and community structure.Some taxonomic/faunistic studies on the decapod fauna in theintertidal zone of the Iranian coasts have been performed (e.g.Pretzmann 1971; Bahmani 1997; Hosseini 2009; Naderloo andSchubart 2009, 2010; Naderloo and Türkay 2009, 2011; Ankeret al. 2010; Naderloo et al. 2011). These mostly deal with asingle species/taxon or focus on a small patchy habitat.

Since October 2007, a research program is being undertak-en along the Iranian coastline of the Persian Gulf, includingQeshm Island, focusing mainly on the intertidal habitat andtheir associated decapod crustacean assemblages. The resultsregarding decapod crustacean communities relating to differ-ent types of habitats of Qeshm Island are presented herewith.

Currently, no detailed data are available on the physical andbiological characteristics of the shoreline of this island.Höpner et al. (2000) presented a very short account on theintertidal area of the Persian Gulf, with brief reference to theintertidal habitats of Qeshm Island. Kavousi et al. (2011)provided basic information on the coral reefs at two locationsalong the south coast of Qeshm Island. The decapod crusta-cean fauna of the island, like inmost parts of themainland coastof Iran, was completely unknown. Naderloo and Türkay (2009)were the first to describe a decapod crustacean from QeshmIsland, Nanosesarma sarii Naderloo and Türkay 2009, and toredescribeN. jousseaumeiNobili, 1905, as a new record for thePersian Gulf. Naderloo and Schubart (2010) described a furthersesarmid crab, Parasesarma persicum, from the Persian Gulfwith records from Qeshm Island, as did Anker et al. (2010) bydescribing a new caridean shrimp (Athanas iranicus) from theisland. Macrophthalmus sinuspersici was recently describedfrom the Persian Gulf and recorded from the island (Naderlooand Türkay 2011). However, these are all isolated taxonomicstudies not dealing with the whole decapod crustacean com-munities and providing no information on their occurrence anddistribution around the island.

The present study provides a basic description of habitatcharacteristics of intertidal regions together with their associ-ated decapod crustacean assemblages. It is intended to informfuture ecological surveys and underpin management decisionsregarding the sustainable use of marine resources and regionalconservation programs.

Materials and methods

The research area is located in the south-western part of thePersian Gulf, a few kilometres (about 1.8 km at the closestposition between Laft Port of Qeshm Island and Pohl Villageon the mainland) off the southern coast of Iran, at the entranceto the Strait of Hormuz. It is the largest island in the Persian

Gulf, with a length of about 110 km, a maximum breadth of34 km (between Laft and Shibderaz) and a minimum breadthof 10 km (between Salakh and Tabl); with an area of1,491 km2, it is twice as large as one of the western coastalstates, Bahrain. Climatic conditions are arid to semi-arid, witha mean annual rainfall of 160 mm (http://www.irimet.net).The largest mangal ecosystem of the Persian Gulf is locatedalong the north coast of the island (26° 40′-27° 00′N, 55° 21′-55° 52′E). It has been assigned in “the Ramsar Conventionlist” under the name of “Khuran straits” with a total protectedarea of 100,000 ha. This ecosystem is now regarded as aBiosphere Reserve of IUCN Management Category IV andIX (Höpner et al. 2000).

Sampling surveys were primarily conducted to investigatethe biophysical structure of the intertidal habitats of QeshmIsland. In a rapid scanning survey, 120 coastal sites around theisland were visited, covering almost the entire coastline of theisland. Habitat structure was defined mainly using four gen-eral substrate categories, comprising: muddy, sandy, rocky/cobble andmangrove. In most cases, the width of the intertidalregion together with the relative slope was measured and theirgeneral profile sketched. The salinity of shore water wasmeasured using a hand refractometer. Air and water tempera-tures were recorded. Accurate site locations were recordedusing a hand-held GPS (Garmin, Olathe, KS, USA), and allsites were photographed. Surveys were usually performedduring low spring tides, when the major extent of the shorewas exposed. A tidal time-table was extracted from http://www.iranhydrography.org. The tidal system of QeshmIsland completely follows the usual tidal regime of thePersian Gulf, which is generally semidiurnal. Tidal range isfairly regular and considerable, varying from about 4.5 m inthe north-eastern end (e.g. Dargahan) to about 3.8 m in north-eastern end (e.g. Basaeedu). Most of the sampling surveyswere performed in the summer months, when two low tidesoccur during daylight, one early in the morning and a succes-sive one in the late afternoon. Mapping of the coastal structurewas performed by Geographic Information Service (GIS)using software ArcGIS 10.

Since 2007, samples have been collected around theQeshm Island, irregularly from October to April. Taxonomicsurveys, assessing the crustacean faunal assemblages, wereperformed at 40 selected sites, covering all representativehabitats (Fig. 1). Our sampling transects ran from land to thesea, i.e. we collected samples along a vertical line. Hence, werefer to the vertical distribution of species along the transectsthroughout the manuscript. Here vertical does not mean ver-tically in the sediment, but vertical to the coast line. Sampleswere collected by a group of at least four biologists, who spent2–3 h at each site, depending on the shore type. Samples werecollected at least twice from all the selected sites in order toensure a high probability that all existing species were col-lected. It must be noted that caridean shrimps received less

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http://www.irimet.net

http://www.iranhydrography.org

http://www.iranhydrography.org

attention than other decapods and that the real species num-bers for this group will therefore be higher than the onespresented here.

Intertidal regions were divided based upon tidal inundationinto 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. Specimensfrom the different zones were collected in separate jars orplastic bags. Material was mainly collected by hand, diggingin muddy substrate and sieving the fine sediments. At somesites, a hand-made pushnet (metal frame, 75×27 cm; length ofnet, 130 cm; mesh size, 1 cm) (Manning 1975) was used to

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

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Table 1 List of families and species collected around Qeshm Island

Familya Species No. sitesb PGc QId Habitat and zonee

Alpheidae (16) Alpheus lobidens De Haan, 1850 27 + Muddy, cobble (mid, low)

Alpheus edamensis De Man, 1888 5 + + Rocky, cobble (mid, low)

Alpheus edwardsi Audouin, 1827 5 + Rocky, cobble (mid, low)

Alpheus macrodactylus Ortmann, 1890 1 + + Muddy (mid)

Alpheus paracrinitus Miers, 1881 1 + + Sandy, cobble (mid)

Alpheus lutosus Anker & De Grave, 2009 2 + Muddy (mid, low)

Alpheus n. sp. 1 5 Rocky, cobble (mid)

Alpheus n. sp. 2 1 Rocky bed (mid)

Athanas dimorphus Ortmann 1894 6 + Sandy, rocky (mid, low)

Athanas djiboutensis Coutiere, 1897 1 + + Sandy, cobble (mid, low)

Athanas iranicus Anker, Naderloo & Marin, 2010 1 Muddy-sand with shells (mid)

Athanas n. sp. 1 Muddy-sand with shells (mid)

Salmoneus gracilipes Miya, 1972 1 + + Cobble (mid, low)

Synalpheus coutierei A. H. Banner, 1953 2 + Cobble (mid, low)

Synalpheus gracilirostris De Man, 1910 2 + + Rocky, cobble (mid, low)

Synalpheus hastilicrassus Coutiere, 1905 1 + + Rocky, cobble (mid, low)

Hippolytidae (5) Latreutes anoplonyx Kemp, 1914 1 + Jellyfish (low)

Lysmata vittata (Stimpson, 1860) 2 + Rocky (mid, low)

Lysmata sp. 1 1 Cobble (mid, low)

Lysmata sp. 2 1 Rocky (sponge) (mid, low)

Thor amboinensis (De Man, 1888) 2 + Rocky, cobble (mid, low)

Palaemonidae (5) Cuapetes elegans (Paulson, 1895) 1 + + Rocky, cobble (mid, low)

Cuapetes grandis Stimpson, 1860 1 + + Rocky, cobble (mid, low)

Palaemon cf. serrifer (Stimpson, 1860) 2 + + Rocky, cobble (mid, low)

Palaemon cf. debilis Dana, 1852 1 + + Muddy-sand (mid, low)

Palaemonetes sp. 1 1 Muddy-sand (mid, low)

Crangonidae (1) Philocheras Parvirostris Kemp, 1916 1 + + Rocky, cobble (mid, low)

Spongicolidae (1) Microprosthema validum Stimpson, 1860 1 + + Cobble (mid, low)

Upogebiidae (4) Upogebia carinicauda (Stimpson, 1860) 3 Muddy-sand (high, mid)

Upogebia pseudochelata Tattersal, 1921 1 + + Rocky bed (high, mid)

Upogebia darwinii (Miers, 1884) 3 + + Rocky (sponge) (mid, low)

Upogebia cf. savignyi (Strahl, 1862) 1 + Rocky (high, mid)

Callianassidae (3) Callianidea typa H. Milne Edwards, 1837 2 Rocky, cobble (mid, low)

Michaelcallianassa indica Sakai, 2002 2 Muddy-sand (mid, low)

Neocallichirus jousseaumei (Nobili, 1904) 5 Rocky (mid, low)

Hippidae (1) Emerita holthuisi Sankoli, 1965 4 + Exposed sandy (mid)

Coenobitidae (1) Coenobita scaevola (Forskål, 1775) 25 Sandy (sup, high)

Diogenidae (8) Calcinus latens (Randall, 1840) 1 Rocky, cobble (mid, low)

Clibanarius signatus Heller, 1861 16 Rocky, cobble (high, mid)

Diogenes avarus Heller, 1865 13 Sandy, muddy-sand, cobble (high, mid)

Diogenes karwarensis Nayak & Neelakantan, 1989 7 Muddy-sand (high, mid)

Diogenes planimanus Henderson, 1893 2 Sandy, cobble (high, mid)

Diogenes tirmiziae Siddiqui & McLaughlin, 2003 3 Sandy, cobble (high, mid)

Areopaguristes perspicax (Nobili, 1906) 2 Sandy, cobble (mid)

Areopaguristes sp. 1 Cobble (mid)

Paguridae (1) Pagurus kulkarnii Sankolli, 1961 1 Cobble (high, mid)

Porcellanidae (11) Pachycheles natalensis (Krauss, 1843) 3 + Rocky, cobble (high, mid)

Pachycheles tomentosus Henderson, 1893 2 + Rocky, cobble (high, mid)

Petrolisthes boscii (Audouin, 1826) 5 + Rocky, cobble (high, mid)

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Table 1 (continued)


Petrolisthes leptocheles (Heller, 1861) 12 Rocky, cobble (high, mid)

Petrolisthes rufescens (Heller, 1861) 18 Rocky, cobble (high, mid)

Pisidia dehaani (Krauss, 1843) 8 + Rocky, cobble (high, mid)

Pisidia gordoni (Johnson, 1970) 3 + Rocky, cobble (high, mid)

Pisidia inaequalis (Heller, 1861) 3 + Rocky, cobble (high, mid)

Polyonyx loimicola Sankolli, 1965 2 + + Muddy (high, mid)

Polyonyx obesulusMiers, 1884 2 + Rocky, (sponge) (mid, low)

Raphidopus indicus Henderson, 1893 1 + Rocky, muddy (mid, low)

Dromiidae (1) Cryptodromia fallax (Lamarck, 1818) 1 + Rocky bed (mid)

Matutidae (1) Matuta victor (Fabricius, 1781) 3 Exposed sandy beach (low)

Leucosiidae (4) Ebalia abdominalis Nobili, 1906 1 + Sandy, cobble (mid, low)

Nursia rubiferaMüller, 1887 6 + Rocky, cobble (mid)

Hiplyra variegata (Rüppel, 1830) 2 + Sandy (mid, low)

Ryphila cancellus (Herbst, 1783) 5 Sandy (mid, low)

Hymenosomatidae (1) Elamena sindensis Alcock, 1900 2 + Rocky, cobble (mid, low)

Majidae (7) Achaeus lacertosus Stimpson, 1907 1 + Rocky (mid, low)

Menaethiops nodulosa (Nobili, 1905) 1 + Rocky bed, cobble (mid, low)

Menaethiops n. sp. 1 1 Rocky bed, cobble (mid, low)

Menaethiops n. sp. 2 2 Rocky bed, cobble (mid, low)

Menaethius monocerus (Latreille, 1825) 1 + Rocky bed, cobble (mid, low)

Micippa platipes (Rüppel, 1830) 2 + Rocky (mid, low)

Schizophrys aspera (H. Milne Edwards, 1834) 2 + Rocky (mid, low)

Portunidae (10) Charybdis helleri (A. Milne-Edwards, 1867) 2 Rocky, cobble (mid, low)

Gonioinfradens paucidentata (A. Milne-Edwards 1861) 1 + Rocky, cobble (mid, low)

Portunus segnis (Forskål, 1775) 14 Sandy, rocky (mid, low)

Scylla serrata (Forskål, 1775) 2 + Mangroves (mid, low)

Thalamita admete (Herbst, 1803) 1 + Rocky, cobble (mid, low)

Thalamita crenata Rüppell, 1830 2 Sandy, cobble (mid, low)

Thalamita poissonii (Savigny, 1817) 1 + Rocky (mid, low)

Thalamita prymna (Herbst, 1803) 4 Rocky, cobble (mid, low)

Thalamita rubridens Apel & Spiridonov, 1998 1 + Rocky, cobble (mid, low)

Thalamita savignyi A. Milne-Edwards, 1861 1 + Cobble (mid, low)

Eriphidae (1) Eriphia smithi MacLeay, 1838 10 Rocky, cobble (high, mid)

Menippidae (1) Menippe rumphii (Fabricius, 1798) 1 + Rocky, cobble (high, mid)

Oziidae (2) Epixanthus frontalis (H. Milne Edwards, 1834) 6 Rocky, cobble (high, mid)

Lydia tenax (Rüppell, 1830) 1 Rocky, cobble (high, mid)

Pseudoziidae (1) Pseudozius caystrus (Adams & White, 1849) 2 + Rocky, cobble (high, mid)

Xanthidae (7) Actaea jacquelinae Guinot, 1976 1 + Oyster bank (low)

Atergatis laevigatus A. Milne-Edwards, 1865 2 + Sandy (low)

Leptodius exaratus (H. Milne Edwards, 1834) 19 Rocky, cobble (high, mid, low)

Medeaops neglectus (Balss, 1920) 2 + Rocky, cobble (mid)

Phymodius drachi Guinot, 1964 3 + Rocky, cobble (mid)

Xanthias sinensis (A. Milne-Edwards, 1867) 2 Rocky, cobble (mid)

Zozymodes xanthoides (Krauss, 1843) 2 + Rocky, cobble (mid)

Pilumnidae (10) Eurycarcinus integrifrons De Man, 1879 6 Muddy (mid, low)

Eurycarcinus orientalis A. Milne-Edwards, 1867 15 Muddy (mid, low)

Heteropanope glabra (Stimpson, 1858) 2 + + Cobble (mid)

Heteropilumnus trichophoroides (De Man, 1895) 2 + Cobble (mid)

Pilumnopeus convexus (Maccagno, 1936) 26 Rocky, cobble, muddy (mid, low)

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sample in shallow subtidal zones of sandy beaches. In a fewcases, shallow subtidal zones of rocky shores were investigatedby snorkelling. Samples were preserved in 70 % ethanol andsome specimens were preserved in 96 % ethanol for furthergenetic analyses. The material was first shipped to the ZoologyMuseum of the University of Tehran (ZUTC), where it wassorted and identified preliminarily. Taxonomic examinations ofthe species were undertaken in the Senckenberg Museum inFrankfurt am Main, Germany (SMF), where they were

compared with specimens deposited in the collection or withloaned material from other museums. The material of thepresent survey is deposited at the ZUTC and SMF.

A multivariate cluster analysis of all 40 taxonomic siteswas conducted for the decapod species composition using thesoftware package Brodgar 2.6.6. Hierarchical clustering basedon Euclidean distance was applied using Ward’s method inorder to group the sites with similar species compositions.Taxa which could not be identified to species level were

Table 1 (continued)


Pilumnopeus vauquelini (Audoun, 1826) 1 + Rocky, cobble, (mid)

Pilumnus longicornis Hilgendorf, 1879 2 + Rocky, cobble (mid, low)

Pilumnus savignyi Heller, 1861 6 + Rocky, cobble (mid, low)

Pilumnus sp. 1 Rocky, cobble (mid)

Pilumnidae sp. 2 Rocky (mid)

Dotillidae (4) Dotilla blanfordi Alcock, 1900 2 Sandy flat (high)

Scopimera crabricauda Alcock, 1900 11 Sandy flat (high)

Ilyoplax frater (Kemp, 1919) 9 + Muddy, mangroves (mid, low)

Ilyoplax stevensi (Kemp, 1919) 13 Muddy, mangroves (high, mid)

Macrophthalmidae (7) Ilyograpsus rhizophorae (Barnard, 1950) 15 Muddy, mangroves (mid, low)

Macrophthalmus sinuspersici Naderloo & Türkay, 2011 4 Rocky, cobble (mid)

Macrophthalmus grandidieri A. Milne-Edwards, 1867 1 Sandy-mud (high, mid)

Macrophthalmus depressus Rüppell, 1830 10 Muddy, mangroves (mid, low)

Macrophthalmus dentipes Lucas, 1836 5 Muddy, muddy-sand (mid, low)

Macrophthalmus laevis A. Milne-Edwards, 1867 5 Muddy-sand (high, mid)

Macrophthalmus sulcatus H. Milne Edwards,1852 3 Muddy-sand, sandy (high, mid)

Camptandriidae (3) Manningis arabicum (Jones & Clayton, 1983) 12 Muddy, mangroves (mid, low)

Nasima dotilliformis (Alcock, 1900) 13 Muddy, mangroves (high)

Opusia indica (Alcock, 1900) 11 Muddy, mangroves (mid, low)

Ocypodidae (4) Ocypode rotundata Miers, 1882 19 Sandy (high)

Uca inversa (Hoffman, 1874) 1 + Sandy-mud (high)

Uca iranica Pretzmann, 1971 13 Muddy-sand, mangroves (high)

Uca sindensis Alcock, 1900 12 Sandy-mud, mangroves (high)

Grapsidae (4) Grapsus albolineatus Lamarck, 1818 15 Rocky (high)

Grapsus granulosus H. Milne Edward, 1853 5 Rocky, cobble (high, mid)

Metopograpsus messor (Forskål, 1775) 28 Rocky, muddy, mangroves

Metopograpsus thukuhar (Owen, 1839) 4 Muddy, mangroves (high, mid)

Sesarmidae (3) Nanosesarma sarii Naderloo & Türkay, 2009 27 Cobble, muddy, mangroves (high, mid)

Nanosesarma jousseaumei (Nobili, 1906) 5 Cobble (high, mid)

Parasesarma persicum Naderloo & Schubart, 2010 14 Muddy, mangroves (high, mid)

Varunidae (2) Metaplax indica H. Milne Edwards, 1852 10 Muddy, mangroves (mid, low)

Thalassograpsus harpax (Hilgendorf, 1892) 7 Rocky, cobble (high, mid)

Pinnotheridae (1) Arcotheres placunae (Hornell & Southwell, 1909) 4 + + Muddy-sandy/sandy flat (high)

a Total number of species in each family is given in parenthesesb Number of sites where species have been foundc Indicating first records from the Persian Gulfd Indicating first records from Qeshm IslandeHigh, mid and low included in parentheses refer to high-intertidal, mid-intertidal and low-intertidal, respectively

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ignored. Species which were found on more than four sitesand also species with ten occurrences at any site (if occurringat less than four sites) were included in the statistical analyses.

Results

Faunistic survey

A total of 131 decapod species were collected along the inter-tidal coast of Qeshm Island (Table 1), of which 11 species couldnot be exactly assigned reliably to hitherto known species. Ofthese, three alpheid shrimps, including two from the genusAlpheus Fabricius, 1798 and one from the genus AthanasLeach, 1814, are currently being described as new to science.Three caridean species from the families Hippolytidae andPalaemonidae are subject of an ongoing study. The remainingfive species in different suborders were unidentifiable, mostlydue to being small-sized juveniles.

Taxonomic composition

The 131 species collected belonged to five different infraorders,i.e. Caridea, Stenopodidea, Thalassinidea, Anomura andBrachyura. The highest number of species (74) or 56 % of allspecies belonged to the Brachyura (Fig. 2). The species withinBrachyura fell into 20 different families, of which Portunidaeand Pilumnidae, each with ten species (14 %), were the bestrepresented along the coast of Qeshm Island. Two familiesMacrophthalmidae (with species characteristics of muddy/sandy habitats) and Xanthidae (mostly occurring on stonysubstrate), each with seven species (11 %), had a consider-able diversity (Fig. 3). Seven families each were representedby one species only, including: Dromiidae, Matutidae,Hymenosomatidae, Eriphidae, Menippidae, Pseudoziidae andPinnotheridae. Of seven rare brachyuran families, two(Dromiidae and Menippidae) occurred at a single site (Table 1).

The infraorder Stenopodidea was represented by a singlespecies (Microprosthema validum). Caridea and Anomuracomprised 27 species (20 %) and 22 species (17 %), respec-tively, which were relatively well represented in the intertidalregion of Qeshm Island. Four families were recorded amongcaridean shrimps. Alpheidae had the highest species richness,comprising 62 % (16 species) of all collected shrimps, witheight species belonged to the well represented genus Alpheus.The families Hippolytidae and Palaemonidae each with fivespecies (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 andCallianassidae, had a relatively low species richness. Withinthe thalassinidian shrimps, the genus Upogebia with four spe-cies, namelyU. carinicauda,U. darwinii, U. pseudochelata and

Upogebia cf. savignyi, was well represented along the coast ofQeshm Island.

Anomuran crabs belonged to five different families, includ-ing: Hippidae, Coenobitidae, Diogenidae, Paguridae andPorcellanidae. The families Hippidae (Emerita holthuisi),Coenobitidae (Coenobita scaevola), and Paguridae (Paguruskulkarnii) were represented by a single species each. The ma-jority of the anomuran species belonged to the two familiesDiogenidae and Porcellanidae, with eight species (36 %) and11 species (50 %), respectively. These families had the highestspecies richness amongst Anomura. Five species of the hermitcrabs of the family Diogenidae belonged to the genusDiogenesDana, 1851, while the generaCalcinusDana, 1851,ClibanariusDana, 1852, and Areopaguristes Thomson, 1899, were repre-sentedwith one, one and two species, respectively. The crab-likeAnomura of the family Porcellanidae were particularly diverseat Qeshm Island, and were represented by five genera. Of these,Petrolisthes Stimpson, 1858 and Pisidia Leach, 1820 eachcomprised three species, and Pachycheles Stimpson, 1858 andPolyonyx Stimpson, 1858 each comprised two species, whileRaphidopus 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 atwhich a taxon was found) and distribution, it is apparent that thePilumnidae (33 of 40 sites), Sesarmidae (32 sites), Grapsidae (31sites), Alpheidae (29 sites) and Ocypodidae (28 sites) were themost widely distributed families around Qeshm Island.Diogenidae (26 sites), Coenobitidae (25 sites), Xanthidae (24sites), Macrophthalmidae (23 sites) Porcellanidae (20 sites), andPortunidae (18 sites) were also common. The familyCoenobitidae was represented by as single species (Coenobitascaevola), but occurred at more than half of the surveyed sites(71 %). Several abundant families, which were not as diverse interms of species richness, were represented by low numbers ofspecies (e.g. Sesarmidae with three species, Grapsidae andOcypodidae with four species each).

The most frequently collected decapod species wereMetopograpsus messor (28 sites), Nanosesarma sarii (27sites), 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 andmangrove) occurred along the intertidal zone of Qeshm Island.The majority of the study sites were a combination of these foursubstrates. Generally, the north coast was sedimentary, mostlysheltered sandy-mud (covering 38 % of the coastline) or muddy(covering 25 % of the coastline), together covering more thanhalf of the northern coastal line (Fig. 4).Mangroves (Tablmangalsystem together with the patchily distributed mangrove stands)were the second most common habitat along the north coastcovering about 33 % of the northern coastline. Rocky/cobble

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coastswere less common and showed amore patchy distribution,their coverage hardly reached 5 %. Pure sandy beaches weretotally lacking on the north coast, while sandy/rockywas the leastrepresented substrate (3 %) (Fig. 4). Over than half of the moreexposed south coast was sandy in combination with rocky andcobbles, which covered about 53 % of the southern coastline. Incontrast, pure sandy beaches were a less common habitat, cov-ering 18 % of the southern coastline. Rocky shores were thesecond most common coastal habitat (22 %), generallyfound in association with cobble. Sedimentary flats oc-curred only in the central part of the south coast (betweenShib Deraz and Nagasheh), where a small stand of plantedmangrove was present (Fig. 4).

Multivariate cluster analyses of the 40 sites with referenceto their species composition (Fig. 5), separated the northernsites from the southern ones. Group A includes mostly north-ern coastal sites plus one site (station 11) from the south coastcomprised sedimentary habitats with muddy, sandy-mud ormuddy-sand substrates. Group A is further divided into twodistinct subgroups: subgroup I includes sedimentary sites withmuch 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 sub-group. 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 closelyrelated to the diverse southern sites and was included withthem in the same group. Amongst these seven diverse sites,four sites were considerably decapod crustaceans diverse [sta-tion 1 (43 species), station 5 (36 species), station 21 (39species), and station 24 (34 species)]. Subgroup IV (Fig. 5)of group B contains sites with rocky/sandy substrate. Withinthis subgroup two smaller groups are distinguishable, a smallgroup (with 6 sites) comprising sandy substrate, and a largergroup (with 13 sites) comprising sandy/rocky substrate.

Main habitats and decapod communities

Mud flat

The north coast of Qeshm Island lays in an extremely shel-tered location without wave action and with very limited tidalenergy, resulting in a wide, gently sloping, muddy-sandy/sandy-mud flat. Sedimentary flats of the northern coastlinewere generally soft with a high proportion of coarse particles(sand and shell fragments) in the high-intertidal zone, usuallyfringed with narrow bands of sandy beach towards thesupralittoral zone. Different intertidal zones of extensive

sedimentary flats are very difficult to determine according tospring and neep tide levels. But the zonation of the muddy flatwas clearly defined by the vertical distribution of decapodspecies belonging to the superfamily Ocypodoidea (Table 2).Burrows ofNasima dotilliformiswere characteristic of the high-intertidal zone, extending normally landwards into thesupralittoral zone, where they burrowed in relatively consolidat-ed substrate among salt-marsh plants. In the high intertidal zone,Uca sindensis was sympatric with N. dotilliformis, which wasalso sympatric with (or sometimes replaced by) Uca iranica inregions with high proportions of coarse particles. Two small-sized species, Ilyoplax stevensi andOpusia indica overlapped intheir occurrences with N. dotilliformis in its lower limit in thehigh-intertidal zone. High-intertidal zones were characterised bymuddy-sandmixedwith shell fragments patchily fringing with athin veneer of limestones in their higher limit. Such zones weredominated by a newly recorded thalassinid shrimp, Upogebiacarinicauda. Chimney-shaped structures of burrows of this mudshrimp with an average height of 4 cm were characteristic in thesummer 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 thishabitat. The mid-intertidal zones were generally marked byburrows of a large macrophthalmid crab (Macrophthalmusdentipes) which tended to burrow in loose sandy-mud sub-strates. The most distinct associate of M. dentipes was themudskipper (Periophthlamus waltoni). Awide variety of deca-pod crustaceans was found sympatric with M. dentipes in themid-intertidal zone (see Table 2).

The low-intertidal zones of mud flats could be determined bythe occurrence of Macrophthalmus depressus. This medium-sized macrophthalmid crab was usually found in a narrow bandof low-intertidal zones, together with some common decapodcrustaceans of the mid-intertidal zone, mainlyMetaplax indica,Eurycarcinus orientalis, Eurycarcinus integrifrons and Opusiaindica, Alpheus lobidens,Michaelcallianassa indica (Table 2).

Mangroves

The Qeshm mangal ecosystem “Harra of Tabl” is the largestmangrove 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 to4–6 m in height and located in a fully sheltered embayment inthe northwestern coast of Qeshm. The muddy banks of the waterchannels in most areas of the Qeshm mangal system wereoccupied by oyster. Presence of the dead shells promote theestablishment of mangroves via the trapping of sediments andhosts 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 themuddy banks of the mangroves (low-intertidal zone), the most

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common decapod species were Macrophthalmus depressus,Metaplax indica, Eurycarcinus orientalis, Ilyograpsusrhizophorae, Opusia indica, Alpheus lobidens and Manningisarabicum (Fig. 6). Among the pneumatophores and trunk zones,usually impenetrable, was a habitat for the most commongrapsoid crabs, Parasesarma persicum and M. messor, withjuveniles distributed on the sediment surface, while adults weregenerally found in the canopy zone of trees. The rare speciesMetopograpsus thukuharwas found at only two sites in syntopywith M. messor. Other decapod species found in this zone arepresented in Table 1. The landward edge of the Qeshm mangal

stand was usually a wide band of muddy flats with sparsehalophytic plants, mainly Salicornia sinus-persica, Arthroc-nemum macrostachyum and Halocnemum strobilaceum. Thiszone was dominated by two ocypodoid crabs, Nasimadotilliformis and Uca sindensis. Remarkable burrow structuresof N. dotilliformis were one of the prominent features of thiszone, which extended to the high-intertidal zone.

There were also patchy mangrove swamps which were arti-ficially planted alongmost parts of the north coast.Most of thesefringing artificial mangroves are exposed to the open sea; themajority of planted mangrove bushes had not established or had

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)

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developed very poorly, most likely due to the loose nature ofmuddy-sand/sandy substrate, and partly perhaps because oftheir immediate exposure to the sea. The single best developedmangrove stand occurred at Holor coast. The substrate wasdominantly muddy-sand, trees reaching up to 2m in height, butnot densely populated. This was inhabited by some mangal-characteristic decapod crustaceans, common fiddler crab, Ucairanica in the N. dotilliformis zone, where it replaced U.sindensis in relatively coarser sediments.

On the southern coast of the island, a small stand ofA.marinaoccurred between Band-Chapi and Naghasheh (Fig. 4), wheremangrove shrubs were planted along banks of artificial tidalchannels running parallel to the sea coast in the sandy ormuddy-sand flat. Sea grasses, dominantly Halodule uninervis,were abundant in the sheltered unconsolidated sandy or muddy-sand substrata.

South coast mangroves had some of the brachyuran crabsthat are also characteristic for the well-developed mangroves ofthe north coast. Avery small population of the mud crab, Scyllaserrata, with medium-sized specimens was found in thismangal ecosystem. Apparently this species has establisheditself in this narrow habitat quite recently. A further rare specieswas the fiddler crab Uca inversa, which occurred in a smallpatch on consolidated substrate along the landward edge.

Sandy beaches

Most of the sandy beaches were found, usually together withrocky patches, along the south coast of Qeshm Island. Pure

sandy beaches were a less common habitat covering about18% of the southern coastline (Fig. 4). To the south-east of theisland, most beaches had a relatively steep profile, with thenarrow intertidal zone sloping down, in most parts, to a rockysubtidal. Rocky structures in the shallow subtidal or lowerintertidal zone of the sandy beaches are exposed only duringlow spring tides in some regions. Supralittoral zones of thesandy beaches were usually characterised by sandy dune belts,vegetated mainly with halophytes (e.g. Suaeda maritima, andSeidlitzia rosmarinus), and bordered mostly along their land-ward fringe with Acacia spp.

The typical decapod crustacean inhabitant of this zone wasa semi-terrestrial anomuran known as the land hermit crab,Coenobita scaevola (Table 3). Footprints of this nocturnallyactive hermit were a main feature of this supralittoral zone,while all collected specimens were found buried in the shadedarea of coastal vegetation during day-time, emerging usuallyat night to scavenge in the intertidal zone. This species wasoften seen crossing the coastal road, and its footprints werefound several hundred metres further landward. The high-intertidal zone of such a steep sandy beach was dominatedby the only ghost crab of the Persian Gulf, Ocypoderotundata. Conical small pyramids of male animals, in partic-ular, were seen during the breeding season (e.g. with a densityof two burrows per 1.0 m2 counted in May in Namakdan).Further down on a steeper slope, the suspension feeding molecrab Emerita holthuisi inhabited a swash active zone. Similarexposed sandy beaches, favourably populated by E. holthuisi,were fully lacking on the north coast. This species was

Fig. 3 Percentages of differentfamilies of the Brachyuracollected from Qeshm Island.Seven families, each with onespecies, are included in Others

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collected only at four sites along the eastern part of the southcoast in a dense population of 25 specimens per 0.25 m2. Mid-intertidal continued to low intertidal and shallow subtidal zoneswith a gentle profile were favourite habitats of a small leucosiidcrab Ryphila cancellus. Matuta victor is a large-sizedbrachyuran crab usually found sympatric with R. cancellus,but distributed slightly deeper than the latter. Two small-sizedbrachyuran species of the family Dotillidae, namely Dotillablanfordi and Scopimera crabicaudawere common inhabitantsof sandy flats of seaward fringe of mangroves along the southcoast. Only the pinnotherid species Arcotheres placunae livingin bivalve molluscs Amiantis umbonella also occurred in suchsheltered sandy flats. The small-sized hermit crab Diogenesavarus usually occurred on all sandy beaches, being particu-larly abundant on sandy flats.

Rocky shore

Rocky shores were merely found along the south coast ofQeshm Island but not as long stretches, rather interruptingsandy beaches in the south-eastern half and cobble-shoresmainly in the southwestern half. Nevertheless, rocky shorescovered about 18 % of the southern coastline (Fig. 4). Very

small patches of rocky substrate occurred in the northeastof the island. These consisted mainly of a thin veneer ofsedimentary limestone rocks, and often flourished with typicalrocky fauna (e.g. in Kuweii). The coastal rocks around QeshmIsland were mainly sedimentary (limestone), composed ofmarine organisms’ fragments, mainly those of Foraminifera,echinoderms and mollusc shells. Rocky substrates were oftencombined with dead corals on the south coast, while no signsof dead corals were found along the north coast. An interestinglimestone rocky substrate with coquina rocks occurred in thenorthwestern part of the island on the Basaeedu headland.Rocky shores of the south coast mainly included rocky cliffsin 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 distri-bution are shown in Table 3. Common species found in therocky beds of the Shib Deraz were Nanosesarma sarii,Leptodius exaratus, Pilumnopeus convexus, Eriphia smithi,Petrolisthes rufescens, and Petrolisthes leptocheles. A narrowband (5-10 m wide) of about a 1-km long rocky bed stretchedfrom about 2 km of Qeshm city westwards along the southcoast. This rocky bed was a very poor habitat for the crusta-cean species. The herbivorous crab Grapsus albolineatus

Fig. 4 Distribution and extent of different habitats along the coastline of Qeshm Island

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occurred in the high-intertidal zones spreading towards themid-intertidal zones of nearly all rocky shores, where thishighly active crab was sympatric withMetopograpsus messor.The scavenger eriphid crab, E. smithi was found exclusivelyin crevices, slightly lower on the shore than G. albolineatus.

The most common anomuran crab Clibanarius signatus wasgenerally present in shaded areas. Grapsus granulosus was aless common species (found at five sites), recorded for the firsttime in the Persian Gulf and coexisting withG. albolineatus inthe mid-intertidal zone.

Table 2 Vertical distribution patterns of the common species of decapod crustaceans in muddy and sandy/muddy shores on Qeshm Island

Fig. 5 Hierarchical clusteranalysis using the SquaredEuclidean distance performedwith Ward’s method in order togroup the sites with similarspecies compositions. Cluster Aincludes sites on the north coastwith mangrove, muddy, sandy-mud or muddy-sand substrate.Cluster B includes all south coastsites together with some from thenorth coast, which were mostlycombination structure of sandy/rocky or exclusively rocky orsandy

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Rocky shores at the south coast, in most cases, supportedgrowth of the brown algae Sargassum sp. in the sublittoralzone. These algae reached up to 60 cm of height in somelocations, particularly during the cold season, and were foundwashed-up on the shore. On the contrary, rocks of the northerncoast supported the growth of a green algal tuft. Sedimentaryrocks were densely covered with a filamentous green lawn(Table 4).

Cobble beaches

Cobble beaches, composed of small limestone rocks, occurredscattered only along the south coast. This kind of habitat wasusually found in the proximity of rocky shores and sometimes incombination with sandy substrate. Such a complex habitat,usually trapping sediments between loose rocks and supportingalgal cover, provided diverse microhabitats allowing differentspecies to occur together in the same locality. Consequently, thisenvironment was the most diverse among the coastal habitatsaround the island. High-intertidal zones were characterised ei-ther by rocks (e.g. Zeytun Park Beach) or by sandy beaches(e.g. Dustku). Cobbles were mainly present in the mid-intertidalzone of the shore, hosting many of the species also occurring atthe rocky shores and rock beds, as well as a few on the sandyflat. The common species of this zone, whichweremainly foundunder stones, are Clibanarius signatus, Leptodius exaratus,Nanosesarma sarii, Petrolisthes leptocheles and P. rufescens.The eriphid species Eriphia smithi and different alpheid shrimpswere also characteristic of this habitat. Several different speciesof the families Xanthidae, Pilumnidae and Upogebiidae were

present as well. Common species of the cobble-beaches andtheir vertical distribution are presented in Table 3, together withrocky shore species.

Discussion

Species richness

The faunistic results of the present survey illustrate the scar-city of the information on the taxonomy and assemblagestructure of the decapod crustaceans along the intertidal hab-itats of the Persian Gulf, particularly along the eastern coast(Iranian coast). Of 131 decapod crustaceans collected, about47 % (61 species) were new records for Qeshm Island. About14% (18 species) of all species collectedwere recorded for thefirst time from the Persian Gulf. These new records includecaridean shrimps:Alpheus edamensisDeMan, 1888,A.macro-dactylus Ortmann, 1890, A. paracrinitusMiers, 1881, AthanasdjiboutensisCoutiere, 1897, Salmoneus gracilipesMiya, 1972,Synalpheus gracilirostris De Man, 1910 and S. hastilicrassusCoutiere, 1905, Cuapetes elegans (Paulson, 1895), Cuapetesgrandis 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 loimicolaSankolli, 1965; Brachyura: Heteropanope glabra (Stimpson,1858), Arcotheres placunae (Hornell and Southwell, 1909).Nine species (7 %) are new to the science, of which four had

Fig. 6 Schematic diagram of the main mangrove system along north-western Qeshm Island in Tabl area. Most common decapod crustacean species areshown together with their relative vertical distribution

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already been described from among the material of the presentsurvey, including: Nanosesarma sarii Naderloo and Türkay,2009, Parasesarma persicum Naderloo and Schubart, 2010,Athanas iranicus Anker et al., 2010 and Macrophthalmussinuspersici Naderloo and Türkay, 2011.

The island surprisingly includes the highest number ofdecapod species known from the Iranian coast, where 150species were recorded (Naderloo and Türkay 2012). Of these,87 % were recorded from Qeshm Island. In terms of speciesnumbers, Qeshm Island is comparable with other regions inthe Indian Ocean. For instance, Socotra Island, located in thetropical Indian Ocean with a total area of 3.796 km2, has 130recorded decapod species (Simões et al. 2001), while QeshmIsland (1,491 km2) with slightly larger than half the Socotraarea has roughly the same number (131) of the decapod

species. This occurs in spite of the generally harsh environ-mental conditions of the Persian Gulf, i.e. high salinity andfluctuating 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 PersianGulf results in an inflow of fresh low-salinity oceanic watersthrough the Gulf of Oman and this makes this island verydifferent from the other parts of the Gulf. The high diversity ofthe island was mainly a result of the high decapod diversity instony habitats along the south coast, whereas the muddysubstrate of the northern coast was about as diverse as othermuddy shores within the Gulf (Apel 1994a, 1996; Clayton1986; Naderloo and Türkay 2012). This is confirmed bycomparing the number of decapod species recorded from arocky/cobble site in Qeshm Island (e.g. 43 species at station

Table 3 Vertical distribution patterns of the common species of decapod crustaceans in exposed sandy shores of Qeshm Island

Table 4 Vertical distribution patterns of the common species of decapod crustaceans in rocky and rocky/cobble shores of Qeshm Island

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1), with similar sites along the mainland coast of Iran (37species from Bushehr, 32 species from Moghdan, Naderlooand Türkay 2012).

Intertidal habitat and associated decapod fauna

The intertidal area of Qeshm Island includes a considerableecosystem diversity, varying from extensive mangroves, gent-ly sloping sedimentary flats, exposed sandy beaches tocobble/rocky shore. Different habitats with varying physicalcharacteristics favour different species communities; hencethe north coast of the island with a predominantly sedimentarystructure hosted completely different assemblages of decapodcrustaceans compared with the south coast. This was shownby results from the cluster analysis (Fig. 5). The sites clusteredaccording to the nature of the physical structures, which werefavoured by different species assemblages. The high decapodspecies diversity of Qeshm Island is thus a consequence of itshigh ecosystem diversity.

The highest species richness was present in stony habitats, inparticular in the mixed substrates of rocky/cobble, rather than issofter sedimentary habitats. This is consistent with results ofother local studies (e.g. Basson et al. 1977; Titgen 1982) andgenerally supported by findings of similar surveys from otherparts of the world (Raffaelli and Hawkins 1996; Simões et al.2001). The high richness is most likely due to the availability ofdifferent microenvironments in stony substrates. Of 131 identi-fied decapod species, 88 (67 %) were found in stony shores(rocky, cobble, rocky/cobble), 76 of which exclusively inhabitedrocky/cobble (Table 1) in the mid-intertidal zone. Four morecommon species of the rocky/cobble habitat found to be alsocommon along muddy shores were: Alpheus lobidens,Metopograpsus messor, Nanosesarma sarii and Pilumnopeusconvexus. Four fairly common species in stony habitats includedMacrophthalmus sinuspersici, Nanosesarma jousseaumei, N.sarii, Thalassograpsus harpax. These species are only knownfrom the Iranian coast of the Gulf and not recorded from theArabian side, despite of the quite extensive samplings conductedalong 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 % (30species) of all collected decapod crustaceans. This is less thanhalf of the rocky/cobble species richness. The brachyurancommunity of the mud flat and the vertical distribution ofdifferent crab species is comparable with those recorded fromthe northern Persian Gulf coast of Kuwait (Clayton 1986),Saudi Arabia (Apel 1994a, 1996) and from other Iranian coastsof the Persian Gulf (Naderloo and Türkay 2012). The onlybrachyuran species which is dominant on the high intertidalzone of mud flats on the northern part of the Gulf (Iran, Iraq andKuwait) is Leptochryseus kuwaitensis (Al-Khayat and Jones1996; Ng et al. 2009; Naderloo and Türkay 2012). This species

is completely missing along the southern coast of the PersianGulf and replaced by Nasima dotilliformis. The latter has beenrecorded 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 diver-sity is relatively low in this habitat, the abundance and biomassis considerably high. Some species (e.g. Uca sindensis, Ucairanica andMacrophthalmus dentipes) had the highest density,which can be attributed to the high content of organic materialin the muddy substrate.

The mangroves of Qeshm Island were not as diverse regard-ing the decapod crustaceans as other intertidal habitats, but stillmanymore species have been recorded from here comparedwithother parts of the Persian Gulf. Of the 23 brachyuran crab speciesrecorded from Qeshm Island, 13 and 14 species have beenrecorded from mangroves in Abu Dhabi (Cooper 1997) andQatar (Al-Khayat and Jones 1999), respectively. The mangrovehabitat shared many specific decapod assemblages with the mudflat. Mangroves ecologically are important because of their highprimary productivity, importance as nursery grounds for shrimpsand fish larvae and as providing feeding ground for manyresident and migratory birds. Two species, Metopograpsusthukuhar and Scylla serrata were exclusively found in man-groves.Metopograpsus messor andParasesarma persicumwerethe two most common crabs amongst the trunks and pneumato-phores in the study area, whereas in east African mangroves twospeciesMetopograpsus thukuhar and Perisesarma guttatum arethe most common and ecologically important species (Vanniniand Valmori 1981; Gillikin and Schubart 2004). An interestingrecord of the present study is the giant mud crab Scylla serrata,which is commercially important. This species is widely distrib-uted in the Indo-West Pacific and has already been recorded fromthe 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 Khor-khalasi (R.N. pers. obs.). Here it is the largest and most commonspecies and is used as a food source by local people. Scyllaserrata has recently been recorded from the southeast PersianGulf in Ras al-Khaimah (UAE) by Hogarth and Beech (2001).This species is thought to have been very common in thepast when extensive mangroves were widely distributedalong the coast of the Persian Gulf (Hogarth and Beech2001). Numbers declined following the disappearance ofmangroves from most parts of the Persian Gulf, but thereare sign of recovery. We found this commercially importantcrab in the planted mangroves along the south coast(Naghasheh), while, despite extensive sampling survey,no individual was recorded in the largest mangal systemalong the north coast (Tabl).

Sandy beaches are the least diverse habitat within the PersianGulf (Jones 1986b) and all over the world (Raffaelli and

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Hawkins 1996). The Decapod community of the sandy beach ofQeshm Island is fully comparable with those of the coast ofKuwait (Jones 1986b) and Saudi Arabia (Basson et al. 1977).In comparison to the adjacent regions, the only genus whichappears 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 ofPakistan (Türkay et al. 1996), four species were recorded fromSocotra Islands (Türkay et al. 1996; Simões et al. 2001) and twospecies from the Red Sea (Türkay et al. 1996). The terrestrialhermit crab Coenobita scaevola is the only hermit crab occur-ring in the supralittoral zone of sandy beaches in the PersianGulf andGulf of Oman. This hermit crabwas a common speciesin 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, especiallywhen comparing sites with different physical and topographicalcharacteristics (English et al. 1997). Many workers believe thatthe zonation should be based on biological elements withdetermined species compositions and distribution of the keyspecies (e.g. Raffaelli and Hawkins 1996). As this study wasthe first to determine the species composition of decapodcrustaceans and their vertical distribution along the coast ofQeshm Island, it was only possible to use water movementto define different zones. Therefore, we advocate that inusing the present data in comparison with data from otherregions, differences in tidal regimes and regional topographyas well as the physical nature of the shores must be takeninto consideration.

The vertical distribution of the different brachyuran speciesin the mud flats of Qeshm Island is relatively similar with thatrecorded from Kuwait (Clayton 1986), Saudi Arabia (Apel1994a, 1996) and Iran mainland (Naderloo and Türkay 2012).Unfortunately, there is no documentation on the intertidalrocky/cobble crustacean assemblage from other parts of thePersian Gulf, except a brief study of Jones (1986b) fromKuwait that deals with the most common rocky fauna andflora and this includes few crustacean decapod species.

Intertidal zone and biodiversity conservation

Monitoring and documenting the biodiversity along the coastsof the Persian Gulf is very important since the area is underextreme pressure due to several environmental and anthropo-genic factors, including oil-related industries, coastal devel-opments, urban sewage, developing tourist industry, immigra-tion of invasive species and, most importantly, globalwarming (e.g. Facey 2008; Khan 2008; Krupp andAbuzinada 2008).

Two recent reviews by Sheppard et al. (2010) and by Saleet al. (2011) clearly address the environmental problems of thePersian Gulf and the necessity for an integrative conservationprogram. Mangroves and reefs of the Persian Gulf are undergreatest threat (Sheppard et al. 2010). Qeshm Island,possessing the largest mangal ecosystem with about a 6,800-ha area within the Gulf, is of major importance for conserva-tion of this environment (Spalding et al. 2010). In the past,mangroves were used as building material, firewood andforage for livestock by traditional users, which is possiblythe main destructive reason for this ecosystem in the world(Saenger et al. 1983). Currently, only the leaves and youngbranches of mangroves are used as livestock fodder by ruralpeople. Camels are sometimes also being seen foragingamong the arid zones of the system, which locally seem tohave a small destructive effect on the mangroves. Illegalshrimp catching is occasionally seen in the Tabl mangalecosystem, which seems to have the second rank indestructing the mangroves’ fauna and flora. Further, Tablmangrove forest is now the most important recreational sitearound the island, and the ecosystem is being opened up totourism, which needs an integrative tourism management.Nonetheless, Qeshm mangroves are in relatively good condi-tion compared with the mangroves of the other Persian Gulfstates like Saudi Arabia and UAE (Spalding et al. 2010).

No published record is available on the oil-related pol-lution of intertidal habitats of Qeshm Island. Consideringthe location of the island in the Strait of Hormoz, which is amain oil transporting passage, it has the potential to besubject to a heavy oil spill. There are some pollution im-pacts along the northern coast due to its vicinity to theBandar-Abbas port.

In general, anthropogenic effects presently cannot be con-sidered as a big threat to the intertidal habitats of the island,mainly because of the small rural population inhabiting thecoastal area. Developing the free economic zone with sub-sequent impacts on the island coastal environments is by farthe greatest threat to the region. Clear evidence for such apossible threat is seen in the coastal situation of Dubai(Sheppard et al. 2010).

Acknowledgments We would like to thank Dr. Ranjbaran (School ofGeology, University of Tehran) for helping with identification of thecoastal rocks and boulders. The help of Mr. Abbas Kazemi (Universityof Tehran) in field works is highly acknowledged. We are grateful toMoritz Sonnewald (Senckenberg Museum, Frankfurt) for his help withstatistical analysis and workingwith Brodgar software. The English of thepaper was checked by “GRAGE Language Service” of University ofFrankfurt, which is especially acknowledged.

This biodiversity project was mainly funded by Iran National ScienceFoundation (INSF) and University of Tehran. Their financial help forfacilitating the biodiversity project (including the present one) in the Iraniancoast of the Persian Gulf is highly appreciated. We are indebted to DAAD(German Academic Exchange Service) for financial support in the form ofa PhD Scholarship for R. Naderloo facilitated extensive data analysis.

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