Chrysymenia tigillis sp. nov. (Rhodymeniales, Rhodophyta) from the Sultanate of Oman, with a census of currently recognized species in the genus Chrysymenia

Chrysymenia tigillis sp. nov. is described on the basis of a few specimens collected from Dhofar, Sultanate of Oman. The new species is known only from southern Oman, a region of the northern Arabian Sea that is strongly impacted by the upwelling from the summertime monsoon. It is distinguished from other species of the genus by the simple nature of the blades, their dimensions (to 75 cm in length and to 21 cm in width), the rough, bumpy surface of the blades, and the presence of internal struts connecting the inner sides of the blades. A census of the currently recognized species in the genus Chrysymenia is provided.     

Turbinaria foliosasp. nov. (Fucales, Phaeophyceae) from the Sultanate of Oman, with a census of currently recognized species in the genus Turbinaria

Turbinaria foliosasp. nov. is described on the basis of several collections made from Dhofar, Sultanate of Oman. The new species is known only from southern Oman, a region of the northern Arabian Sea that is strongly impacted by the upwelling from the summertime monsoon. It is distinguished from other species of the genus by the shape of the leaves and by their loose, non‐congested arrangement. Air vesicles, embedded in the leaves, may be present, but more often are absent. A census of the currently recognized species in the genus Turbinaria is provided. Reference is made to Sargassum turbinarioides and Sargassum turbinati‐folium. These species possess discrete blades and spherical vesicles and thus conform to Sargassum.     

Further connections between the benthic marine algal floras of the northern Arabian Sea and Japan

Identifications made on some recent collections of benthic marine algae from the Sultanate of Oman, northern Arabian Sea, have revealed several new records for this region. A total of eight species, representing five species of Rhodophyceae, two species of Phaeophyceae, and a single species each of Chlorophyceae, are newly reported for Oman. Of particular interest is the fact that these species represent new records for the Indian Ocean, some of which had previously been known from Japan and environs.     

Stock differentiation of the greater lizardfish Saurida tumbil (Teleostei: Synodontidae) collected along the western coast of the Arabian Gulf and Sea of Oman using meristic characters

Meristic variation among stocks of greater lizardfish Saurida tumbil through the western coasts of the Arabian Gulf and Sea of Oman was examined using meristic characters. Statistical analysis of meristic traits proposed that there is constrained migration of populations of greater lizardfish along the western coast of the Arabian Gulf and Sea of Oman. Overlapping of the two samples from the northern part of the Arabian Gulf (Iraq-Kuwait waters), three samples from the middle region of the Arabian Gulf (Bahrain-Qatar-Saudi Arabia) and two samples from the southern part of the Arabian Gulf/Sea of Oman (United Arab Emirates–Sultanate of Oman) suggested that there are three self-recruiting populations in the studied area. Inspection of the role of each meristic trait variable to Canonical discriminant analysis showed that changes among samples appeared to be linked with the pattern of distribution of water temperature and configuration of current in both the Arabian Gulf and Sea of Oman areas.     

Sea cucumbers of the Arabian Peninsula and Iran – A review of historical and current research trends

Sea cucumbers are benthic marine invertebrates with immense ecological and commercial value. Processed sea cucumbers known as “Beche-de-mer” are a delicacy in southeast Asian countries with an ever-increasing demand depleting wild stocks on a global scale. Aquaculture techniques are well developed for commercially important species (e.g. Holothuria scabra) to aid in conservation and trade. In the Arabian Peninsula and Iran, where the major land mass is surrounded by marginal seas (Arabian Gulf, Gulf of Oman, Arabian Sea, Gulf of Aden, and Red Sea), studies on sea cucumbers are rather limited and its economic value is underestimated. Historical and current research trends indicate impoverished diversity (82 species) due to environmental extremes. Artisanal fisheries exist for the sea cucumbers of Iran, Oman, and Saudi Arabia, with Yemen and United Arab Emirates (UAE) playing a key role in collection and export to Asian countries. Stock assessment and data on export indicates depletion of natural stocks in Saudi Arabia and Oman. Aquaculture trials of high value species (H. scabra) were successful in Saudi Arabia, Oman and Iran with prospects for further expansion. Research on ecotoxicological properties and bioactive substances conducted in Iran demonstrates an immense research potential. Molecular phylogeny, biology, use in bioremediation, and characterisation of bioactive compounds were identified as potential gaps in research. Expanding aquaculture operations could revive exports and recuperate damaged stocks through sea ranching. Furthermore, regional cooperation, networking, training, and capacity building could help fill the gaps in sea cucumber research, which will aid in its effective conservation and management.     

First geolocator tracks of Swedish red‐necked phalaropes reveal the Scandinavia‐Arabian Sea connection

We studied migration and wintering patterns of a wader with a pelagic lifestyle during the non‐breeding period, the red‐necked phalarope Phalaropus lobatus. Using light‐level geolocation, we obtained three full annual tracks and one autumn migration track of male red‐necked phalaropes caught during breeding in Scandinavia. These tracks confirmed expectations that individuals from the Scandinavian population winter in the Arabian Sea. Migration was accomplished in two to four migration leaps, staging for a few days in the Gulf of Finland (autumn) or the southern Baltic Sea (spring) and for up to a month in or near the Black and Caspian Sea (autumn and spring). In addition, travel speeds suggested that only the flights between the Baltic and Black/Caspian Sea are non‐stop, and thus the birds seem to make additional short stops during the other flights. Stopover time in the Black/Caspian Sea is only 8–10 d in spring but up to 36 d in autumn, which is longer than expected if only used for pre‐migratory fattening to cover the ca 2000 km to the Gulf of Oman. After entering the Arabian Sea via the Gulf of Oman, birds dispersed over the entire presumed winter range. Winter movements appear to correspond to the spatio‐temporal patterns in primary production linked to seasonally changing monsoon winds. These are not only the first tracks of Scandinavian red‐necked phalaropes, but also the first seabird tracks in the Arabian Sea, one of the most productive and dynamic marine areas on the planet.     

TEMPERATURE THRESHOLD AS A BIOGEOGRAPHIC BARRIER IN NORTHERN INDIAN OCEAN MACROALGAE1

The most eastern point of the Arabian Peninsula, Ras Al Hadd, marks the boundary between the Arabian Sea and the Gulf of Oman. This geographic landmark coincides with an abrupt floristic turnover, probably one of the sharpest biotic transitions known in marine biogeography. The floras of different Arabian localities across this floristic break were compared using macrophyte distribution data throughout the Indian Ocean and seasonal sea‐surface temperature (SST) data. The localities from the Arabian Gulf and Gulf of Oman differ significantly from those of the Arabian Sea based on their species richness, species composition, average distribution range per species, general temperature affinity of the composing species, and seasonal temperature data of the coastal waters. Pooling the temperature data into two groups (SST_3avg, average SST of the three warmest seasons; SST_min, minimum of the seasonal SSTs) revealed a temperature limit at 28°C using both the temperature affinity data of the floras and the seasonal temperatures recorded for the specific Arabian localities, which significantly separates the Arabian Sea from localities of both Gulfs. Finally, SST data of the Indian Ocean were analyzed using this upper temperature threshold of macrophytes at 28°C and the lower temperature limit of corals at 25°C, revealing general macrophyte diversity patterns.     

Sea cucumbers of the Arabian Peninsula and Iran – A review of historical and current research trends

Sea cucumbers are benthic marine invertebrates with immense ecological and commercial value. Processed sea cucumbers known as “Beche-de-mer” are a delicacy in southeast Asian countries with an ever-increasing demand depleting wild stocks on a global scale. Aquaculture techniques are well developed for commercially important species (e.g. Holothuria scabra) to aid in conservation and trade. In the Arabian Peninsula and Iran, where the major land mass is surrounded by marginal seas (Arabian Gulf, Gulf of Oman, Arabian Sea, Gulf of Aden, and Red Sea), studies on sea cucumbers are rather limited and its economic value is underestimated. Historical and current research trends indicate impoverished diversity (82 species) due to environmental extremes. Artisanal fisheries exist for the sea cucumbers of Iran, Oman, and Saudi Arabia, with Yemen and United Arab Emirates (UAE) playing a key role in collection and export to Asian countries. Stock assessment and data on export indicates depletion of natural stocks in Saudi Arabia and Oman. Aquaculture trials of high value species (H. scabra) were successful in Saudi Arabia, Oman and Iran with prospects for further expansion. Research on ecotoxicological properties and bioactive substances conducted in Iran demonstrates an immense research potential. Molecular phylogeny, biology, use in bioremediation, and characterisation of bioactive compounds were identified as potential gaps in research. Expanding aquaculture operations could revive exports and recuperate damaged stocks through sea ranching. Furthermore, regional cooperation, networking, training, and capacity building could help fill the gaps in sea cucumber research, which will aid in its effective conservation and management.     

Scales and otoliths as identity cards of the Indian oil sardine Sardinella longiceps (Teleostei: Clupeiformes) populations: Ultrastructure and ornamentation characteristics using light and scanning electron microscopy

Scale and otolith morphology and morphometry of Indian oil sardine Sardinella longiceps (Clupeidae) were investigated and described using light and scanning electron microscopy from eight different body regions for scales and the right and left otoliths. Scales of the Indian oil sardine show general characteristics of the other studied clupeids sand that are easily distinguishable from other fish groups, by having striae in the posterior field. The studied cycloid scales of S. longiceps were classified into three types based on the overall shape including circular (e.g. true circular and cordate), pentagonal and quadrilateral in the different body regions. The circular shape was the most common shape (87.5%), while the quadrilateral and pentagonal forms constituted 6.25% each. The results also showed that the relative scale size (J-index) plays a desirable contribution in separating the examined populations. The results showed that the mean (or relative) scale size for all the eight regions in the Oman Sea population is larger than the Arabian Sea population. Also, another scale variable, the scale shape index (Si index), demonstrated variation (a mean of 0.86 to 1.1) in different regions of both populations from the Oman and Arabian Seas. Interestingly, here, we found that scale characters of S. longiceps not only differ from its other congeneric species, but also differ in the populations from both sides of the Oman Sea (Iran and Oman) and the Arabian Sea. It shows a positive signal for the presence of different taxonomic and management unit in the Oman and Arabian Seas. The idea should be approved by using integrated molecular and morphological traits. The otolith morphology of S. longiceps from the Oman and Arabian Seas was more conservative than the scales, which can be due to its function actin primarily as a balance organ and also enhancing hearing. The overall shape of S. longiceps otolith was lanceolate, with an elongated morphology and a well-developed rostrum, an ostial sulcus acusticus that opens to the anterior/ dorsal margin. These morphological characters are also found in the Iranian population of S. longiceps. However, otolith displayed variation in biometric parameters among two populations and left and right otoliths and the RRL parameter were important characters to discriminate the Oman and Arabian Sea populations. Thus, the structural/biometrical variability of the otoliths may be used for population distinctness, especially in water bodies with various environmental factors, and the otolith has turned out to be a useful tool to track the life history of teleostean fishes in environments with physicochemical gradients.     

Comparative phylogeography of reef fishes from the Gulf of Aden to the Arabian Sea reveals two cryptic lineages

The Arabian Sea is a heterogeneous region with high coral cover and warm stable conditions at the western end (Djibouti), in contrast to sparse coral cover, cooler temperatures, and upwelling at the eastern end (southern Oman). We tested for barriers to dispersal across this region (including the Gulf of Aden and Gulf of Oman), using mitochondrial DNA surveys of 11 reef fishes. Study species included seven taxa from six families with broad distributions across the Indo-Pacific and four species restricted to the Arabian Sea (and adjacent areas). Nine species showed no significant genetic partitions, indicating connectivity among contrasting environments spread across 2000 km. One butterflyfish (Chaetodon melannotus) and a snapper (Lutjanus kasmira) showed phylogenetic divergences of d = 0.008 and 0.048, respectively, possibly indicating cryptic species within these broadly distributed taxa. These genetic partitions at the western periphery of the Indo-Pacific reflect similar partitions recently discovered at the eastern periphery of the Indo-Pacific (the Hawaiian and the Marquesan Archipelagos), indicating that these disjunctive habitats at the ends of the range may serve as evolutionary incubators for coral reef organisms.

     

Chloroplast DNA variation in the shrub Justicia areysiana (Acanthaceae) endemic to the monsoon affected coastal mountains of the southern Arabian Peninsula

The monsoon affected mountains of the southern Arabian Peninsula harbour in climatically favoured refugia vegetation elements of palaeo-African origin. To understand better the temporal and spatial differentiation of these refugia, chloroplast variation in Justicia areysiana Deflers (Acanthaceae), a shrub species endemic to the Yemeni and Omani mountains close to the Arabian Sea, was studied using PCR-RFLP and chloroplast microsatellite diversity. Eleven haplotypes were characterized and show a distinct geographical distribution pattern with a deep split between populations from south Yemeni fog oases and those from Hawf Mountains/Dhofar region in east Yemen and south Oman. Very limited haplotype diversity within populations (h_S = 0.15) and a high level of population differentiation (G_ST = 0.81) demonstrate the strong genetic isolation of populations from each other. Past oscillations between humid and arid periods connected with glacial and interglacial episodes in the Pleistocene and Holocene are considered responsible for the observed patterns of genetic variation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 437–444.     

A NEW SPECIES OF ABYSSOCHRYSOS (GASTROPODA: LOXONEMATOIDEA) FROM THE OMAN MARGIN

A new species of Abyssochrysos, A. xouthos n. sp. is describedfrom south-west Oman in the northern Arabian Sea at a depth of3150 metres. This represents the first record of this genusfrom the northern Indian Ocean. Of the five previously describedextant species of Abyssochrysos A. xouthos is most similar to A.melanioides from which it differs in radular and shell characters. (Received 22 February 1999; accepted 20 May 1999)     

Living Neogloboquadrina pachyderma sin and its distribution in the sediments from Oman and Somalia upwelling areas

Microfaunal analysis of plankton nets collected offshore Oman/Yemen and sediment traps offshore Somalia shows that Neogloboquadrina pachyderma sin is abundant during the SW monsoon upwelling. The sediment traps recorded the highest flux (136 specimens m⁻² day⁻¹) and the highest relative frequency (3.55% of the total foraminifera assemblage) of the species during this period. During the intermonsoon it became less abundant and decreased in size, and only very few N. pachyderma sin were found in the water column during the NE monsoon. Sediment trap and Recent sediment data collected along a downslope transect off Somalia show that the species frequency decreases offshore. The highest concentrations of N. pachyderma sin in plankton nets off Oman/Yemen were found at a depth of 300–500 m. However, the oxygen isotope compositions of N. pachyderma sin at the depth of 300–500 m and from the surface 8 m are identical, and also similar to that of Globigerina bulloides which reflects properties of the upper 25 m of the water column. The carbon isotope values are relatively consistent within the studied samples from both areas except for the specimens from the surface water samples offshore Oman/Yemen. The data show that the δ¹⁸O ratio of N. pachyderma sin is in equilibrium with the ambient water while the δ¹³C values are at least 0.8‰ lower. We infer that N. pachyderma sin in the Arabian Sea reproduces, grows and calcifies in the upper 25 m of the water column, but possibly descends into deeper waters later during its life cycle.     

Phytogeography of upwelling areas in the Arabian Sea

Aim The identification of the marine plant communities of two islands from different upwelling areas of the Arabian Sea, with a similar diversity in biotopes. A comparison of the species composition of these macroalgal communities and their biogeographical affinities within the Indian Ocean should give insight into the biogeographical position of the Arabian Sea within the larger Indian Ocean. The incorporation of environmental parameters in the analysis is instructive in understanding their importance in shaping the diverse marine assemblages of the Arabian Sea. Location Arabian Sea: (1) the Socotra Archipelago (Yemen; 12.47° N, 53.87° E) in the Somali upwelling area, (2) Masirah Island (Oman; 20.42° N, 58.79° E) in the upwelling area of the southern Arabian Peninsula. Methods The marine flora of different biotopes around both islands were examined by means of qualitative assessments. Ordination analysis [detrended correspondence analysis (DCA)] was used to identify the different plant communities and to correlate these with environmental parameters. The species composition of the identified communities were compared (tripartite similarity index) and their biogeographical affinity with nations bordering the Indian Ocean was determined. Indicator species analyses were performed to identify the characteristic species of the different plant communities and their biotopes. Results The DCA analysis shows a clustering of sites (plant communities) corresponding with their geographical position, linked in turn to the prevailing environmental conditions of the different coastal areas. The combined interpretation of the ordination, similarity and biogeographical analyses results in the aggregation of similar plant communities of both upwelling areas into four biotopes. Main conclusions The north coast communities of Socotra and the west coast communities of Masirah can be grouped into three biotopes related to the degree of exposure (to upwelling) and sedimentation. These biotopes are typified by indicator species, characteristic for specific substrata, and have a high biogeographical affinity with the East African coast. The plant communities of Socotra's south coast and Masirah's east coast constitute a fourth biotope, being diverse and species rich, typified by a large proportion of red macroalgae including the characteristic species of the unique Arabian Sea flora. This biotope has a pronounced biogeographical affinity with distant regions (disjunctly distributed taxa) as South Africa's East Coast and Western Australia. Within the different biotopes, the communities of Masirah are more divergent from an East African flora in comparison to Socotra, the latter being a stepping stone between the East African and Arabian Sea flora.     

Zoogeography of the coral reef fishes of the Socotra Archipelago

Fish communities and habitats were studied at the Socotra archipelago (Gulf of Aden, ≈12°N 54°E). Extensive and unexpected hermatypic coral communities were recorded, at the centre of a 2200 km gap in knowledge of species and habitat distributions which coincides with a change from a western Indian Ocean coral reef fauna to an Arabian one. The fish assemblage associated with the Socotra archipelago corals is predominantly south Arabian. An east African influence, minimal on the mainland coasts of Arabia, is more evident here, and results in previously unrecorded sympatry between Arabian endemic species and their Indian Ocean sister taxa. A study of distributions of Chaetodontidae (butterflyfishes) in the north-western Indian Ocean reveals a number of distinct patterns, with a trend for species replacement along a track from the northern Red Sea to the Indian Ocean. A major feature of the reef fish zoogeography of the region is found to be a distinct south Arabian area, characterized by a 'pseudo-high latitude effect' which results from seasonal cold water upwelling along the Arabian sea coasts of Yemen and Oman and the Indian Ocean coast of Somalia. This south Arabian feature is consistent across a wide range of fish families. It is most pronounced in Oman and Yemen, and although it is the dominant influence at Socotra it is slightly 'diluted' here by the east African influence. The south Arabian area wholly or partly accounts for most of the major marine zoogeographic features around Arabia, and is the principal feature fragmenting Arabian coastal fish assemblages, and separating them from those of the wider Indo-west Pacific.     

Pedobesia simple (Kützing) comb. nov. (Chlorophyta), a new name for P. lamourouxii and its first report from the Indian Ocean

The green alga Pedobesia lamourouxii (J. Agardh) Feldmann et al. (Bryopsidales) is reported from the northern Arabian Sea on the basis of collections from the Sultanate of Oman and Socotra Island (Yemen). Vegetative and sporangial plants as well as the ecology are described. These collections constitute the first record of this widely distributed species for the Indian Ocean. In the course of this study we discovered that Bryopsis simplex Kützing is an older taxonomic synonym, and thus we propose Pedobesia simplex (Kützing) comb. nov. as the correct name for this species. A specimen of Bryopsis simplex in the Leiden Herbarium (L) is designated as lectotype.     

Consequences of marine barriers for genetic diversity of the coral‐specialist yellowbar angelfish from the Northwestern Indian Ocean

Ocean circulation, geological history, geographic distance, and seascape heterogeneity play an important role in phylogeography of coral‐dependent fishes. Here, we investigate potential genetic population structure within the yellowbar angelfish (Pomacanthus maculosus) across the Northwestern Indian Ocean (NIO). We then discuss our results with respect to the above abiotic features in order to understand the contemporary distribution of genetic diversity of the species. To do so, restriction site‐associated DNA sequencing (RAD‐seq) was utilized to carry out population genetic analyses on P. maculosus sampled throughout the species’ distributional range. First, genetic data were correlated to geographic and environmental distances, and tested for isolation‐by‐distance and isolation‐by‐environment, respectively, by applying the Mantel test. Secondly, we used distance‐based and model‐based methods for clustering genetic data. Our results suggest the presence of two putative barriers to dispersal; one off the southern coast of the Arabian Peninsula and the other off northern Somalia, which together create three genetic subdivisions of P. maculosus within the NIO. Around the Arabian Peninsula, one genetic cluster was associated with the Red Sea and the adjacent Gulf of Aden in the west, and another cluster was associated with the Arabian Gulf and the Sea of Oman in the east. Individuals sampled in Kenya represented a third genetic cluster. The geographic locations of genetic discontinuities observed between genetic subdivisions coincide with the presence of substantial upwelling systems, as well as habitat discontinuity. Our findings shed light on the origin and maintenance of genetic patterns in a common coral reef fish inhabiting the NIO, and reinforce the hypothesis that the evolution of marine fish species in this region has likely been shaped by multiple vicariance events.     

Reconstruction of Indian monsoon variability over the past 230,000 years: Planktic foraminiferal evidence from the NW Arabian Sea open-ocean upwelling area

We determined the faunal composition and total number of tests (#/g) of planktic foraminifera (> 125 μm) in core KH00-05 GOA 6 near Oman in order to decipher monsoon-induced variability of oceanographic productivity in the open-ocean upwelling area in the northwest Arabian Sea. The core contains a continuous record of sedimentation over the last 230 kyr, with the age model based on oxygen isotope and accelerator mass spectrometry ¹⁴C dates. We focused on species (Globigerina bulloides and Globigerinita glutinata) typical for SW monsoonal upwelling and species typical for NE monsoon conditions (Neogloboquadrina incompta, Neogloboquadrina dutertrei, Globigerinoides ruber, and Globigerinoides sacculifer). The changes in relative abundance of these monsoonal indicators suggest that the open-ocean upwelling area was dominated by the SW monsoon during interglacial periods, but by the NE monsoon during glacial periods.Increases in total test abundance during glacial periods confirmed that the NE monsoon rather than SW monsoon contributes largely to planktic foraminiferal productivity in this area. We argue that three types of circumstances resulted in high productivity, with nine high productivity events occurring at a 23-kyr frequency. The first type caused high productivity events at 102 and 199 ka (interglacial periods), characterized by the dominance of upwelling species, indicating high productivity during strong SW monsoons, correlated with high July insolation at 45° N. An exceptional high productivity event occurred at 37 ka during interglacial marine isotope stage (MIS) 3, with contributions from both SW and NE monsoons. The second type of high productivity event occurred at 61, 147, and 175 ka, during glacial periods, characterized by dominance of NE monsoon species, and correlated with low January insolation at 45° N. In addition, a high productivity event at 85 ka (interglacial period) also was induced by enhanced NE monsoons. The last two high productivity events occurred during transitional periods from glacial to interglacial (MIS 6/5.5 and 2/1), were characterized by the replacement of NE monsoon species with upwelling species, and corresponded to abrupt climate warming, suggesting that they are related to both accelerated SW monsoon systems and reduced NE monsoon systems.     

Upper Triassic reefs of the Oman Mountains: Data from the South Tethyan margin

Summary The Upper Triassic reefal limestones of the Oman Mountains were investigated with respect to their microfacies, palaeontology and community structure. The reef fauna described and figured for the first time occurs in parautochthonous slope deposits of the Arabian platform (Sumeini Group) and in allochthonous reefal blocks (‘Oman Exotics’, Hawasina Complex). The ‘Oman Exotics’ are tectonically dislocated blocks, derived from isolated carbonate platforms on seamounts in the Hawasina basin or in the South Tethys Sea. The lithofacies and fauna of these blocks comprise a cyclic platform facies with megalodonts, reef and reef debris facies. The reefal limestones are dated as Norian/Rhaetian by benthic foraminiferal associations (Costifera, Siculocosta, Galeanella) and typical encrusting organisms (Alpinophragmium, Microtubus). Some small ‘Oman Exotics’ are of Carnian age. The shallow-marine organisms include scleractinian corals of different growth forms, ‘sphinctozoans’, ‘inozoans’ chaetetids, spongiomorphids, disjectoporids and solenoporacean algae as the main reef builders, various encrusters like microbes, foraminifers, sponges and many different problematical organisms for the stabilisation of the reef framework and a group of dwellers including benthic foraminifers, gastropods, bivalves and a few dasycladacean algae. The reef communities are characterized by the coverage of organisms and distributional pattern. Analogies with the coeval reef deposits from the European part of the Tethys have been recognized. Some species, now collected in Oman, were also reported from American and Asian localities.