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.     

Cleome socotrana (Capparaceae) and allied species in the Horn of Africa region

Four species of Cleome with small flowers, 6 stamens, petals without appendages, spreading to erect capsules, and hairy seeds, are recognized in the Horn of Africa region: C. socotrana in the Socotra archipelago (Yemen), C. hadramautica , sp. nov., in southern Yemen, C. omanensis , comb, nov., in the Mahrah Region of Yemen and in Oman, and C. albescens in northern Somalia. C. socotrana is lectotypified and a key to the species treated is provided.     

The phylogenetic affinities of the Socotra Bunting Emberiza socotrana

The monophyly of the African ‘brown bunting’ complex was corroborated by a recent molecular study. However, the little-known Socotran endemic Emberiza socotrana, which is morphologically similar to the other taxa in this complex, was not included. Here we present a hypothesis of the phylogenetic relationships of the Socotra Bunting based on one mitochondrial gene and one nuclear intron. We found the Socotra Bunting to be deeply nested within the African ‘brown bunting’ complex and, although morphologically most similar to E. capensis, it proved to be more closely related to the E. striolata/sahari and E. tahapisi/goslingi species groups. The phylogenetic uniqueness of the Socotra Bunting underpins once more the evolutionary importance of the Endemic Bird Area of Socotra, which is often considered the ‘Galápagos of the Indian Ocean’.     

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.     

New record of the butterfly Euchrysops cnejus (Fabricius) from Oman,with notes about phylogeographic patterns of E. cnejus and E. osiris (Hopffer) (Lepidoptera: Lycaenidae)

We document the occurrence of the Oriental butterfly species Euchrysops cnejus in northern Oman. It is very similar to E. osiris whose distribution extends from the Afrotropical region into Dhofar, southern Oman. Oman is thus the only country where both species co-occur. The two species differ in genitalia and in molecular data. We show that based on the COI marker, E. cnejus and E. osiris are sister species and the Oman populations of E. osiris are closely related to the population from continental Africa. Haplotype diversity is lower in E. cnejus, than in E. osiris. Genetic differences were found between the Australian, Oriental and Oman populations of E. cnejus but available data were not sufficient to clarify the origin of the Oman population.     

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.     

New or noteworthy species of Leguminosae in NE tropical Africa

The following new species are described: Chamaecrista dunensis (Somalia), Acacia densispina (Somalia), A. flagellaris (Somalia), Tephrosia karkarensis (Somalia), T. scopulata (Somalia), Indigofera ancistrocarpa (Kenya), I. curvirostrata (Ethiopia), I. hiranensis (Somalia), I. karinensis (Somalia), I. kucharii (Somalia), I. nugalensis (Somalia), I. pellucida (Kenya, Somalia), I. sabulosa (Ethiopia, Somalia), I. sessilis (Somalia), I. ienuirostris (Somalia), Rhynchosia megalocalyx (Somalia), Crotalaria allophylla (Somalia), C. intricata (Somalia), C. warfae (Somalia), and Coronilla somalensis (Somalia). 2n = 22 is reported in Tephrosia karkarensis. Cassia somalensis, Acacia puccioniana, Sophora somalensis , and Crotalaria sennii are reduced to synonyms of respectively Senna hookeriana, Dichrostachys kirkii, Millettia usaramensis and Crotalaria boranica subsp. trichocarpa . Thirty-seven taxa are reported from Somalia for the first time, including Ptycholobium biflorum , known previously only from Southern Africa. Crotalaria pieropoda , known previously only from Socotra, is reported from Somalia and Oman. Neptunia oleracea and Vatovaea pseudolablab are new generic records for respectively Ethiopia and Oman. The fruits of Indigofera boranica are described. Vigna somaliensis and an allied species are discussed. Rhynchosia gansole is lectotypified and its flowers and fruits are described.     

A taxonomic revision of Socotran Indigofereae (Leguminosae—Papilionoideae) with insights into the phytogeographical links of the Socotran Archipelago

This revision of Socotran Indigofereae (Leguminosae) treats two genera, Microcharis and Indigofera , and 16 species, one with two varieties. Indigofera coerulea var. coerulea and I. nugalensis are new records for the archipelago. Indigofera nephrocarpoides, I. marmorata (lectotypified here) and I. socotrana are endemic, whilst Microcharis disjuncta vat. fallax and Indigofera nugalensis are near-endemic to the Socotran islands. This study indicates that the closest relatives of the Socotran Indigofereae are to be found in African lineages. The strongest affinities are between Socotra and extreme NE Somalia ( Indigofera pseudointricata, I. nugalensis ) and, to a lesser extent, with SW Oman ( Microcharis disjuncta var. fallax ), SW Pakistan ( I. nephrocarpa ) and Southern Yemen. Socotran Indigofereae are mainly derived from Tertiary African palaeotropical and drought-adapted lineages. We also suggest that following the separation of the islands from mainland Africa some Indigofereae would have reached the archipelago as a result of recent, long-distance dispersal events.     

Erythroxylon (Erythroxylaceae) on Socotra

Erythroxylon socotranum , sp. nov., is described from limestone rocks on southwestern Socotra (Yemen). The species is the first representative of Erythroxylaceae to be known from Socotra and from South-West Asia in general.     

A new arborescent Euphorbia (Euphorbiaceae) from northern Somalia

Euphorbia spissa, sp. nov., a tree known from a single population on a rocky limestone slope in northern Somalia, is described and illustrated. The species is compared with the widespread E. tirucalli, as well as with E. cameronii and E. bariensis (endemics in Somalia), E. arbuscula (endemic on Socotra), and E. uzmuk (endemic in northern Yemen).     

A new species and a new record of Tephrosia (Leguminosae) from Socotra

Tephrosia socotrana sp. nov., a woody species growing on limestone cliffs in north-central Socotra (Yemen), is described and illustrated. strigosa is recorded from Socotra and Yemen for the first time.     

Diversity and composition of estuarine and lagoonal fish assemblages of Socotra Island,Yemen

Estuarine and lagoonal surveys of Socotra Island and selected sites on the Hadhramout coast of Yemen were conducted with the objective of documenting and analysing fish diversity and assemblage structure. A total of 74 species in 35 families were recorded, among which 65 species in 32 families were from Socotra and 20 species in 17 families were from mainland Yemen. Twenty‐one species represent new faunal records for Socotra. Including historic records re‐examined in this study, the total fish species richness of estuaries and lagoons of Socotra Island reaches 76, which is relatively high compared to species inventories of well‐researched coastal estuaries in southern Africa. Five species dominate the occurrence and abundance frequencies: Terapon jarbua, Hyporhamphus sindensis, Aphanius dispar, Ambassis gymnocephala and Chelon macrolepis. Rarefaction and extrapolation analyses suggest that the actual number of fish species inhabiting some of those estuaries might be higher than the one observed. Thus, additional sampling at specific sites should be conducted to record other less conspicuous species. Ordination and multivariate analyses identified four main distinct assemblage clusters. Two groups are geographically well structured and represent northern Socotra and mainland Yemen, respectively. The other two assemblage groups tend to be determined to a greater extent by the synchrony between physical (e.g. estuary opening periods) and biological (e.g. spawning and recruitment periods) variables than by geographical location. Finally, the single intertidal lagoon of Socotra represents by itself a specific fish assemblage. The high proportion of economically important fish species (38) recorded underscores the paramount importance of these coastal water bodies as nursery sites, and for sustaining vital provisioning ecosystem services.     

Forgotten in the ocean: systematics,biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago

Sindaco, R., Metallinou, M., Pupin, F., Fasola, M. & Carranza, S. (2012). Forgotten in the ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago. —Zoologica Scripta, 41, 346–362. The Socotra Archipelago, in the north‐west Indian Ocean, is considered to be one of the most remote and most biodiversity rich and distinct islands in the world. Often referred to as the ‘Galapagos of the Indian Ocean’, it was designated a UNESCO World Heritage Natural site in 2008. Despite having a very rich and bizarre fauna and flora with a high level of endemicity at both species and generic levels, the taxonomy of most of the groups is still not clear, and their origin and evolution remain unknown. Reptiles constitute the most relevant vertebrate fauna of the Socotra Archipelago, with 90% of the 30 species and 45% of the 12 genera being found nowhere else in the world. The skinks of the endemic species Trachylepis socotrana are the only reptile species in the Archipelago distributed across all four islands (Socotra, Darsa, Samha and Abd Al Kuri). Although the species is very well known from Socotra Island, it was not discovered on Samha until 1999 and on Darsa until 2000, whereas only a few citations and one single Museum specimen exist for the population from Abd Al Kuri. To clarify the systematics, biogeography and evolution of Trachylepis socotrana, we assembled a dataset for Mabuya sensu lato including 904 base pairs (bp) of sequence (392 bp from the 12S and 512 from the 16S rRNA mitochondrial genes) for 115 individuals, including specimens of T. socotrana from all four island populations, numerous representatives of the genus Trachylepis from the Middle East, Africa and Madagascar, plus some individuals from each of the other three genera of Mabuya sensu lato (Chioninia, Eutropis and Mabuya). The results of the phylogenetic analyses indicate that, contrary to what was previously thought, members of the genus Trachylepis have colonized the Socotra Archipelago in two independent events, with the first giving rise to the populations from Socotra, Samha and Darsa and the second to the Trachylepis from Abd Al Kuri Island. According to the calibrations, both colonization events occurred within the last fourteen million years, when the Socotra Archipelago had already drifted away from Arabia, thus ruling out vicariance. Both morphological and genetic data show that the Trachylepis from Abd Al Kuri is a distinct taxon, which is herein described as a new species belonging to the T. brevicollis species complex. On the basis of this evidence, the terrestrial herpetofauna from Abd Al Kuri is composed exclusively of endemic species (one of which, the gecko Pristurus abdelkuri, was introduced into some parts of Socotra).     

New species of Euphorbia subgenus Tirucalli (Euphorbiaceae) from Somalia and Oman

A small shrub, Euphorbia bariensis related to E. arbuscula , a sturdy shrub, E. pachyclada related to E. consobrina and E. nubica , and one with conspicuous bracts, E. papilionum related to E. cameronii , are described as distinctive new endemic-species from northeast Somalia. A new endemic, E. dhofarensis related to E. larica , is also described from the Dhofar region of Oman.     

Origin and in situ diversification in Hemidactylus geckos of the Socotra Archipelago

The Socotra Archipelago is an ancient continental fragment of Gondwanan origin and one of the most isolated landforms on Earth and a biodiversity hot spot. Yet, the biogeography and evolutionary history of its endemic fauna still remain largely overlooked. We investigate the origin, tempo and mode of diversification in the Hemidactylus geckos of the Socotra Archipelago. Concatenated and multilocus species coalescent analyses of Hemidactylus from Arabia and North Africa indicate that the Hemidactylus from Socotra do not form a monophyletic group and branch as three independent and well-supported clades instead. Both the chronogram inferred using the gene tree approach of BEAST and the age-calibrated multilocus species tree obtained using *BEAST suggest that the origin of Hemidactylus from Socotra may have involved a first vicariance event that occurred in the Early Miocene, followed by two independent transoceanic dispersal events that occurred more recently, during the Pliocene. Within Socotra, we analysed patterns of genetic diversity, the phylogeography and the demographic history in all seven nonintroduced species of Hemidactylus. Results based on two mitochondrial and two nuclear loci from 144 individuals revealed complex patterns of within-island diversification and high levels of intra-species genetic divergence. The interplay of both historical and ecological factors seems to have a role in the speciation process of this group of geckos. Interestingly, the case of H. forbesii and H. oxyrhinus, which inhabit the island of Abd al Kuri with an area of 133 km(2), may represent one of the most extreme cases of intra-island speciation in reptiles ever reported.     

Chromosome numbers of Umbelliferae (Apiaceae) from Africa south of the Sahara

All previously reported chromosome counts for Umbelliferae of tropical and southern Africa are summarized, two from Socotra are included. New counts on 35 taxa in 14 genera are provided, mostly from Malawi or South Africa. Alepidea, Baumiella, Diplolophium, Lefebvrea , and Lichtenskinia are included for the first time.     

Myxobdella socotrensis sp. nov., a new parasitic leech from Socotra Island,with comments on the phylogeny of Praobdellidae (Hirudinida: Arhynchobdellida)

The Socotra Archipelago (Yemen) is remarkable for unique ecosystems and high endemism, for which it is often referred to as the “Galápagos of the Indian Ocean”. Here we describe a new parasitic leech Myxobdella socotrensis sp. nov. from Socotra, the largest island of the archipelago. The new species was found in a freshwater spring attached to the endemic crab Socotrapotamon socotrensis (Hilgendorf, 1883) (Crustacea: Potamidae). Based on its morphology, ecology and a phylogenetic analysis of DNA sequence data (12S, 18S, 28S and COI gene markers), the new leech species is classified into the highly diversified family Praobdellidae (Hirudinida: Arhynchobdellida), distributed in tropical regions of Africa, Asia and Central and South America. The praobdellid leeches are known to infest mammalian mucous membranes, however, some taxa are associated with other hosts, namely crabs, amphibians and birds. By its morphology, the new species fits quite well in the current concept of the genus Myxobdella Oka, 1917. However, the monophyly of Myxobdella was not supported here by molecular data, pointing at the need of a more comprehensive systematic revision of the genus and family. As far as known, Myxobdella socotrensis sp. nov. is endemic to the Socotra Island, but more data are needed to understand its evolutionary origin, biology and distribution.     

Seascape genetics along environmental gradients in the Arabian Peninsula: insights from ddRAD sequencing of anemonefishes

Understanding the processes that shape patterns of genetic structure across space is a central aim of landscape genetics. However, it remains unclear how geographical features and environmental variables shape gene flow, particularly for marine species in large complex seascapes. Here, we evaluated the genomic composition of the two‐band anemonefish Amphiprion bicinctus across its entire geographical range in the Red Sea and Gulf of Aden, as well as its close relative, Amphiprion omanensis endemic to the southern coast of Oman. Both the Red Sea and the Arabian Sea are complex and environmentally heterogeneous marine systems that provide an ideal scenario to address these questions. Our findings confirm the presence of two genetic clusters previously reported for A. bicinctus in the Red Sea. Genetic structure analyses suggest a complex seascape configuration, with evidence of both isolation by distance (IBD) and isolation by environment (IBE). In addition to IBD and IBE, genetic structure among sites was best explained when two barriers to gene flow were also accounted for. One of these coincides with a strong oligotrophic–eutrophic gradient at around 16–20?N in the Red Sea. The other agrees with a historical bathymetric barrier at the straight of Bab al Mandab. Finally, these data support the presence of interspecific hybrids at an intermediate suture zone at Socotra and indicate complex patterns of genomic admixture in the Gulf of Aden with evidence of introgression between species. Our findings highlight the power of recent genomic approaches to resolve subtle patterns of gene flow in marine seascapes.