One of the S. ramani specimens is nested within the central Indian Ocean clade, but the other is sister to the Australia clade. Despite the commercial importance of S. pharaonis, very little has been published on the phylogeography or population genetics of this species, presumably due in part to its broad geographic distribution. Widespread Mediterranean, Indo-West Pacific Video by Japan Ethological Society & Springer Japan. They are also able to shoot a cloud of black ink at predators when threatened. Size: 18 inches (45 cm) ... Genus: Sepia. Tissue samples from two additional specimens of S. pharaonis and two specimens of S. ramani were collected in Tuticorin, India, in October 2007. Our central Indian Ocean subclade may be Norman's S. pharaonis III, although we found that S. pharaonis individuals collected along the west coast of India from as far north and west as Veraval are also members of this clade (i.e. It is commonly hunted in the Philippines, India, and Persia for food. PCRs were performed using a Perkin-Elmer 9700 thermal cycler. Furthermore, the cuttlebone of S. pharaonis has a distinctive cuplike extension covering the striated zone of the posterior inner cone (Khromov et al., 1998; Norman, 2000), which may allow fossil members of the S. pharaonis complex to be identified. An outer shell once covered the cuttlefish's … in 80–100% EtOH as part of an earlier study (Anderson et al., 2007). Sepia ramani is so far only known from the Gulf of Mannar in southeastern India (Neethiselvan, 2001). However, two of the four specimens collected in the Gulf of Oman (N Gulf of Oman 3* and N Gulf of Oman 5) showed discordance between clade membership and collection locality. Sometimes the boys of these cephalopods will fight each other over a girl they like, but they won't touch each other, they just flash different patterns of colors. 3), and the distinction between this subclade and all other subclades in the complex is the only distinction that is supported by the rhodopsin sequence data. The best (lowest) AICc and BIC scores are in bold text. Sign in Sign up for FREE Prices and download plans Not Reef Tank Suitable. At present, we have no divergence time information for clades within the S. pharaonis complex. Recovery of cuttlebones attributable to the S. pharaonis complex might allow estimation of the age of the complex and divergence times within the complex, allowing phylogeographic comparisons of S. pharaonis with other neritic species in the Indian Ocean. Sepia pharaonis. Sepia pharaonis Ehrenberg, 1831 Pharaoh Cuttlefish. Preliminary analyses suggested that the default temperature (T = 0.2) resulted in very few state swaps between chains, and some analyses were succumbing to the ‘long tree’ problem, in which estimated branch lengths were unreasonably long, as described by Marshall (2010) and Brown et al. (2005) noted that S. pharaonis is the most common species of cuttlefish caught in the Persian Gulf, the Gulf of Oman, the Andaman Sea, the Gulf of Thailand, the Philippines and along the southern coast of China, and Nesis (1987) wrote that “[Sepia pharaonis] is the most important object of the cuttlefish fishery in the northern part of the Indian Ocean and southeastern Asia”. They feed by catching their prey by two powerful tentacles which shoot out from beneath the creatures eyes. COI and rhodopsin sequences obtained in this study were combined with all available sepiid 16S rRNA, COI and rhodopsin sequences in GenBank (http://www.ncbi.nlm.nih.gov/) as of 17 February 2009. Norman (2000) suggested that S. pharaonis comprises three forms: S. pharaonis (s. s.) (found in the western Indian Ocean from the Red Sea to the Persian Gulf; the eastern limit is unknown); S. ‘pharaonis II’ (Japan to the Gulf of Thailand, Philippines and north Australia) and S. ‘pharaonis III’ (Maldives to Andaman Sea coast of Thailand). One group of closely related individuals (the central Indian Ocean subclade) is distributed across the central Indian Ocean along the east and west coasts of India and the Andaman Sea coast of Thailand; in contrast, another group seems to be restricted to the Persian Gulf and northern Gulf of Oman (the Iranian subclade). In light of this, we suggest that the binomen S. pharaonis be restricted to the western Indian Ocean subclade. Substitution model abbreviations are as follows: GGI = GTR = G = I, HG = HKY85 = G, HI = HKY85 = I, K = K2P, KG = K2P = G; see Anderson & Swofford (2004) for more information on model abbreviations and original citations for each model. Created to help individuals around the world identify tropical fish - WHATSTHATFISH.com. Four Bayesian analyses, each consisting of one cold and three heated Metropolis-coupled Markov chains, were run simultaneously in MrBayes v. 3.1.1, with random starting trees and trees sampled every 1,000 generations. When these different estimates of sample size caused MrDT-ModSel to select different models for a given data partition, we chose the model with fewer parameters. We suggest that ‘S. It is possible that Pleistocene glaciations also played a role in the divergence between the central Indian Ocean and western Pacific clades of S. pharaonis, though the current lack of divergence time information for the S. pharaonis complex limits our ability to test hypotheses of causality. Sepia pharaonis is a neritic demersal species so direct dispersal across the Gulf of Oman seems unlikely. Males can only produce once and the females die shortly after laying their eggs. These calculations required estimation of model likelihoods. As shown by Anderson et al. Allcock et al. Genus: Sepia . *Rerun for 50 million generations. (2007): 1, Red Sea; 2, Gulf of Aden; 3, Persian Gulf (Iran); 4, northern Gulf of Oman (Iran); 5, southern Gulf of Oman (Oman); 6, Veraval; 7, Kochi; 8, Tuticorin; 9, Vishakapatanam; 10, Phuket; 11, Prachuap; 12, Chumphon; 13, Taiwan; 14, Gulf of Carpentaria; 15, northeast Queensland. The taxonomic status of S. ramani (as well as usage of the binomen S. pharaonis itself) hinges on the taxonomic status of the five unnamed subclades, and this cannot be fully addressed without detailed morphological and morphometric work, preferably coupled with additional genetic data collection to provide a link with our study. A total of 141 out of 684 sites for COI were variable and 109 of these were parsimony-informative within Sepia pharaonis. Comparisons with other studies are somewhat compromised by the fact that although numerous phylogeographic studies of Indian Ocean species have been published, the Malay Archipelago, South Africa and Australia have received substantially more attention than the northern Indian Ocean (i.e. Abbreviation: ∼, approximate values denoting collections from several sites in close proximity to one another. Customise filters (scroll to see full list) Taxon. The cuttlefish isn’t a fish at all – it is a cephalopod. The deepest divergence within the S. pharaonis complex is between the western Indian Ocean clade and the rest of the complex. There appear to be consistent reproductive differences among these three forms. Found in shallow waters over sand and seagrass beds of coral and rocky reefs. to compare only substitution models that are available in MrBayes v. 3.1.1 (Perl script available upon request to F.E.A.). Welcome to FishSource, an online information resource about the status of fish stocks and fisheries. DNA markers indicate that distinct spawning cohorts and aggregations of Patagonian squid, Subtle population structuring within a highly vagile marine invertebrate, the veined squid, Widely distributed Pacific plate endemics and lowered sea-level, Molecular phylogeny of coleoid cephalopods (Mollusca: Cephalopoda) using a multigene approach; the effect of data partitioning on resolving phylogenies in a Bayesian framework, PAUP*. Only 16S rRNA sequences are available from the two specimens of S. pharaonis from Taiwan. Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae. Furthermore, phylogeographic studies of Indian Ocean marine fauna encompass taxa of differing ages, which may have been impacted by different vicariant events or paleooceanographic phenomena (Page, 1990, 1991). Photography By: Ramin Ketabi Editor:Shahla Jamili (IFSRI) The. Sepia ramani is a neritic demersal southeastern Indian species that is morphologically very similar to S. pharaonis, and there has been some controversy regarding the status of S. ramani as a distinct species. Flamboyant Cuttlefish: This species is well-named for the rather bright and exuberant pattern of colors on … All of these sequences appear to be protein-coding sequences rather than pseudogenes; translation into amino acids using the flatworm mitochondrial genetic code (COI sequences) or the universal genetic code (rhodopsin sequences) in MacClade v. 4.08 (Maddison & Maddison, 2005) revealed no premature stop codons. əl‚fish] (invertebrate zoology) An Old World decapod mollusk of the genus Sepia; shells are used to manufacture dentifrices and cosmetics. This time, when 33 cuttlefish … The Bayesian consensus phylogram does not clearly support monophyly of the S. pharaonis complex; sequences from Metasepia tullbergi and S. lycidas are part of a polytomy that includes all S. pharaonis specimens sampled in this study, although one resolution of this polytomy would have the S. lycidas/M. Neethiselvan (2001) also noted that some morphometric characters (cuttlebone width, inner cone length and tentacular club length) could be useful for distinguishing between the species, but there is some overlap between the two species in all of these characters. For full access to this pdf, sign in to an existing account, or purchase an annual subscription. Our only specimens from the Persian Gulf are members of the Iranian subclade, but we do not know if members of the western Indian Ocean subclade are also found in the Persian Gulf, so we cannot discern whether the boundary between these two subclades is in the Gulf of Oman or the Persian Gulf (or both). Multiple data partitioning schemes were tested for Bayesian analyses of the separate and combined datasets. They usually … The strong morphological similarities suggest a close relationship between S. pharaonis and S. ramani, but the nature of this relationship is unknown. Have a photo you want identified? Partition abbreviations are as follows: C, COI; R, rhodopsin; C1, COI position 1; C2, COI position 2; C3, COI position 3; R12, rhodopsin position 1 + position 2; R3, rhodopsin position 3. Sepia (genus) - WikiMili, The Free Encycloped The Pharaoh Cuttlefish is found in the Mediterranean, Indo-West Pacific region growing up to 33cm in length. The geographic regions in question are adjacent to one another; one member of the western Indian Ocean subclade (S Gulf of Oman 1) was collected from the southern coast of the Gulf of Oman, while the Iranian specimens (N Gulf of Oman 2, 3, 4 and 5) were collected about 230 km to the northeast, on the opposite side of the Gulf of Oman. To our knowledge, no fossil cuttlebones attributable to S. pharaonis have been found. These likelihood scores were used to select a best-fitting substitution model using ‘MrDT-ModSel’, a modification of DT-ModSel (Minin et al., 2003) developed by F.E.A. (2007), five strongly supported geographically delimited clades are evident on both the mtDNA and three-gene phylogenies. Ten such analyses were run, with bootstrap support values for each node averaged across all 10 runs. Scientific name Scientific name (unprocessed) Subspecies Species Genus Family Order Class Phylum Kingdom Scientific name (unprocessed) Subspecies Species Genus … There is some morphological and behavioural evidence that S. pharaonis may be a complex of closely related species. 1) and shipped to the first author (F.E.A.) While mating, S. pharaonis s. s. males show zebra lines on the third arm pair, while S. pharaonis II males have broken lines and S. pharaonis III males have spots (Norman, 2000). The pharaoh cuttlefish Sepia pharaonis Ehrenberg, 1831 (Sepiidae) is a broadly distributed neritic demersal cephalopod species found from East Africa to southern Japan. The combined mtDNA phylogeny is shown in Figure 2. Given that there are over 100 described species within Sepiidae (Khromov et al., 1998; Lu, 1998), our inference of monophyly for the S. pharaonis complex must be considered provisional pending sampling of additional sepiid species. Depth - 0-130m Such data could be particularly important for S. pharaonis. 5 kg, and for females 50 cm and 2 kg in … (2010). The average longevity of the species is between 1-2 years. Genetic divergence between Indian and Pacific populations of marine species has been attributed to reductions in gene flow during repeated periods of glaciation over the last 140,000 years, which resulted in lower sea levels, reduced transfer of warm surface water between the Indian and Pacific Ocean basins and increased cold-water upwelling as recently as 18,000 years ago (Potts, 1983, 1984; Fleminger, 1986; Springer & Williams, 1990; Williams et al., 2002). The three-gene phylogeny is shown in Figure 3. FYI: Shown here only for identification only. For the protein-coding gene datasets, the data were either not partitioned or partitioned by codon position (with a separate substitution model for each codon position and model parameters estimated separately for each partition). The Gulf of Oman ranges from 60 km (at the Strait of Hormuz) to 370 km wide (from Ras Al Hadd, Oman to Gwadar Bay, Pakistan) and the Gulf of Oman basin is about 3,400 m deep (Uchupi, Swift & Ross, 2002). Members of the class Cephalopoda are gonochoric. A pharaoh cuttlefish pretends to be a hermit crab, raising its front legs to look like eyestalks and appearing to walk on the bottom of the tank. File:Partes de la sepia.ogv. Upload image Colour changes to match surroundings. Cuttlefishes. Sequences for S. madokai have been removed from GenBank subsequent to the completion of the analyses described in this paper. Rhodopsin sequences for E. scolopes, M. tullbergi and S. officinalis are from different individuals than the mitochondrial sequences. Due to the size and composition of the dataset, analyses of each MP bootstrap pseudoreplicate resulted in thousands of equally parsimonious trees. Pharaoh Cuttlefish: This is a large species of cuttlefish that inhabits the Pacific region between Japan and Australia and as far west as the Red Sea. The number of parameters, run length (‘length’) in millions of generations, best-fitting models, and AICc and BIC values for different partitioning schemes for the combined mtDNA (16S rRNA + COI) and three-gene (16S rRNA + COI + rhodopsin) datasets. Numbers above branches are clade posterior probability (BPP) estimates; numbers below branches are MPBS values. only one substitution model was used), partitioning by gene, or partitioning by gene and codon position. Sepia pharaonis. Maturation, fecundity and seasonality of reproduction of two commercially valuable cuttlefish, The preservation of the shells of Sepia in the middle Miocene of Malta, Proceedings of the Geologists’ Association, Molecular and morphological analyses of the cuttlefish, A synopsis of Sepiidae outside Australian waters (Cephalopoda: Sepioidea), A synopsis of Sepiidae in Australian waters (Cephalopoda: Sepioidea), Phylogenetic systematics and biogeography of hummingbirds: Bayesian and maximum likelihood analyses of partitioned data and selection of an appropriate partitioning strategy, MacClade: analysis of phylogeny and character evolution, Version 4.08, Sunderland, Massachusetts, USA, Cryptic failure of partitioned Bayesian phylogenetic analyses: lost in the land of long trees, Performance-based selection of likelihood models for phylogeny estimation, First multi-generation culture of the tropical cuttlefish, Enlightenment of old ideas from new investigations: more questions regarding the evolution of bacteriogenic light organs in squids, Bayesian phylogenetic analysis of combined data, Temporal congruence and cladistic analysis of biogeography and cospeciation, Clocks, clades and cospeciation: comparing rates of evolution and timing of cospeciation events in host-parasite assemblages, Testing hypotheses of population structuring in the Northeast Atlantic Ocean and Mediterranean Sea using the common cuttlefish, Selecting the best-fit model of nucleotide substitution, Model selection and model averaging in phylogenetics: advantages of Akaike information criterion and Bayesian approaches over likelihood ratio tests, Evolutionary disequilibrium among Indo-Pacific corals, Generation times and the Quaternary evolution of reef-building corals, Cephalopods of the world. Ref: https://en.wikipedia.org/wiki/Pharaoh_cuttlefish. We have also added sequence data from two specimens of Sepia ramaniNeethiselvan, 2001, collected in southeastern India. When this value reached 0.01, the runs were terminated. Collection locality and GenBank accession data for all specimens of Sepia pharaonis complex. 49, Paris, France, DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates, Molecular Marine Biology and Biotechnology. Cuttlefish, Octopus & Squid. (2007). A similar pattern has been found in Lunella coronata, a gastropod found on rocky shores from southeastern Africa through the Gulf of Oman to the western Pacific (Williams et al., in review). As Anderson et al. (2007) and confirmed here corresponds quite well to Norman's S. pharaonis s. s., although we found evidence of a genetic break between the southern and northern Gulf of Oman (see below). In analyses of the partitioned datasets, all model parameters except topology and branch lengths were unlinked across partitions. Want to share your pictures? The relationships among the S. pharaonis subclades are still not fully resolved, but some inferences can be made. By comparison, the rhodopsin data showed very low levels of variation. Estimated Bayesian posterior probabilities (BPPs) of clades on inferred trees were interpreted as measures of support. Another type of cephalopod is the Pharaoh Cuttlefish. Pharaoh cuttlefish are cephalopods related to cuttlefish, squid, octopus and chambered nautilus. The cuttlefish's flat body allows it to live and hover near the ocean bottom where it finds its favorite food. Anderson et al. Males are larger than females, the maximum recorded size for males is 80 cm and. They feed on crustaceans and small fish. Though less work has been done on western Indian Ocean marine populations, some studies have found evidence of a phylogeographic break between the eastern and western Indian Ocean (Ridgway & Sampayo, 2005). For each dataset or partition, models were evaluated by using all sites or only variable sites as estimates of sample size (Posada & Buckley, 2004). All hope is not lost, however, because sepiids possess a calcified structure that would seemingly be amenable to fossilization – the cuttlebone. Cuttlefish possess the ability to swim in different manners, usually gently rippling their side fins. Cephalopod researcher Dr. James Wood sums it up well; “Octopuses, squids, cuttlefish and the chambered nautilus belong to class Cephalopoda, which means ‘head foot”. During the southwestern (summer) monsoon, the Ras Al Hadd jet (a continuation of the Somali and Oman Coastal Currents that flows eastward from the eastern tip of Oman; Schott & McCreary, 2001) and the cyclonic eddy it produces in the Gulf of Oman could promote occasional dispersal of S. pharaonis juveniles across the Gulf. Furthermore, additional specimens from as-yet-unsampled parts of the range of the S. pharaonis complex must be evaluated, as there may be additional subclades (or species) waiting to be discovered; regions of particular interest are Madagascar, the Philippines, the Yellow Sea (Hwang Hai) and Indonesia. This specimen could be misidentified, or it could be a hybrid (or backcross) between S. ramani and S. pharaonis that exhibits S. ramani morphology but carries a S. pharaonis mtDNA haplotype. Also included in this clade are the lesser-known cuttlefish (Sepioidea), the Ram's Horn Squid, which has an internal coiled shell and floats head down in the water, and an enigmatic deep water genus called the "vampire squid" (Vampyromorpha). The phylogeny reveals five strongly supported subclades within S. pharaonis: a western Indian Ocean clade (Red Sea, Gulf of Aden and the northeast coast of Oman), a northeastern Australia clade (with a representative of ‘S. unpubl. Several phylogenetic analyses were performed for the COI and rhodopsin data individually and for three combinations of data – one consisting of the combined mtDNA data only (i.e. Sepia pharaonis is a commercially harvested species, and it is a significant component of cephalopod fisheries throughout its range (Nesis, 1987; Reid, Jereb & Roper, 2005). Sepia. tullbergi clade as sister to a monophyletic S. pharaonis complex (an S. pharaonis sequence from GenBank is distantly related to the S. pharaonis sampled in our study, suggesting that the GenBank specimen was misidentified; this sequence was excluded from the three-gene dataset prior to analysis). Though we did not obtain samples from the type localities, we did obtain samples from the Yemeni Red Sea coast (340 km east of Massawa) and found that these specimens were members of our western Indian Ocean subclade. The pharaoh cuttlefish, Sepia pharaonis Ehrenberg, 1831, is a commercially fished species found from Japan to East Africa. Sepia ramani is a member of the S. pharaonis species complex, though one of our S. ramani samples may represent an additional, previously unsampled subclade within the complex. However when in danger, the cuttlefish sucks water into their body cavity and expels it through a funnel like extension on the underside of the body, causing a backward propulsion enabling the cuttlefish to escape from predators. Customise filters × Customise filters (scroll to see full list) (scroll to see full list) a single substitution model was used for the dataset). (2007) noted, the type localities of S. pharaonis are both in the Red Sea (near El-Tor in the Sinai in the northern Red Sea and near Massawa in Eritrea along the west coast of the Red Sea). A fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene and a fragment of the rhodopsin gene were amplified using universal metazoan COI PCR primers (Folmer et al., 1994) and cephalopod-specific rhodopsin PCR primers (Strugnell et al., 2005), and HotStar Master Mix (QIAGEN) following manufacturer's protocols (half-reactions). ramani’ weakly supported as sister to this clade), an Iranian clade (northeastern Persian Gulf and northern Gulf of Oman), a western Pacific clade and a broadly distributed central Indian Ocean clade (west and east coasts of India and the Andaman Sea coast of Thailand). For both the combined mtDNA dataset and the three-gene dataset, the AICc and BIC values were lowest for the ‘by gene and codon position’ partitioning scheme, indicating that this was the best-fitting partitioning scheme of those evaluated for these data (Table 3).