Early otolith development in the critically endangered tooth-carp, <Emphasis Type="Italic">Aphanius farsicus</Emphasis> (Teleostei: Cyprinodontidae)

Otolith morphology in the tooth-carp/killifish genus Aphanius is a source of informative taxonomic characters at both the species and population level. Most work on otoliths has focused on adult specimens, while evidence of ontogenetic variation is rarely provided. In this study we describe the development of otolith morphology during the early life stages of an endangered and endemic species, the Fars tooth-carp Aphanius farsicus from southern Iran. The study material comprises 34 larvae and early juveniles representing nine different developmental stages (0–120 days post hatching), all reared under the same laboratory conditions. The results reveal (i) a significant correlation between standard length and otolith size (length) in larval and early juvenile stages, (ii) clear differences in otolith morphology between larvae/early juveniles and adults, and (iii) a temporal link between the appearance of the sulcus on the otolith’s inner face and the emergence of the dorsal and anal fins. Our results indicate that otoliths of Aphanius can be recognized as originating from larval or early juvenile fish based on their short rostrum and antirostrum lengths and wide excisura, in addition to their small size. These immature otoliths are, however, not diagnostic at the species level in A. farsicus, nor most probably in other species of tooth-carp. The outcome of our study is also of interest to palaeontologists working with fossil killifish otoliths, as it can help avoid misinterpretation of ancient species diversity.     

A new and unique species of the genus Aphanius Nardo, 1827 (Teleostei: Cyprinodontidae) from Southern Iran: A case of regressive evolution

A primarily vicariance-based speciation has been suggested for the killifish genus Aphanius Nardo, 1827, but ecological factors are likely to have promoted the speciation processes in addition. Here, we report on the discovery of a unique Aphanius species from Southern Iran and show that also regressive evolution has shaped the present-day diversity of Aphanius. The species is characterized by complete absence of scales and reduction in the biomineralization of hard structures, particularly of the caudal skeleton and jaw teeth. Based on mt-DNA sequences, morphometric and meristic data, osteology, jaw teeth and otoliths, it is described as Aphanius furcatus sp. n. The new species is sympatric with A. dispar (Rüppell, 1829) in salty rivers and hot sulphuric springs in the Hormuzgan Basin (Southern Iran), and is sister taxon to this species plus A. ginaonis Holly, 1929. Based on geological data we estimate that the divergence between the lineages of A. furcatus and A. dispar is about 12–14 million years old. We conclude that the reductive phenomena observed in A. furcatus have evolved as an evolutionary response to the extreme habitat conditions in order to save energy (because storage of Ca²⁺ is not necessary), and to transport oxygen efficiently. The results confirm that regressive evolution is an important factor in speciation and occurs independently in separate lineages.     

Ontogenetic pattern,morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae)

The relationships between ontogenetic and sexual factors with otolith morphology are poorly studied in killifish genus Aphanius Nardo, 1827. We studied the ontogeny, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus inhabiting a high‐salinity environment in Southern Iran. The results highlighted growth‐dependent variability in the otolith of A. furcatus, which might be useful to define its certain developmental stages. We underlined those characters that were consistent during fish development, which could be valuable to identify the species. Considering the ontogenetic pattern of otolith in A. furcatus, it was found that the otoliths of young individuals have largely been influenced by sexes and that it was reduced during the fish development. Morphological side dimorphism in the otolith of A. furcatus tends to be increased during fish development. The outcomes of this study are essential since they have provided new information on ontogenetic changes and the sexual and side dimorphism of Aphanius otolith.     

Geographic variation in otolith morphology among freshwater populations of Aphanius dispar (Teleostei,Cyprinodontiformes) from the southeastern Arabian Peninsula

Aphanius Nardo is a large genus of teleost fishes in the Old World, with 19 described species. Several of these species have only recently been recognized and additional species can be expected from isolated populations in remote areas. We show here that otolith morphology and statistical analyses of otolith variables can contribute to the detection of genetic differentiation in Aphanius. We studied samples of eight Aphanius dispar populations from the southeastern part of the Arabian Peninsula. Two populations originate from freshwater habitats far inland and probably have been isolated since the late Holocene some 4,000 years ago, three populations come from freshwater habitats with occasional connections to the sea, one population originates from a coastal site, and two were artificially introduced populations. The coastal population is interpreted as possessing the basic otolith type of A. dispar. The basic otolith type also occurs in fishes from the freshwater habitat which is located closest to the coast. Otoliths from the two other freshwater populations with occasional connections to the coast differ slightly from the basic type. However, the two populations from the long‐term isolated freshwater habitats far inland show distinct morphological changes. Our results are consistent with the hypotheses that i) otolith morphology is primarily genetically determined and is little influenced by physical parameters of the habitat, and ii) isolated A. dispar populations may be capable of evolving into new species within short periods of time. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Microstructural characterization of the body key scale morphology in six Iranian endemic <Emphasis Type="Italic">Aphanius</Emphasis> species (Cyprinodontidae): Their taxonomic and evolutionary significance

The microstructural characteristics of the body key scales were described for six Iranian endemic Aphanius Nardo, 1827 species. Among them, five species genetically belong to the “Iranian inland and inland-related Aphanius species, IIRAS” group, while A. ginaonis (Holly, 1929) belongs to the “brackish water species” group. General scale shape in the studied Aphanius species was cycloid with the exception of A. ginaonis, which has cycloid scales with few spinous-like structures present in the posterior edge of the scales. Considering phylogenetic relationships of the studied taxa, the most likely explanation for presence of the spinous scales in Aphanius species is the primary existence of these scales in this group. In addition, among the IIRAS group, the most different lepidont was recognized in scales of the A. isfahanensis Hrbek, Keivany and Coad, 2006 (pointed), which is similar to lepidonts in the A. ginaonis. The scales of these species are polygonal with similar typology. The similarity in lepidont morphology between A. ginaonis and A. isfahanensis can explained by (i) the primary existence of the rounded lepidont in this group, and (ii) since A. ginaonis and A. isfahanensis belong to two separate phylogenetic lineages, therefore, similarity in their lepidont morphology could be results of convergent evolution.     

Phylogenetic analysis of Aphanius from the endorheic Kor River Basin in the Zagros Mountains,South‐western Iran (Teleostei: Cyprinodontiformes: Cyprinodontidae)

Morphologically similar populations of Aphanius that are currently considered as A. sophiae inhabit the endorheic Kor River Basin in the Zagros Mountains. Using genetic analysis based on mtDNA (cytochrome b), combined with examination of morphology (morphometry, meristics, otoliths), we discovered that what is thought to be A. sophiae is actually two distinct species, one of which is described as A. shirini sp. n. The males of the new species can be distinguished from those of all other Iranian inland Aphanius species by having only 7–10 clearly defined white flank bars, which is the lowest number of flank bars among the Iranian inland Aphanius species. Both males and females differ from all other Iranian inland Aphanius species by having a significantly longer caudal peduncle and a smaller dorsal fin depth. Based on the PhyML and Bayesian likelihood trees, A. shirini is sister to A. vladykovi from the Karoun Basin in the Zagros Mountains. Our results indicate that an ancient exorheic Kor River Basin existed in the Late Miocene and Pliocene. The close phylogenetic relationship between A. shirini and A. vladykovi suggests that the pre‐Pliocene drainage in the ancient Kor River Basin was directed to the north‐west (to the Karoun Basin), and not to the south‐east as in the present‐day Kor Basin. Both A. shirini and A. vladykovi represent the highest altitude records for Aphanius. We conclude that the splits of A. shirini and A. vladykovi can be linked to tectonic events in the Middle to Late Miocene, which created the highest altitudes (>3000 m) in the Zagros Mountains, and led to isolation of populations. The present‐day endorheic Kor Basin is known to have formed in the Late Pleistocene or Early Holocene, and the ‘young’ age of A. sophiae is clearly related to this history. Our results contribute to elucidate the link between geological history and the present‐day species diversity in the tectonically still active Zagros Mountains of Iran.     

Molecular confirmation of the occurrence of Elysia cf. tomentosa (Mollusca: Heterobranchia) in the Persian Gulf

The molluscan fauna of the Persian Gulf has recently been relatively well documented, yet there are few records of heterobranch sea slugs (opisthobranchs) from the Arabian parts and no report from the Iranian waters. Here we report for the first time the occurrence of one of these molluscs in the northern Persian Gulf (Bandar Abbas, Iran). Sacoglossan specimens were collected in association with the seaweed, Caulerpa sertularioides. Since morphological attributes were not adequately reliable for species identification, molecular approaches were carried out. Maximum-likelihood and Bayesian Inference analysis of partial DNA sequences of the mitochondrial cytochrome c oxidase subunit I (COI) locus were used for DNA barcoding of large-bodied specimens of Elysia. All Persian Gulf specimens were genetically confirmed as Elysia cf. tomentosa sp. 5, one of at least five morphologically similar but genetically distinct species in the taxonomically challenging and unresolved E. tomentosa complex. This species has previously been recorded only from Australia and Thailand and our finding adds another distant point to the geographic distribution of this species.     

Hidden taxonomic characters in otoliths of blenniid fishes (Teleostei: Blenniidae) from the Iranian coasts of the Persian Gulf and Oman Sea ecoregions: A scanning electron microscopy approach

In the present work, the morphological characteristics of sagitta were described and analysed for 10 species of combtooth blennies (Blenniidae) inhabiting the intertidal region of the Persian Gulf and Makran zone of Oman Sea to see the taxonomic usefulness of otoliths, especially their hidden characters for species identification. Based on the obtained results, sagitta morphology could be considered as diagnostic characters distinguishing the genera Alticus, Antennablennius, Istiblennius, Omobranchus and Parablennius showing differences related to general shape, that is, elliptic/oval to triangular; shape of antirostrum and its degree of development; the degrees of convexity of ventral and dorsal margins, and shape of ostium. The common/general otolith characters shared in the all studied combtooth blennies, is the heterosulcoid condition, which is a shared character with Perciformes and Gobiiformes. The type of sulcus is ostial, and it can be considered as a plesiomorph character shared with a basal group of actinopterygian fishes. The obtained results could be useful in taxonomical, paleontological, ecological and food and feeding habits studies.     

Comparative otolith morphology of clupeids from the Iranian brackish and marine resources (Teleostei: Clupeiformes)

The diagnostic features of otolith morphology were provided for the clupeid fishes of the Iranian brackish and marine resources to be used as diagnostic features for the identification of clupeids diversity in these regions. Fish individuals belong to 20 species belong to 13 genera, and four families were collected from the Caspian Sea, the Persian Gulf and the Makran zone of the Oman Sea. Overall, seven otolith morphotypes were distinguished, that is lanceolated (45%), fusiform (20%), boot-like (15%) and clamp-like, pentagonal, elliptic and rectangular (each 5%, respectively). The univariate analysis showed that all variables except shape index [sulcus area (SS)/otolith area (OS)] and ROx (roundness) were significantly different among the clupeid species. The interspecific relationships of the otoliths were analysed based on the data of otolith morphology and otolith shape indices. Both dendrograms showed somehow an obvious separation among the studied species. However, the interspecific relationships in the dendrogram based on the otolith morphology have a better resolution. These phenotypic relationships based on otolith morphology among the studied clupeids are largely consistent with the previous hypothesis on the systematics of these fishes and emphasized that the morphological and morphometric features of the otolith, which are highlighted in this study, could be adequately used as diagnostic features for the identification of clupeids diversity.     

Geographical differentiation of Aphanius dispar (Teleostei: Cyprinodontidae) from Southern Iran

The Arabian cyprinodontid Aphanius dispar (Rüppell, 1829) is known to show considerable morphological variation. It has remained unknown, however, whether this variation is a result of environmental differences or allopatric divergence owing to geographical isolation. In this study, 11 populations of A. dispar from three geographically separated basins were analysed, that is, the Makran Basin (I, one river system), the Hormuzgan Basin (II, five rivers and three hot springs) and the Helleh Basin (III, two hot springs) in southern Iran. Statistical analyses do not indicate significant differences between the fishes from river and hot spring habitats (T‐test, p < 0.05), which is also supported by the Canonical Discriminant Analysis (CDA). Nevertheless, morphometric and meristic characters of the fishes, as well as otolith morphology and morphometry, demonstrate that six phenotypic characters discriminate the A. dispar populations of the three basins, that is, (1) predorsal distance (Prdd.SL), (2) head length (HL.SL), (3) pelvic fin length (Lplf.SL), (4) number of pelvic fin rays, as well as relative length of both the (5) medial part and (6) rostrum of the otolith. In addition, these characters display a consistent pattern of variation, thus providing support for the assumption that the phenotypically different A. dispar populations are a result of geographical isolation and not related to environmental differences. It is likely that the geological history of the drainage systems caused isolation event(s) that may date back to the Pleistocene (1.8 million years before present). The high phenotypic differences might suggest that the A. dispar populations from the three studied basins represent separate subspecies or even species.     

Genetic connectivity and phenotypic plasticity in the cyprinodont Aphanius farsicus from the Maharlu Basin,south‐western Iran

Meristic and morphometric characteristics, including otolith data, of the Farsi tooth‐carp Aphanius farsicus, which is endemic to the endorheic Maharlu Basin in south‐western Iran, were analysed for a sample of 92 individuals from four spring‐streams; DNA sequence data (cytochrome b gene) are presented for 29 specimens. Some phenotypic variation was detected but the genetic data clearly indicate connectivity between the populations. Possible links between phenotypic variation and environmental variables such as water temperature, habitat size and absence or presence of predators and competitors are discussed. Based on a literature survey and the new data, it is concluded that population connectivity is maintained during times of droughts via large aquifers that formed during the late Pliocene to early Pleistocene, when the extant endorheic Maharlu Basin was created. Based on new data presented here and previous work, it is apparent that plastic and constant characteristics are present in Aphanius species, and that, if a population becomes isolated, a given trend of evolution may give rise to a taxonomically useful characteristic.     

Length‐weight relationships of six fishes from intertidal and coastal waters in the northern Persian Gulf

The length–weight relationships (LWRs) for six little‐known fish species collected in the northern Persian Gulf (south of Iran) are presented, namely, Plicofollis dussumieri, Pseudotriacanthus strigilifer, Cynoglossus bilineatus, Cynoglossus arel, Solea elongata, and Pseudorhombus elevates.     

Using otolith shape and morphometry to identify four Alburnus species (A. chalcoides,A. escherichii,A. mossulensis and A. tarichi) in Turkish inland waters

Asteriscus otolith shapes as well as their morphometry and shape contours were investigated in order to identify four allopatric Alburnus species: A. chalcoides (Güldenstädt, 1772) (Ordu), A. escherichii Steindachner, 1897 (Eski?ehir), A. mossulensis Heckel, 1843 (Tunceli), and A. tarichi (Güldenstädt, 1814) (Van) in Turkish inland waters. These were compared using the shape indices (form factor, roundness, circularity, ellipticity, rectangularity and aspect ratio), and the morphological characters [otolith weight (OWE), otolith length (OL), otolith width (OW), otolith perimeter (OP), and otolith area (OA)]. The overall canonical discriminant analysis (CDA) classification score was 93.8%, with the lowest score for A. escherichii (82.5%) and the highest for A. chalcoides (100%). The otolith shapes, morphology and shape contours of all sampled fish were a clear species differentiator, thereby demonstrating that the otolith shape is species‐specific. The current study presents for the first time comprehensive variation information on interspecific left‐right asteriscus otoliths in males and females of each Alburnus species: A. chalcoides from Ordu, A. escherichii from Eski?ehir, A. mossulensis from Tunceli and A. tarichi from Van, based on a total of 307 individuals. Scanning electron microscopy (SEM) images, shape contours and other otolith characters vary within the same genus; these differences should be investigated not only in other freshwater fish species or genera but also in the same species living in different habitats. In addition, further investigation is required not only with respect to the morphometry, biometry, shape, geometry, and shape contours of the otoliths, but also regarding the genetic methods for robust identification of various sympatric and allopatric fish populations.     

Morphology and microchemistry of abnormal otoliths in the ayu, <Emphasis Type="Italic">Plecoglossus altivelis</Emphasis>

The sagittal otolith morphology and microchemistry of reared juvenile ayu, Plecoglossus altivelis, were examined to describe the occurrence and microchemical characteristics of the abnormal otoliths in this species. Juvenile ayu (N = 31) were collected in June 2004 at three different locations, Wakayama, Kumamoto, and Biwa Lake in Japan, where they were being reared in freshwater aquaculture ponds after having been collected in the wild as larvae. Otolith abnormality was found in the sagittae of 26% (N = 8) of the individuals examined, of which five fish had abnormal otoliths only on one side, while the otolith on the other side was normal. Abnormal otoliths were more transparent and crystalline in appearance with irregular shapes compared to normal ones that were more opaque and less irregular. Abnormal otoliths were divided into two types, semi-abnormal (Type 1) with a normal part in the center, and fully-abnormal (Type 2) that were completely crystalline in appearance. The line transects and whole otolith concentration maps showed that the contents of Sr, Na and K were lower in the abnormal otolith regions compared to the normal ones, while those of Ca and S were almost constant in both. The appearance and microchemical properties of the abnormal ayu otoliths were similar to the abnormal otoliths in other species in which vaterite replaces the aragonite. Abnormal formation of otoliths occurred in ayu from Biwa Lake (30%) and Kumamoto (45%), while the Wakayama samples had no abnormality. The microchemical analyses of the normal and abnormal otoliths indicated that some abnormal otoliths had formed before the fish were captured and transferred to the hatchery, so the possible causes of otolith abnormality in ayu are discussed.     

Length–weight and length–length relationships for three Sillago species (Sillaginidae) from the Persian Gulf

This study investigates length–length and length–weight relationships of three Sillago species [Sillago arabica McKay & McCarthy (Memoirs of the Queensland Museum, 27, 1989, 551), Sillago attenuata McKay, 1985 and Sillago sihama (Forsskål, 1775)] captured in the Persian Gulf, Iran. A collection of 220 specimens was captured from August 2015 to January 2016 by cast net (mesh size: 10 mm) and beach seine (mesh size: 15 mm). The relationships of total length (TL) and standard length (SL) and the relationships between total length and body weight are given. This study presents the first reference on LWRs for two of the species (S. arabica and S. attenuata) in their range of distribution.     

Validation of yellowedge grouper, <Emphasis Type="Italic">Epinephelus flavolimbatus</Emphasis>, age using nuclear bomb-produced radiocarbon

Age validation and estimates of longevity of yellowedge grouper (Epinephelus flavolimbatus) from the Gulf of Mexico (GOM) are needed to inform fishery management decisions. Yellowedge grouper sagittal otoliths (n = 100) were collected, aged using conventional means, and cores were submitted for radiocarbon (¹⁴C) measurement. Radiocarbon values of yellowedge grouper otoliths were compared to established radiocarbon chronologies in the region to validate the age and ageing methodology of this species. The yellowedge grouper chronology displayed a similar sigmoidal trend as previously published chronologies. In addition to the core analysis, multiple areas on otolith sections from eight specimens were analyzed for Δ¹⁴C to validate age estimates for fish born prior to the ¹⁴C increase. Our results indicate that yellowedge grouper live longer than previously reported (minimum of 40 years based on radiocarbon measurements). The validated ageing methodology supported an estimated maximum longevity of 85 years and established that yellowedge grouper have the longest lifespan currently known for any species of grouper in the GOM. Results also indicate a depth-age interaction in that material extracted from adult otolith sections assigned to post-bomb dates exhibited lower Δ¹⁴C values than cores (juvenile material) assigned to the same post-bomb dates. This finding is likely explained by lower ¹⁴C levels reported from water masses at deeper depths (>100 m) which are inhabited by adults.     

Length–weight relationships of five commercial fish species from the Strait of Hormuz in the Persian Gulf

Length‐weight relationships are described for five fish species from the coast of the Hormuz Strait in the Persian Gulf, Iran. Specimens were collected between August 2010 and July 2011 using shrimp beam trawls, gill nets and intertidal fishing weirs. Relationships for three of the species had no previous data in the international science global databank on fishes: www.FishBase.org . A new maximum length was recorded for Acanthopagrus latus.     

Length–weight relationships for five stingray species from the Persian Gulf

Length–weight relationships (LWR) of fishes are necessary for various purposes such as stock/ecological assessment, monitoring and management (Froese, 2006). In this study we report length–weight relationships for five species of stingrays found in the Persian Gulf, Iran. Only one LWR was previously reported in the scientific literature for Rhinoptera javanica (Raje, 2000).     

Length–weight relationships of four fish species caught in the Northern Persian Gulf (Horomzgan waters,Iran)

Length–weight relationships (LWRs) for four little‐known fish species collected in the northern Persian Gulf (south of Iran) are presented, namely, Leiognathus lineolatus, Grammoplites suppositus, Eupleurogrammus muticus and Acanthocepola abbreviate. Thus far, these are the first LWRs for these species in the international scientific literature.