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Molecular Systematics, Zoogeography, and Evolutionary Ecology of the Atlantic Parrotfish Genus Sparisoma
Abstract
Parrotfishes of the genus Sparisoma (Scaridae) are ecologically important tropical reef fishes restricted to the Atlantic Ocean. We investigated phylogenetic relationships among the eight extant species within this genus using mitochondrially encoded 12S and 16S ribosomal genes. Our molecular data support the view that (i) Sparisoma originated approximately 14-35 million years ago (mya), probably in the tropical western Atlantic, off Brazil; (ii) there have been at least four discrete bouts of cladogenesis within the genus, with the most recent one ( approximately 2.8-5.6 mya) involving four events in both the east and the west Atlantic and across the Atlantic; and (iii) the genus invaded the eastern Atlantic on two different occasions, probably by at least two different routes. The data also offer support for Bellwood's ideas concerning the evolutionary changes in adult feeding patterns and habitat use within Scarids. Specifically, they support the evolutionary position of the ecological traits of Sparisoma as intermediate within the family.
Supplementary resources (16)
... The evolutionary history of parrotfishes has been previously studied using molecular (Bernardi et al., 2000;Streelman et al., 2002;Smith et al., 2008;Cowman et al., 2009;Price et al., 2010;Choat et al., 2012;Siqueira et al., 2019a) and morphological information (Bellwood, 1994). These studies have identified a clear break between two monophyletic groups within the subfamily, the "reef " clade scarinine (Scarus, Chlorurus, Bolbometopon, Hipposcarus, and Cetoscarus), and the "seagrass" clade sparisomatine (Sparisoma, Nicholsina, Leptoscarus, Calotomus, and Cryptotomus) (Streelman et al., 2002;Westneat and Alfaro, 2005;Smith et al., 2008;Choat et al., 2012;Aiello et al., 2017;Siqueira et al., 2019a). ...
... The "reef " clade (scarinine) is more taxonomically diverse (∼75 species) with two main genera accounting for the majority of species (Scarus 52 and Chlorurus 18); species of this group feed either by scraping or excavating and are distributed in all tropical oceans. The "seagrass" clade (sparisomatine) has far fewer species (∼25 species) with one genus accounting for the majority of species (Sparisoma 15; Froese and Pauly, 2017); in sparisomatine, species are performing each of the three feeding modes, occurring in all tropical oceans with most of the species distributed in the western Atlantic (Bernardi et al., 2000;Taylor et al., 2018). ...
... The separation between Scarus and Sparisoma is strongly supported, with a posterior probability (PP) of 1. Our phylogenetic reconstruction of the evolutionary relationships of 52 parrotfish species (Figure 3) concurs almost completely with the topology and divergence of previous works (Bellwood, 1994;Bernardi et al., 2000;Smith et al., 2008;Choat et al., 2012;Aiello et al., 2017;Siqueira et al., 2019a). The recovered estimated age for the genus Scarus was 8.02 mya ago (HPD: 5.01-10.9), ...
In functional ecology, morphology is expected to reflect function; however, occasional decoupling of these two can be found. In the case of feeding ecology, the diversity of the diet or diversity of the feeding modes within a clade is expected to be positively related to the diversity of the morphological traits involved in the feeding performance. Parrotfishes are separated into two main groups, the “reef” clade and the “seagrass” clade. Both groups have important differences in their evolutionary history. Still, more interestingly, they have important morphological and ecological differences. The genera Scarus and Sparisoma are the most specious genera of parrotfishes. They belong to each of those main groups, respectively. All Scarus species have the same feeding mode, while in Sparisoma, there are three different feeding modes. We want to test if the morphological jaw diversity of these genera corresponds with the diversity in their feeding modes. Using a disparity analysis of feeding traits within a phylogenetical framework, we did not find a relationship between functional feeding morphology and feeding modes of the American parrotfishes of the genera Scarus and Sparisoma. Interestingly we found that some muscular traits are the source of the high disparity in the genus Scarus. We explore some possible morpho-functional reasons for this phenomenon and reappraise the parrotfishes’ scraper feeding mode’s functional diversity. We also consider that there could be more ecological differentiation between Scarus species that we are aware of. Using an ancestral reconstruction of feeding modes of 52 species of parrotfishes, we found that the scraping feeding mode exhibited by all Scarus species is an evolutionary convergence with the scraping feeding modes performed by some Sparisoma species. Different selective pressures or ecological conditions may have shaped the differences in the feeding ecology and the feeding morphology of these two genera. Probably, key novel structures and muscular properties found in the Scarus species’ jaw played an essential role in this genus’s morpho-functional diversification. Finally, we propose that feeding modes may not fully capture the complexity of feeding ecology in parrotfishes.
... According to this phylogeny, Bellwood described a progressive emergence of coalesced dentitions from non-coalesced dentitions. Sparisoma dentitions are viewed as a transitional form between non-coalesced dentitions and the group constituted by Cetoscarus, Bolbometopon, Hipposcarus, Chlorurus and Scarus Bernardi et al., 2000). The transition to coalesced dentitions is described as (1) a flattening of the anterior caniniform teeth present in Cryptotomus to form teeth similar to Calotomus, (2) progressive coalescence in Leptoscarus, (3) increase of the depth of dental plates and teeth as in Sparisoma, (4) enlargement of teeth in Bolbometopon and Cetoscarus, (5) entire coalescence in most coalesced dental plates are formed by an anterior dental fusion i.e. by teeth homologous to the anterior teeth of non-coalesced dentitions. ...
... We found equivalent results on the tooth structure with a strong similarity between the teeth of Calotomus carolinus and other Labridae (Berkovitz & Shellis, 2017) while teeth of tightly-organized dentitions present reduced dentine. With his phylogeny based on morphological characters, proposed that dental plates progressively evolved from non-coalesced dentitions, with Sparisoma species being considered as transitional forms (Bernardi et al., 2000). On the contrary, the phylogenetic analyses based on molecular data Kazancıoğlu et al., 2009;Choat et al., 2012;Baliga & Law, 2016) confirmed the separation of parrotfishes proposed by Schultz (1958) into two tribes: the Sparisomatini and the Scarini. ...
... Sparisoma are classified as excavators or scrapers Bernardi et al., 2000). The tight organization of teeth thus appears as an adaptation for feeding on hard substrate and arises independently in Scarini and Sparisoma. ...
Parrotfishes, divided into the tribes Scarini and Sparisomatini , are labrid fishes that arouse from carnivorous ancestors. One of the main factors that explain this radiation is the exploitation of microorganisms as a food source. While most Sparisomatini browse macroalgae to get epiphytic microorganisms, grazing parrotfishes (all Scarini and some Sparisoma) scrap or excavate hard substrate to obtain epilithic and endolithic microorganisms. Changes in feeding behaviour are associated with dentition specialization with notably dental plates in grazing but also some browsing parrotfishes. Gene-based phylogenies contradicted the previous evolutionary scenario about parrotfish dentition evolution, which states the progressive emergence of dental plates from non-coalesced dentitions. This thesis manuscript aims to re-examine the evolution of parrotfish dentitions in the light of the new phylogenies by using X-ray 3D microtomography, and to link this evolution with diet and ecology.
... As a result, many authors have recommended the combination of different, but complementary sampling methods to accurately study fish assemblages inhabiting seagrass meadows (Harmelin-Vivien and Francour 1992;Bobsien and Brandelberger 2006). The parrotfish Sparisoma cretense (Linnaeus, 1758) is one of the few temperate species of the Scaridae family (Bernardi et al. 2000). This species is distributed in the eastern coasts of the Mediterranean Sea, the Macaronesian archipelagos (Azores, Madeira, Canaries and Cape Verde) and the northwest coast of Africa (González 1993). ...
... This is supported by, firstly, the presence of adult parrotfish inhabiting shallow-water rocky reefs in the study region (Tuya et al. 2006b). Secondly, a similar pattern has also been detected in the Mediterranean, where S. cretense is found on rocky reefs, feeding on macroalgae, yet occasional displacements towards seagrass meadows dominated by Posidonia oceanica can also occur to feed on seagrass epiphytes (Bernardi et al. 2000;Kalogirou et al. 2010). In tropical areas, distinct parrotfish species inhabit coral reefs; intermittent displacements of adults towards neighboring seagrasses are common to feed (Randall 1965;Ogden and Zieman 1977;Tribble 1981;Macintyre et al. 1987). ...
Seagrass habitats are relevant for numerous nearshore fish species, particularly as nursery grounds. Seagrass meadows are often interspersed with other habitats, what can alter the distribution and abundance of seagrass ichthyofauna. This research aimed to determine whether there is a change in the abundance and biomass of the parrotfish, Sparisoma cretense, in seagrass meadows (Cymodocea nodosa) with varying proximity from rocky reefs, specifically seagrass interiors (>200 m away from reefs) vs. seagrass adjacent to reefs (<10 m away). Sampling was undertaken using a seine net and underwater visual census through an entire annual cycle. Adults were predominantly observed in seagrass adjacent to reefs, which seem to be restricted to incursions of large-sized parrotfish from adjacent reefs. Juvenile abundance did not significantly differ between seagrass interiors and seagrass adjacent to reefs; however , juvenile biomass was greater in seagrass meadows adjacent to reefs compared to those far away from reefs. This pattern was consistent through times for both sampling techniques. These results suggest a transition of juvenile parrotfish from seagrass interiors to seagrass near reefs, so juveniles are located in the vicinity of their adult habitat, i.e. rocky reefs.
... Biogeographic history (e.g. Bernardi et al. 2000, Choat et al. 2012, and more recently human exploitation (Mumby 2006, Rasher et al. 2013, Taylor 2014, has shaped the considerable regional variation in the taxonomic and functional composition of parrotfishes. The most pronounced division in the parrotfish assemblage structure is between the Indo-Pacific and Atlantic Oceans, with assemblages in the Atlantic and Indo-Pacific being dominated by seagrass associated (i.e. ...
... Scraping parrotfishes are represented primarily by Scarus and Hipposcarus in the Indo-Pacific (46 of 75 species), and Scarus in the Atlantic (eight of 25 species), and supplemented by smaller individuals of excavating species in both regions. Further, Sp. aurofrenatum has also been recorded to function as a scraper on Caribbean reefs (McAfee and Morgan 1996, Bernardi et al. 2000). The relatively high species diversity within scraping parrotfishes, together with their preference for feeding on EAM covered substratum, suggest the loss of one or two species would not greatly impact the overall process. ...
... El pez loro o 'vieja', S. cretense, es una de las pocas especies templadas de la familia Scaridae (Bernardi et al., 2000). Esta especie se distribuye en las costas orientales del mar Mediterráneo, los archipiélagos de la Macaronesia (Azores, Madeira, Canarias), en las Islas de Cabo Verde, y en las costas del noroeste de África (González, 1993). ...
... Además, la presencia de adultos de S. cretense en praderas de fanerógamas marinas parece estar restringida a incursiones desde los arrecifes adyacentes, probablemente para alimentarse (ver Figura 4.7.a-c). En primer lugar, esta idea es apoyada por la presencia de adultos de peces loro que habitan los arrecifes rocosos someros en la región de estudio (Tuya et al., 2006b); segundo, un patrón similar ha sido detectado en el mar Mediterráneo, donde S. cretense se encuentra en arrecifes rocosos, alimentándose de macroalgas, pero también pueden ocurrir desplazamientos ocasionales hacia las praderas de fanerógamas marinas constituidas por P. oceanica, para alimentarse de los epífitos de las fanerógamas (Bernardi et al., 2000;Kalogirou et al., 2010). En áreas tropicales, los individuos adultos de distintas especies de peces loro que habitan los arrecifes de coral, realizan, de forma común, desplazamientos intermitentes hacia las vecinas praderas de fanerógamas marinas para alimentarse (Randall, 1965;Ogden y Zieman, 1977;Tribble, 1981;Macintyre et al., 1987). ...
El objetivo de la tesis es investigar y conocer los patrones de variabilidad espacio-temporal de las poblaciones de peces asociados a praderas constituidas por la fanerógama marina Cymodocea nodosa. En el Capítulo 1, se realiza un análisis macro-ecológico de las comunidades de peces asociadas a praderas de C. nodosa en toda su área de distribución, excepto en las cotas del norte de África, donde no existen datos; en este trabajo se analiza, por primera vez, la variación en la composición de la ictiofauna a escala biogeográfica; los resultados muestran que la composición de la comunidad de peces en las Islas Canarias difiere significativamente de las demás eco-regiones. En el Capítulo 2, se estudia la variabilidad espacial en la estructura, abundancia y diversidad de la comunidad de peces asociada a praderas canarias de C. nodosa a tres escalas espaciales: entre islas (100s km), entre praderas dentro de islas (10s km) y entre sitios dentro de praderas (100s m); la mayor variabilidad en la estructura de la comunidad se produce entre sitios, seguida por la variación entre praderas, mientras que la menor se registró entre islas. En el Capítulo 3, se estudia la composición y estructura general de la comunidad íctica en tres islas del Archipiélago Canario, así como la función de ‘guardería’ que desempeña este ecosistema en las islas; los resultados muestran que las praderas son un hábitat de cría para numerosas especies de peces litorales, muchas de ellas de interés comercial. En el Capítulo 4, se analizan las variaciones espaciales y temporales de la abundancia y biomasa de una especie emblemática en las Islas Canarias, la ‘vieja’ (Sparisoma cretense (Linnaeus, 1758)), en praderas marinas adyacentes y lejanas a arrecifes rocosos; los resultados sugieren una transición de los ejemplares juveniles desde la zona interior de las praderas hacia la zona adyacente a los arrecifes rocosos que constituirán su hábitat de adulto. En el Capítulo 5, se estudia la influencia de las características del hábitat de mosaicos de pradera-arena sobre la estructura poblacional del ‘pejepeine’ (Xyrichtys novacula (Linnaeus, 1758)); los resultados muestran que la variación espacial de la abundancia y biomasa de esta especie se explica por un equilibrio entre la protección provista por la bóveda foliar y la protección derivada de su comportamiento de enterramiento. En el Capítulo 6, se realiza una estimación del valor económico de los stocks de peces de interés pesquero en praderas de C. nodosa de Gran Canaria, el valor económico de los stocks pesqueros asociados fue relativamente alto comparado con los obtenidos en otras áreas del mundo. Por último, en el Capítulo 7, se establecen las relaciones talla-peso para diez especies de peces teleósteos que habitan las praderas de fanerógamas en las Islas Canarias; los resultados de este trabajo contribuyen a mejorar los conocimientos sobre la ecología de estas especies y la gestión de sus stocks poblacionales.
... The lineage that led to the two eastern Atlantic species, S. cretense (presently reported from the Mediterranean Sea and from the eastern Atlantic shores of mainland Portugal, the Azores, Madeira, Canaries, Cabo Verde and Senegal; cf. Table 2) and S. strigatum (Günther, 1862) (endemic to Saint Helena and Ascension islands; Table 2), originated at c. 10 Ma from a west Atlantic ancestor (Bernardi et al. 2000). The oldest known fossil specimens of S. cretense were collected in two Pliocene outcrops from Santa Maria Island (Pedra-que-pica and Ponta dos Frades) and dated as 4.78 ± 0.13 to 4.13 ± 0.19 Ma in age (Table 1); our data are therefore in agreement with the estimates of Bernardi et al. (2000). ...
... Table 2) and S. strigatum (Günther, 1862) (endemic to Saint Helena and Ascension islands; Table 2), originated at c. 10 Ma from a west Atlantic ancestor (Bernardi et al. 2000). The oldest known fossil specimens of S. cretense were collected in two Pliocene outcrops from Santa Maria Island (Pedra-que-pica and Ponta dos Frades) and dated as 4.78 ± 0.13 to 4.13 ± 0.19 Ma in age (Table 1); our data are therefore in agreement with the estimates of Bernardi et al. (2000). ...
Fossil fishes are among the rarest in volcanic oceanic islands, their presence providing invaluable data for the understanding of more general (palaeo)biogeographical patterns and processes. Santa Maria Island (Azores Archipelago) is renowned for its palaeontological heritage, with representatives of several phyla, including the Chordata. Herein, we report on the fossil fishes, resulting in an increase of the number of Pliocene fishes from the Azores to 11 taxa: 7 Chondrichthyes and at least 4 Actinopterygii. The genus Sparisoma is reported for the first time in the fossil record. The presence of fossil remains of the parrotfish Sparisoma cretense in Last Interglacial outcrops is significant, because it posits a setback for the theory that most of the present-day Azorean marine species colonized the area after the last glacial episode. Our multidisciplinary approach combines palaeontological data with ecological and published genetic data, offering an alternative interpretation. We suggest that most of the Azorean shallow-water subtropical and temperate marine species living in the archipelago during the Last Interglacial were not affected by the decrease in sea surface temperatures during the last glacial episode. We also predict low genetic diversity for fish species presently living in the Azores and ecologically associated with fine sediments, due to the remobilization and sediment transport to abyssal depths, during the Last Glacial episode; these are viewed as post-glacial colonizers or as ‘bottleneck’ survivors from the Last Glaciation.
... A number of tropical reef fish phylogenies have been inferred from molecular data (e.g. Bernardi and Crane 1999;Tang et al. 1999;Bernardi et al. 2000;Craig et al. 2001;Tang 2001;Wang et al. 2001;Lo Galbo et al. 2002;Streelman et al. 2002;Clements et al. 2003;Ruber et al. 2003;Thacker 2003). Most of these examined high-level relationships using mtDNA (sequencing 1 -4 gene regions) rather than species-level relationships. ...
... Some of the studies (e.g. Bernardi et al. 2000;Streelman et al. 2002;Ruber et al. 2003) examined specific features, such as foraging modes, in relation to the phylogeny obtained, an issue also examined and addressed in this study. Two genus-level molecular phylogenies for the family Acanthuridae (surgeonfishes) have been published (Tang et al. 1999;Clements et al. 2003). ...
... Nonetheless, the color patterns commonly used to a precise identification of Sparisoma species are best observed in fresh specimens only, and they might vary according to sex and ontogenetic state (Bernardi et al., 2000). Thus, the utilization of genetic markers should represent an efficient and reliable tool to identify these species, independent of the morphological aspects. ...
... In spite of the scarcity of cytogenetic data about Scarinae, the currently available reports suggest that repeated pericentric inversions and centric fusions have taken place during the evolutionary history of this subfamily. Such a divergent karyotypic trend in relation to other marine Perciformes might have been favored by vicariance events combined with habitat differentiation and specialization of parrotfishes (Bernardi et al., 2000;Robertson et al., 2006). To test this hypothesis, additional cytogenetic analyses in other Scarinae representatives are highly encouraged. ...
Parrotfishes (Labridae, Scarinae) comprise a large marine fish group of difficult identification, particularly during juvenile phase when the typical morphology and coloration of adults are absent. Therefore, the goal of this study was to test cytogenetic markers and DNA barcoding in the identification of bucktooth parrtotfish Sparisoma radians from the northeastern coast of Brazil. Sequencing of cytochrome c oxidase subunit I (COI) confirmed all studied samples as S. radians, and all showed high similarity (99-100%) with Caribbean populations. The karyotype of this species was divergent from most marine Perciformes, being composed of 2n = 46 chromosomes. These consisted of a large number of metacentric and submetacentric pairs with small amounts of heterochromatin and GC-rich single nucleolar organizer regions (NORs) not syntenic to 5S rDNA clusters. These are the first data about DNA barcoding in parrotfish from the Brazilian province and the first refined chromosomal analysis in Scarinae, providing useful data to a reliable genetic identification of S. radians.
... These genera are among the most representative groups of the Scaridae, aggregating almost 70% of all species. Sparisoma, recorded only from the Atlantic, is the most speciose scarid genus in the Atlantic (Bernardi et al., 2000;Streelman et al., 2002). The species presently know from Brazil are S. amplum, S. axillare and S. frondosum (Moura et al., 2001), and more recently a new species, S. tuiupiranga (Gasparini et al., 2003). ...
... In spite of the importance of the Scaridae, there are no karyotypic data on this group. The phylogenetic position of the Sparisoma genus has been considered intermediary within the family, based on analyses of mitochondrial DNA with ribosomal gene probes 12S and 16S (Bernardi et al., 2000). To date about 8% of the species in the Perciformes have been karyotyped, revealing a modal diploid number of 2n = 48 chromosomes (Ohno, 1970;Le Grande and Fitzsimons, 1988;Affonso et al., 2001). ...
The parrotfishes (family Scaridae) are comprised of the subfamilies
Sparisomatinae and Scarinae. They are important agents of marine
bioerosion, which rework the substrate with their beaklike jaws. Despite
their importance, there are no published cytogenetic data on this group. We
made cytogenetic analyses of Sparisoma axillare (Sparisomatinae) and Scarus
coelestinus (Scarinae) from the Brazilian coast. Differentiation in the
diploid number in S. axillare compared to the basal karyotype of the Perciformes
apparently occurred due to a Robertsonian fusion, combined with
pericentric inversions. S. coelestinus presented a conserved diploid number,
but showed considerable structural karyotypic changes, resulting mainly
from pericentric inversions. The Ag-NOR sites were unique and located on
the short arm of the 1st subtelocentric pair in both species (possibly homeologous),
corresponding to the 11th pair in S. axillare and the 9th pair in
S. coelestinus. The constitutive heterochromatin is reduced in these species
and is distributed in centromeric and pericentromeric regions in most of thechromosomes. The low fundamental number compared to the Scarus genus
suggests a more basal condition for Sparisoma. The chromosome formula
in S. coelestinus was more diversified, deriving from large-scale pericentric
inversions. Karyotypic evolution patterns observed for these representatives
of the Sparisomatinae and Scarinae subfamilies, added to new data from a
larger number of species, would allow us to determine if there is a tendency
among the Sparisomatinae for centric fusion events.
... Moura, Figueiredo & Sazima 2001]. These fishes are of tropical origin (Bernardi et al., 2000;Streelman et al., 2002), encompass both scraper and excavator feeding modes (Bellwood & Choat, 1990;Choat et al., 2002), and do not present any characteristics of a fermenting-like species [e.g. acid stomach, long gut or a hindgut fermentation chamber: Lobel (1981); Horn (1989)]. ...
... Contrasts among the assemblages of herbivores observed in this study are similar to the described latitudinal variations in species distributions along the Brazilian coast (Ferreira et al., 2004;Floeter et al., 2005). On the Brazilian coast, fishes with tropical origins from the genera Sparisoma, Scarus and Acanthurus (Bernardi et al., 2000;Streelman et al., 2002) decrease in abundance towards higher latitudes (Ferreira et al., 2004). Only Kyphosus spp. ...
Assemblages of roving herbivores were consistently different between eastern, warmer, sheltered sites and western, colder, more wave-exposed sites. At eastern sites, detritivorous-herbivorous species dominated while omnivores had the highest biomass and were dominant at western sites. Macroalgivores did not show any trends related to location. These distributional patterns, at relatively small spatial scales of a few kilometres, mirror large-scale latitudinal patterns observed for the studied species along the entire Brazilian coast, where cold water associated species are abundant on south-eastern rocky reefs (analogous to the western sites of this study), and tropical species are dominant on north-eastern coral reefs (analogous to the eastern sites). Species-level analyses demonstrated that depth was an important factor correlated with biomasses of Diplodus argenteus, Sparisoma axillare and Sparisoma tuiupiranga, probably due to resource availability and interspecific competition. Herbivorous fish assemblages in the study area have been historically affected by fishing, and combined with the variation in assemblage structure, this is likely to have important, spatially variable effects on the dynamics of benthic communities.
... Yet, there are no studies on the behavior of these species on other reef habitats. Parrotfishes of the Sparisoma genus are recorded only in the Atlantic Ocean, being the most speciose scarid genus in this ocean basin (Bernardi et al. 2000;Streelman et al. 2002). This genus is ecologically diverse, with a wide range of feeding modes and patterns of habitat use (Bernardi et al. 2000;Streelman et al. 2002;Bonaldo et al. 2006). ...
... Parrotfishes of the Sparisoma genus are recorded only in the Atlantic Ocean, being the most speciose scarid genus in this ocean basin (Bernardi et al. 2000;Streelman et al. 2002). This genus is ecologically diverse, with a wide range of feeding modes and patterns of habitat use (Bernardi et al. 2000;Streelman et al. 2002;Bonaldo et al. 2006). ...
Structural complexity is a key parameter for fish on reef habitats. Several studies have investigated the influence of this variable on aspects of reef fish population and community dynamics. However, there is a lack of knowledge on the influence of structural complexity on antipredator behavior. Here we studied the effect of habitat type and structural complexity on flight initiation distance (FID) and the escape behaviors of four labrid fishes (Halichoeres brasiliensis, H. penrosei, H. poeyi and Sparisoma axillare) on two different reef habitats (coral and rocky reefs). Habitat type influenced the FID of three of the studied species (H. brasiliensis, H. penrosei and S. axillare), and structural complexity negatively influenced the FID of two species (H. brasiliensis and S. axillare). The frequency of escape behaviors varied between species. All of them showed high frequency of the ‘run away’ behavior and low frequency of the ‘leave the habitat’ behavior. On coral reefs, structural complexity influenced the ‘fled to the holes’ for S. axillare only. Reef ecosystems worldwide are being modified by anthropogenic activities. Our results suggest that if such activities reduce structural complexity, then ‘seascapes of fear’ for labrid fishes will become more widespread, which could lead to negative consequences in the reef ecosystems.
... Nonetheless, the color patterns commonly used to a precise identification of Sparisoma species are best observed in fresh specimens only, and they might vary according to sex and ontogenetic state (Bernardi et al., 2000). Thus, the utilization of genetic markers should represent an efficient and reliable tool to identify these species, independent of the morphological aspects. ...
... In spite of the scarcity of cytogenetic data about Scarinae, the currently available reports suggest that repeated pericentric inversions and centric fusions have taken place during the evolutionary history of this subfamily. Such a divergent karyotypic trend in relation to other marine Perciformes might have been favored by vicariance events combined with habitat differentiation and specialization of parrotfishes (Bernardi et al., 2000;Robertson et al., 2006). To test this hypothesis, additional cytogenetic analyses in other Scarinae representatives are highly encouraged. ...
Parrotfishes (Labridae, Scarinae) comprise a large marine fish group of difficult identification, particularly during juvenile phase when the typical morphology and coloration of adults are absent. Therefore, the goal of this study was to test cytogenetic markers and DNA barcoding in the identification of bucktooth parrtotfish Sparisoma radians from the northeastern coast of Brazil. Sequencing of cytochrome c oxidase subunit I (COI) confirmed all studied samples as S. radians, and all showed high similarity (99-100%) with Caribbean populations. The karyotype of this species was divergent from most marine Perciformes, being composed of 2n = 46 chromosomes. These consisted of a large number of metacentric and submetacentric pairs with small amounts of heterochromatin and GC-rich single nucleolar organizer regions (NORs) not syntenic to 5S rDNA clusters. These are the first data about DNA barcoding in parrotfish from the Brazilian province and the first refined chromosomal analysis in Scarinae, providing useful data to a reliable genetic identification of S. radians.
... As a result, many authors have recommended the combination of different, but complementary sampling methods to accurately study fish assemblages inhabiting seagrass meadows (Harmelin-Vivien and Francour 1992;Bobsien and Brandelberger 2006). The parrotfish Sparisoma cretense (Linnaeus, 1758) is one of the few temperate species of the Scaridae family (Bernardi et al. 2000). This species is distributed in the eastern coasts of the Mediterranean Sea, the Macaronesian archipelagos (Azores, Madeira, Canaries and Cape Verde) and the northwest coast of Africa (González 1993). ...
... This is supported by, firstly, the presence of adult parrotfish inhabiting shallow-water rocky reefs in the study region (Tuya et al. 2006b). Secondly, a similar pattern has also been detected in the Mediterranean, where S. cretense is found on rocky reefs, feeding on macroalgae, yet occasional displacements towards seagrass meadows dominated by Posidonia oceanica can also occur to feed on seagrass epiphytes (Bernardi et al. 2000;Kalogirou et al. 2010). In tropical areas, distinct parrotfish species inhabit coral reefs; intermittent displacements of adults towards neighboring seagrasses are common to feed (Randall 1965;Ogden and Zieman 1977;Tribble 1981;Macintyre et al. 1987). ...
Seagrass habitats are relevant for numerous nearshore fish species, particularly as nursery grounds. Seagrass meadows are often interspersed with other habitats, what can alter the distribution and abundance of seagrass ichthyofauna. This research aimed to determine whether there is a change in the abundance and biomass of the parrotfish, Sparisoma cretense, in seagrass meadows (Cymodocea nodosa) with varying proximity from rocky reefs, specifically seagrass interiors (>200 m away from reefs) vs. seagrass adjacent to reefs (<10 m away). Sampling was undertaken using a seine net and underwater visual census through an entire annual cycle. Adults were predominantly observed in seagrass adjacent to reefs, which seem to be restricted to incursions of large-sized parrotfish from adjacent reefs. Juvenile abundance did not significantly differ between seagrass interiors and seagrass adjacent to reefs; however , juvenile biomass was greater in seagrass meadows adjacent to reefs compared to those far away from reefs. This pattern was consistent through times for both sampling techniques. These results suggest a transition of juvenile parrotfish from seagrass interiors to seagrass near reefs, so juveniles are located in the vicinity of their adult habitat, i.e. rocky reefs.
... Echinocardium meteorense populations from Great Meteor, Seine seamounts and the southern Mid-Atlantic Ridge differ conspicuously from each other in the form of rostrate pedicellariae (Mironov, present volume). Recent molecular studies show significant genetic differentiation between populations from the components of the Macaronesian archipelagos (Azores, Canaries, Madeira) (Benzie, 1999;Bernardi et al., 2000;Ó Foighil et al., 2001, and other). For example, the parrotfish genus Sparisoma and the bivalve genus Lasaea exhibit heterogeneous patterns in which previously unsuspected genetic discontinuities are apparent (Bernardi et al., 2000;Ó Foighil et al., 2001). ...
... Recent molecular studies show significant genetic differentiation between populations from the components of the Macaronesian archipelagos (Azores, Canaries, Madeira) (Benzie, 1999;Bernardi et al., 2000;Ó Foighil et al., 2001, and other). For example, the parrotfish genus Sparisoma and the bivalve genus Lasaea exhibit heterogeneous patterns in which previously unsuspected genetic discontinuities are apparent (Bernardi et al., 2000;Ó Foighil et al., 2001). ...
... Echinocardium meteorense populations from Great Meteor, Seine seamounts and the southern Mid-Atlantic Ridge differ conspicuously from each other in the form of rostrate pedicellariae (Mironov, present volume). Recent molecular studies show significant genetic differentiation between populations from the components of the Macaronesian archipelagos (Azores, Canaries, Madeira) (Benzie, 1999;Bernardi et al., 2000;Ó Foighil et al., 2001, and other). For example, the parrotfish genus Sparisoma and the bivalve genus Lasaea exhibit heterogeneous patterns in which previously unsuspected genetic discontinuities are apparent (Bernardi et al., 2000;Ó Foighil et al., 2001). ...
... Recent molecular studies show significant genetic differentiation between populations from the components of the Macaronesian archipelagos (Azores, Canaries, Madeira) (Benzie, 1999;Bernardi et al., 2000;Ó Foighil et al., 2001, and other). For example, the parrotfish genus Sparisoma and the bivalve genus Lasaea exhibit heterogeneous patterns in which previously unsuspected genetic discontinuities are apparent (Bernardi et al., 2000;Ó Foighil et al., 2001). ...
This study considers the distribution patterns of the benthic fauna of the Meteor Seamounts. In total, 301 species of macro/megafaunal benthic invertebrates, 94 species of meiofaunal invertebrates and 115 fish species were included. Most of the macro/megafaunal invertebrates (73%) and fishes (96.5%) have a panthalassic distri-bution, i.e. they known both from near-continental and open oceanic areas, whereas most of meiofaunal inverte-brates (95%) have open-oceanic distribution (confined to open oceanic seamounts and islands). Of the pan-thalassic macro/megafaunal invertebrates, 102 have trans-Atlantic distribution, 96 species are not found near the American continents (panthalassic east-Atlantic distribution), and 21 species are not found near European and African continents (panthalassic west-Atlantic distribution). The panthalassic fish of these three groups number 71, 35 and 5, respectively. The group of open oceanic species consists of 48 macro/megafaunal endemics of the Meteor Seamounts (16% of the total macro/megafauna), 88 meiofaunal endemics (91%), 1 fish endemic (1%) and 41 non-endemic species. The Meteor Seamount fauna is much closer to the European and African faunas than to the American fauna. Most macro-and megafaunal invertebrates (66%) are common to the northeastern Atlantic mainland areas, and numerous species (44%) are endemics to the north-eastern Atlantic. The number of species of gorgo-nians, madreporarians and echinoids shared with the Lusitanian region is larger than that shared with the Mauri-tanian region. An opposite situation is observed with cirripedes, brachiopods and fishes. The fishes of the Mete-or Seamounts are more typical of the Mauritanian Province, than are the invertebrates. A cluster analysis shows three main groups of the northern and northeastern Atlantic areas, however the composition of each group varies from taxon to taxon. The Meteor Seamounts are in association with either the Madeiran and Lusitanian areas (madreporarians and stylasterins), or with the Madeiran and Mediterranean areas (echinoids), or with the Madei-ran and Azores areas (brachiopods). Most endemics of the Meteor Seamounts are rather autochthonous descendants of immigrants than geo-graphic relics. Biogeographical and paleontological data suggest that the recent Meteor Seamounts fauna is mainly a result of late (Pliocene-Pleistocene) colonization from the Indo-West Pacific around the southern tip of Africa, and through the eastern and western Atlantic. The dispersal route through the northeastern Atlantic areas disagrees with the overall eastward surface flow in the northern Atlantic. However, a great variability of hydro-dynamics in three-dimensional space and in time made dispersal possible in any direction. The idea of dispersal through the western Atlantic is supported by meridional asymmetry in the distribution patterns of the panthalas-sic species. The panthalassic western Atlantic species are distributed more widely in the open oceanic regions than are the panthalassic eastern Atlantic species. Participants in the Trans-Arctic interchange are not found among the Meteor Seamount species. A parallel between the biogeographies of the Meteor Seamounts and the Azores is drawn throughout the paper. The Meteor Seamounts as well as the Azorean slopes can be classified as the recipient areas where the faunas of different origins are mixing. The events leading to faunistic similarity with the Indo-West Pacific and western Atlantic are generalised in a conception of the centres of redistribution of the marine fauna, which unites the conception of centres of origin and the conception of centres of accumulation.
... In addition, the A1 and A2 components of the adductor mandibulae are fused in Hipposcarus and Scarus (Bellwood 1994). Interestingly, some members of the genus Sparisoma exhibit several characteristic features of both scraping (S. aurofrenatum) and excavating parrot shes (S. amplum and S. viride) from the reef-associated lineage (Bruggemann et al. 1994b, Bernardi et al. 2000. These similarities appear to have arisen as a result of functional convergence (Streelman et al. 2002, Robertson et al. 2006. ...
... There is considerable regional variation in the taxonomic and functional composition of parrot shes that has been shaped by a complex biogeographic history (Bellwood 1994, Bernardi et al. 2000, Streelman et al. 2002) and, more recently, by human exploitation of these sh and their habitats , Hawkins & Roberts 2003, Mumby 2006. The most pronounced division in the composition of parrot sh assemblages occurs between the Indo-Paci c and Atlantic Oceans, with no overlap in species and limited overlap in genera between these two regions (Parenti & Randall 2000, Streelman et al. 2002. ...
Global reductions in biodiversity and the accelerating loss and degradation of many of the world's ecosystems have intensi research into the roles of species in ecosystem processes. Within coral-reef systems, the parrot (Labridae) are widely viewed as a key functional group in facilitating the recovery of reefs from recurrent disturbances. Although parrot are commonly viewed as herbivores, exerting top-down control of algal communities, their unique jaws allow them to feed on almost all coral-reef substratum types. Consequently, parrot are the primary agents responsible for a number of key ecological processes on coral reefs, namely, bioerosion, sediment production and transport, provision of space for coral settlement, and predation of live coral colo-nies. The parrot however, cannot be considered a uniform group.Their functional roles are highly dependent on species feeding mode (scrapers, excavators, and browsers) and body size, with larger individuals having a disproportionately greater effect on the dynamics of benthic communi-ties than smaller conspeci Parrot are ubiquitous on tropical reefs worldwide, yet there is strong spatial structuring in the taxonomic and functional composition of the group. This spatial variation has been shaped by their biogeographic history, the productivity of their environment, and the habitat requirements of individual taxa. Over recent decades, increasing pressure and habitat destruction have had a dramatic impact on the structure of parrot assemblages, and as a consequence, on many reefs, normal ecosystem processes have been disrupted. Indeed, reef systems with severely depleted parrot communities are less resilient to anthropogenic or natural environmental perturbations. Management strategies for the protection of this unique and critical functional group are urgently needed if we are to maintain the diversity, resilience, and structure of coral-reef ecosystems.
... This is a good starting point for examining the status of Hemiurids, especially those belonging to the genera Lecithocladium, documented in FishBase. The majority of phylogenetic connections are based on morphological features and molecular data [17]. The evidence given here supports the idea that all major Helminthes parasite groups are monophyletic [18]. ...
A total of 55 specimens of Bloch: Apolectus niger (Pisces: Carangidae) were obtained during the current inquiry from the sea beach of Puri, Orissa, India. It is a commercially significant marine fish, since it is a highly valued source of nutrition for human consumption. One type of the digenetic trematodes was collected from the gut of the affected fish, which belong to the genus, Lecithocladium, of the family Hemiuridae. Partial nucleotide sequences of the 18S ribosomal gene were acquired from this trematode species and deposited in the GenBank. Molecular analysis of the parasites using 18s small subunit rDNA was undertaken to evaluate their phylogenetic position with respect to previously known species in GenBank, which strikingly confirmed their molecular phylogeny revealed here. The hemiurid species from the genus, Lecithocladium are found to be L. excisum. A comparison of 18S rDNA sequences indicated that they exhibited 98.0 percent sequence similarity with Dinurus longisinus.
... Following these common classifications, we grouped herbivorous fish species into five functional groups (croppers, browsers, farmers, scrapers and excavators) based on the literature (Green & Bellwood, 2009;Choat et al., 2012;Adam et al., 2018;Supporting Information Table S1). Parrotfish of the genera Scarus and Sparisoma that were not identified by Adam et al. (2018) were assigned functional groups guided by the closest related species (for further details, see Supporting Information Table S1; Bellwood & Choat, 1990;Bernardi et al., 2000;Choat et al., 2012). Species from the parrotfish genus Cryptotomus did not appear in work of Adam et al. (2018) and ...
Aim
Biodiversity loss is impacting essential ecosystem functions and services across the globe. Recently, our interest in the benefits of biodiversity for ecosystem function has shifted focus from measurements of species richness to functional diversity and composition. However, the additional importance of other community characteristics, such as species evenness and co‐occurrence, for diversity‐driven ecosystem function is less known. We used herbivorous coral reef fish as a model system to investigate how co‐occurrence of different functional groups, rather than purely functional diversity, within an assemblage might affect the coral reef benthic state.
Location
Western Atlantic.
Time period
2007–2017.
Major taxa studied
Herbivorous reef fish.
Methods
We analysed benthic and fish assemblage data from 601 sites across 12 countries in the western Atlantic. Using diversity–interaction models, we investigated how the composition and relative abundances of reef fish functional groups were correlated with benthic cover and estimates of coral calcification rates. We used statistical interactions to explore the importance of co‐occurrence of herbivorous fish functional groups for the coral reef benthic state.
Results
We found that co‐occurrence of herbivorous fish functional groups, in addition to functional diversity, was correlated with reduced algal cover and increased coral accretion. Moreover, pairwise statistical interactions between functional groups were significantly correlated with an improvement in the coral reef benthic state.
Main conclusions
Our results support the idea that functional group co‐occurrence, in addition to functional diversity, within herbivorous fish offers additional benefits to the coral reef benthic state. We identify farming damselfish and excavating parrotfish as potential key determinants of the coral reef benthic state and highlight that co‐occurrence of cropping and scraping herbivores might promote coral accretion. Our findings support the argument that protecting herbivore abundance without regard to the species and functional groups present is not enough to preserve coral reef health and that fine‐scale community composition must be considered.
... Scarines are currently divided into the two tribes: Sparisomatini and Scarini (Baliga & Law, 2016;Schultz, 1958;Smith et al., 2008;Streelman et al., 2002). The tribe Sparisomatini (Figure 1a) includes the genera Leptoscarus, Calotomus, Nicholsina, Cryptotomus, and Sparisoma, which are generally associated with herbal reef environments as they feed primarily on marine angiosperms or macroalgae (Bellwood, 1994;McClanahan et al., 1999;Nakamura et al., 2006), except for some species of the genus Sparisoma (Bellwood, 1994;Bernardi et al., 2000). In contrast, the tribe Scarini (Figure 1a) includes the genera Cetoscarus, Bolbometopon, Hipposcarus, Scarus, and Chlorurus, which feed on hard substrates, including dead corals and rubbles (Bellwood & Choat, 1990;Bonaldo et al., 2014). ...
Parrotfish play important ecological roles in coral reef and seagrass communities across the globe. Their dentition is a fascinating object of study from an anatomical, functional and evolutionary point of view. Several species maintained non‐interlocked dentition and browse on fleshy algae, while others evolved a characteristic beak‐like structure made of a mass of coalesced teeth that they use to scrape or excavate food off hard limestone substrates. While parrotfish use their highly specialized marginal teeth to procure their food, they can also develop a series of large fangs that protrude from the upper jaw, and more rarely from the lower jaw. These peculiar fangs do not participate in the marginal dentition and their function remains unclear. Here we describe the morphology of these fangs and their developmental relationship to the rest of the oral dentition in the marbled parrotfish (Leptoscarus vaigiensis), the star‐eye parrotfish (Calotomus carolinus), and the palenose parrotfish (Scarus psittacus). Through microtomographic and histological analyses, we show that some of these fangs display loosely folded plicidentine along their bases, a feature that has never been reported in parrotfish. Plicidentine is absent from the marginal teeth and is therefore exclusive to the fangs. Parrotfish fangs develop a particular type of simplexodont plicidentine with a pulpal infilling of alveolar bone at later stages of dental ontogeny. The occurrence of plicidentine and evidence of extensive tooth wear, and even breakage, lead us to conclude that the fangs undergo frequent mechanical stress, despite not being used to acquire food. This strong mechanical stress undergone by fangs could be linked either to forced contact with congeners or with the limestone substrate during feeding. Finally, we hypothesize that the presence of plicidentine in parrotfish is not derived from a labrid ancestor, but is probably a recently evolved trait in some parrotfish taxa, which may even have evolved convergently within this subfamily. Some parrotfish display fangs that exhibit a particular type of simplexodont plicidentine with a pulpal infilling of alveolar bone at later stages of dental ontogeny. The extensive tooth wear and the breakage of certain fangs lead us to conclude that they undergo frequent mechanical stress, although they are not used to acquire food. This strong mechanical stress undergone by fangs could be linked either to forced contact with congeners or with the limestone substrate during feeding. We hypothesize that the presence of plicidentine in parrotfish is not derived from a labrid ancestor, but is probably a recently evolved trait in some parrotfish taxa, which may even have evolved convergently within this subfamily.
... While many species are shared between Ascension/St. Helena and the Brazilian or Tropical Eastern Atlantic Provinces (Manning and Chace 1990;Zibrowius et al. 2014;Brown et al. 2016;Padula et al. 2017), several taxa show strong genetic isolation of Ascension populations, such as in Sparisoma parrotfish (Bernardi et al. 2000), Eucidaris sea urchins (Lessios et al. 1999) and Grapsus crab (Freire et al. 2021). The haplotype networks for both loci showed that all individuals from Ascension had at least six fixed mutational differences from the nearest haplotypes in other biogeographic regions. ...
Understanding how populations are inter-connected can help identify species that are vulnerable or those that are likely to persist over ecological and evolutionary time scales. Favia fragum and Favia gravida are sister species of brooding corals common in the Atlantic Ocean that are secondary reef-builders that contribute to habitat complexity in intertidal and shallow-water coral reefs. To better understand population connectivity in these two congeneric species with broad and presumably non-overlapping distributions, one single-copy nuclear marker (masc1) and one ribosomal marker (ITS) were used to estimate population genetic parameters for Favia in four biogeographic regions: the Caribbean, Brazilian, Ascension and Tropical Eastern Atlantic provinces. Strong and significant levels of population structure were found among all four biogeographic regions, with each province being characterized by nearly all private alleles. Phylogenetic analyses based on each gene and on a combined marker analysis showed strong support for three sister clades composed of individuals from either the Caribbean, South Atlantic (Brazilian + Tropical Eastern Atlantic), or Ascension provinces. However, low genetic diversity was observed at each location, and no differentiation was observed among populations along the entire coast of Brazil (~ 1600 km). Furthermore, the Tropical Eastern Atlantic was likely colonized by at least two independent founder events, one from the Caribbean and another from the Brazilian provinces. Life-history traits of the genus might explain the different patterns found. Favia fragum and F. gravida release zooxanthellae larvae monthly and may be capable of rafting, which could favor occasional long-distance dispersal. On the other hand, they are capable of self-fertilization and commonly disperse over a few meters which may explain low genetic variability locally. Our results show that while long-distance migration events are possible for Favia corals, they are insufficient to maintain connectivity across biogeographic regions. Lastly, the Ascension Island population was more divergent than the sister clades composed of Caribbean or South Atlantic Favia, indicating a possible cryptic lineage, but further work may clarify species boundaries within the Atlantic coral genus Favia.
... The Mediterranean parrotfish is a coastal reef fish inhabiting shallow rocky bottoms and seagrass beds throughout the Macaronesian archipelagos (Azores, Madeira, Canary, and Cape Verde) and the warmer south Mediterranean Sea (Bernardi et al. 2000), although some studies suggest a climate change related expansion to the north (Guidetti and Boero 2001;Yapici et al. 2016) and out of the western Mediterranean (Abecasis et al. 2008). This species has two distinct mating strategies: (1) a single-male territorial strategy whereby usually larger males hold female harems in small areas and (2) a multi-male (usually smaller < 35 cm) group strategy (Afonso et al. 2008a, b). ...
Aggressive behaviour in fishes, particularly in territorial species, is a common trait used to defend resources such as food or mates. Territorial males of the Mediterranean parrotfish Sparisoma cretense have been described to chase away conspecifics yet other aggressive behaviour repertoire has not been reported for this species. We describe, for the first time, an extreme aggressive behaviour between two male Mediterranean parrotfish which includes biting and prolonged mouth locking.
... Scaridae identification is based on skeletons of 15 fish representing seven species of Red Sea parrotfish (Randall, 1986) and the single species endemic to the Mediterranean Sea (Sparisoma cretense; Mediterranean parrotfish) (Bernardi et al., 2000, Golani and Bogorodsky, 2010, Golani et al., 2006 (Table S1). We used the Micro CT to scan two Table S4) (after Bellwood, 1994;Gregory, 1959;Nanami, 2016), and of Sparisoma cretense from the Mediterranean Sea (photographs by Roee Shafir, Haifa University). ...
Recent archaeological excavations in the Negev desert of the southern Levant have yielded a surprising and unprecedented amount of fish remains, found in the landfills at Byzantine (4th–7th century CE) and Early Islamic (7th–9th century CE) sites. The significant economic and dietary role of Red Sea fish, especially parrotfish (Scaridae), in addition to fish originating from freshwater habitats, including the Nile, as well as from the
Mediterranean Sea, attest to the importance of fish in sophisticated trade networks, which facilitated the transport of fish from different aquatic habitats to the distant provinces. The discovery of Red Sea parrotfish at all three studied sites is of particular interest. Analysis of the Scaridae remains revealed their role as a flagship species of the Red Sea coral-reef fishery and as an essential commodity in the Byzantine and Early Islamic economic systems; as well as the presence of more than one Scaridae species. The study of skeletal element representation indicated that complete fish, larger than 23 cm in length (SL), were transported to the sites. The bone fragmentation patterns reflected typical traditional butchering during the desiccation processes. The fish remains recovered from the desert provide new insights into the infrastructure of commercial networks that
enabled the transport of fish over long distances during periods of high demand for fish products.
... Scaridae identification is based on skeletons of 15 fish representing seven species of Red Sea parrotfish (Randall, 1986) and the single species endemic to the Mediterranean Sea (Sparisoma cretense; Mediterranean parrotfish) (Bernardi et al., 2000, Golani and Bogorodsky, 2010, Golani et al., 2006 (Table S1). We used the Micro CT to scan two Table S4) (after Bellwood, 1994;Gregory, 1959;Nanami, 2016), and of Sparisoma cretense from the Mediterranean Sea (photographs by Roee Shafir, Haifa University). ...
Recent archaeological excavations in the Negev desert of the southern Levant have yielded a surprising and unprecedented amount of fish remains, found in the landfills at Byzantine (4th–7th century CE) and Early Islamic (7th–9th century CE) sites. The significant economic and dietary role of Red Sea fish, especially parrotfish (Scaridae), in addition to fish originating from freshwater habitats, including the Nile, as well as from the Mediterranean Sea, attest to the importance of fish in sophisticated trade networks, which facilitated the transport of fish from different aquatic habitats to the distant provinces. The discovery of Red Sea parrotfish at all three studied sites is of particular interest. Analysis of the Scaridae remains revealed their role as a flagship species of the Red Sea coral-reef fishery and as an essential commodity in the Byzantine and Early Islamic economic systems; as well as the presence of more than one Scaridae species. The study of skeletal element representation indicated that complete fish, larger than 23 cm in length (SL), were transported to the sites. The bone fragmentation patterns reflected typical traditional butchering during the desiccation processes. The fish remains recovered from the desert provide new insights into the infrastructure of commercial networks that enabled the transport of fish over long distances during periods of high demand for fish products.
... The phenotypic characterization of parrotfish is challenging due to the presence of a series of coloration changes. For instance, Sparisoma species color pattern varies by sex and developmental stage and is only clearly visible in fresh specimens (Bernardi et al., 2000). ...
Cheilinus trilobatus, Cheilinus quinquecinctus, and Chlorurus sordidus specimens from Saudi Arabia's Farasan Islands were collected and genotyped using inter simple sequence repeats (ISSRs) and start codon targeted (SCoT) primers. Mitochondrial cytochrome C oxidase subunit I (COI) gene fragments were used for DNA barcoding. ISSRs and SCoT primers showed moderate polymorphisms: expected heterozygosity (H exp) of 0.470 and 0.435 and average polymorphism information contents (PICs) of 0.359 and 0.339 for ISSRs and SCoT markers were observed, respectively. Cheilinus quinquecinctus had the highest genetic diversity from ISSRs (70%) and SCoT (73%). Chlorurus sordidus and C. trilobatus showed similar genetic diversities of 29% and 39% based on ISSRs, respectively, and 60.32% for both species based on SCoT. Cheilinus quinquecinctus had the lowest nucleotide diversity (π) of 0.003, while C. sordidus and C. trilobatus had π values of 0.065 and 0.103, respectively. Analysis of molecular variance (AMOVA) revealed greater genetic variation among species rather than within them using ISSRs (65% and 67%, respectively) and SCoT (35% and 33%, respectively). COI-based AMOVA showed similar genetic variation among (51.11%) and within (48.89%) species. The current study highlighted outperformance of COI compared to ISSRs and SCoT markers in differentiating among parrotfish species. Also, ISSR outperformed SCoT since it was able to clearly show three distinct groups in principal component analysis. This study also confirmed the presence of three distinct parrotfish species, which will provide an insight into parrotfish diversity. Moreover, the results will contribute to monitoring parrotfish migration between Farasan Islands and different geographic locations which significantly affect species conservation.
... This occurs on reefs with early successional-stage algae. Bruggemann et al. 1994cAdam et al. 2015 McAfee & Morgan 1996 Bernardi et al. 2000 Reducing competition between macroalgae and corals Sparisoma spp. ...
Parrotfishes are caught primarily by fish traps and spearfishing (targeted and incidental) for personal and commercial consumption. Over half (65 percent) of the responding countries and overseas territories indicated that they have either a complete harvesting ban or fishing regulations in place to promote the protection of parrotfishes to some extent. About half (56 percent) of the responding countries have fishery-independent surveys for parrotfishes, but the metrics evaluated by these surveys are not the same. Similarly, parrotfish-related outreach and education was varied throughout the Caribbean. Twelve countries (32 percent) report having specific parrotfish outreach campaigns or materials, while in other countries campaigns and materials were geared towards marine conservation in general. Most countries agree that long-term protection for parrotfishes is critical, but consensus about the specific measures required to achieve this is lacking.
... Although congeneric sponge-feeding fishes tend to eat the same sponge species, other members of their genera and families do not eat sponges. Two small-bodied Caribbean Sparisoma species are browsers in seagrass habitats (Bernardi et al. 2000). And although the large Pacific puffer Arothron mappa has sometimes been found with its gut packed with sponges (Hiatt and Strasburg 1960), and A. hispidus focuses on sponges when they are available, a third congener, A. meleagris, primarily eats corals (Glynn et al. 1972, Guzm an andRobertson 1989). ...
Defenses that target particular consumers often influence community organization, ecosystem function, and diversity maintenance. In coral reef, mangrove, and seagrass ecosystems, sponges affect substratum stability, water clarity, diversity of associated species, and survival of habitat‐providing organisms, key roles not duplicated by other organisms. Whether and how predators control sponges are much disputed. Substantial ecosystem consequences of losing or gaining sponges motivated definitive experiments on how predators control sponge distribution and abundance. Caribbean sponges of 94 species representing 13 taxonomic orders and three linked habitats (coral reefs, mangroves, and seagrass meadows) were exposed to seven predator species representing different habitats and degrees of spongivory in 4,493 in situ trials. The resulting data force reassessment of popular interpretations of several patterns and processes. Contrary to extract pellet assays that declare most sponges deterrent, 78% of these 94 species were eaten by at least one predator. But “palatability” is consumer dependent: a sponge species eaten by one predator can be rejected by other predators, and predator species differed in what sponges they ate in 55.4% (214/392) of pairwise comparisons between predators. Because spongivore species are usually restricted to particular habitats, they impose abrupt boundaries on sponges’ habitat distributions, reflecting inverse relationships between accessibility and palatability to each predator. Thus a seagrass‐dwelling starfish eats only 9% of seagrass sponge species, but 70% of coral reef species, and 78% of mangrove species. Reef‐dwelling angelfishes completely consume only 13% of reef species, but 29% of seagrass species, and 63% of mangrove species. Defenses that target specific predators reveal that spongivore influence on community organization cannot be inferred from extract pellet/omnivore assays that assume defenses target all predators equally. In fact, pellet data wrongly predicted actual consumption of living sponges of that pellet's species in 43% of field experiments with spongivores. In contrast with herbivore–plant interactions, opportunistic spongivory is at least as important as routine spongivory for community organization and ecosystem function. Potential for loss of key functional roles of sponges, if opportunistic predators gain access to sponge species that lack defenses against them, must inform conservation plans for coral reef, mangrove, and seagrass ecosystems.
... Bellwood 1994) and some authors also categorizing Sp. chrysopterum and Sp. rubripinne as excavators (Bernardi et al. 2000). Within the lineage that possesses the intramandibular joint there is wide diversity in feeding mode, from superficial scraping to excavating. ...
... The topology dependent test (t-PTP; Faith 1991) has been criticized for problems dealing with the formulation of the null hypothesis, and the fact that the test is sensitive to the number of taxa and characters (the parameters that determine the number of possible trees) (see Harshman 2001 for references on critics and replies from Faith and collaborators). Despite those critics, this test has been used in different studies to compare the topology obtained to a prioiri hypotheses (see for instance Alvarez et al. 2000, Bernardi et al. 2000. Alternative tests have been proposed to compare both a priori and a posteriori hypotheses: a nonparametric test, the Shimodaira-Hasegawa test Hasegawa 1999, Buckley et al. 2001), and two parametric tests, the parametric bootstrap test (Huelsenbeck et al. 1996) and the SOWH test . ...
The systematics of the babblers (Timaliidae) and related members of the Old World insectivorous passerines have been particularly difficult. To clarify our understanding of this group, phylogenetic relationships were constructed using sequences of three mitochondrial genes (cytochrome b, rRNA 12S and 16S). The results indicated that several species traditionally placed among babblers, the shrike babblers (Pteruthius) and the Gray-chested Thrush Babbler (Kakamega poliothorax), are not related to the Timaliidae, but belong to other passerine groups. Furthermore, the phylogenetic hypotheses inferred from molecular data suggest that the babblers assemblage includes two other oscine taxa traditionally considered to be distantly related, Sylvia (Sylviidae) and Zosterops (Zosteropidae). The polyphyly of several babbler genera is discussed, with particular attention to the laughingthrushes (genera Garrulax and Babax) for which the phylogeny is compared to previous hypotheses of relationships. Results from different tests under the maximum-parsimony and maximum-likelihood criteria indicate the rejection of the hypothesis of monophyly for the laughingthrushes group. Thus, the molecular phylogeny challenges the traditional classification of the Timaliidae.
... rubripinne and Sp. aurofrenatum, have been variously described as macroalgal browsers, excavators, or scrapers (Bellwood 1994, Bruggemann et al. 1996, Bernardi et al. 2000, Bellwood et al. 2004, Cardoso et al. 2009, Adam et al. 2015b), but it is unlikely that they serve all of these roles. Developing a better understanding of the functional roles of different parrotfishes will give us a clearer picture of the patterns of functional diversity and redundancy on Caribbean reefs. ...
Feeding behavior of coral reef fishes often determines their species‐specific ecological roles. We studied the two most common Caribbean surgeonfishes (Acanthurus coeruleus and Acanthurus tractus) to examine their species‐specific grazing rates and feeding preferences and how these differed with environmental context. We quantified the feeding activity of both surgeonfishes at four spur and groove reefs in the Florida Keys, USA, that varied in fish abundance, rugosity, algal community composition, and sediment loading. Overall, A. tractus fed twice as fast as A. coeruleus. Both species selected for turf algae but avoided feeding on turf algae that had become laden with sediment. Selectivity for upright macroalgae was more complex with A. tractus targeting Dictyota spp., while A. coeruleus avoided Dictyota spp. relative to the alga's abundance. Both species selected for epiphytes growing on other organisms such as macroalgae and sponges. However, several of these feeding patterns changed with ontogeny. For example, larger individuals of both species fed more frequently on long, sediment‐laden algal turf and less frequently on Dictyota spp. compared to smaller sized individuals. In addition, A. tractus also increased its preference for upright calcareous algae as they attained larger sizes. Overall, the disparity in feeding preferences of surgeonfishes likely indicates subtle differences in species‐specific ecological roles. Both A. coeruleus and A. tractus likely prevent development of turf algae and thus maintain algal communities in the early stages of succession. Additionally, A. tractus may also help reduce macroalgal abundance by targeting common macroalgal species.
... In this study, the genus Sparisoma, which only occurs in the Atlantic (Bernardi et al. 2000), was 5 times more abundant than the genus Scarus. Unlike in the Caribbean, the genus Sparisoma predominates over the genus Scarus in the southwestern Atlantic , which may be related not only to dispersion potential, but also to habitat requirements. ...
Parrotfishes may affect the structure of benthic communities and reef ecosystem functioning. Despite extensive studies worldwide, parrotfishes in the southwestern Atlantic are relatively understudied, limiting our ability to propose effective management strategies. We assessed abundance, biomass and size class distribution of parrotfish assemblages in northeastern Brazil and identified habitat preferences based on reef attributes. Outer-shelf reefs sustained larger individuals and higher biomasses for all species (except Sparisoma radians). In contrast, inner-shelf reefs supported higher abundances of small individuals. Even though most species occurred across all areas, their abundances, biomass and size-class distributions were variable and related to their respective feeding modes and reef attributes. Benthic cover, reef structural complexity, depth and distance from the coast affected the composition of parrotfish assemblages, but had different effects on each species. The endemic greenbeak parrotfish Scarus trispinosus was more abundant on calcareous substrates and higher-complexity reefs. Sc. zelindae and Sp. amplum were more common in deeper biogenic reefs further from the coast, which were characterized by high abundances of sponges, stony corals and cyanobacterial mats. Sp. axillare and Sp. radians were more abundant on reefs that had high cover of large-bladed macroalgae, while Sp. frondosum was ubiquitous across all studied reefs. Such heterogeneity in habitat use is suggestive of functional complementarity rather than functional redundancy among parrotfish assemblages. Conservation of Brazilian endemic parrotfishes requires protecting reefs with diverse attributes and a better understanding of habitat connectivity and the role of different habitats in parrotfish reproduction and life cycle.
... rubripinne and Sp. aurofrenatum, have been variously described as macroalgal browsers, excavators, or scrapers (Bellwood 1994, Bruggemann et al. 1996, Bernardi et al. 2000, Bellwood et al. 2004, Cardoso et al. 2009, Adam et al. 2015b), but it is unlikely that they serve all of these roles. Developing a better understanding of the functional roles of different parrotfishes will give us a clearer picture of the patterns of functional diversity and redundancy on Caribbean reefs. ...
Parrotfishes are a diverse group of herbivores that can influence benthic community dynamics and ecosystem function on coral reefs. Different species and size classes of parrotfishes vary in their feeding ecology and can impact reef ecosystems in distinct ways. We documented differences in the feeding ecology of 9 species of parrotfishes in the Florida Keys National Marine Sanctuary (FKNMS). Many of the key differences can be summarized by assigning species to functional groups (e.g. scrapers, excavators, croppers, macroalgae browsers), which are differentially responsible for carrying out specific ecological processes. For example, we found that Sparisoma viride, Scarus coelestinus, Sc. guacamaia, Sc. taeniopterus, and Sc. vetula feed on short turfs with few sediments, while Sp. aurofrenatum, Sp. chrysopterum, and Sp. rubripinne feed on longer sediment-laden turfs in addition to macroalgae. Further, parrotfishes use distinct bite types that indicate contrasting impacts on the benthos. Species that feed on short turfs scrape and excavate epilithic and endolithic algae, while species that feed on longer turfs and macroalgae tend to tear or crop algae from the reef. These distinct feeding behaviors result in different rates of algae removal, carbonate erosion, and sediment production. Recognizing that different species of parrotfishes interact with the benthos in fundamentally different ways will enable scientists and managers to better predict how changes in the structure of parrotfish assemblages may affect benthic communities and ecosystem processes.
... Sp. aurofrenatum, Sparisoma rubripinne and Sp. chrysopterum are considered 'grazers' (the term grazer is used because these species are not obligate browsers and also scrape the epilithic algal matrix), although Sp. aurofrenatum does frequently bite live corals (Miller and Hay 1998) and in some classifications is considered a scraper (Bernardi et al. 2000, Streelman et al. 2002. Sp. viride is a 'bioeroder' or 'excavator' (removes both coralline rock and live coral when feeding but feeds extensively on algal turfs and several macroalgae including Dictyota spp., Mumby 2006a), and Scarus vetula is a 'bioeroder/scraper' (removes coralline rock but feeds primarily on turf algae). ...
... Parrotfish evolved in tropical ocean waters prior to the mid-Miocene (Bellwood andSchultz 1991, Bernardi et al. 2000), with the earliest known fossil a Calotomus species (Bellwood and Schultz 1991). In terms of habitat associations, the phylogeny suggests that parrotfish have shifted over time from non-reef (seagrass) habitats to a more exclusive association with coral reefs, with the reef association being only a relatively recent one (Bellwood 1994, Bernardi et al. 2000, Streelman et al. 2002, Choat et al. 2012. It is argued by some that coral reefs have facilitated the diversification and speciation of the Labroidei, however, given the historical associations with non-reef habitats, it seems possible that parrotfish could persist in alternative environments, albeit at depaurperate levels of species diversity, or potentially with a loss over evolutionary time of some of the oral jaw modifications that have evolved within the reef clade (Streelman et al. 2002). ...
... However, in some species, it seems to be particularly affected by ecological differences among regions (Rocha et al., 2005). Vicariant events, combined with habitat differentiation (Bernardi et al., 2000), have been shown to be promoters of genetic diversification in some marine groups. In this respect, analysis based on the linear distances, especially among insular environments, may disregard physical and biological parameters that may act on regional chromosomal differentiations. ...
The distribution patterns of marine biodiversity are complex, resulting from vicariant events and species dispersion, as well as local ecological and adaptive conditions. Furthermore, the wide geographic distribution of some species may be hindered by biogeographical barriers that can interfere in the gene flow. Cytogenetic analyses in marine fishes, especially those involving populations in small remote insular environments, remain scarce. In the Western Atlantic, species of wrasses from the genera Halichoeres and Thalassoma occur in biogeographic arrangements that make it possible to analyze cytogenetic patterns between coastal and widely separated island populations. Species of these genera were punctually analyzed in some Atlantic regions. In this study, we compared several chromosomal features, such as karyotype macrostructure, heterochromatic patterns, patterns of base-specific fluorochromes, Ag-NORs, and 18S and 5S ribosomal sites in Thalassoma noronhanum, Halichoeres poeyi, and Halichoeres radiatus individuals from distinct coastal or insular regions
of Atlantic. Notably, all of them are characterized by multiple 18S and 5S rDNA sites with syntenic arrangements in some chromosome pairs. Individuals of T. noronhanum (between the insular regions of Rocas
Atoll and Fernando de Noronha Archipelago - FNA) and H. poeyi (coastal areas from Northeastern Brazil) show no detectable differences among their cytogenetic patterns. On the other hand, H. radiatus from FNA and São Pedro and São Paulo Archipelago exhibit differences in the frequency of rDNA sites that could suggest some level of population structuring between these insular regions. Interregional cytogenetic inventories of marine species with wide geographic distribution need to be rapidly expanded. These data will allow a better understanding of the level of chromosomal stability between vast oceanic spaces, which may be less than previously thought.
... All the species were found genetically distinct from each other and got delineated into three groups without any of their barcodes sharing or overlapping. The study in addition to authenticating the species, validated their taxonomical status as established based on morphological characters by [13], like earlier studies in fishes [8,25] along with the baseline information regarding intra specific variation which might be important for future studies leading to their conservation. The sequences of COI region were simple and unambiguous with well defined peaks. ...
Authentic identification of fish species is essential for conserving them as a valuable genetic resource in our environment. DNA barcoding of living beings has become an important and ultimate tool for establishing their molecular identity. Among cyprinids, Barilius is an important genus having nearly 23 species in Indian region whose morphological identification is often difficult due to minute differences in their features. Five species collected from Indian waters and primarily identified as Opsarius bakeri (syn. Barilius bakeri), B. gatensis, B. vagra, B. bendelisis and B. ngawa were authenticated by their DNA barcoding based on mitochondrial COI gene sequences. Five individuals of each species were taken for barcode preparation by COI gene sequencing which yielded one barcode for B. ngawa, two barcodes each for O. bakeri, B. gatensis, B. bendelisis and three barcodes for B. vagra. The order of inter and intra-specific variation was estimated to know a preliminary status of variation prevailing in these cold stream fish species significant for evolution and conservation of these valued species of our ichthyofauna. Average variation within genera was found to be 13.6% with intra-specific variation ranging from 0.0% (B. ngawa) to 0.6% (B. gatensis). These distance data are in the same order found by various researchers globally using COI barcode sequences in different fish species. Phylogenetic relatedness among Barilius species and some other cyprinids validate their status of individual species as established by conventional taxonomy.
... All the species were found genetically distinct from each other and got delineated into three groups without any of their barcodes sharing or overlapping. The study in addition to authenticating the species, validated their taxonomical status as established based on morphological characters by [13], like earlier studies in fishes [8,25] along with the baseline information regarding intra specific variation which might be important for future studies leading to their conservation. The sequences of COI region were simple and unambiguous with well defined peaks. ...
Authentic identification of fish species is essential for conserving them as a valuable genetic resource in our environment. DNA barcoding of living beings has become an important and ultimate tool for establishing their molecular identity. Among cyprinids, Barilius is an important genus having nearly 23 species in Indian region whose morphological identification is often difficult due to minute differences in their features. Five species collected from Indian waters and primarily identified as Opsarius bakeri (syn. Barilius bakeri), B. gatensis, B. vagra, B. bendelisis and B. ngawa were authenticated by their DNA barcoding based on mitochondrial COI gene sequences. Five individuals of each species were taken for barcode preparation by COI gene sequencing which yielded one barcode for B. ngawa, two barcodes each for O. bakeri, B. gatensis, B. bendelisis and three barcodes for B. vagra. The order of inter and intra-specific variation was estimated to know a preliminary status of variation prevailing in these cold stream fish species significant for evolution and conservation of these valued species of our ichthyofauna. Average variation within genera was found to be 13.6% with intra-specific variation ranging from 0.0% (B. ngawa) to 0.6% (B. gatensis). These distance data are in the same order found by various researchers globally using COI barcode sequences in different fish species. Phylogenetic relatedness among Barilius species and some other cyprinids validate their status of individual species as established by conventional taxonomy.
... Despite the limited information available, S. trispinosus has been described as a species with a more restricted distribution, establishing populations mostly near the coast. The positive relationship between S. trispinosus and primary production found in this study may explain the more restricted distribution of this species due to its specialized feeding habits (Bernardi et al., 2000;Streelman et al., 2002), which results in intense grazing that enhances the productivity of benthic algal assemblages by removing large algae with low specific productivity (Hatcher, 1988). In addition, the distribution prediction shown in this study supports the patterns of a decreasing occurrence at higher latitudes, which is explained by the fact that the tropics sustain higher primary productivity (Carpenter, 1986;Hatcher, 1988). ...
In coral reef environments, there is an increasing concern over parrotfish (Labridae: Scarini) due to their rising exploitation by commercial small-scale fisheries, which is leading to significant changes in the reefs' community structure. Three species, Scarus trispinosus (Valenciennes, 1840), Sparisoma frondosum (Agassiz, 1831) and Sparisoma axillare (Steindachner, 1878), currently labeled as threatened, have been intensively targeted in Brazil, mostly on the northeastern coast. Despite their economic importance, ecological interest and worrisome conservation status, not much is known about which variables determine their occurrence. In this study, we adopted a hierarchical Bayesian spatial-temporal approach to map the distribution of these three species along the Brazilian coast, using landing data from three different gears (gillnets, spear guns, and handlines) and environmental variables (bathymetry, shore distance, seabed slope, Sea Surface Temperature and Net Primary Productivity). Our results identify sensitive habitats for parrotfish along the Brazilian coast that would be more suitable to the implementation of spatial-temporal closure measures, which along with the social component fishers could benefit the management and conservation of these species.
Copyright © 2015 Elsevier Ltd. All rights reserved.
... However, despite COI being a robust evolutionary marker, it has been shown to have a higher mutation rate in several organisms. We therefore selected the mitochondrial 16SrRNA gene, which has also been shown to be a good marker for species differentiation, and has been of great use in studies of intergeneric and interspecific comparisons in several Perciform families (Ritchie et al., 1997;Bernardi et al., 2000;Streelman et al., 2000). ...
This study addresses variations of mtDNA and otolith morphometrics in Scomberomorus concolor and Scomberomorus sierra collected along the Mexican Pacific Coast and Gulf of California (GC) in 2002, 2007 and 2008. The catches of GC are reported without distinguishing species, serious problem account that S. concolor is included in the red list by the IUCN. For molecular identification a 592 bp fragment of the 16SrRNA gene was used, and a 410 bp fragment from the mtDNA control region was analysed. The 16SrRNA analysis gene generated two clades, corresponding to each species, which could only be
distinguished from two nucleotide sites. The mtDNA sequences maintained the detected lineages, even when both species presented elevated polymorphism in relation to their levels of divergence. For morphometric analysis, a first analysis included samples of fish obtained in 2007 that were genetically identified. A second analysis included fish captured in the central GC that were identified using meristic criteria. Both shape analysis, generated statistically significant classification, the allocation per species was 100%, whereas the allocation of the fish identified using meristic characters was 95%. The deformation grids identified the dorsal zone of the otolith as the origin of the discrimination between species and the criterion for separation.
... The observed anti-adenine bias in stems, and anti-guanine as well as antitymine bias in loops was similar to a study on piranhas (Ortí et al., 1996). The average ti/tv ratio of 3.85 was close to values of serranid fishes (Craig et al., 2001) as well as parrotfishes ( Bernardi et al., 2000). Even tough the number of phylogenetically informative sites was lower than in the cyt b sequences, a strong phylogenetic signal in the 16S rDNA sequences was indicated by g 1 statistics. ...
Aim of this thesis is the study of biogeography and ecology, genetic population structure, and molecular phylogeny of fishes on coral reefs in the Gulf of Aqaba and northern Red Sea. Ecological and genetic pattern are compared on different spatial scales and molecular markers add a temporal scale to study of evolutionary processes.Biogeographic analysis supported the differentiation of the Arabian sub-province from the Indian Ocean, but the affiliation of the Arabian Gulf is not clear.The analysis of the genetic population structure based on the mtDNA control region revealed homogenity between populations of Pterois miles in the Gulf of Aqaba and northern Red Sea and indicate high levels of gene flow.Investigations on interrupted gene flow in the evolutionary history of lionfishes (Scorpaenidae) revealed a differentiation between the Indian Ocean and Western Pacific. Phylogenetic analysis of cyt b and 16S rDNA sequences of the siblings P. miles and P. volitans suggest a divergence time of 2.4-8.3 million years. This coincide with tectonic events and sea level changes in Southeast Asia during the glacial maxima. Additionally, this genetic study gave indications for taxonomic revision of the genera Pterois and Dendrochirus.Ecological studies on the shore fishes off the Jordanian coast showed that fish species richness was positively correlated with hard substrate cover and benthic diversity. Abundance of corallivores was positively linked to live coral cover. The multivariate analysis of the fish community has revealed several associations of fishes in different habitats, such as deep and shallow reef slope.The northern tip of the Gulf of Aqaba and its western shores are particularly subject to human disturbances. Fish abundance at an industrial site was 50% lower than on an undisturbed reef and the trophic community structure was different.
... Furthermore, some Scaridae predominantly ingest detritus (Wilson et al., 2003). Scaridae are known to have diverse habitat use and migration patterns, and this behaviour implies a wide range of feeding ecology (Bernardi et al., 2000;Streelman et al., 2002). Fatty acids generally cannot be synthesized in higher trophic levels and are incorporated into the tissues of predators (Sargent et al., 1987). ...
To understand the trophic ecology and habitat use of coral reef fish, the fatty acid composition and levels were examined in three species of parrotfish, Scarus psittacus: Scarus quoyi and Scarus rivulatus, which were collected at the Bidong Island of the Malaysian South China Sea. The proportions of saturated fatty acids (SAFA) are 57.6–64.9%, where fatty acids have the highest proportion, monounsaturated fatty acids (MUFA) (25.3–25.9%) have the second highest proportion, and polyunsaturated fatty acids (PUFA) (9.2–17.1%) have the lowest proportion. Palmitic acid (C16:0) is the most common acid in SAFA, palmitoleic acid (C16:1) is the dominant acid in MUFA, and eicosatrienoic acid (C20:3n3) and eicosapentaenoic acid (EPA, C20:5n3) have the highest proportions in PUFA. The composition of each fatty acid does not differ among species, which suggests that the feeding ecology and habitat use of the fish in the coral reef ecosystem are similar. The fatty acid concentrations, particularly that of PUFA, increase with the fish growth, which suggests that the diet and habitat shift during the life history of the fish. The diet shifts revealed by the fatty acid composition and levels suggest the similarity in habitat use and feeding ecology of those types of parrotfish.
... These corallivorous excavating fishes play a major role in reef dynamics and sedimentation while rasping algal communities (e.g. Bellwood & Choat 1990, Bernardi et al. 2000, Comeros-Raynal et al. 2012, Bonaldo et al. 2014). According to Choat et al. (2012), the Guinean parrotfish Scarus hoefleri (Fig. 1E) is 'basal' in the tropical Atlantic clade and the sole Scarus species that ranges from Mauritania to Congo (Wirtz et al. 2007). ...
The zoogeographic composition of the coastal ichthyofauna of the Cape Verde archipelago shows a predominance of Guinean species, followed by tropical-subtropical (amphi-Atlantic) fishes and several endemic species. Recent taxonomic revisions, new species descriptions, new fish records in the area and recent ecological surveys of intra-specific fish interaction as well as emergent research on seamounts in national waters warrant an up-to-date review of research on the ichthyofauna of Cape Verde. An updated percentage of endemic coastal reef fish of 10.2% attest significant speciation in Cape Verde waters. Almost half of the total cryptobenthic fish species richness in Cape Verde comprises endemic species, some of them newly discovered. Examples of endemism are discussed, notably white seabream Diplodus sargus lineatus, bulldog dentex Dentex (Virididentex) acromegalus, blackfish drummer Girella stuebeli, Lubbock's chromis Chromis lubbocki, Cape Verde damselfish Similiparma hermani and Guinean parrotfish Scarus hoefleri. Cape Verde diverges considerably from the other Macaronesian island groups in terms of its ichthyofauna and its uniqueness warrants special attention by governmental agencies and conservation organizations.
... Moreover, warmer regions were also characterised by greater abundance of the parrotfish Sparisoma cretense (an EAM consumer) which has a distribution largely restricted to the warmest eastern and southern parts of the Mediterranean basin (Bernardi et al., 2000). ...
A striking example of climate‐mediated range shifts in marine systems is the intrusion of tropical species into temperate areas world‐wide, but we know very little about the ecological consequences of these range expansions.
In the Mediterranean Sea, the range expansion of tropical rabbitfishes that first entered the basin via the Suez Canal provides a good example of how tropical herbivorous fish can impact the structure of rocky bottoms in temperate seas. Two species of rabbitfishes have now become a dominant component of total fish biomass in the southernmost part of the eastern Mediterranean. Experimental evidence shows these species can profoundly transform benthic communities, turning algal forests into ‘barrens’, but the specific mechanisms that facilitate this shift have not been established.
We surveyed ˜1000 km of coastline in the eastern Mediterranean and identified two clearly distinct areas, a warmer group of regions with abundant tropical rabbitfish and a colder group of regions where these consumers were absent/ extremely rare. In regions with abundant rabbitfish, canopy algae were 65% less abundant, and there was a 60% reduction of overall benthic biomass (algae and invertebrates) and a 40% decrease in total species richness.
Video‐recorded feeding experiments showed that the extensive barrens characteristic of regions with abundant rabbitfish were not due to greater rates of herbivory by these tropical consumers, but rather by functional differences among the herbivores. Temperate herbivorous fish displayed the greatest macroalgae consumption rates overall, but they fed exclusively on established adult macroalgae. In contrast, in regions with abundant rabbitfishes, these consumers fed complementarily on both established macroalgae and on the epilithic algal matrix, which typically contains macroalgal recruits.
Synthesis . Range‐shifting tropical rabbitfish can severely reduce the biomass and biodiversity of temperate reefs at a scale of hundreds of kilometres. A shift from macroalgal dominance to barrens is mediated by the addition of functionally diverse herbivores that characterize tropical reefs. This work highlights the importance of assessing the functional traits of range‐shifting species to determine potential mechanisms of impact on ecological communities.
The parrotfish Sparisoma cretense, a marine species native to the eastern and southern
coastal areas of the Mediterranean, has extended its distribution northward. Here, we provide
an update on its distribution based on currently published data and two new records from the
coastline of Sardinia, Italy (central-western Mediterranean). The survey methods were scuba diving
and spearfishing: one specimen of S. cretense was caught along the Argentiera coastline (northwest
Mediterranean) and the others were photographed in the Gulf of Orosei, Osalla Bay (central-eastern
Mediterranean). A literature update, together with new records, documents the distribution of this
species in the northernmost areas of the Mediterranean. Probably a result of global warming, the
ongoing northward expansion of S. cretense highlights the need for sampling campaigns to obtain
timely updates on population and distribution of this thermophilic species
Nibblers (family Girellidae) are reef fishes that are mostly distributed in the Indo-Pacific, with one exception: Girella stuebeli, which is found in the Cabo Verde Archipelago, in the Atlantic Ocean. We capitalized on this unusual distribution to study the evolutionary history of the girellids, and determine the relationship between G. stuebeli and the remaining nibbler taxa. Based on thousands of genomic markers (RAD sequences), we identified the closest relatives of G. stuebeli as being a clade of three species endemic to the northwestern Pacific, restricted to the Sea of Japan and vicinity. This clade diverged from G. stuebeli approximately 2.2 Mya. Two alternative potential routes of migration may explain this affinity: a western route, from the Tropical Eastern Pacific and the Tropical Western Atlantic, and an eastern route via the Indian Ocean and Southern Africa. The geological history and oceanography of the regions combined with molecular data presented here, suggest that the eastern route of invasion (via the Indian Ocean and Southern Africa) is a more likely scenario.
In the present study we used a molecular approach to investigate
the species‐level diversity of the parrotfishes of the genus
Sparisoma, applying the GMYC, bPTP, and ABGD approaches to the
delimitation of species from the Atlantic Ocean and Mediterranean
Sea, using a single locus. The different species delimitation procedures produced distinct results. While the GMYC model recovered 12 species‐level entities, the ABGD approach identified only 10, and the bPTP model, only 11. The GMYC analysis indicated potential species‐level diversity in all four Sparisoma species analyzed here. We agree with the number of lineages established by GMYC,
which corresponded to the method that generated the largest number
of specific entities, or ESUs (Evolutionary Significant Units), in
Sparisoma, followed by bPTP and ABGD.
Corals and seaweeds fulfill important ecological functions in tropical reef ecosystems. In an environment where space is a limiting factor, competition between both players is critical in defining the structure of coral reef communities. Dramatic shifts from coral- to macroalgal-dominated reefs have highlighted competitive interactions between macroalgae and corals. Defining the nature of the interaction between corals and reef algae, however, has been challenging, and it was questioned whether algae are the cause or the consequence of coral reef shifts. Although it is commonly accepted that macroalgae may outcompete corals under conditions of reduced herbivory or enhanced nutrient levels, there is also evidence that algae may have a negligible or even a positive effect on corals in undisturbed reefs. Interactions between macroalgae and corals date back to the Paleocene, when ‘modern’ coralgal reefs became established. Macroalgae and corals share a long evolutionary history. A combination of abiotic and biotic interactions shaped coral reef ecosystems as we presently know them, reaching stable ecological dynamics. However, natural and anthropogenic disturbances may rupture this equilibrium resulting in unbalanced population dynamics. Intensified competition between macroalgae and corals is therefore symptomatic of damaged reefs, and usually results from decrease in herbivory as well as coral morbidity and mortality.
Tub gurnard is one of the most abundant and commercially important Triglidae species in Marine waters of Turkey found in the Black, Marmara, Aegean and Mediterranean Seas. In this study, it’s aimed to investigate genetic and morphological structure of C. lucerna populations. Morphometric and meristic charecters were used for morphological analysis, and 16S rRNA gene of mtDNA was used for DNA sequencing analysis.
As a result of sequencing, the average nucleotide composition of 16S rRNA gene was determined the ratio of T, C, G, A as 21.5%, 26.6 %, 31.4 %, 20.5 % respectively. 16S rRNA region was found as 809 base pair and compused 796 base pair as conserved sites and 13 base pair as variable sites. 6 base pair seen as a indicator between populations. The average value of genetic diversity within populations was 0.0016. The lowest genetic diversity was found in Iskenderun population, while the highest genetic diversity was found Marmara population. A total of 14 haplotypes were obtained from all populations. The highest haplotype diversity was found in Marmara populations, whereas the lowest haplotype diversity was found in Iskenderun population. Akcakoca and Marmara populations showed the least genetic divergence (0.0012). The highest level of genetic divergence was found between Antalya-Izmir populations and between Iskenderun-Izmir (0.0021). According to Neighbour Joining tree, Antalya and Iskenderun populations were found to be close to each other. Izmir population was found to be far away from the other populations.
Discriminant function analysis of morphometric characters showed that Akcakoca populationis completely different from the other populations. It was observed that the other populations are not significantly different from each otherin statistical terms. According to morphological affinity matrix, Antalya and Akcakoca populations were the most distant populations to each other. Izmir and Iskenderun populations were the closest populations to each other. Izmir and Iskenderun populations showed morphological similarity. However, Akcakoca population was found to be highly separated from the other populations.
An important part of our planet’s biodiversity is the genetic diversity, which is the basis of evolution. The loss of genetic diversity entails the risk that species can no longer adapt to (human induced) changes in the environment and become extinct. Additionally, these genetic resources are essential for industries, such as fisheries and aquaculture. The loss of genetically distinct populations within a species is almost as critical as the loss of an entire species. Biotechnology provides molecular tools for the detection and analysis of genetic diversity, which can aid the assessment and monitoring of biodiversity. Molecular genetic key technologies for the assessment and monitoring of biodiversity include polymerase chain reaction (PCR), DNA sequencing, microsatellite markers, and microarrays. This molecular toolbox opens a window to several levels of genetic diversity, such as (1) genetic population structure and gene flow, (2) delineation of species boundaries and evolutionary relationships, and (3) genetic identification of species. This work comprises several case-studies that illuminate, how molecular genetic tools can reveal processes that shape biodiversity on a micro-evolutionary (population genetics) as well as macro-evolutionary (phylogenetics) scale, and how they can be utilised in environmental research and provide baseline data for conservation.
Natural feeding on freshly dead spotted sea hares Aplysia dactylomela by reef fish Sparisoma frondosum at wild was registered and described for the first time, from two different observations made on the northeastern coast of Brazil, South Atlantic.
With a standard set of primers directed toward conserved regions, we have used the polymerase chain reaction to amplify homologous segments of mtDNA from more than 100 animal species, including mammals, birds, amphibians, fishes, and some invertebrates. Amplification and direct sequencing were possible using unpurified mtDNA from nanogram samples of fresh specimens and microgram amounts of tissues preserved for months in alcohol or decades in the dry state. The bird and fish sequences evolve with the same strong bias toward transitions that holds for mammals. However, because the light strand of birds is deficient in thymine, thymine to cytosine transitions are less common than in other taxa. Amino acid replacement in a segment of the cytochrome b gene is faster in mammals and birds than in fishes and the pattern of replacements fits the structural hypothesis for cytochrome b. The unexpectedly wide taxonomic utility of these primers offers opportunities for phylogenetic and population research.
MATERIALS AND METHODS Global and regional differences in the structure of Recent coral reef fish assemblages are eXamined and compared with the fossil record. Family-level comparisons between Atlantic and Indo-Pacific sites show more sparid and haemulid species in the Atlantic but fewer species in characteristic reef families (Labridae, Pomacentridae, Apogonidae, Chaetodontidae) and the absence of several widespread Indo-Pacific reef fish taxa (Caesionidae, Lethrinidae, Nemipteridae, Siganidae). These differences appear to be primarily the result of faunal loss in the Atlantic during the mid-late Cenozoic. The role of non-reef habitats in the evolution of reef fish assemblages is examined based on an analysis of the structure of Recent tropical and temperate fish assemblages and phylogenetic analyses incorporating habitat associations. Throughout their 50 Ma history, reef fish assemblages appear to have maintained strong links with non-reef habitats.
With 5 figures and 4 plates) Manuscript received August 31 st 1990 Summary The status of 7 fossil species and 6 unidentified fossil fragments, currently placed in the family Scaridae are reviewed. Of these, two fragments are considered to be scarid material, both belonging to the genus Bolbometopon SMITH, 1956. The remainder include fragments of Diodontidae, Labridae and Oplegnathidae. Most are of uncertain affinity. A new species, Calotomus preisli n. sp., from the Upper Badenian (Middle Miocene) of the Bay of Eisenstadt, Austria, is described. This new species is the first unequivocal fossil representative of the Scaridae from Europe and provides new evidence for tropical conditions in the Central Paratethys during the Upper Badenian. Zusammenfassung Es erfolgt die Überprüfung der systematischen Zugehörigkeit von 7 fossilen Arten und weiteren 6 fossilen Fragmenten, die bisher der Familie Scaridae (Papageifische) zugeordnet wurden. Nur zwei der Fragmente sind aber eindeutige Belege für Scaridae; es handelt sich um Belege für die Gattung Bolbometopon SMITH, 1956. Einige der übrigen Fossilreste stammen von Diodontidae, Labridae und Oplegnathidae, aber die meisten lassen nicht einmal eine Zuordnung zu einer Familie zu. Ein Neufund aus der gebankten Fazies des Leithakalkes (Oberes Badenien, Mittel-Miozän) der Eisenstädter Bucht (St. Margarethen, Burgenland, Österreich) wird als neue Art beschrieben: Calotomus preisli n. sp. Diese neue Art ist der erste und einzige unzweifelhafte Beleg für fossile Scaridae in Europa und ein weiterer Beleg für tropische Verhältnisse in der Zentralen Paratethys zur Zeit des Oberen Badeniens.
Bootstrapping is a common method for assessing confidence in phylogenetic analyses. Although bootstrapping was first applied in phylogenetics to assess the repeatability of a given result, bootstrap results are commonly interpreted as a measure of the probability that a phylogenetic estimate represents the true phylogeny. Here we use computer simulations and a laboratory-generated phylogeny to test bootstrapping results of parsimony analyses, both as measures of repeatability (i.e., the probability of repeating a result given a new sample of characters) and accuracy (i.e., the probability that a result represents the true phylogeny). Our results indicate that any given bootstrap proportion provides an unbiased but highly imprecise measure of repeatability, unless the actual probability of replicating the relevant result is nearly one. The imprecision of the estimate is great enough to render the estimate virtually useless as a measure of repeatability. Under conditions thought to be typical of most phylogenetic analyses, however, bootstrap proportions in majority-rule consensus trees provide biased but highly conservative estimates of the probability of correctly inferring the corresponding clades. Specifically, under conditions of equal rates of change, symmetric phylogenies, and internodal change of less-than-or-equal-to 20% of the characters, bootstrap proportions of greater-than-or-equal-to 70% usually correspond to a probability of greater-than-or-equal-to 95% that the corresponding clade is real. However, under conditions of very high rates of internodal change (approaching randomization of the characters among taxa) or highly unequal rates of change among taxa, bootstrap proportions >50% are overestimates of accuracy.
Food selection by the Caribbean stoplight parrotfish Sparisoma viride was investigated on a fringing coral reef of Bonaire, Netherlands Antilles. For different reef zones, the diet composition for each life phase was determined by description of randomly selected bites, and compared to the availability of food resources, as determined with the aid of chain-link transects. S. viride employs an excavating grazing mode, and feeds almost exclusively on algae associated with dead coral substrates. Preferred food types are large and sparse turfs growing on carbonate substrates inhabited by endolithic algae. Crustose corallines, with or without algal turfs, are not preferred. Feeding forays were longer on the preferred food types. Foraging preferences are related to nutritional quality of the food types and their yield, i.e. the amounts of biomass, protein and energy that can be ingested per bite, as calculated from the size of grazing scars and the biochemical composition of the algae. In spite of selective foraging, a large proportion of bites is taken on inferior food types. Endolithic algae constitute an important food resource for scraping herbivores, such as S. viride, These algae have relatively high energetic value, and allow a high yield as a result of weakening the carbonate matrix by their boring filaments. The yield of algal resources also depends on the skeletal density of the limestone substrates. On deeper reef parts (> 3.5 m depth), low-density substrates predominate, resulting in higher yields of algae per bite than are attained from high-density substrates that predominate on shallower reef parts. The increased availability of high-yield food and substrate types coincides with the occurrence of haremic territorial behaviour in S. viride males on the deeper reef parts. Territories are defended against conspecifics and have an important function as spawning sites. It is argued that the access to superior food resources on the deeper reef makes territorial defence feasible for S. viride.
On the fringing reef of Bonaire, Netherlands Antilles, a comparative study was made of habitat use, diet selection, foraging behaviour and food acquisition of the parrotfish species Scarus vetula and Sparisoma viride. The species are sympatric and live in the same reef habitats (depth zones). Both species show similar foraging selectivity, but exploit algal resources differently. Preferred food items are turf algae on substrates infested with endolithic algae, whereas crustose corallines are avoided. Foraging preferences are related to yield, i.e. the amount of AFDW (ash-free dry weight), protein and energy that can be harvested per bite. Foraging behaviour differs between the species. S. vetula takes more bites in long forays, has higher bite rates (no, of bites s(-1)), and makes fewer and smaller scars on grazing substrates than S. viride. Furthermore, S. vetula prefers flat substrate surfaces while adult S. viride graze by preference on concave surfaces. Species-specific differences in preference and utilization of grazing substrates are related to feeding mode. S. vetula employs a scraping feeding mode by which mainly epilithic algae are ingested. In contrast, S. viride is an excavating grazer that ingests large amounts of endolithic and crustose algae. Intake and assimilation of algal AFDW, protein and energy were quantified through a combination of laboratory feeding trials and field observations. S. vetula has lower food intake (mg AFDW bite(-1)) than S. viride (0.8 x 10(-3) x fish wet wt, FWW, and 2.3 x 10(-3) x FWW respectively), resulting from smaller (shallower) bites. Assimilation efficiencies of total AFDW, protein and energy by S. vetula were higher than in S. viride grazing on the same dead coral substrates, In spite of different feeding modes and different fractions of the primary production harvested, daily amounts of assimilated nutrients and energy are similar for both species, resulting from higher feeding rates (no. of bites h(-1)) and higher assimilation efficiency in S. vetula.
The reproductive biology and social structure of the protogynous stoplight parrotfish, Sparisoma viride, were investigated from 1985-1987 in shallow-water areas (<10 m) at Grand Turk, Turks and Caicos Islands, B.W.I. Over 225 fish were speared, and size at maturation and sex reversal, number of eggs in ripe females, stage of sexual development and variation in reproductive condition were determined. Sex reversal occurred from 150 to 250 mm standard length (SL). Females matured between 170 and 270 mm SL, but ripe eggs were found only in females >260 mm SL indicating a period of growth between maturation and reproduction. Gonadal indices for females >260 mm SL varied among sites, with higher gonadal indices found at the more exposed sites. Egg counts among ripe females varied widely (5,543-68,606 eggs˙female−1) and showed no relationship to size. Visual surveys indicated that abundance and sex ratio varied according to habitat, with males predominating in the lagoon and females off the ends of the island. Overall, stoplight parrotfish were more abundant off the ends of the island where reproductively active individuals appeared to congregate. Results indicate that S. viride at Grand Turk maintains some sort of haremic mating system.
SummaryA total of 399 parrot fishes (Sparisoma cretense) were collected by trammel bottom nets between April 1985 and October 1986 off Kastellorizo Island (Dodecanese, Greece). The fishes were measured, sexed and aged, to determine some population parameters, such as growth, mortality and exploitation rate. One check mark per year occurring during spring was found on the scales. The relationship between scale radius and fish total length was plotted and the linear description was chosen for the back-calculation of length. No significant difference in length was found between sexes. Most growth takes place in the first three years of life, after which it proceeds slowly. Weight is proportional to the cubic power of length. Females mature in the first year, males in the third; the breeding season covers the summer months (July to September). The male to female ratio was almost 1:1.
Daily food intake by the herbivorous parrotfish Sparisoma v~nde, as well as assimdation efficiencies of algal food, protein and energy, were quantified through a combination of laboratory feeding tnals and field observations. The intake of algal ash-free dry wt (AFDW) per bite increases hnearly with fish wet wt (FWW, g) and algal biomass (mg AFDW cm-2), and is further determined by the skeletal density of the limestone substrate. Low-density substrates yield higher amounts of AFDW per bite than do high-density substrates. The percentage of the total food intake that is derived from endolithic and crustose coralline algae increases with the size of the flsh, and can be 9 0 % depending on the biomass of epilithic algae. The daily pattern of foraging activity is positively correlated with diurnal changes in food quality, while seasonal daylength variations result in 13 % variation in total daily bites taken. Daily number of bites of S. viride in the field decreases wlth fish size, and is further dependent on life phase and foraging depth. In experiments, fish attained an assimilation efficiency of ca 20% from a natural diet of low algal biomass and high-density dead coral substrates that predomi-nates in the shallow reef. Assimilation efficiency was ca 70 % from a diet of high algal biomass and low-density substrates that predominates on the deeper reef parts. In spite of lower daily foraging effort, territorial fish, living in deeper parts of the reef, ingest and assimilate higher amounts of AFDW, pro-tein and energy per day than non-territorial fish foraging on the shallow reef. The difference is caused by increased availability of high-yield food and substrate types inside territories compared to the situ-ation on the shallow reef. Daily assimilated energy (kJ d-') is 0.85 X FWW'.~'~ for fish foraging in the shallow reef zone, and 1.22 X FWW0854 for S. viride foraging inside territories on the deeper reef.
Sediment produced by parrotfishes (family Scaridae) may comprise new bioeroded material and/or reworked sediment. The relative contribution of the two components was examined in two bioeroding Chlorurus species, C. gibbus and C. sordidus, from Lizard Island in the Northern Great Barrier Reef. The relative importance of reworked sediment was determined based on direct estimates of sediment ingestion. In C. gibbus, 2.4% of the sediment produced is reworked. In C. sordidus, reworking contributes 27.2%. Comparisons of sediment size-distributions in epilithic algal communities, gut contents and defaecation sites suggest that both C. gibbus and C. sordidus markedly decrease the particle size of sediment as a result of ingestion and trituration in the pharyngeal apparatus.
Resource use by five sympatric species of parrotfish was quantified in the San Blas Archipelago of the Republic of Panama from March to August 1987. Detailed observations of parrotfishes on patch reefs and surrounding seagrass beds showed that they partition resources with respect to habitat, food and size, but not time. Although parrotfishes shared resources, the proportions of each resource used differed significantly among species. Scarus iserti (Bloch) scraped filamentous microalgae that grew from eroded coral pavement on lower slopes of patch reefs and in halos, the area of sparse vegetation surrounding reefs. Sparisoma viride (Bonnaterre) foraged on upper slopes of patch reefs where they mostly took bites from dead coral and associated algae. S. aurofrenatum (Cuvier and Valenciennes) had the broadest diet, which consisted mostly of seagrasses and macro- and microalgae that were attached to dead coral on lower reef slopes and in halos. Although S. chrysopterum (Bloch and Schneider) commonly occurred on patch reefs, it primarily foraged in seagrass beds that surround them. S. rubripinne (Cuvier and Valenciennes) was distributed most widely, ranging from seagrass beds to reef crests, where it took bites from seagrasses, dead coral and macroalgae. Juveniles of all species occurred on lower slopes or in halos where they scraped filamentous microalgae from coral pavement. As they matured, parrotfishes moved into other habitats changing access to different types of food. All of these parrotfishes fed throughout the daytime, and resource use did not differ between morning and afternoon.
The foraging behavior of the bucktooth parrotfish Sparisoma radians was studied in seagrass beds off St. Croix, US Virgin Islands and in laboratory preference tests. Thalassia testudinum was the dominant item in the field diet with the epiphytized distal portion of the blades most favored. Other seagrasses, Syringodium filiforme and Halodule wrightii, were taken in relation to their abundance. Several algae were also eaten, particularly Halimeda spp. and Penicillus spp. Although abundances of the algae varied, the fish maintained a steady mixture of plant species in their diet. Laboratory feeding tests for various plants presented in pairs showed clear preferences in nearly all cases. The preference hierarchy was (1) T. testudinum with epiphytes, (2) H. wrightii, (3) T. testudinum without epiphytes, (4) S. filiforme, and the algae, (5) Dictyota divaricata, (6) Enteromorpha flexuosa, (7) Caulerpa mexicana, (8) Halimeda incrassata and (9) Penicillis pyriformis. Preference did not vary significantly with satiation. The catch per unit effort (kilocalories absorbed per bite) was calculated for each plant, using (a) absorption values calculated for S. radians fed the different plants, (b) the calorific values for tested plants, and (c) an estimate of the amount of material taken per bite. The ranking of catch per unit effort closely paralleled the preference hierarchy with the exception of C. mexicana which has a toxin. Fish fed diets of single plants, mixed plant diet, and starved controls showed differential survival which paralleled the preference hierarchy, the most preferred plants leading to longest survival. Comparison of laboratory results with field feeding behavior shows that inclusion of plants in the diet is not related directly to preference rank, availability or survival value, but that the fish deliberately eat a variety of plants presumably which maintains balanced diet. This result indicates that models of optimal foraging for herbivores should include nutrient constraints and avoidance of toxins in order to predict accurately the behavior of an animal in the field.
The assumption that parrotfishes represent a single group of grazing herbivores is addressed by morphological, functional and ecological analyses. This assumption is rejected. The 24 scarine parrotfishes from the Great Barrier Reef, Australia, are divided into two functional groups: excavators and scrapers. The osteology and myology of the jaws of all 24 species were examined, and a detailed description of a representative species from each group is provided. The relative weights of the major jaw structures of five representative species were quantified. Morphological differences between species were interpreted in a functional context and were assessed based on observations of feeding in the field. Species in the two functional groups show major group-related differences in their bite speeds, microhabitat utilization patterns, and form and extent of substratum excavation during grazing. Group-related differences are also apparent in feeding rates, foray sizes and bite rates. The division of the species into two functional groups is supported by both morphological and behavioural observations. The ecological significance of two functional groups within the family is discussed in relation to the role of parrotfishes on reefs, particularly in terms of bioerosion.
The major agents of biological destruction of coral reefs can be divided into grazers, etchers and borers. Each of these groups is reviewed on a world wide basis, together with the mechanisms by which they destroy the coral substrate. Rates of bioerosion attributed to major agents of grazers, etchers and borers are given, together with limitations of some of the measurements. Recent work is highlighting the variability in rates of bioerosion both over time and space. Factors which may be responsible for this variability are discussed. Bioerosion is a major factor influencing reef morphology and the ways in which this is achieved is discussed in some detail. Although the review concentrates mainly on present day reefs, some attempt is made to consider the impact of bioerosion on older reefs.
A maximum likelihood method for inferring evolutionary trees from DNA sequence data was developed by Felsenstein (1981). In evaluating the extent to which the maximum likelihood tree is a significantly better representation of the true tree, it is important to estimate the variance of the difference between log likelihood of different tree topologies. Bootstrap resampling can be used for this purpose (Hasegawa et al. 1988; Hasegawa and Kishino 1989), but it imposes a great computation burden. To overcome this difficulty, we developed a new method for estimating the variance by expressing it explicitly.The method was applied to DNA sequence data from primates in order to evaluate the maximum likelihood branching order among Hominoidea. It was shown that, although the orangutan is convincingly placed as an outgroup of a human and African apes clade, the branching order among human, chimpanzee, and gorilla cannot be determined confidently from the DNA sequence data presently available when the evolutionary rate constancy is not assumed.
A description of the daily feeding patterns is provided for the two most abundant bioeroding parrotfish species on the Great Barrier Reef: Chlorurus gibbus and C. sordidus. This study was based at Lizard Island in the northern section of the GBR. Dawn to dusk observations were used to estimate feeding rates and the length of the feeding day. The effects of location and season on these values were also examined. These data are used to estimate erosion rates by the two species. A direct estimate of erosion rates by the two species is provided based on estimated feeding rates and bite sizes. In both species, feeding was characterized by an abrupt initiation and termination with relatively consistent feeding rates throughout the day. Both species spend 83.6 to 91.0% of daylight hours feeding. The proportion of daylight hours spent feeding increases in winter, although the actual length of time spent feeding decreases. Overall, the two species are similar in their general feeding patterns but differ in their seasonal response to time of day effects. C. gibbus is conservative with limited site, time of day or seasonal changes in feeding rates. In contrast, C. sordidus displayed marked seasonal changes in daily feeding rates and diel feeding patterns, with higher feeding rates in the afternoon and in the summer. Seasonally adjusted estimates of annual erosion rates per individual of the two species were 1017.7186.3 kg yr-1 (0.410.07 m3 yr-1) for C. gibbus and 23.63.4 kg yr-1 (9.7 10-31.3 10-3 m2yr-1) for C. sordidus. The likely impact of location and season effects on erosion rates appear to be limited.
With a standard set of primers directed toward conserved regions, we have used the polymerase chain reaction to amplify homologous segments of mtDNA from more than 100 animal species, including mammals, birds, amphibians, fishes, and some invertebrates. Amplification and direct sequencing were possible using unpurified mtDNA from nanogram samples of fresh specimens and microgram amounts of tissues preserved for months in alcohol or decades in the dry state. The bird and fish sequences evolve with the same strong bias toward transitions that holds for mammals. However, because the light strand of birds is deficient in thymine, thymine to cytosine transitions are less common than in other taxa. Amino acid replacement in a segment of the cytochrome b gene is faster in mammals and birds than in fishes and the pattern of replacements fits the structural hypothesis for cytochrome b. The unexpectedly wide taxonomic utility of these primers offers opportunities for phylogenetic and population research.
Las migraciones internas en España experimentan en la década de los ochenta importantes cambios que afectan tanto al volumen de los flujos, que se reducen claramente, como a su estructura interna (edad, profesiones, etc.) y a la matriz de origen-destino de los migrantes. Junto a la creciente importancia de los movimientos a corta distancia se distinguen tres procesos simultáneos: periurbanización, retornos de antiguos migrantes, y consolidación de nuevas áreas de atracción; el peso de cada uno de éstos varía a lo largo de la década en función de la coyuntura económica.
The bootstrap is a statistical method for obtaining a nonparametric estimate of error (Efron 1979, 1982). Felsenstein ( 1985) was the first to apply this method to phylogeny estimation, and his approach is now widely used. Taxa are held constant, and the characters (for sequence data, nucleotide or amino acid sites) are resampled randomly with replacement. A phylogeny is constructed from each replication of the data, and the frequency of appearance of particular phylogenetic groups (groups of alleles or taxa) among all of the trees constructed by this resampling is the bootstrap confidence limit, or bootstrap P value (BP). The BPS of different nodes within a tree can be used to assess the relative stability of those phylogenetic groups or, if applied strictly, to test their statistical significance (e.g., at the 95% or 99% level). The appli- cation of bootstrapping to phylogeny estimation is a tradeoff between the maximum number of replications that can be performed by the researcher in a reasonable amount of time and the minimum number of replications needed for accurate estimation of the BP. The purpose of the present report is to explore the variance (and hence the accuracy) of the phylogenetic BP and to establish guidelines for efficient bootstrap sampling. BP is the proportion of trees containing a particular phylogenetic group. It there- fore follows the binomial distribution, which has a variance of o2 = (P( 1 - P)/n), where P is the BP and II is the number of replications. Although Li and Gouy ( 1990) recently suggested that more replications are needed for larger numbers of taxa, the accuracy of the BP is a function only of P and n. If the interval containing 95% of the samples (+ 1.96 standard deviations) is used as a measure of accuracy, then the application of the above formula shows that 1,825 replications ( =0.95 X 0.05( 1.96/ 0.0 1 )2) are needed to attain + 1% accuracy at a BP of 0.95 (fig. 1). This is more than an order of magnitude higher than the number of replications (50-100) normally used in phylogenetic analyses. Based on this, a practical guideline for efficient and accurate bootstrap sampling can be made: If one wishes the expectation to be that the 95% confidence range is + 1% of the BP, then one must perform 2,000 bootstrap replications (if BP = 0.95) or 400 replications (if BP = 0.99) in phylogenetic analyses, unless the computational time is prohibitive; additional replications are unnecessary, and fewer replications may sacrifice statistical accuracy. Moreover, statistical testing at the 95% level cannot be made using ~73 replications, even if the group is supported by a BP of 1 .OO. This is because the inaccuracy at a mean BP of 0.95 is greater than f5% (fig. 1) when ~73
The subfamily of fishes Trematominae is endemic to the subzero waters of antarctica and is part of the longer notothenioid radiation. Partial mitochondrial sequences from the 12S and 16S ribosomal RNA (rRNA) genes and a phylogeny for 10 trematomid species are presented. As has been previously suggested, two taxa, Trematomus scotti and T. newnesi, do not appear to be part of the main trematomid radiation. The genus Pagothenia is nested within the genus Trematomus and has evolved a unique cyropelagic existence, an association with pack ice. Using a mitochondrial rRNA molecular clock rate of 0.14% transversion changes per million years, the average age of the trematomids is estimated at 3.4 million years (MY). If the age of the trematomids is approximately 3.4 MY, this group could have speciated during the period of deglaciation in Antarctica 2.5-4.8 million years ago. This era was marked by significant changes on the Antarctic shores, such as the opening of fjords, which might have provided a stimulus for specification.
Forty-nine complete 12S ribosomal RNA (rRNA) gene sequences from a diverse assortment of mammals (one monotreme, 11 marsupials, 37 placentals), including 11 new sequences, were employed to establish a "core" secondary structure model for mammalian 12S rRNA. Base-pairing interactions were assessed according to the criteria of potential base-pairing as well as evidence for base-pairing in the form of compensatory mutations. In cases where compensatory evidence was not available among mammalian sequences, we evaluated evidence among other vertebrate 12S rRNAs. Our results suggest a core model for secondary structure in mammalian 12S rRNAs with deletions as well as additions to the Gutell (1994: Nucleic Acids Res. 22) models for Bos and Homo. In all, we recognize 40 stems, 34 of which are supported by at least some compensatory evidence within Mammalia. We also investigated the occurrence and conservation in mammalian 12S rRNAs of nucleotide positions that are known to participate in the decoding site in E. coli. Twenty-four nucleotide positions known to participate in the decoding site in E. coli also occur among mammalian 12S rRNAs and 17 are invariant for the same base as in E. coli. Patterns of nucleotide substitution were assessed based on our secondary structure model. Transitions in loops become saturated by approximately 10-20 million years. Transitions in stems, in turn, show partial saturation at 20 million years but divergence continues to increase beyond 100 million years. Transversions accumulate linearly beyond 100 million years in both stems and loops although the rate of accumulation of transversions is three- to fourfold higher in loops. Presumably, this difference results from constraints to maintain pairing in stems.
The use of molecular phylogenies to examine evolutionary questions has become commonplace with the automation of DNA sequencing
and the availability of efficient computer programs to perform phylogenetic analyses. The application of computer simulation
and likelihood ratio tests to evolutionary hypotheses represents a recent methodological development in this field. Likelihood
ratio tests have enabled biologists to address many questions in evolutionary biology that have been difficult to resolve
in the past, such as whether host-parasite systems are cospeciating and whether models of DNA substitution adequately explain
observed sequences.
The hypotheses that grazing losses and/or ambient inorganic nitrogen concentrations control the standing crop of the epilithic algal community were tested in two habitats at One Tree Reef (Great Barrier Reef, Australia). Short (12-15 days) and long (167-306 days) multifactorial experiments using grazer exclusion and nitrogen fertilization treatments were used to partition variance in algal community biomass on portable segments of natural reef substratum during 1980. On outer reef slopes, inorganic nitrogen limited algal community growth, but the standing crop was determined by grazing losses. In the subtidal lagoon inorganic nitrogen and grazing alternated seasonally in controlling standing crop. The recolonization of cleared natural substratum was followed at two additional sites. The algal standing crop in subtidal habitats reached control levels within 4 months, while that in an intertidal reef habitat took up to 14 months. The standing crop of benthic algae on natural reef substrata was monitored in all habitats over 2 yr. In shallow and intertidal habitats, the standing crop was three to five times higher than in deeper areas, and showed a spatial and seasonal variation apparently controlled by factors other than grazing intensity, despite high levels of yield to grazers. Seasonal variation was much less in subtidal habitats. It is concluded that only within limited temporal and spatial scales is grazing intensity alone an adequate predictor of benthic algal standing crop.
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data, In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Abstract — Commonly used permutation tail probability (PTP) and topology dependent permutation tail probability (T-PTP) tests incorporate an inappropriate treatment of designated outgroup taxa, and for that reason are biased either for (PTP) or for or against (T-PTP) rejection of the null hypothesis. A modified test is proposed, in which this source of bias is eliminated.
Based on a critical review of the literature, and on abundant material recently collected along the coast of Brazil, a synthesis of the occurrence and distribution of the Brazilian species of seagrasses is presented. Two species of Hydrocharitaceae, namely Halophila baillonii Aschers. and H. decipiens Ostenf., and three species of Potomogetonaceae, Halodule emarginata den Hartog, H. wrightii Aschers. and Ruppia maritima L. s.s., were found. Ruppia maritima was found all along the Brazilian coast, from about 3 to 32°S latitude, in brackish water ponds and lakes, with salinities varying from 0.3 to 28ℵ.. The other species are restricted to normal seawater (±35ℵ.), although Halodule wrightii was also found in hypersaline waters (45ℵ.). This last species is the most common seagrass in Brazil. The species of Halophila are restricted to warmer waters and were not found further south than Rio de Janeiro State, being especially common on the NE coast at depths down to 62 m. During the course of these studies, Halophila baillonii was found for the second time in Brazil since its original collection in 1888. A critical comparison of a large number of plants leads us to the conclusion that Halodule brasiliensis Lipkin cannot be distinguished from H. wrightii, and H. lilianae den Hartog cannot be distinguished from H. emarginata. Reproductive structures of H. emarginata are described for the first time.
In the protogynous stoplight parrotfish (Sparisoma viride), large males defend territories that encompass the home-ranges of several mature females. However, high-quality habitat is in short supply, such that smaller, competitively inferior males do not defend territories. We investigated the role of 11-ketotestosterone (11KT) and testosterone (T) in the regulation of territorial behavior in a wild population of a protogynous reef fish, the stoplight parrotfish, at Glover's Reef, Belize. Radioimmunoassay of plasma samples from individuals of known social status revealed that nonterritorial males have lower levels of T and 11KT than territorial males. Nonterritorial males allowed access to vacant territories underwent pronounced increases in T and 11KT. When sampled 1 week after territory acquisition, levels of T and 11KT in new territorial males were significantly higher than the levels in established territorial males, but by 3 weeks after territory acquisition, there was no significant difference. We further investigated the hypothesis that such short-term increases in androgen levels are a response to intense male-male interactions during territory establishment. Simulated territorial intrusion promoted increased plasma levels of both T and 11KT while access to vacant territories without neighboring territorial males did not. These findings suggest that the endocrine system plays a role in fine-tuning the levels of territorial aggression exhibited by male stoplight parrotfish. We discuss these results in light of recent theory in behavioral endocrinology.
Publisher Summary This chapter examines the range of the variation of secondary structure among the 16-S-like rRNAs. This brings into a larger structural context a recent detailed analysis of the individual helical elements and provides a basis for an accurate alignment of the corresponding regions of different primary structures. Computer-assisted comparative is used in the analysis of aligned sequences to describe the pattern of phylogenetic conservation for each nucleotide position in 16-S rRNA. A search for matching patterns among unpaired positions in the RNA chain then produces a list of candidates for potential base–base tertiary interactions. The completion of nucleotide sequences for 34 16-S-like rRNAs includes 4 eubacteria, 4 chloroplasts, 12 mitochondria, 4 archaebacteria, and 10 eukaryotes. Secondary structure models for these molecules have been developed in the course of refinement of the E. coli model, and have been used to arrive at improved sequence alignments for the 16-S-like rRNAs. Schematic drawings of (1) eubacterial, (2) archaebacterial, (3) eukaryotic cytoplasmic, (4) plant mitochondrial, (5) fungal mitochondrial, and (6) mammalian mitochondrial structures are shown in the chapter.
including seagrass beds, rocky reefs, and coral reef-fringe and back-reef areas of coral rubble and sand, as well as zones dominated by live coral
- Gonzalez
Species of Sparisoma are found in a particularly broad range of tropical marine habitats (Bellwood, 1994), including seagrass beds, rocky reefs, and coral reef-fringe and back-reef areas of coral rubble and sand, as well as zones dominated by live coral (e.g., Robertson and Warner, 1978; Randall, 1996; Petrakis and Papacon-stantinou, 1990; Gonzalez et al., 1993; Bellwood, 1994; McAfee and Morgan, 1996).
Reef fishes: Their history and evolutionThe Ecology of Fishes on Coral Reefs
- Academic Press
- London
- J H Choat
- D R Bellwood
Academic Press, London. Choat, J. H., and Bellwood, D. R. (1991). Reef fishes: Their history and evolution. In ''The Ecology of Fishes on Coral Reefs'' (P. F. Sale, Ed.), pp. 39–68.
6 are species of Sparisoma that are ecologically important components of coral reef communities in that area. Species of Sparisoma have been the subject of a wide variety of ecological, evolutionary, and bio-chemical studies during the past 30 years (e.g., Robert-son and Warner
- Bruggemann
Of the 14 scarids found in the Caribbean (Randall, 1996), 6 are species of Sparisoma that are ecologically important components of coral reef communities in that area. Species of Sparisoma have been the subject of a wide variety of ecological, evolutionary, and bio-chemical studies during the past 30 years (e.g., Robert-son and Warner, 1978; Lobel and Ogden, 1981; Cardwell and Liley, 1991; Koltes, 1993; Bruggemann et al., 1994).