Asymmetric conspecific sperm precedence in relation to spawning times in the Montastraea annularis species complex (Cnidaria: Scleractinia)

In broadcast spawners, prezygotic reproductive isolation depends on differences in the spatial and temporal patterns of gamete release and gametic incompatibility. Typically, gametic incompatibility is measured in no‐choice crosses, but conspecific sperm precedence (CSP) can prevent hybridization in gametes that are compatible in the absence of sperm competition. Broadcast spawning corals in the Montastraea annularis species complex spawn annually on the same few evenings. Montastraea franksi spawns an average of 110 min before M. annularis, with a minimum gap of approximately 40 min. Gametes are compatible in no‐choice heterospecific assays, but it is unknown whether eggs exhibit choice when in competition. Hybridization depends on either M. franksi eggs remaining unfertilized and in proximity to M. annularis when the latter species spawns or M. franksi sperm remaining in sufficient viable concentrations when M. annularis spawns. We found that the eggs of the early spawning M. franksi demonstrate strong CSP, whereas CSP appears to be lacking for M. annularis eggs. This study provides evidence of diverging gamete affinities between these recently separated species and suggests for the first time that selection may favour CSP in earlier spawning species when conspecific sperm is diluted and aged and is otherwise at a numeric and viability disadvantage with heterospecific sperm.     

Reproductive Cycle and Developmental Processes During Embryogenesis of Clavularia hamra (Cnidaria,Octocorallia)

Clavularia hamra Gohar, 1948 is a common octocoral on the reefs of the Gulf of Eilat (northern Red Sea). Reproductive biology of C. hamra was studied for two years. Direct observations of spawning were conducted in situ and in aquaria. Cleavage of eggs and further embryo metamorphosis into mature planulae were examined by scanning electron microscopy and histological sections. Clavularia hamra is dioecious. Young oocytes appear annually in September, gradually grow in size and attain maturity within 11 months. The main spawning event of the population is highly synchronized, occurring on a single night each year between the last quarter and the new moon in mid summer. The released orange eggs are held together by mucus and remain attached amongst the polyps on the outside of the female colonies. Twelve hours after spawning, the young embryos have blastomeres of equal size with numerous microvilli on their external surface. Due to unequal cleavage, bizarre embryos are also formed. By 48 h a blastopore is visible, indicating that a gastrula is developing. Eight days after spawning mature planulae are observed. The external mode of embryo development on the surface of the parent colony reduces dispersal of the planulae. However, this reproductive feature enhances formation of locally dense populations of C. hamra, with distinct habitat preferences.     

Lipid content of Favia fragum larvae: changes during planulation

Lipid content of planula larvae of the brooding scleractinian coral Favia fragum was analysed through a monthly planulation period. The average lipid content per dry weight of F. fragum planulae was 39%, which is low compared with other scleractinian coral species. Lipid content of planulae was significantly affected by the day of release and decreased during the planulation period.     

Estimating the spawning activity of fish species using nuclear and mitochondrial environmental DNA concentrations and their ratios

1. During spawning activity, fish release large amounts of sperm and eggs into the water, which has been assumed to cause an increase in environmental DNA (eDNA) levels and nuclear DNA/mitochondrial DNA ratios. To test whether these assumptions are valid and whether nuclear and mitochondrial eDNA analysis can be used to monitor the spawning activity of freshwater fish, we conducted field eDNA surveys and traditional surveys using common carp (Cyprinus carpio), largemouth bass (Micropterus salmoides) and bluegill sunfish (Lepomis macrochirus) as model species.
2. Fish spawning periods were estimated based on age, as estimated using the body lengths of juveniles collected in the Miharu reservoir in Fukushima, Japan. The results showed that the main spawning periods of largemouth bass and bluegill sunfish were from April to July and from July to August, respectively.
3. Field eDNA surveys were conducted in the Hebisawagawa front reservoir, which is connected to the Miharu reservoir. From March to August 2019 and 2020, weekly eDNA sampling was conducted at three sites, and daily sampling was conducted at six sites from 23 June to 3 July 2020. The eDNA concentrations of the nuclear internal transcribed spacer 1 (ITS1) and mitochondrial cytochrome B (CytB), as well as the ITS1/CytB ratio, were measured for each of the three fish in each water sample. Water temperature had a statistically significant effect on eDNA concentration, probably reflecting the relationship between water temperature and spawning.
4. We created generalised additive mixed models to estimate spawning activity periods based on weekly eDNA data. The estimated periods of spawning activity for common carp, largemouth bass and bluegill sunfish were March to May, May to July, and May to August, respectively. The estimated spawning periods coincided with known fish ecology or the results of traditional methods. This method also has been applied to daily eDNA samples, showing the feasibility of high-resolution estimation of spawning activity.
5. For common carp and bluegill sunfish, we were able to estimate the spawning period using this method. Although the method is affected by biomass and the diffusion and degradation of eDNA, it has the potential to accurately estimating spawning activities. These then can be estimated without conducting laborious traditional surveys, facilitating the monitoring of reproduction by rare, invasive or important fishery species. Further research on the diffusion distance and degradation time of the eDNA concentration peak caused by fish spawning activity may improve the accuracy of monitoring.

     

Reproductive characteristics and gamete development of the soft coral Sclerophytum cf. heterospiculatum in Okinawa Island,Japan

Sexual reproduction data are important to understand how organisms can replenish their populations and proliferate on coral reefs. Despite the importance of such data, the reproductive characteristics of most soft coral species are still unknown. Here, we examined the reproductive strategies of a species from the often-dominant genus Sclerophytum in a coral reef on subtropical Okinawa Island, Japan. DNA barcoding and histological examinations of the tissues were conducted to confirm colony conspecificity and identify reproductive characteristics, respectively, between March 2020 and March 2021. Results indicated that the studied species, identified as Sclerophytum cf. heterospiculatum, exhibits gonochorism with longer oogenesis and shorter spermatogenesis. Female colonies produced immature oocytes throughout the year, with mature oocytes observed from late July to early September, and thus, extended spawning is likely characteristic of this species. In male colonies, spermatogenesis took place over ~5 months, with spermaries present from April through August. Mature spermaries were noted beginning in July and the inferred peak of sperm release was between late August and early September, which suggests that spermatogenesis duration was ~5 months. The largest mean oocyte and spermary sizes (628.45 ± 61.36 and 240.04 ± 49.49 μm, respectively) were both recorded in August. Gamete spawning presumably occurred during the summer season, which suggests seasonality in reproduction as influenced by changes in seawater temperature. However, the proximate cue for exact dates of spawning could be the lunar period because the inferred release of spawning materials seemed to occur between full moon and last-quarter moon phases in both the months of August and September. The results of this study represent the first detailed report of reproductive characteristics of the genus Sclerophytum in Japan.     

Polymorphism in a common Atlantic reef coral (Montastraea cavernosa) and its long-term evolutionary implications

Recent advances in morphometrics and genetics have led to the discovery of numerous cryptic species in coral reef ecosystems. A prime example is the Montastraea annularis scleractinian coral species complex, in which morphological, genetic, and reproductive data concur on species boundaries, allowing evaluation of long-term patterns of speciation and evolutionary innovation. Here we test for cryptic species in the Atlantic species, Montastraea cavernosa, long recognized as polymorphic. Our modern samples consist of 94 colonies collected at four locations (Belize, Panamá, Puerto Rico in the Caribbean; S?o Tomé in the Eastern Atlantic). Our fossil samples consist of 78 colonies from the Plio-Pleistocene of Costa Rica and Panamá. Landmark morphometric data were collected on thin sections of 46 modern and 78 fossil colonies. Mahalanobis distances between colonies were calculated using Bookstein coordinates, revealing two modern and four fossil morphotypes. The remaining 48 of the 94 modern colonies were assigned to morphotype using discriminant analysis of calical measurements. Cross-tabulation and multiple comparisons tests show no significant morphological differences among geographic locations or water depths. Patterns of variation within and among fossil morphotypes are similar to modern morphotypes. DNA sequence data were collected for two polymorphic nuclear loci (β-tub1 and β-tub2) on all 94 modern colonies. Haplotype networks show that both genes consist of two clades, but morphotypes are not associated with genetic clades. Genotype frequencies and two-locus genotype assignments indicate genetic exchange across clades, and ϕst values show no genetic differentiation between morphotypes at different locations. Taken together, our morphological and genetic results do not provide evidence for cryptic species in M. cavernosa, but indicate instead that this species has an unusually high degree of polymorphism over a wide geographic area and persisting for >25 million years (myr).     

Larval development and survivorship in the corals Favia favus and Platygyra lamellina

Two Red Sea faviid species, Favia favus and Platygyra lamellina spawn eggs and sperm once a year, during the summer. External fertilization occurs 0.5 h after spawning, and mobile gastrulae appear 20 h later. Four stages in the early ontogenesis of these corals are described. The slow development (2–3 months) to the polyp stage in broadcasting species is attributed to the lack of zooxanthellae in their planulae and their appearance in the primary polyp only at a later stage. Survivorship of one-month-old primary polyps is ca 0.21% and 0.25% in F. favus and P. lamellina respectively, from the populations of 2–9-day-old planulae. Despite these low rates of survival, both species form dense populations in the Gulf of Eilat.     

Analysis of complete mitochondrial DNA sequences of three members of the Montastraea annularis coral species complex (Cnidaria, Anthozoa, Scleractinia)

Complete mitochondrial nucleotide sequences of two individuals each of Montastraea annularis, Montastraea faveolata, and Montastraea franksi were determined. Gene composition and order differed substantially from the sea anemone Metridium senile, but were identical to that of the phylogenetically distant coral genus Acropora. However, characteristics of the non-coding regions differed between the two scleractinian genera. Among members of the M. annularis complex, only 25 of 16,134 base pair positions were variable. Sixteen of these occurred in one colony of M. franksi, which (together with additional data) indicates the existence of multiple divergent mitochondrial lineages in this species. Overall, rates of evolution for these mitochondrial genomes were extremely slow (0.03–0.04% per million years based on the fossil record of the M. annularis complex). At higher taxonomic levels, patterns of genetic divergence and synonymous/nonsynonymous substitutions suggest non-neutral and unequal rates of evolution between the two lineages to which Montastraea and Acropora belong.     

Synchronous spawning in a recently established population of the zebra mussel,Dreissena polymorpha,in western Lake Erie,USA

Seasonal patterns of reproduction in a newly established population of Dreissena polymorpha are described for a site in the western basin of Lake Erie. Reproduction was monitored by histological examination of gonads, analysis of shell length-dry weight relationships, and following abundance of planktonic and settling larvae. Patterns of planktonic larval abundance and settling showed a distinct bimodal pattern in July and August, but conflicted with histological and length-dry weight data that showed spawning was a brief, highly synchronous event occurring in late August. Differences between histological data and abundance of larvae in the plankton can be explained as resulting from drift of larvae from disjunct populations of D. polymorpha which spawned earlier, into the study area. Veliger larvae were present in the plankton at low densities throughout the warm months before and after periods of peak abundances. The presence of these larvae can be explained by a combination of asynchronous spawning among local populations and postponed settlement by planktonic larvae.     

Metamorphosis and acquisition of symbiotic algae in planula larvae and primary polyps of Acropora spp.

Coral planulae settle, then metamorphose and form polyps. This study examined the morphological process of metamorphosis from planulae into primary polyps in the scleractinian corals Acropora nobilis and Acropora microphthalma, using the cnidarian neuropeptide Hym-248. These two species release eggs that do not contain Symbiodinium. The mode of acquisition of freshly isolated Symbiodinium (zooxanthellae) (FIZ) by the non-symbiotic polyp was also examined. Non-Hym-248 treated swimming Acropora planulae did not develop blastopore, mesenteries or coelenteron until the induction of metamorphosis 16 days after fertilization. The oral pore was formed by invagination of the epidermal layer after formation of the coelenteron in metamorphosing polyps. At 3 days after settlement and metamorphosis, primary polyps exposed to FIZ established symbioses with the Symbiodinium. Two–four days after exposure to FIZ, the distribution of Symbiodinium was limited to the gastrodermis of the pharynx and basal part of the polyps. Eight–ten days after exposure to FIZ, Symbiodinium were present in gastrodermal cells throughout the polyps.     

Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

The relationship between corallite morphology and colony shape in some massive reef-corals

Multivariate statistical tests are performed on three coral species to determine if a relationship exists between colony shape and corallite morphology. Hemispherical to branched colony shapes are emphasized. The material consists of specimens of Montastraea annularis and M. cavernosa from modern reef habitats in Jamaica. Colonies of the more branched M. limbata from the Neogene of the Dominican Republic are analyzed for comparison. The characters studied consist largely of linear measurements of colony dimensions and corallite structures in transverse thin-sections. The results show that different complexes of corallite characters vary within each species. With the possible exception of a complex describing theca thickness in M. annularis, none of these corallite complexes is related to variation in colony shape within species. Colony shape variation within species is related to upward colony growth rate. These results indicate that corallite morphology and colony shape can be represented by two different vectors of variation within species. Comparisons of intraspecific with interspecific patterns suggest that a strong genetic component explains colony shape variation in corals.     

Seasonal and interannual variation of the ichthyoplankton off the coast of Northern Morocco

Changes in the qualitative and quantitative composition of the ichthyoplankton off the coast of Northern Morocco (36–28° N) are analyzed in different seasons of 1994–2008. Pelagic eggs and larvae of over 80 fish species belonging to 57 families are recorded. In cold periods at the Morocco coast, there is maximum spawning of fish belonging to the subtropical (Mediterranean-Atlantic) fauna, the dominant of which are European pilchard Sardina pilchardus, common scad Trachurus trachurus, and Atlantic mackerel Scomber japonicus. In warm periods, in addition to the aforementioned species, the ichthyopankton often comprises eggs and larvae of fish form the tropical fauna. Among the latter are round sardinella Sardinella aurita and West-African mackerel T. trecae dominate. The spawning biomass of principal commercial fish is also calculated by the quantity of eggs laid by females.     

Larval development of certain gamete-spawning scleractinian corals

Embryogenesis and larval development were documented in 19 species of hermatypic scleractinians which release gametes during the summer coral spawning season on the Great Barrier Reef. Cleavage of fertilized eggs began approximately 2 h after spawning in all species, and gave rise to blastulae after 7–10 h. Endoderm formation in Platygyra sinensis was by invagination, and this appeared to occur in all species studied. All species observed at 36 h after spawing were mobile and full mobility was reached by 48 h. Settlement of planulae placed in aquaria occurred between 4 and 7 days after fertilization. These results suggest that larval corals produced by most gamete-releasing coral species are likely to be dispersed away from the parent reef.     

The spawning behaviour of the endangered freshwater fish Ladigesocypris ghigii (Gianferrari, 1927)

Ladigesocypris ghigii (Gianferrari, 1927), commonly called gizani, is an endangered freshwater fish endemic to the Greek island of Rhodes. The spawning behaviour of gizani was studied in aquaria, to develop an artificial breeding technique for this endangered species. Spawning in captivity was recorded over 41 spawning days between 15 February and 14 July 2001, under constant temperature (18 or 21°C) and photoperiod 14L/10D. The number of spawning acts in a day was 82 ± 28. No aggressiveness and no territoriality were developed. The mating system of the fish was polygamous, with females spawning more than once. In captivity, eggs are laid either on aquatic plants or on gravel, while in nature fish spawn mainly on algae and plants. The latter suggests that, as far as breeding substrate is concerned, gizani is an opportunistic species using different substrates when necessary.     

Reproductive ecology of the deep-sea scleractinian coral <Emphasis Type="Italic">Fungiacyathus marenzelleri</Emphasis> (Vaughan, 1906) in the northeast Atlantic Ocean

The reproductive biology and its seasonality were examined in the deep-water, solitary coral Fungiacyathus marenzelleri from 2,200 m depth in the NE Atlantic, using histological techniques. A total of 186 corals were collected using either an Otter trawl (semi-balloon) or Agassiz trawl from the research vessel RRS Challenger between 1979 and 1991. All individuals were gonochoric, with all 48 mesenteries being fertile. A high fecundity was recorded (2,892 ᇀ.4 oocytes per polyp) with strong size-dependency. This species is a quasi-continuous reproducer with overlapping gametogenesis for both spermacysts and oocytes. No planulae were observed and broadcasting of gametes is inferred, with the large egg size (max. 750-µm diameter) suggesting lecithotrophic development. There is a non-significant trend towards a synchronous spawning event during June/July. Asexual fission was observed, though in low numbers.     

New records of rare derogenids (Digenea: Hemiuroidea) from Mediterranean sparids,including the description of a new species of <Emphasis Type="Italic">Magnibursatus</Emphasis> Naidenova, 1969 and redescription of <Emphasis Type="Italic">Derogenes adriaticus</Emphasis> Nikolaeva, 1966

Records of derogenid digeneans in the Mediterranean and Black Sea region are scarce and tend to be restricted to a small number of host-groups, but especially to sparid fishes. This work reports on the presence of derogenine and halipegine derogenids from two sparids, Diplodus annularis (L.) and D. sargus (L.), from off the Mediterranean coast of Spain. Five derogenid forms were recovered. Derogenes adriaticus Nikolaeva, 1966 is redescribed from Diplodus annularis, and Derogenes sp. is described from the same host but differentiated from the former species. Magnibursatus barretti n. sp. is described from Diplodus sargus and distinguished from other species of the genus especially by its smaller body size and smaller eggs. M. bartolii Kostadinova, Power, Fernández, Balbuena, Raga & Gibson, 2003 is redescribed from D. sargus, a new host for this species. A single specimen from D. sargus, somewhat similar to M. minutus Kostadinova, Power, Fernández, Balbuena, Raga & Gibson, 2003, is also described, as it exhibits some morphometric differences from the latter species.     

Reproductive biology of three sponge species of the genus Xestospongia (Porifera: Demospongiae: Petrosida) from the Great Barrier Reef

The reproductive development of three species of the Petrosida, Xestospongia bergquistia, X. exigua, and X. testudinaria, was monitored for four years on a fringing reef at Orpheus Island, Great Barrier Reef, Australia. All three species were oviparous and female reproductive activity began prior to males becoming active. X. bergquistia and X. testudinaria were gonochoric and broadcast eggs in spawning events that were synchronous within species. Egg development occurred over more than five months in X. bergquistia and X. testudinaria and two months in X. exigua. Spawning was during periods of warm temperature and occurred in October or November for X. bergquistia and X. testudinaria, and January or February for X. exigua. Lunar phase was implicated in timing of spawning of X. testudinaria. Diel timing of spawning in X. testudinaria and X. bergquistia was consistently a morning event.     

Corallite wall and septal microstructure in scleractinian reef corals: Comparison of molecular clades within the family Faviidae

Recent molecular phylogenies conflict with traditional scleractinian classification at ranks ranging from suborder to genus, challenging morphologists to discover new characters that better agree with molecular data. Such characters are essential for including fossils in analyses and tracing evolutionary patterns through geologic time. We examine the skeletal morphology of 36 species belonging to the traditional families Faviidae, Merulinidae, Pectiniidae, and Trachyphylliidae (3 Atlantic, 14 Indo‐Pacific, 2 cosmopolitan genera) at the macromorphological, micromorphological, and microstructural levels. Molecular analyses indicate that the families are not monophyletic groups, but consist of six family‐level clades, four of which are examined [clade XV = Diploastrea heliopora; clade XVI = Montastraea cavernosa; clade XVII (“Pacific faviids”) = Pacific faviids (part) + merulinids (part) + pectiniids (part) + M. annularis complex; clade XXI (“Atlantic faviids”) = Atlantic faviids (part) + Atlantic mussids]. Comparisons among molecular clades indicate that micromorphological and microstructural characters (singly and in combination) are clade diagnostic, but with two exceptions, macromorphologic characters are not. The septal teeth of “Atlantic faviids” are paddle‐shaped (strong secondary calcification axes) or blocky, whereas the septal teeth of “Pacific faviids” are spine‐shaped or multidirectional. Corallite walls in “Atlantic faviids” are usually septothecal, with occasional trabeculothecal elements; whereas corallite walls in “Pacific faviids” are usually trabeculothecal or parathecal or they contain abortive septa. Exceptions include subclades of “Pacific faviids” consisting of a) Caulastraea and Oulophyllia (strong secondary axes) and b) Cyphastrea (septothecal walls). Diploastrea has a diagnostic synapticulothecal wall and thick triangular teeth; Montastraea cavernosa is also distinct, possessing both “Pacific faviid” (abortive septa) and “Atlantic faviid” (paddle‐shaped teeth) attributes. The development of secondary axes is similar in traditional Atlantic faviids and mussids, supporting molecular results placing them in the same clade. Subclades of “Pacific faviids” reveal differences in wall structure and the arrangement and distinctiveness of centers of rapid accretion. J. Morphol. 272:66–88, 2011. © 2010 Wiley‐Liss, Inc.