Early signs of recovery of Acropora palmata in St. John,US Virgin Islands

Since the 1980s, diseases have caused significant declines in the population of the threatened Caribbean coral Acropora palmata. Yet it is largely unknown whether the population densities have recovered from these declines and whether there have been any recent shifts in size-frequency distributions toward large colonies. It is also unknown whether colony size influences the risk of disease infection, the most common stressor affecting this species. To address these unknowns, we examined A. palmata colonies at ten sites around St. John, US Virgin Islands, in 2004 and 2010. The prevalence of white-pox disease was highly variable among sites, ranging from 0 to 53 %, and this disease preferentially targeted large colonies. We found that colony density did not significantly change over the 6-year period, although six out of ten sites showed higher densities through time. The size-frequency distributions of coral colonies at all sites were positively skewed in both 2004 and 2010, however, most sites showed a temporal shift toward more large-sized colonies. This increase in large-sized colonies occurred despite the presence of white-pox disease, a severe bleaching event, and several storms. This study provides evidence of slow recovery of the A. palmata population around St. John despite the persistence of several stressors.     

A dynamic size-structured population model: does disturbance control size structure of a population of the massive coral Gardineroseris planulata in the Eastern Pacific?

Using a long-term data set of Gardineroseris planulata (Dana) on Uva Island reef, Panamá, we developed a simulation model that relates size-specific schedules of growth and partial mortality to predation by Acanthaster planci (Linnaeus) and El Niño–Southern Oscillation (ENSO)-related elevation of water temperature. We compared the model predictions to field observations for both the subpopulation of colonies that was used for model development and for the entire population. No statistically significant differences in the size-frequency distributions of the real and modeled coral populations were found for the subpopulation during any of 9 yr or for the entire population during 4 yr. These results suggested that the model relationships were reflecting field conditions. Longer-term (100 yr) simulations were conducted to assess the relative importance of predation and ENSO-related colony losses in determining the size structure of the coral population. Predation by A. planci was of overwhelming importance due to both stronger effects of predation (larger transitions) and the frequency of predation (yearly) compared to ENSO (episodically). Even the least-frequent predation scenario skewed the distribution toward smaller colonies, while simulations where populations were subjected to frequent ENSO events (≥3 yr) still maintained colonies in the largest size class. The model results suggested that sea star predators may not have been in the present abundance on this reef prior to the last 30 yr; with predators present, the model predicts that the distribution would be skewed toward smaller colonies.     

Size-dependent reproduction in the Caribbean gorgonian Pseudoplexaura porosa

Reproductive success among clonal taxa is often portrayed as a simple function of clone size, but reproduction in Pseudoplexaura porosa (Houtuyn), a common Caribbean gorgonian, reflects a more complex relationship between size and gamete production. Tagged colonies were sampled at two reefs in the San Blas Islands, Panama during the 1995 spawning season to determine the colony size at first reproduction, size-dependent polyp fecundities, and whole colony gonad production, and to follow the gametogenic cycle. Additional data were obtained during non-spawning months in 1984 and 1985. Of 120 colonies ranging in height from 20 to 250 cm, only colonies taller than 50 cm were reproductive. An average of 4.33 oocytes polyp⁻¹, 560 to 800 μm in diameter, were released during monthly synchronous spawning events from June to September. Oocytes released each month developed from a group of intermediate size oocytes present at the end of the preceding month. Almost all oocytes >560 μm were released during each spawning event. Spermatogenesis was a shorter process than oogenesis. During each spawning month, mature spermaries developed from a group of <125 μm spermaries. Virtually all spermaries >190 μm were released each month, and most of the spermary volume in mature male polyps was generated anew each month of spawning. Among reproductive colonies, polyp fecundity increased with colony height from 50 to 200 cm. Per capita fecundity of >200 cm tall female colonies was lower than for 150 to 200 cm colonies, but whole colony fecundity of large colonies was greater due to the exponential increase in the number of polyps that occurs with increasing colony height. In male colonies, both polyp and colony reproductive output increased with colony height. The large amounts of sperm produced by large male colonies probably contributes to the high rates of in situ fertilization observed in P. porosa. Differences in fecundity as a function of colony size and sex suggest differences in some combination of the cost of reproduction and/or allocation of resources to reproduction. Received: 9 September 1998 / Accepted: 7 June 1999     

Shifts in coral community structures following cyclone and red tide disturbances within the Gulf of Oman (United Arab Emirates)

This study documents the effects of two consecutive disturbances on coral community structures in the Gulf of Oman (United Arab Emirates); Cyclone Gonu in June 2007 and the Cochlodinium polykrikoides harmful algal bloom (HAB) that persisted from August 2008 until May 2009. Coral cover, colony densities, size class frequency distributions, and geometric growth rates derived from size class transition probability matrices were used to assess the post-Gonu and post-HAB recovery trajectories at four sites. The net effects of these disturbances were fourfold: (i) storm damage caused >50% losses of live branching and tabular coral cover by fragmentation and dislodgment of pocilloporid and acroporid colonies; (ii) Pocillopora damicornis colonies that survived the cyclone experienced mass mortality during the first 3?months of the HAB, resulting in localized extirpation of this species; (iii) variable Acropora mortality during the HAB indicated individual colony, rather than taxa-wide, susceptibility; and (iv) massive colony coral taxa were resistant to both disturbances.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Almost royal: incomplete suppression of host worker ovarian development by a social parasite wasp

Insect social parasites, like other parasites, may benefit from inhibiting their host from reproducing (complete or partial parasitic castration) because they can then exploit more of the host’s resources for their own reproduction. In particular, social parasites that kill or expel the host queen need to prevent host workers from reproducing; this is a common worker response to the absence of their queen. Indeed, host workers would benefit from detecting the presence of the parasite and investing in direct and indirect fitness. Studying whether and how social parasites control host worker reproduction can provide information about the degree of integration of the parasite in the host colony and help identify factors regulating workers’ reproductive decisions in social insects. We investigated whether the paper wasp social parasite, Polistes sulcifer, suppresses Polistes dominula (host) worker reproduction as efficiently as the dominant host female does in queen-right colonies by comparing worker reproductive efforts in parasitized and non-parasitized (control) colonies. Our results show that 6 weeks after usurpation of their colony by the social parasite, parasitized workers (1) had more developed ovaries than control workers and (2) laid more eggs as soon as the opportunity arose. This reproductive readiness of parasitized workers was not apparent 2 weeks after colony usurpation. This suggests that P. dominula workers have evolved means to react to social parasitism, as occurs in some ants, and that the parasite has only limited control over host reproduction.     

Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. I: Population structure of snails and corals

Despite their potential importance in structuring reef communities, invertebrate corallivores and their population structures are poorly understood. We found distinct differences in the population structures (length-frequency distribution and sex ratio) of the corallivorous gastropod Coralliophila abbreviata residing on two coral-host taxa, Montastraea spp. and Acropora palmata, in the Florida Keys. In each of two survey years, around 50% of the Montastraea spp. colonies were infested, with a mean snail density of eight snails per infested colony (range 1–45), while around 20% of A. palmata colonies harbored three snails per infested colony (range 1–23). Variation in patterns of snail occurrence was also observed within a host taxon. A. palmata occurred in low- and high-density stands (0.4 and 1 colony m^–2, respectively, at the initial survey) at different sites. Hurricane Georges struck the area in September 1998. When resurveyed in 1999, density of colonies in low-density stands had decreased by 75% to 0.1 colonies m^–2. This decrease was accompanied by a doubling in the proportion of colonies infested with snails (from 19% to 46%) and an increase in snail density per infested colony (from 3.7±3.3 SD to 5.4±4.6 SD) as snails apparently concentrated on surviving A. palmata. In contrast, sites with high density A. palmata stands (thickets) retained colony densities of about ~1 colony m^–2 among years, while snail infestation increased only from 9% to 14% of colonies surveyed and snail density essentially remained unchanged (from 2.7±1.8 to 2.9±1.9 snails per infested colony). Snails collected from Montastraea spp. were shorter than those from A. palmata in low-density stands and were longest on A. palmata in thickets. On both host taxa, female snails were longer than males. The sex ratio of snails on Montastraea spp. hosts was even (1:1), while that of snails on A. palmata was skewed (70% males). Factors that could explain observed differences in size structure and sex ratio between Coralliophila populations on the two coral host taxa include: differential susceptibility to predators, influence of host tissue nutritional quality and/or secondary metabolite content, and genetic differences (cryptic species). The host-specific characteristics of C. abbreviata populations imply that the impact of gastropods on reef communities will vary with the coral species composition.     

Long-term changes in coral colony size distributions on Kenyan reefs under different management regimes and across the 1998 bleaching event

Colony size is an important life-history characteristic of corals and changes in colony size will have significant effects on coral populations. This study summarizes ∼21,000 haphazard colony size measurements of 26 common coral taxa (mostly coral genera) collected annually between 1992 and 2006 in seven Kenyan reef lagoons. There was a major coral bleaching and mortality event in early 1998 and all seven reefs were affected. The seven locations include two long-protected Marine National Parks (Malindi and Watamu), one relatively recently established park (Mombasa), and four unprotected locations (Vipingo, Kanamai, Ras Iwatine, and Diani). They span about 150 km and represent three distinct fishery management regimes: old protected (OP), newly protected (NP), and unprotected (UP). Seventeen taxa had statistically significant different sizes for comparisons of the management regimes, with only one genus, Pavona, having larger sizes in the unprotected reefs. The size of eight coral genera showed a significant time and management interaction, and size frequency differences that existed in management areas prior to 1998 were further increased after the bleaching event. Time alone was a significant factor for eleven genera, and in all cases colonies were smaller after 1998. For most taxa, colony size distributions were significantly skewed and had right-tailed distributions. After 1998, the right-tailed distributions of Acropora, Hydnophora, and Montipora were significantly reduced. Most taxa had peaky distributions and only Acropora experienced a statistically significant change from peaky to flat. The mean sizes of taxa were not related to their mortality across 1998, which indicates that the size effect was within rather than between taxa. Astreopora and Platygyra were well-sampled taxa that did not show an effect of management, but had reduced median sizes across 1998. Consequently, no taxa were tolerant of both fishing and bleaching disturbances and the combined effect was to reduce the size of all corals.     

When fragmented coral spawn? Effect of size and timing on survivorship and fecundity of fragmentation in <Emphasis Type="Italic">Acropora formosa</Emphasis>

In order to determine competency of sexual reproduction and survival rate after fragmentation, the branching coral Acropora formosa was fragmented and fragments in three different sizes (ca. 5, 10 and 20 cm long) and three different stages of gametogenesis were transplanted on coral pavement. Their oocyte development and fecundity, as well as spawning were monitored for a 3-year period. The oocyte development was affected by both fragment size and by the developmental stage of oocytes when fragmented. In small fragments, the oocytes were resorbed while in large fragments they continued development. Oocytes in the early vitellogenic stage at the time of fragmentation were resorbed, whereas those in the late stage continued developing. Smaller fragments showed a lower survival rate and histological observations of their gonads revealed resorption of oocytes, suggesting that there was a trade-off of energy between reproduction and survival. Transplanted fragments often spawned one month earlier than the donor colonies in the first year, but spawning occurred in the same month as the donors or did not occur at all in the second year and none spawned in the third year. The risk of colony death may cause the fragments to re-allocate energy for sexual reproduction.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and the Galápagos Islands (Ecuador). VI. Agariciidae, <Emphasis Type="Italic">Pavona clavus</Emphasis>

The reproductive ecology of the zooxanthellate reef coral Pavona clavus was investigated at several sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador) over the period 1985–2009. Pavona clavus displayed stable gonochorism as only five hermaphrodites were found in 590 samples. At four of five locations, sex ratios were skewed toward female dominance; however, at Saboga Island (Panamá) male colonies predominated. In Panamá, sexual maturity was observed in an estimated eight-year-old female colony, and several colonies of 10–20 years of age demonstrated gametogenesis. Sexual activity was observed at all study sites, but gamete development occurred in only 14–31% of colonies sampled sporadically. Seasonality of gametogenic activity occurred predominantly during the warm/wet season, June to August, at mainland sites (Caño Island, Costa Rica, and Gulfs of Chiriquí and Panamá, Panamá). This pattern was repeated in the Galápagos Islands, but mainly from March to May when seasonally high sea temperatures and rainfall prevailed there. Histological sampling and field observations indicated that spawning was centered around the full moon, most frequently on lunar day 17, and near sunset (1,800 h). Mean fecundity (mature ova cm^?2 live tissue) estimates were significantly different for two sites and ranged from ~1,780 (Saboga Island, Gulf of Panamá, seasonally upwelling) to ~4,280 (Uva Is, Gulf of Chiriquí, non-upwelling). Assuming three annual spawning events colony^?1 (August, September, October), extrapolation of minimum and maximum fecundities yield 5,340 and 12,840 ova cm^?2 year^?1. Seasonal, lunar, and diel spawning patterns in nine zooxanthellate species at Uva Island indicate asynchronous coral community spawning.     

Polymorphism in soft coral larvae revealed by amplified fragment-length polymorphism (AFLP) markers

The dioecious Red Sea soft coral Parerythropodium fulvum fulvum breeds its nonsymbiotic planula larvae on the surface of female colonies for less than a week. After completing their development, larvae crawl and settle near maternal colonies. Here we study the genetic polymorphism of developing larvae by the use of amplified fragment-length polymorphism markers. Four reproductive colonies from shallow water populations (two from a dense population and two from a less densely populated area 100 m away) were chosen, and ten larvae were randomly collected from each colony. DNA was analyzed by using three different primer combinations producing 61, 63, 63 polymorphic markers, respectively. All larvae exhibited different banding patterns from one another, illustrating the prominent role of sexual reproduction for the production of larvae. Nei's mean genetic distances for all 12 possible pair-wise combinations for larval origins revealed, in most cases, that sister larvae are genetically closer than larvae from different colonies and that larvae may be grouped into three statistical clusters in accordance with colony origin and population studied. The usefulness of molecular methodologies in coral population genetics is discussed. Received: 26 March 1999 / Accepted: 19 October 1999     

The effects of temperature on the growth of juvenile scleractinian corals

Tropical reef corals are well known for their sensitivity to rising temperature, yet surprisingly little is known of the mechanisms through which temperature acts on intact coral colonies. One such mechanism recently has been suggested by the association between the growth of juvenile corals and seawater temperature in the Caribbean, which suggests that temperature causes a transition between isometric and allometric growth scaling in warmer versus cooler years, respectively (Edmunds in Proc R Soc B 273:2275–2281, 2006). Here, this correlative association is tested experimentally for a cause-and-effect relationship. During April and May 2006, juvenile colonies (8–35 mm diameter) of massive Porites spp. from Moorea, French Polynesia, were incubated at warm (27.8°C) and cool (25.7°C) temperatures for 15 days, and their response assessed through the scaling of growth (change in weight) with colony size. The results reveal that the scaling of colony-specific growth (mg colony⁻¹ day⁻¹) was unaffected by temperature, although growth absolutely was greater at the cool compared to the warm temperature, regardless of colony size. This outcome was caused by contrasting scaling relationships for area-specific growth (mg cm⁻² day⁻¹) that were negatively allometric under warm conditions, but independent of size under cool conditions. In April 2007, a 22 days field experiment confirmed that the scaling of area-specific growth in juvenile Porites spp. is negatively allometric at a warm temperature of 29.5°C. Based on strong allometry for tissue thickness, biomass, and Symbiodinium density in freshly collected Porites spp., it is hypothesized that the temperature-dependency of growth scaling in these small corals is mediated by the interaction of temperature with biomass.     

Seasonal cycle of sexual reproduction of the Mediterranean soft coral Alcyonium acaule (Anthozoa, Octocorallia)

Alcyonium acaule (Cnidaria, Octocorallia) is a common, hard-bottom soft coral in the northwestern Mediterranean Sea. This study describes sexual reproduction and the gamete development cycle of this soft coral. A population at 15–18 m depth in the Marine Protected Area of the Medes Islands (42º02′55″ N, 3º13′30″ E) was sampled from July 1994–August 1995. A. acaule is gonochoristic and a surface brooder, spawning once a year in early summer. The mean diameter of ripe spermatic sacs was 400 ± 91 (SD) μm, and the mean diameter of mature oocytes was 473 ± 37 (SD) μm. There were 30 spermatic sacs polyp^?1 in males and 14 oocytes polyp^?1 in females. Different phases of gametogenesis in female and male colonies were examined separately with respect to seasonal changes in bottom temperature and solar irradiance. The data suggest that the relatively constant temperatures in January–April are probably not related to oocyte maturation, but that rising temperatures in May could affect sperm maturation. Rapidly increasing solar irradiance in March may be the trigger for vitellogenesis and oocyte maturation, although the mechanism for this in anthozoans is not understood.     

Alternate coral–bryozoan competitive superiority during coral bleaching

Bleaching of corals results from the loss of their symbiotic algae (zooxanthellae) and/or pigments. The supply of photoassimilates provided by the zooxanthellae to the coral declines during bleaching and reduces the ability to activate energy-costly processes such as maintenance, growth and reproduction. In the present study we compared the competitive outcomes, expressed as overgrowth and changes in colony sizes of Oculina patagonica (an encrusting Mediterranean stony coral) and the bryozoan Watersipora sp., growing in contact with each other, during and between bleaching events. Year-round observations of tagged colonies showed alternating competitive outcomes: O. patagonica wins over Watersipora sp. between bleaching events, but loses during bleaching events. Using the ¹⁴C-point-labeling technique on coral tissue, we examined intra-colonial translocation of photosynthetic products from the point-tissue labeling towards interaction zones. In non-bleached O. patagonica, competition resulted in preferentially oriented translocation of ¹⁴C products to the interaction zone located up to 8 cm away from the tissue-labeling site. Sites opposite the interaction zone received significantly less labeled photoassimilates compared to the interaction zone. In bleached colonies (40-85% bleached surface area), such translocation did not occur, probably explaining the failure to compete with the encrusting neighbor Watersipora sp. Our findings demonstrate the importance of colonial integration and resource orientation for the competitive superiority of O. patagonica.     

Porites and the Phoenix effect: unprecedented recovery after a mass coral bleaching event at Rangiroa Atoll,French Polynesia

The 1997/1998 El Niño Southern Oscillation (ENSO) was the most severe coral bleaching event in recent history, resulting in the loss of 16 % of the world’s coral reefs. Mortality was particularly severe in French Polynesia, where unprecedented mortality of massive Porites was observed in lagoonal sites of Rangiroa Atoll. To assess the recovery of massive Porites 15 years later, we resurveyed the size structure and extent of partial mortality of massive Porites at Tivaru (Rangiroa). Surveys revealed an abundance of massive Porites colonies rising from the shallow lagoonal floor. Colony width averaged 2.65 m, reaching a maximum of 7.1 m (estimated age of ~391 ± 107 years old). The relative cover of recently dead skeleton within quadrats declined from 42.8 % in 1998 to zero in 2013, yet the relative cover of old dead skeleton increased only marginally from 22.1 % in 1998 to 26.1 % in 2013. At a colony level, the proportion of Porites dominated by living tissue increased from 34.9 % in 1998 to 73.9 % in 2013, indicating rapid recovery of recent dead skeleton to living tissue rather than transitioning to old dead skeleton. Such rapid post-bleaching recovery is unprecedented in massive Porites and resulted from remarkable self-regeneration termed the ‘Phoenix effect’, whereby remnant cryptic patches of tissue that survived the 1997/1998 ENSO event regenerated and rapidly overgrew adjacent dead skeleton. Contrary to our earlier predictions, not only are large massive Porites relatively resistant to stress, they appear to have a remarkable capacity for recovery even after severe partial mortality.     

Coral architecture affects the habitat choice and form of associated gobiid fishes

Gobiid fishes of the genus Gobiodon live in strong association with certain reef-building corals that vary considerably in size and architecture. These fishes hence are excellent model systems for studying evolutionary adaption to specific microhabitats. Using a sample of Gobiodon histrio and G. rivulatus and their most important host corals (Acropora digitifera and A. gemmifera) from the northern Red Sea, we assess (1) how corals that are occupied by gobies differ in their architecture from colonies that are not occupied and (2) how fish body shape is associated with the architecture of their host coral. Fish body shape was assessed by geometric morphometric techniques. Coral measurements included colony size, branch length (BL), and interbranch as well as branch tip distance of adjacent branches, for which we applied a new and non-destructive measurement technique based on casts of two-component epoxy resin. The most important factor influencing the occupation of corals was a BL of more than 5 cm. The distance between coral branches was clearly related to the width of the fishes and hence constrained overall fish size. G. histrio and G. rivulatus differ in adult body shape as well in their allometric development of lateral body compression, resulting in different maximum body sizes attainable in the restricted interbranch space of corals. The strong dependence of coral-associated fishes on large coral colonies with specific architectures increases the extinction risk of fishes within deteriorating coral reefs.     

Coral mortality following extreme low tides and high solar radiation

Extreme tidal events are one of the most predictable natural disturbances in marine benthic habitats and are important determinants of zonation patterns in intertidal benthic communities. On coral reefs, spring low tides are recurrent disturbances, but are rarely reported to cause mass mortality. However, in years when extremely low tides coincide with high noon irradiances, they have the potential to cause widespread damage. Here, we report on such an event on a fringing coral reef in the central Great Barrier Reef (Australia) in September 2005. Visual surveys of colony mortality and bleaching status of more than 13,000 corals at 14 reef sites indicated that most coral taxa at wave-protected sites were severely affected by the event. Between 40 and 75% of colonies in the major coral taxa (Acropora, Porites, Faviidae, Mussidae and Pocilloporidae) were either bleached or suffered partial mortality. In contrast, corals at wave-exposed sites were largely unaffected (<1% of the corals were bleached), as periodic washing by waves prevented desiccation. Surveys along a 1–9 m depth gradient indicated that high coral mortality was confined to the tidal zone. However, 20–30% of faviid colonies were bleached throughout the depth range, suggesting that the increase in benthic irradiances during extreme low tides caused light stress in deeper water. Analyses of an 8-year dataset of tidal records for the area indicated that the combination of extended periods of aerial exposure and high irradiances occurs during May–September in most years, but that the event in September 2005 was the most severe. We argue that extreme low-tide, high-irradiance events are important structuring forces of intertidal coral reef communities, and can be as damaging as thermal stress events. Importantly, they occur at a time of year when risks from thermal stress, cyclones and monsoon-associated river run-off are minimal.     

Colony life history and demography of a swarm-founding social wasp

Colonies of social insects are sometimes viewed as superorganisms. The birth, reproduction, and death of colonies can be studied with demographic measures analogous to those normally applied to individuals, but two additional questions arise. First, how do adaptive colony demographies arise from individual behaviors? Second, since these superorganisms are made up of genetically distinct individuals, do conflicts within the colony sometimes modify and upset optima for colonies? The interplay between individual and superindividual or colony interests appears to be particularly complex in neotropical, swarm-founding, epiponine wasps such as Parachartergus colobopterus. In a long-term study of this species, we censused 286 nests to study colony-level reproduction and survivorship and evaluated individual-level factors by assessing genetic relatedness and queen production. Colony survivorship followed a negative exponential curve very closely, indicating type II survivorship. This pattern is defined by constant mortality across ages and is more characteristic of birds and other vertebrates than of insects. Individual colonies are long-lived, lasting an average of 347 days, with a maximum of over 4.5 years. The low and constant levels of colony mortality arise in part from colony initiation by swarming, nesting on protected substrates, and an unusual expandable nest structure. The ability to requeen rapidly was also important; relatedness data suggest that colonies requeen on average once every 9–12 months. We studied whether colony optima with respect to the timing of reproduction could be upset by individual worker interests. In this species, colonies are normally polygynous but new queens are produced only after a colony reaches the monogynous state, a result which is in accord with the genetic interests of workers. Therefore colony worker interests might drive colonies to reproduce whenever queen number happens to cycled down to one rather than at the season that is otherwise optimal. However, we found reproduction to be heavily concentrated in the rainy season. The number of new colonies peaked in this season as did the percentages of males and queens. Relatedness among workers reached a seasonal low of 0.21–0.27, reflecting the higher numbers of laying queens. This seasonality was achieved in part by a modest degree of synchrony in the queen reduction cycle. Worker relatedness reached peaks of around 0.4 in the dry season, reflecting a decrease to a harmonic mean queen number of about 2.5. Thus, a significant number of colonies must be approaching monogyny entering the rainy season. Coupled with polygynous colonies rearing only males (split sex ratios), this makes it possible for a colony cycle driven by selfish worker interests to be consistent with concentrating colony reproduction during a favorable season.     

Morphology and growth of the deep-water gorgonians <Emphasis Type="Italic">Primnoa resedaeformis</Emphasis> and <Emphasis Type="Italic">Paragorgia arborea</Emphasis>

The morphology of the gorgonian corals Paragorgia arborea and Primnoa resedaeformis was studied from video records and colonies collected from different locations in Atlantic Canada, at depths between 200 and 600 m. Growth was studied by relating colony height to age (number of growth rings) in P. resedaeformis, and from a photographic time-series of a P. arborea colony in a Norwegian fjord. The highest P. resedaeformis and P. arborea colonies were 86 and 180 cm, respectively. The height of P. arborea seemed to be restricted by the size of the boulder it was attached to. When the coral exceeds a critical height (approximately twice the stone size), the drag of strong currents can turn the coral and its substrate over. No limiting factors for the height of P. resedaeformis colonies were identified. P. arborea occurred in three colour varieties: red, salmon red, and white. The red and white contributed 41% to the population each, while 18% of the colonies were salmon red. On average the salmon red P. arborea were taller than the red and white. P. arborea colonies >50 cm were mainly concave fan shaped. The orientation of these indicated a near-bottom current pattern similar to what is known from previous current measurements in the area. P. resedaeformis occurred mainly on the up-current side of boulders, but its bushy morphology does not indicate influence by unidirectional current to the same degree as P. arborea. The different height, morphology, and position on boulders of the two species indicate that they utilize different food sources. P. resedaeformis seems to be adapted to a near-bottom environment with turbulent currents, whereas P. arborea utilize uni- or bidirectional currents higher above bottom by developing planar colonies perpendicular to the current. The oldest P. resedaeformis colony was 61 years. The relationship between height and age indicated an average growth of 1.7 cm year^–1 for P. resedaeformis. X-ray images of skeletal sections of P. arborea showed clear growth bands with a maximum band width of 1.3 cm. It is not clear what time scales these bands represent, and they could therefore not be used for indicating age. The limited previously reported data on age and growth of P. arborea indicate an average growth rate of 1 cm year^–1. This gives an age of about 180 years for the largest colony in this study. The time-series photographs, however, indicated a much higher growth rate (varying between 2 and 6 cm year^–1 within the colony), which may be more representative for colonies of an intermediate size.Communicated by R.J. Thompson, St. Johns     

Assessing the effectiveness of marine reserves on unsustainably harvested long-lived sessile invertebrates

Although the rapid recovery of fishes after establishment of a marine reserve is well known, much less is known about the response of long-lived, sessile, benthic organisms to establishment of such reserves. Since antiquity, Mediterranean red coral (Corallium rubrum) has been harvested intensively for use in jewelry, and its distribution is currently smaller than its historical size throughout the Mediterranean Sea. To assess whether establishment of marine reserves is associated with a change in the size and number of red coral colonies that historically were not harvested sustainably, we analyzed temporal changes in mean colony diameter and density from 1992 to 2005 within red coral populations at different study sites in the Medes Islands Marine Reserve (established in 1992) and in adjacent unprotected areas. Moreover, we compared colony size in the Medes Islands Marine Reserve, where recreational diving is allowed and poaching has been observed after reserve establishment, with colony size in three other marine protected areas (Banyuls, Carry-le-Rouet, and Scandola) with the enforced prohibition of fishing and diving. At the end of the study, the size of red coral colonies at all sampling sites in the Medes Islands was significantly smaller than predicted by growth models and smaller than those in marine protected areas without fishing and diving. The annual number of recreational dives and the percent change in the basal diameter of red coral colonies were negatively correlated, which suggests that abrasion by divers may increase the mortality rates of the largest red coral colonies within this reserve . Our study is the first quantitative assessment of a poaching event, which was detected during our monitoring in 2002, inside the marine reserve. Poaching was associated with a loss of approximately 60% of the biomass of red coral colonies.