Multi-year, seasonal genotypic surveys of coral-algal symbioses reveal prevalent stability or post-bleaching reversion

This report documents the extent to which coral colonies show fluctuations in their associations with different endosymbiotic dinoflagellates. The genetic identity of Symbiodinium from six coral species [Acropora palmata (Lamarck), A. cervicornis (Lamarck), Siderastrea siderea (Ellis and Solander), Montastrea faveolata (Ellis and Solander), M. annularis (Ellis and Solander), and M. franksi (Gregory)] was examined seasonally over five years (1998 and 2000–2004) in the Bahamas and Florida Keys at shallow (1 to 4 m) fore-reef/patch reef sites and at deeper fore-reef (12–15 m) locations. Symbionts were identified genetically using denaturing gradient gel electrophoresis (DGGE) fingerprinting of the internal transcribed spacer region 2 (ITS2) of ribosomal RNA gene loci. Repetitive sampling from most labeled colonies from the Bahamas and the Florida Keys showed little to no change in their dominant symbiont. In contrast, certain colonies of M. annularis and M. franksi from the Florida Keys exhibited shifts in their associations attributed to recovery from the stresses of the 1997–1998 El Niño southern oscillation (ENSO) event. Over several years, a putatively stress-tolerant clade D type of Symbiodinium was progressively replaced in these colonies by symbionts typically found in M. annularis and M. franksi in Florida and at other Caribbean locations. Greater environmental fluctuations in Florida may explain the observed changes among some of the symbioses. Furthermore, symbiotic associations were more heterogeneous at shallow sites, relative to deep sites. The exposure to greater environmental variability near the surface may explain the higher symbiont diversity found within and between host colonies.     

Experimental evidence that workers recognize reproductives through cuticular hydrocarbons in the ant Odontomachus brunneus

Eusociality is characterized by a reproductive division of labor, wherein workers respond to the presence of reproductive individuals by refraining from reproduction themselves and restricting the reproductive efforts of others. Our understanding of how eusociality is maintained therefore depends on characterizing the mechanism by which workers detect the presence of a reproductive. Variations in cuticular hydrocarbons correspond to changes in reproductive ability in ants, and experimental studies are beginning to reveal the function of hydrocarbons as signals. In this study, we compare the cuticular hydrocarbon profiles of dominant and reproductive workers and queens of the ant Odontomachus brunneus with profiles of non-reproductive workers. Using split/reunification tests we document the existence of worker policing in both queenless and queenright colonies; supernumerary reproductives were treated aggressively by nestmates. Finally, we induce aggression and replicate queen-like submissive nestmate responses by supplementing the hydrocarbon profile of workers with (Z)-9-nonacosene, a compound that was significantly more abundant on the cuticles of reproductives. In three bioassays, we compare this manipulation to various control manipulations of the hydrocarbon profile and demonstrate that workers gauge the reproductive activity of nestmates through changes in their cuticular hydrocarbon profiles.     

Condition-dependent response to changes in pollen stores by honey bee (Apis mellifera) colonies with different parasitic loads

The impact of a parasitic infestation may be influenced by nutritional state, in both individuals and colonies. This study examined the interaction between pollen storage and the effects of an infestation by the mite, Varroa jacobsoni Oudemans, in colonies of the honey bee, Apis mellifera L. We manipulated the pollen storage and mite infestation levels of colonies, and measured pollen foraging and brood rearing. Increased pollen stores decreased both the number of pollen foragers and pollen load size, while initially at least foragers from colonies with moderate infestations carried smaller pollen loads than those from lightly infested colonies. Over the course of the experiment, all colonies significantly increased pollen-foraging rates and pollen consumption, which was presumably a seasonal effect. Lightly infested colonies exhibited a larger increase in pollen forager number than moderately infested colonies, suggesting that more intense mite infestations compromised forager recruitment. Brood production was not affected by the addition of pollen, but moderately infested colonies were rearing significantly less brood by the end of the experiment than lightly infested colonies. Furthermore, the efficiency with which colonies converted pollen to brood decreased as the pollen storage level decreased and the infestation level increased. The results of this study may indicate that honey bee colonies adaptively alter brood-production efficiency in response to parasitic infestations and seasonal changes. Received: 3 May 1999 / Received in revised form: 14 September 1999 / Accepted: 25 September 1999     

Oligogyny by unrelated queens in the carpenter ant, Camponotus ligniperdus

Multilocus DNA fingerprinting and microsatellite analysis were used to determine the number of queens and their mating frequencies in colonies of the carpenter ant, Camponotus ligniperdus (Hymenoptera: Formicidae). Only 1 of 61 analyzed queens was found to be double-mated and the population-wide effective mating frequency was therefore 1.02. In the studied population, 8 of 21 mature field colonies (38%) contained worker, male, or virgin queen genotypes which were not compatible with presumed monogyny and therefore suggested oligogyny, i.e., the cooccurrence of several mutually intolerant queens within one colony. Estimated queen numbers in oligogynous colonies ranged between two and five. According to the results of the genetic analysis, most of the queens coexisting in oligogynous colonies were not closely related. Pleometrosis is very rare and queenless colonies adopt mated queens both in the laboratory and field. Therefore, the most plausible explanation for the origin of oligogynous colonies in C. ligniperdus is the adoption of unrelated queens by orphaned mature colonies. The coexistence of unrelated, but mutually intolerant queens in C. ligniperdus colonies demonstrates that oligogyny should be considered as a phenomenon distinct from polygyny. Received: 18 December 1997 / Accepted after revision: 20 June 1998     

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.     

Effects of prey quantity on predatory wasps (<Emphasis Type="Italic">Polistes dominulus</Emphasis>) when patch quality differs

To determine the effects of prey quantity on central-place foraging of predatory wasps ( Polistes dominulus), prey of varying quality were distributed in patches. A field experiment was conducted, which controlled the amount and quality of prey available. 'Low-fed' colonies were provided with one-third the quantity of prey as that of 'high-fed' colonies. Both were provided with a 1:1 ratio of palatable:unpalatable prey. Experienced wasps of the high-fed colonies never selected unpalatable prey as their first choice of the day, whereas those of the low-fed colonies selected unpalatable prey as their first choice about 1:4 times. In general, wasps from the high-fed colonies reduced palatable patches to zero prey before exploiting unpalatable patches. Foundresses of high-fed colonies captured disproportionately more palatable prey than those of low-fed nests, but there was no correlation between ratio of palatable to unpalatable prey taken and the number of offspring produced. Wasps from low-fed colonies attacked unpalatable prey sooner, but not without considerable effort to avoid use of those patches. Foundresses of low-fed colonies also spent a greater proportion of time overall in unsuccessful search, which may explain why only wasps from low-fed colonies foraged on cool days. High-fed colonies produced more cells and more and heavier offspring than low-fed colonies. But the productivity of the low-fed colonies was greater than that of the 'natural' colonies, which had to find their own prey in a field-woodland area. These results indicate that prey scarcity changes foraging behavior and affects prey choice. These changes may not totally alleviate negative effects of unpalatable prey on colony development and offspring production. The results of this study increase understanding of the central-place foraging behavior of paper wasps, which are important biocontrol agents in natural and agricultural settings.     

Racial mixing in South African honeybees: the effects of genotype mixing on reproductive traits of workers

To test the hypothesis that the honeybee hybrid zone in South Africa is a tension zone due to increased reproductive conflict in colonies that contain both Apis mellifera capensis and Apis mellifera scutellata worker genotypes, we constructed mixed subspecies and hybrid colonies via a combination of artificial and natural matings. We measured emergence weight, ovary activation, and the presence/absence of a spermatheca on workers of different genotypes. We show that the measured characteristics were all affected by genotype with some traits also affected by the social environment in which the worker was reared. Workers with both an A. m. capensis mother and father had the highest emergence weight. When workers had an A. m. capensis mother, paternity affected emergence weight with A. m. capensis fathers producing heavier workers. When the queen was A. m. scutellata, paternity had less effect on weight. Presence of spermatheca was highest in mixed colonies irrespective of maternity and colonies containing pure A. m. capensis workers only. Paternity had a significant effect on the presence of a spermatheca within mixed colonies, with workers that had an A. m. capensis father being more likely to possess a spermatheca. Rates of ovary activation were highest in colonies with an A. m. scutellata queen mated to drones of both genotypes, suggesting that mixed subspecies colonies likely suffer increased reproductive strife among workers. Our results provide support for the hypothesis that the South African honeybee hybrid zone is a tension zone arising from reduced fitness of genetically mixed colonies.     

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.     

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.     

The effects of partial mortality on the fecundity of three common Caribbean corals

The recent intensification of human disturbances in the Caribbean has increased the prevalence of partial mortality on coral colonies. Partial mortality can change colony size by directly shrinking colonies or by splitting colonies into fragments. A reduction in colony size can also adversely affect fecundity and fitness as internal resources shift away from reproduction toward colony maintenance. This study aimed to determine whether three Caribbean coral species, Siderastrea siderea, Montastraea faveolata, and Diploria strigosa, along the reef tract in Puerto Morelos, Mexico (20^o52′N, 86^o51′W), continued to dedicate resources to reproduction when colonies were fragmented to pre-maturation size. Contrary to expectations, eggs were found in colonies that were smaller than the maturation size and had been subjected to partial mortality. The continued dedication of resources toward reproduction, even in the smallest colonies, suggests that resource trade-offs away from reproduction are not as rigid as previously suggested in stressed corals.     

Social context influences cue-mediated recruitment in an invasive social wasp

Social insects often serve as model systems for communication and recruitment studies, and yet, it remains controversial whether social vespid wasps can reliably communicate resource information to nestmates. In this study, I present empirical evidence that foraging strategies depend on the initial assessment of resource size and potential competition by foraging yellowjackets. The context dependent foraging behavior of Vespula pensylvanica provides a potential explanation for the inconsistent reports of the existence of recruitment communication in vespid wasps. Furthermore, life history traits may influence yellowjacket foraging behavior; annual V. pensylvanica colonies, whose foragers routinely patrol near the nest, exhibited increased bait visitation in response to the return of successful foragers, whereas perennial colonies did not. These behavioral disparities provide insight into how foraging strategies and search patterns may shift with colony size and longevity. In experiments that investigate the effects of visual cues of conspecifics and bait dispersion, foraging decisions corresponded with expectations of yellowjackets integrating resource quantity and access into a perception of demand. When resource competition could be assessed as high, V. pensylvanica foragers quickly exploited the bait closest to their colony regardless of occupation by other wasps; however, foragers preferred visiting unoccupied baits in situations where competition could be perceived as low. Moreover, a meta-analysis revealed that context-dependent, cue-mediated recruitment was widespread in Vespidae, where such foraging behaviors changed with habitat and the potential for resource competition. Such plastic foraging strategies may contribute to the invasion success of some vespid wasps.     

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.     

Transitioning from unstable to stable colony growth in the desert leafcutter ant Acromyrmex versicolor

Like organisms, cohesive social groups such as insect colonies grow from a few individuals to large and complex integrated systems. Growth is driven by the interplay between intrinsic growth rates and environmental factors, particularly nutritional input. Ecologically inspired population growth models assume that this relationship remains constant until maturity, but more recent models suggest that it should be less stable at small colony sizes. To test this empirically, we monitored worker population growth and fungal development in the desert leafcutter ant, Acromyrmex versicolor, over the first 6 months of colony development. As a multitrophic, symbiotic system, leafcutter colonies must balance efforts to manage both fungus production and the growth of the ants consuming it. Both ants and fungus populations grew exponentially, but the shape of this relationship transitioned at a size threshold of 89?±?9 workers. Above this size, colony mortality plummeted and colonies shifted from hypometric to hypermetric growth, with a distinct stabilization of the relationship between the worker population and fungus. Our findings suggest that developing colonies undergo key changes in organizational structure and stability as they grow, with a resulting positive transition in efficiency and robustness.     

Spatial distribution patterns of the soft corals Alcyonium acaule and Alcyonium palmatum in coastal bottoms (Cap de Creus, northwestern Mediterranean Sea)

Current knowledge on the abundance and distribution patterns of different soft coral species is relatively limited when compared to other benthic suspension feeders such as gorgonians and hard coral species. To overcome this scarcity of information, the distribution patterns of the soft corals Alcyonium acaule and Alcyonium palmatum were investigated in northwestern Mediterranean benthic communities over a wide geographical (60 km of coastline) and bathymetrical (0–70 m depth) extent using a remotely operated vehicle. A. acaule was the most abundant species in the study area with highest recorded density of 18 col m^?2 found at depths of 35–45 m in areas that are directly exposed to strong near-bottom currents. Conversely, A. palmatum was only found as scattered solitary colonies at greater depths in soft bottoms, with maximum density of 2.4 col m^?2. Medium and large colonies of A. acaule were preferentially found on sloping and vertical rocky bottoms where they form dense patches. High-density patches of A. acaule were preferentially found on vertical rocky bottom, while isolated colonies were preferentially observed on coralligenous substrata as well as on flat soft and maërl substrates. A. acaule biomass distribution showed highest values between 40 and 45 m depth, and between 60 and 65 m depth. This suggests that deeper populations are formed by colonies that are bigger than the equivalent shallower ones. Although both species are almost genetically identical, ecologically they are very different. For this reason, conservation plans should consider the differential ecological traits shown by these two soft coral species.     

Context-dependent acceptance of non-nestmates in a primitively eusocial insect

The benefits of cooperation are essential in driving group formation. However, an individual can gain significant benefits by acting selfishly at a substantial cost to others in the group. Thus, group members must find a balance between accepting and rejecting potential new members. Here, I explore the factors that mediate acceptance of non-related individuals during the period of group establishment in the primitively eusocial wasp Mischocyttarus mexicanus. In this species, group composition changes during establishment, with non-related females (non-nestmates) sometimes accepted into a foreign colony. By experimentally introducing non-nestmates to newly established colonies, I test the hypothesis that acceptance threshold of nestmates towards non-nestmates shifts depending on the ecological context, as predicted by the Optimal Acceptance Threshold Model. I explored how non-nestmate age (young vs. old), stage of colony establishment (early vs. late), initial behavior of the non-nestmates (non-aggressive vs. aggressive), and the behavioral response by nestmates (non-aggressive vs. aggressive) affected the rates of acceptance. My results show an effect of both non-nestmate age and stage of colony development on non-nestmate acceptance. Young non-nestmates were more frequently accepted in early than in late colonies. Late colonies more frequently rejected both young and old non-nestmates, suggesting a cost of accepting potential usurpers into late colonies. Surprisingly, non-nestmate aggressive behavior did not have a direct effect on their acceptance, but it triggered an aggressive response from nestmates. These findings reveal a shift in the acceptance threshold, suggesting an effect of the social context and the specific needs of a colony on non-nestmate acceptance.     

Seasonality in reproduction of the deep-water pennatulacean coral Anthoptilum grandiflorum

The deep-sea pennatulacean coral Anthoptilum grandiflorum exhibits a cosmopolitan distribution and was recently determined to serve as habitat for other invertebrates and fish larvae in the northwest Atlantic. Colonies collected at bathyal depths between 2006 and 2010 in eastern Canada were analysed to determine their fecundity and characterize spatial and temporal trends in their reproductive cycle. Anthoptilum grandiflorum is a gonochoric broadcast spawner with a sex ratio that does not differ significantly from equality (although one hermaphrodite colony was observed). In male colonies, all the spermatocysts synthesized become mature over the annual cycle, while only ~21 % of the initial production of oocytes reaches maturity in female colonies. Female potential fecundity based on mature oocytes just before spawning was on average 13 oocytes polyp^?1; male potential fecundity was 48 spermatocysts polyp^?1. The spawning period of A. grandiflorum differs between geographic regions, from April (in southern Newfoundland) to July (in Labrador), closely following regional spring phytoplankton blooms after accounting for the deposition of planktic detritus. Release of oocytes by a live colony held in the laboratory was recorded in April 2011, coincident with field data for similar latitudes. Seawater temperatures at the time of spawning were around 3.6–4.8 °C in all regions and in the laboratory. Early stages of gametogenesis were detected in colonies collected shortly after the spawning season, and early and late growth stages occurred successively until December. Mature colonies were observed between April and July (depending on latitude). The diameter of mature oocytes (~1,100 μm maximum diameter) is consistent with lecithotrophic larval development.     

How is activity distributed among and within tasks in Temnothorax ants?

How social insect colonies behave results from the actions of their workers. Individual variation among workers in their response to various tasks is necessary for the division of labor within colonies. A worker may be active in only a subset of tasks (specialist), perform all tasks (elite), or exhibit no particular pattern of task activity (idiosyncratic). Here we examine how worker activity is distributed among and within tasks in ants of the genus Temnothorax. We found that workers exhibited elitism within a situation, i.e., in particular sets of tasks, such as those associated with emigrations, nest building, or foraging. However, there was weak specialization for working in a particular situation. A few workers exhibited elitism across all situations, i.e., high performance in all tasks in all situations. Within any particular task, the distribution of activity among workers was skewed, with few ants performing most of the work and most ants performing very little of the work. We further found that workers persisted in their task preference over days, with the same individuals performing most of the work day after day. Interestingly, colonies were robust to the removal of these highly active workers; they were replaced by other individuals that were previously less active. This replacement was not short-lived; when the removed individuals were returned to the colony, not all of them resumed their prior high activity levels, and not all the workers that replaced them reduced their activity. Thus, even though some workers specialize in tasks within a particular situation and are persistent in performing them, task allocation in a colony is plastic and colonies can withstand removal of highly active individuals.     

Two independent mechanisms of egg recognition in worker <Emphasis Type="Italic">Formica fusca</Emphasis> ants

Informational constraints can be an important limitation on the accuracy of recognition. One potential constraint is the use of recognition information from the same sources in multiple discriminatory contexts. Worker wood ants, Formica fusca, discriminate eggs based on their maternal sources of origin in two main contexts: recognition of eggs laid by nestmate versus non-nestmate queens and recognition of worker-laid versus queen-laid eggs. We manipulated the experience of F. fusca workers in laboratory colonies to both worker-laid and queen-laid eggs by transferring eggs between colonies in order to investigate whether these two contexts of egg discrimination are independent. Experience of non-nestmate queen-laid eggs significantly increased worker acceptance of both familiar (18% accepted) and unfamiliar (10%) queen-laid eggs compared to control workers without experience of eggs other than those laid by their own colony’s queen (2%). In contrast, worker acceptance of worker-laid eggs was not affected by variation in the egg experience of workers (14% in workers from control colonies exposed only to eggs from their own colony’s queen versus 19% and 17% in workers from colonies which had received eggs laid by either a non-nestmate queen or nestmate workers, respectively). Our results suggest that these two recognition contexts do not strongly constrain each other and are different in their ontogeny. In particular, worker-laid eggs are universally discriminated against by workers from colonies with a queen whatever the egg experience of the workers, while non-nestmate queen-laid eggs are strongly discriminated against only by workers without experience of eggs laid by more than one queen.     

Pink-line syndrome, a physiological crisis in the scleractinian coral Porites lutea

Coral diseases are one of the major factors that alter coral cover and their diversity. We have earlier reported the “Pink-line syndrome” (PLS) in the scleractinian coral Porites lutea wherein a colored band appears between the dead and healthy tissue of a colony. About 20% of the P. lutea colonies were affected in Kavaratti of the Lakshadweep Islands in the Arabian Sea during April 1996 and the incidence increased fourfold within the next 4 years. Fungi were associated in both PLS-affected and healthy specimens, whereas the cyanobacterium Phormidium valderianum occurred exclusively in the PLS-affected specimens. There was an increased expression of a 29 kDa protein without any significant increase in total protein content in the PLS-affected colonies. A reduced number of zooxanthellae and an increase in zooxanthellae size, mitotic index, and chl a concentrations were some of the characteristics of the PLS-affected colonies. PLS induction experiments conducted using selected fungi and the cyanobacterium P. valderianum isolated from the affected colonies and abiotic factors, such as CO₂ enrichment and the effect of cyanobacterial photosynthesis inhibition, indicated that the CO₂ build-up around the host tissue caused the pink coloration. We hypothesize that these physiological changes disturb the mutualism between the zooxanthellae and the host. When the symbiosis is disturbed by the external CO₂, the host loses control over the zooxanthellae, causing their uncontrolled division. This process may lead to a break in photosynthate transfer to the host, thereby resulting in starvation and finally leading to partial mortality. We further hypothesize that these degenerative processes are triggered by the CO₂ produced by P. valderianum through its carbon concentration mechanism. In this context, any opportunistic cyanobacteria or other agents having potential to interfere with the physiology of the host or the symbiont can cause such a physiological disorder. The mechanism of PLS formation is an early warning to protect corals as the increasing atmospheric CO₂ could induce PLS-like physiological disorder in corals.