Attempts to model the bloom dynamics of Pyrodinium, a tropical toxic dinoflagellate

For the first time, several models have been used to aid in the understanding of the bloom dynamics of Pyrodinium bahamense var. compressum, the major causal organism of toxic algal blooms in Manila Bay and several areas in the tropical world. The complex life cycle of Pyrodinium includes the formation of cysts that settle at the sediments, which can serve as the inoculum for the next bloom.The seasonal variation of temperature and salinity reflects the combined effects of convection and water column stability, which can control vertical movement of plankton and other parameters essential to its growth. The significance of wind forcing appears to be related to the potential to resuspend cysts. In the absence of wind, tidal currents in the inner part of the bay may be too weak to induce resuspension. The addition of wind results in a significant increase in bottom current velocity. Off Cavite at the southeast, bottom velocity is enhanced by orbital motion due to waves, one of the reasons why sediments off this area are dominated by sandy material. The strong vertical mixing of the water column at depths of less than 10 m may influence nutrient and consequently, plankton populations.The wave field during the southwest monsoon indicates that its contribution to the bottom velocity dominates in this area of the bay.Bloom simulations using combined bio-physical parameters show that direction of advection is almost always along wind direction. The dispersal distances increases if the Pyrodinium cells are found higher in the water column. For cells originating from southeastern (Cavite) sources, the direction of transport is slightly towards the north. In either case, the formation of cysts after a bloom is adjacent to the northern area (Pampanga) for blooms originating from the western side (Bataan) and along the eastern side (Parañaque–Manila) for blooms originating from the southeastern side (Cavite). Comparison with a few records of bloom occurrences in Manila Bay shows some consistent features. Reports of these blooms also showed that they occurred almost always during spring tides. There appears to be two main systems for bloom formation: one fed by cyst beds in the west (Bataan) which is advected along the west–northwest coast (Bataan–Bulacan) while the other one is fed by the southeast (Cavite) cyst beds that dominates in the east-southeast (Parañaque–Cavite) area.     

Bacterial Endosymbionts of Pyrodinium bahamense var. compressum

The study presents evidence in support of the bacterial theory associated with the toxicity of Pyrodinium bahamense var. compressum. Bacterial endosymbionts from Philippine P. bahamense var. compressum strain Pbc MZRVA 042595 were isolated and identified via 16S rDNA sequence analysis. Taxonomic diversity of the identified culturable intracellular microbiota associated with Philippine P. bahamense var. compressum was established to be limited to the Phyla Proteobacteria, Actinobacteria, and Firmicutes. Major endosymbionts identified included Moraxella spp., Erythrobacter spp., and Bacillus spp., whereas Pseudomonas putida, Micrococcus spp., and Dietzia maris were identified as minor isolates. All identified strains except D. maris, P. putida, and Micrococcus spp. were shown to contain either saxitoxin or neo saxitoxin or both at levels ≤73 ng/10⁷ bacterial cells based on high-performance liquid chromatography analysis. Paralytic shellfish poisoning-like physiologic reactions in test animals used in the mouse assay were recorded for the endosymbionts except for P. putida. The study is the first to elucidate the possible contribution of bacterial endosymbionts in the toxicity of P. bahamense var. compressum isolated in the Philippines.     

Morphology of Pyrodinium bahamense Plate (Dinoflagellata) near Isla San José, Gulf of California, Mexico

The morphology of Pyrodinium bahamense south of Isla San José in the Gulf of California, Mexico was described using light and scanning electron microscopy. Morphological and dimensional examination of whole cells and thecal patterns were conducted to discriminate between P. bahamense var. compressum and P. bahamense var. bahamense. Microscopic examinations confirmed that specimens of P. bahamense from Isla San José are closely related to P. bahamense var. bahamense. The main average dimensions were: cells 41.9 μm long, 43.8 μm wide, 55.6 μm amplitude, apical horn 7.2 μm long, and the left antapical spine 19.5 μm long. The highest cell concentration was 240 cells l⁻¹ and occurred with a bloom of Pseudo-nitzschia spp. The presence of this dinoflagellate off the east coast of the Baja California Peninsula had not been previously reported. Recently, P. bahamense from Florida tested positive for saxitoxins. Paralytic shellfish poisoning from this species has occurred in Mexican waters. Here, significant aspects of taxonomy, toxicology, life cycle, and ecology are discussed.     

Responses of Pyrodinium bahamense var. compressum and associated cultivable bacteria to antibiotic treatment

Pyrodinium bahamense var. compressum is a toxic dinoflagellate that produces paralytic shellfish poisoning toxins. It is responsible for the chronic toxicity of shellfish in many coastal areas of the Philippines and other South East Asian countries. For the purpose of using antibiotic treatment to possibly generate axenic cultures and understand their growth requirements, the antibiotic tolerances of two local P. bahamense var. compressum isolates and their associated bacteria were determined. The antibacterial compounds ampicillin, chloramphenicol, ciprofloxacin, kanamycin, neomycin, penicillin G, and streptomycin were tested, as well as two antifungals, amphotericin B, and nystatin. All except chloramphenicol, amphotericin B, and nystatin were generally well-tolerated. An antibiotic mixture composed of ciprofloxacin, kanamycin, neomycin, and streptomycin completely inhibited the cultivable bacteria associated with P. bahamense var. compressum MZRVA, although epifluorescence microscopy revealed that residual bacteria were still present. From long-term tests with this antibiotic mix, it was observed that survival of isolate MZRVA post-antibiotic treatment appeared to be associated with re-growth of heterotrophic bacteria and that excess vitamins could potentially enhance dinoflagellate survival. These results suggest that the associated heterotrophic bacterial populations help support the growth of P. bahamense var. compressum MZRVA in culture and possibly in nature. This is the first report on the growth responses of P. bahamense var. compressum and associated cultivable bacteria to a variety of single and combinations of antibiotics.     

Temperature mediates secondary dormancy in resting cysts of Pyrodinium bahamense (Dinophyceae)

High‐biomass blooms of the toxic dinoflagellate Pyrodinium bahamense occur most summers in Tampa Bay, Florida, USA, posing a recurring threat to ecosystem health. Like many dinoflagellates, P. bahamense forms immobile resting cysts that can be deposited on the seafloor—creating a seed bank that can retain the organism within the ecosystem and initiate future blooms when cysts germinate. In this study, we examined changes in the dormancy status of cysts collected from Tampa Bay and applied lessons from plant ecology to explore dormancy controls. Pyrodinium bahamense cysts incubated immediately after field collection displayed a seasonal pattern in dormancy and germination that matched the pattern of cell abundance in the water column. Newly deposited (surface) cysts and older (buried) cysts exhibited similar germination patterns, suggesting that a common mechanism regulates dormancy expression in new and mature cysts. Extended cool‐ and warm‐temperature conditioning of field‐collected cysts altered the cycle of dormancy compared with that of cysts in nature, with the duration of cool temperature exposure being the best predictor of when cysts emerged from dormancy. Extended warm conditioning, on the other hand, elicited a return to dormancy, or secondary dormancy, in nondormant cysts. These results directly demonstrate environmental induction of secondary dormancy in dinoflagellates—a mechanism common and thoroughly documented in higher plants with seasonal growth cycles. Our findings support the hypothesis that a seasonal cycle in cyst germination drives P. bahamense bloom periodicity in Tampa Bay and point to environmentally induced secondary dormancy as an important regulatory factor of that cycle.     

Structures of Two Paralytic Shellfish Toxins,Gonyautoxins V and VI,Isolated from a Tropical Dinoflagellate,Pyrodinium bahamense var. compressa

Two paralytic shellfish toxins, gonyautoxin V and gonyautoxin VI, isolated from a tropical dinoflagellate, Pyrodinium bahamense var. compressa, were identified respectively to be derivatives of saxitoxin and neosaxitoxin with a sulfonatocarbamoyl moiety.     

The occurrence of potentially toxic dinoflagellates and diatoms in a subtropical lagoon, the Indian River Lagoon, Florida, USA

The occurrence of potentially toxic phytoplankton species was examined over a 5-year period in a region of the Indian River Lagoon in Florida that has recently been subject to ecologically significant events, putatively related to algal toxins. The results of the study reveal a significant presence of two species of phytoplankton that have been shown to be toxic in Florida or other regions of world, the dinoflagellate Pyrodinium bahamense var. bahamense and the diatom Pseudo-nitzschia pseudodelicatissima. Concentrations of the former species reached 638,000 cells l⁻¹ and concentrations of the latter reached 23.9 million cells l⁻¹. In addition, the abundance of one of these species, P. bahamense var. bahamense appears to have increased over the 5-year study period from 1997 to 2002. It may be hypothesized that rainfall events following a regional drought period resulted in a flushing of bioavailable phosphorus and nitrogen into the Indian River Lagoon that stimulated P. bahamense var. bahamense blooms. The significance of these results is discussed within the context of the ecology of this flow-restricted lagoon.     

Sterols of the Toxic Marine Dinoflagellate,Pyrodinium bahamense

Pyrodinium bahamense is a dinoflagellate of concern in subtropical and tropical coastal environments. To date, there is only a single published study on its fatty acids, but no published data on its sterol composition. Sterols, which are membrane‐reinforcing lipids in eukaryotes, display a great diversity of structures in dinoflagellates, with some serving as chemotaxonomic markers. We have examined the sterol compositions of two isolates of P. bahamense from Indian River Lagoon and Tampa Bay, Florida, and have found both to produce three sterols: cholesterol, dinosterol, and 4α‐methylgorgostanol. All three sterols are found in closely related, armored taxa.     

Development,morphological characteristics and viability of temporary cysts of Pyrodinium bahamense var. compressum (Dinophyceae) in vitro

Pellicle or temporary cysts of Pyrodinium bahamense var. compressum (Böhm) Steidinger, Tester & F.J.R. Taylor and their role in bloom dynamics have not yet been adequately characterized and understood. We investigated the role of temperature- and nutrient-mediated stress as factors that could induce pellicle formation in batch cultures. Cellular features and their implications for temporary cyst viability were examined using confocal laser scanning microscopy (CLSM). Our data suggest that temperature change is one of the key factors influencing pellicle formation, preserving viability at low temperature (i.e. 13°C). Hypnocysts (resting cysts) were not observed. During pellicle formation, motile cells generally undergo ecdysis, extrusion of cytoplasmic materials and bacteria, compaction of the nucleus and non-motility. The outermost covering of the temporary cysts shows red autofluorescence and it contains lower concentrations of chlorophyll (chl) a and no detectable chl c. The nuclear region is surrounded by transitional red bodies and other unidentified cellular structures. Temporary cysts can immediately revert back to the motile state upon exposure to optimum conditions. This is accompanied by the expansion of the nuclear region, regeneration of the chloroplasts and enlargement of the cell. Developmental changes during reversal of temporary cysts to motile forms were also observed to cause breaks in the cell covering that could serve as sites for bacterial entry. Though observed in vitro, such behaviour may also be occurring in nature especially as a response to drastic short-lived environmental changes. This is the first detailed report on the characteristics of temporary cysts of P. bahamense var. compressum.     

Occurrences of harmful algal blooms (HABs) associated with ocean environments in the South China Sea

Harmful algal blooms (HABs) occur frequently in the South China Sea (SCS), causing enormous economic losses in aquaculture. We analyzed historical HAB records during the period from 1980 to 2003 in SCS. We found that HABs-affected areas have expanded and the frequency of HABs varied during this period. The seasonal and annual variations, as well as causative algal species of HABs are different among the four regions. Areas with frequent HABs include the Pearl River Estuary (China), the Manila Bay (the Philippines), the Masinloc Bay (the Philippines), and the western coast of Sabah (Malaysia). HABs occurred frequently during March–May in the northern region of SCS, May–July in the eastern region, July in the western region, and year-round in the southern region. Among the species that cause HABs, Noctiluca scintillans dominated in the northern region, and Pyrodinium bahamense in the southern and eastern regions. Causative species also varied in different years for the entire SCS. Both P. bahamense and N. scintillans were the dominant species during 1980–2003. Some species not previously recorded formed blooms during 1991–2003, including Phaeocystis globosa, Scrippsiella trochoidea, Heterosigma akashiwo, and Mesodinium rubrum. Variations in HABs are related to various regional conditions, such as a reversed monsoon wind in the entire SCS, river discharges in the northern area, upwelling in Vietnam coastal waters during southwest winds and near Malaysia coastal waters during northeast winds, and eutrophication from coastal aquaculture in the Pearl River estuary, Manila Bay, and Masinloc Bay. Handling editor: D. Hamilton     

Alexandrium (Dinophyceae) species in Malaysian waters

A study was carried out to determine the presence of paralytic shellfish poisoning (PSP) toxin-producing dinoflagellates in the coastal waters of Peninsula Malaysia. This followed first ever occurrences of PSP in the Straits of Malacca and the northeast coast of the peninsula. The toxic tropical dinoflagellate Pyrodinium bahamense var. compressum was never encountered in any of the plankton samples. On the other hand, five species of Alexandrium were found. They were Alexandrium affine, Alexandrium leei, Alexandrium minutum, Alexandrium tamarense and Alexandrium tamiyavanichii. Not all species were present at all sites. A. tamiyavanichii was present only in the central to southern parts of the Straits of Malacca. A. tamarense was found in the northern part of the straits, while A. minutum was only found in samples from the northeast coast of the peninsula. A. leei and A. affine were found in both the north and south of the straits. Cultured isolates of A. minutum and A. tamiyavanichii were proven toxic by the receptor binding assay for PSP toxins but A. tamarense clones were not toxic. Mean toxin content for the A. tamiyavanichii and A. minutum clones were 26 and 15 fmol per cell STX equivalent, respectively. This study has provided evidence on the presence of PSP toxin-producing Alexandrium species in Malaysian waters which suggests that PSP could increase in importance in the future.     

Seasonal change of dinoflagellates cyst flux collected in a sediment trap in Omura Bay, West Japan

Sediment trap samples were harvested bimonthly from 1998 to2000 and examined to better understand the species compositionand seasonal variation of dinoflagellate cyst flux in OmuraBay in Japan. Samples’ living cyst flux clearly showedseasonal variation with the higher flux number between autumnand winter. In total, 43 different cyst taxa were recorded,and these were composed of two different ecological groups.The first group included Protoperidinium compressum and Protoperidiniumsubinerme, which increased every autumn to winter. The secondgroup included Gonyaulax spp. and Pheopolykrikos hartmanniiand was trapped throughout the year. These two groups manifestedthe different flux patterns and were, respectively, heterotrophicand autotrophic in nutrition. In the heterotrophic group, protoperidinioidcysts were dominant. Vegetative cells of protoperidinioid areknown to feed mainly diatoms. Sample diatom flux also increasedfrom autumn to winter. Therefore, the increase of protoperidinioidcysts in autumn to winter was observed to correlate with diatomblooms. In contrast, the autotrophic group mostly consistedof Gonyaulacoid cysts and were generally observed throughoutthe year, although occurrence varied between species most likelyresponding to favorable environmental conditions. The resultsindicate that cyst production is closely related to differentnutritional modes.     

Organic-walled dinoflagellate cyst production in relation to upwelling intensity and lithogenic influx in the Cape Blanc region (off north-west Africa)

Fossil dinoflagellate cyst assemblages are increasingly used in paleoclimatic research to establish paleoenvi‐ronmental reconstructions. To obtain reliable reconstructions, it is essential to know which physical factors influence the cyst production. Most information about the relationship between variations in physical parameters and cyst production is known from middle and higher latitudes. Information from the (sub)tropics is rare. To increase this information, the temporal variation in cyst assemblages from the upwelling area off north‐west Africa (off Mauritania) has been compared to environmental conditions of the upper water column. Samples were collected by the sediment trap CB9, off north‐west Africa (Cape Blanc, 21°15′2″N, 20°42′2″W) between 11 June 1998 and 7 November 1999 at 27.5‐day intervals. Off Cape Blanc, upwelling occurs throughout the year with variable intensity. This region is also characterized by frequently occurring Saharan dust storms. Seasonal variations in dust input, upwelling intensity and sea surface temperature are reflected by the production of organic‐walled dinoflagellate cyst assemblages. Several cyst taxa are produced throughout the sampling interval, with the highest fluxes at times of strongest upwelling relaxation and/or dust input (Echinidinium aculeatum Zonneveld, Echini‐din ium delicatum Zonneveld, Echinidinium granulaturn Zonneveld, Echinidinium spp., Impagidinium aculeatum (Wall) Lentin et Williams, Impagidinium sphaeri‐cum (Wall) Lentin et Williams, Protoperidinium americanum (Gran et Braarud) Balech, Protoperidinium stellatum (Wall in Wall et Dale) Rochon etal., Protoperidinium spp., Selenopemphix nephroides (Benedek) Benedek et Sarjeant and Selenopemphix quanta (Bradford) Matsuoka). Species such as, for example, Bitectatodinium spongium (Zonneveld) Zonneveld et Jurkschat and Impagidinium patulum (Wall) Stover et Evitt do not show any production pattern related to a particular season of the year or to specific environmental conditions in the upper water column. The production of cysts of Protoperidinium monospinum (Paulsen) Zonneveld et Dale is restricted to intervals with increased nutrient concentrations in upper waters when sea surface temperatures at the sampling site is below approximately 24°C.     

Impact of grazing and bioturbation of marine benthic deposit feeders on dinoflagellate cysts

The impact of benthic deposit feeders on marine dinoflagellate cysts was studied by adding a concentrated natural Swedish cyst assemblage to sediment with different deposit feeders in replicate 4-l aquaria. The deposit feeders used were the bivalve Abra nitida, the echinoderm Amphiura filiformis, and the polychaetes Melinna cristata and Nereis diversicolor. These species occur naturally near the Swedish west coast and were selected to represent different ways of feeding. The results showed a significant relative decrease of unfossilizable cyst species; whereas, the common fossilizable species Lingulodinium polyedrum significantly increased in the cyst assemblage after grazing. This work suggests that differences in dinoflagellate cyst compositions can in part be caused by different animal grazing behaviors.     

A fish kill of massive proportion in Kuwait Bay, Arabian Gulf, 2001: the roles of bacterial disease, harmful algae, and eutrophication

In August and September 2001, Kuwait Bay, a semi-enclosed embayment of the Arabian Gulf, experienced a massive fish kill involving over >2500 metric tons of wild mullet (Liza klunzingeri), due to the bacterium Streptococcus agalactiae. In the Bay, this event was preceded by a small fish kill (100–1000 dead fish per day) of gilthead sea bream (Sparus auratus) in aquaculture net pens associated with a bloom of the dinoflagellate Ceratium furca. Sea bream were found to be culture positive for S. agalactiae, but did not show any visible signs of disease. Unusually warm temperatures (up to 35 °C) and calm conditions prevailed during this period. As the wild fish kill progressed, various harmful algae were observed, including Gymnodinium catenatum, Gyrodinium impudicum, and Pyrodinium bahamense var. compressum. Cell numbers of G. catenatum and G. impudicum exceeded 10⁶ l⁻¹ in some locations. All fish tested below the limits of detection for paralytic shellfish poisoning (PSP) and brevetoxins. Clams (Circe callipyga) were positive for PSP but at levels below regulatory limits. Nutrient concentrations, both inorganic and organic, were highly variable with time and from site to site, reflecting inputs from sewage outfalls, the aquaculture operations, a high biomass of decomposing fish, and other sources. It is hypothesized that many factors contributed to the initial outbreak of the bacterial disease, including unusual warm and calm conditions. The same factors, as well as enriched nutrient conditions, also apparently were conducive to the subsequent HAB outbreaks. The detection of PSP, while below regulatory limits, warrants further monitoring to protect human health.     

An Alexandrium Spp. Cyst Record from Sequim Bay,Washington State,USA, and its Relation to Past Climate Variability1

Since the 1970s, Puget Sound, Washington State, USA, has experienced an increase in detections of paralytic shellfish toxins (PSTs) in shellfish due to blooms of the harmful dinoflagellate Alexandrium. Natural patterns of climate variability, such as the Pacific Decadal Oscillation (PDO), and changes in local environmental factors, such as sea surface temperature (SST) and air temperature, have been linked to the observed increase in PSTs. However, the lack of observations of PSTs in shellfish prior to the 1950s has inhibited statistical assessments of longer‐term trends in climate and environmental conditions on Alexandrium blooms. After a bloom, Alexandrium cells can enter a dormant cyst stage, which settles on the seafloor and then becomes entrained into the sedimentary record. In this study, we created a record of Alexandrium spp. cysts from a sediment core obtained from Sequim Bay, Puget Sound. Cyst abundances ranged from 0 to 400 cysts · cm^?3 and were detected down‐core to a depth of 100 cm, indicating that Alexandrium has been present in Sequim Bay since at least the late 1800s. The cyst record allowed us to statistically examine relationships with available environmental parameters over the past century. Local air temperature and sea surface temperature were positively and significantly correlated with cyst abundances from the late 1800s to 2005; no significant relationship was found between PDO and cyst abundances. This finding suggests that local environmental variations more strongly influence Alexandrium population dynamics in Puget Sound when compared to large‐scale changes.     

Dinoflagellate cyst distribution in marine surface sediments off West Africa (17–6°N) in relation to sea-surface conditions, freshwater input and seasonal coastal upwelling

An organic-walled dinoflagellate cyst analysis was carried out on 53 surface sediment samples from West Africa (17–6°N) to obtain insight in the relationship between their spatial distribution and hydrological conditions in the upper water column as well as marine productivity in the study area.Multivariate analysis of the dinoflagellate cyst relative abundances and environmental parameters of the water column shows that sea-surface temperature, salinity, marine productivity and bottom water oxygen are the factors that relate significantly to the distribution patterns of individual species in the region.The composition of cyst assemblages and dinoflagellate cyst concentrations allows the identification of four hydrographic regimes; 1) the northern regime between 17 and 14°N characterized by high productivity associated with seasonal coastal upwelling, 2) the southern regime between 12 and 6°N associated with high-nutrient waters influenced by river discharge 3) the intermediate regime between 14 and 12°N influenced mainly by seasonal coastal upwelling additionally associated with fluvial input of terrestrial nutrients and 4) the offshore regime characterized by low chlorophyll-a concentrations in upper waters and high bottom water oxygen concentrations.Our data show that cysts of Polykrikos kofoidii, Selenopemphix quanta, Dubridinium spp., Echinidinium species, cysts of Protoperidinium monospinum and Spiniferites pachydermus are the best proxies to reconstruct the boundary between the NE trade winds and the monsoon winds in the subtropical eastern Atlantic Ocean. The association of Bitectatodinium spongium, Lejeunecysta oliva, Quinquecuspis concreta, Selenopemphix nephroides, Trinovantedinium applanatum can be used to reconstruct past river outflow variations within this region.     

Red tide phenomena in Brunei Darussalam — some implications for fisheries

Since the first recorded occurrence in 1976, the coastal waters of Brunei Darussalam have been sporadically subjected to the effects of red tide. The 1976 incident overwhelmed the nation which had no previous experience of red tides. Subsequently, a routine was established to monitor the red tide phenomenon. This included plankton monitoring and shellfish toxicity testing, measures which helped to prevent or mitigate adverse human impacts and economic losses to the fishing industry.The second red tide occurrence was in 1980. However, this time there was greater awareness and with the experience gained from the 1976 incident, the situation was handled effectively.A subsequent incident in 1988 was discovered in a slightly different manner. High densities of Pyrodinium bahamense var. compressum, the causative organism, were found in the stomach contents of a pelagic fish, Sardinella spp. Cats which had fed on contaminated Sardinella suffered sickness and mortality. This led to a ban on the import and sale of Sardinella spp. and the closure of a local shellfish farm for almost a year.Red tide occurrences have had some impact on the fisheries of Brunei Darussalam, mainly due to the need to impose bans on the import, sale and consumption of certain species of fish and shellfish.It is suggested that the effects of red tides on finfish capture fisheries, especially planktivorous fish can be minimised by implementing simple precautions. Shellfish under culture would invariably face losses in the event of a prolonged red tide occurrence and it is proposed that an insurance scheme be implemented to cover such losses.A Red Tide Action Plan is currently in force in Brunei Darussalam. It is a contingency plan for red tide vigilance, monitoring and management, and will serve to reduce the negative impacts of red tides.     

THE “HYSTRICHOSPHAERID” RESTING SPORE OF THE DINOFLAGELLATE PYRODINIUM BAHAMENSE,PLATF, 19062

Germination experiments demonstrate that the “hystrichosphere” called Hemicystodinium zoharyi, which previously has been found only as a microfossil organism, is the resting spore stage in the life history of Pyrodinium bahamense, a modern bioluminescent, thecate dinoflagellate. The morphology of this spore, together with new details of the thecal structure and ontogeny of P. bahamense, is described, and it is concluded that Pyrodinium is closely related to Gonyaulax but worthy of retention as a discrete genus. The geological history of P. bahamense is traceable to the Eocene through fossil occurrences of its spore, and it is suggested that additional pyrodinioid dinoflagellates which now are extinct were represented in Lower Tertiary seas by another hystrichosphere genus, called Homotryblium. Selected aspects of the physiology and ecology of modern dinoflagellate resting spores are discussed briefly with special reference to Pyrodinium.