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Alexandrium monilatum

Figure 1. A 6-cell chain of living A. monilatum.

Figure 2. Single cell of A. monilatum captured using SEM. Image courtesy of Florida Fish & Wildlife Conservation Commission.

Figure 3. Schematic apical view of A. monilatum, showing ventral pore (vp, blue arrow), isolation of first apical plate (1’)  from apex, and the apical pore plate (Po, green arrowhead) with anterior attachment pore (aap, red arrow and gray).

Figure 4.Antapical view of A. monilatum, showing posterior sulcal plate (light gray) and attachment pore (red arrow and dark gray).

SPECIES NAME: 
Alexandrium monilatum (Howell) Balech

COMMON NAME:
Dinoflagellate

SYNONOMY:
Gonyaulax monilata Howell
Gessnerium mochimaensis Halim
Gessnerium monilatum (Halim) Loeblich III
Pyrodinium monilatum (Howell) Taylor

The original description by Howell (1953) was based on samples from the IRL, so the type locality of A. monilatum is the IRL.  Halim (1967) established the genus Gessnerium, into which Loeblich placed this species.  However, Halim’s description was based on an optically reversed image, so that genus name is not accepted.  The name Gessnerium is now frequently used as a subgenus name within Alexandrium, encompassing several species including A. monilatum. Taylor (1976) later transferred the species to Pyrodinium, and Balech (1995) included it in Alexandrium.

Cells are joined in chains by exudates from the apical attachment pore (Figure 3, red arrow) and the antapical attachment pore, centrally located and surrounded by several raised radial markings, in the large and concave posterior sulcal plate (Figure 4, red arrow).  This attachment is not firm and, though 16-cell chains are quite common, 8- and 4-cell chains are also frequent. In quiet waters, 32-cell chains can be found.  In the original description, Howell (1953) found up to 40 individuals per colony.  Despite this, when grown in stationary culture, there is a tendency for the colonies to degenerate into unicells.

Alexandrium monilatum is photosynthetic and filled with large number of small brown or yellow-brown chloroplasts that radiate from the centrally located nucleus, which is somewhat lunate. At least some strains are bioluminescent (Latz et al. 2008).

GenBank has limited information on the molecular genetics of this species http://www.ncbi.nlm.nih.gov/.

II.  HABITAT & DISTRIBUTION

HABITAT & REGIONAL OCCURENCE:
Alexandrium monilatum is a coastal and estuarine planktonic species of warm temperate and tropical environments, and is known from both eastern Pacific and western Atlantic Oceans.  It is widespread in the Gulf of Mexico and Caribbean Sea, and has been recorded from the Chesapeake Bay.

INDIAN RIVER LAGOON DISTRIBUTION & ABUNDANCE:
In the IRL, A. monilatum is most often found in summer and early fall, particularly south of Fort Pierce, rarely reaching concentrations of more than a few thousand cells per liter.  However, Norris (1983) reported a bloom of 106 cells per liter at Melbourne Beach.  It has been rare in the Banana River and Mosquito Lagoon.

III. LIFE HISTORY & POPULATION BIOLOGY

SIZE:
The width of individuals is 33-60 µm (Hallegraeff 2003) or 39-67 µm (Balech 1995), with a length of 28-52 µm.

REPRODUCTION:
The life cycle has been studied by Walker and Steidinger (1979), and appears to be typical for dinoflagellates (see Figure 9 of Pyrophacus steinii for schematic life cycle stages): fusion of isogametes form a planozygote which then encysts. The cysts are round to ovoid and filed with darkened yellow-brown cytoplasm and lipids. Asexual reproduction is by binary fission.

TOXICITY:
Although A. monilatum has rarely been implicated in marine mortalities in the IRL since its original description, it was the cause of fish kills due to ichthyotoxins along the Texas coast (Gates & Wilson 1960; Ray & Aldrich 1967) when cell concentrations approached 106 cells per liter.  An unusual bloom in a tributary of the Chesapeake Bay had over 107 cells per liter (May et al. 2010).  It was earlier reported to produce PSP toxins (saxitoxin and gonyautoxin) with hemolytic and neurotoxic properties, but the primary toxin is apparently goniodomin-A (Hsia et al. 2006).  In addition to killing fish, A. monilatum reduces adult feeding activity and increases larval mortality in commercially valuable shellfish, as well as a variety of other invertebrates (May et al. 2010).

IV. REFERENCES

Balech, E. 1995. The genus Alexandrium Halim (Dinoflagellata). Sherkin Island Marine Station, Cork, Ireland. 151pp.

Gates, JA & WB Wilson. 1960. The toxicity of Gonyaulax monilata Howell to Mugil cephalus. Limnol. Oceanogr. 5: 171-174.

Halim, Y. 1967. Dinoflagellates of the South East Caribbean Sea (East Venezuela). Internationale Revue des gesamten Hidrobiologie 52: 701-755.

Hallegraeff, GM. 2003. Taxonomic Principles. In: Hallegraeff, GM, Anderson, DM & AD Cembella (Eds.). 383-432. Manual on Harmful Marine Microalgae. UNESCO Publishing, Paris. 793pp.

Hsia, MH, Morton, SL, Smith, LL, Beauchesne, KR, Huncik, KM & PDR Moeller. 2006. Production of goniodomin A by the planktonic, chain-forming dinoflagellate Alexandrium monilatum (Howell) Balech isolated from the Gulf coast of the United States. Harmful Algae 5: 290-299.

Howell, JF. 1953. Gonyaulax monilata sp. nov.,the causative dinoflagellate of a red tide in the east coast of Florida in August-September 1951. Trans. Amer. Microsc. Soc. 72: 153-15.

Latz, MI, Bovard, M, VanDelinder, V, Segre, E, Rohr, J & A Groisman. 2008.  Bioluminescent response of individual dinoflagellate cells to hydrodynamic stress measured with millisecond resolution in a microfluidic device.  J. Exp. Biol. 211: 2865-2875.

Loeblich III, AR. 1970. The amphiesma or dinoflagellate cell covering. In: Yochelson, EL (Ed.). Proceedings of the North American Paleontological Convention, Chicago, September 1969, Part 2. G. Allen Press, Lawrence, KS. 867-929.

May, SP, Burkholder, JM, Shumway, SE, Hegaret, H, Wikfors, GH & D Frank. 2010. Effects of the toxic dinoflagellate Alexandrium monilatum on survival, grazing and behavioral response of three ecologically important bivalve mollusks.  Harmful Algae 9: 281-291.

Norris, DR. 1983. The occurrence of a toxic dinoflagellate in the Indian River system, Florida. Fla. Sci. 46: 150-153.

Ray, SM & DV Aldrich. 1967. Ecological interactions of toxic dinoflagellates and mollusks in the Gulf of Mexico. 75-83. In: Russell, FE & PR Saunders [Eds.]. Animal Toxins, First International Symposium on Animal Toxins. Pergamon Press, NY.

Taylor, FJR. 1976. Dinoflagellates from the international Indian Ocean Expedition. Bibliothec. Botan. 132: 1-234 + 46 Plates.

Unless otherwise noted, all images and text by PE Hargraves
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