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Gametogenic Cycle of <I>Rangia Cuneata</I> (Mactridae, Mollusca

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Gametogenic Cycle of <I>Rangia Cuneata</I> (Mactridae, Mollusca BULLETIN OF MARINE SCIENCE, 45(1): 130-138, 1989 GAMETOGENIC CYCLE OF RANGIA CUNEATA (MACTRIDAE, MOLLUSCA) IN MOBILE BAY, ALABAMA, WITH COMMENTS ON GEOGRAPHIC VARIATION M. C. Jovanovich and K. R. Marion ABSTRACT Stages of the gametogenic cycle of the brackish water clam Rangia cuneata were investigated in Mobile Bay, Alabama, by histological examination of the gonads. Water temperature and salinity measurements were related to the gametogenic cycle and compared to those made in other studies on the same species in different locations. Clams in the early active phase were found from February through April, and those in the late active phase predominated in May. Gonads were ripe from July through September and were partially spawned from September through October. Clams became spent between November and January. Changes in meat, gonad, and total wet weight reflected the stages of the gametogenic cycle, while length of the shells remained nearly constant year round. The gametogenic cycle of R. cuneata in Mobile Bay followed a similar pattern to that observed in clams studied in other locations, except that gametogenesis began earlier, during late winter, and clams became spent earlier in the fall. It is suggested that the interactive effects of temperature, salinity, and nutrients can account for the differences in the timing and length of the stages of the gametogenic cycle between locations. Rangia cuneata (Gray) is a brackish water clam (Family Mactridae) abundant in the estuaries of the Gulf of Mexico. It is also found throughout the coastal areas of the eastern United States as far north as the upper Chesapeake Bay (Woodburn, 1962; Pfitzenmeyer, 1970). R. cuneata inhabits the upper layer of sediment in areas where the salinity ranges between 0 and 150/00and where water temperatures vary seasonally between 0.5 and 35°C (Hopkins et aI., 1973). In the James River, Virginia, Cain (1975) reported that this species made up 95% of the total benthic biomass. As a result of such high population densities and because R. cuneata converts detritus to biomass that can then be used as a food source for many bottom-dwelling and bottom-feeding animals (Odum and Copeland, 1969), the species is considered to be an important link in the food chains of estuaries. This clam has also been used in recent years as a biological indicator of pollution in the upper reaches of estuaries, because of its ability to accumulate organic pollutants, particularly the polycyclic aromatic hydrocarbons (PAH) (Fu- cik et aI., 1977; Neffand Anderson, 1981). During the course of a study conducted by the authors on the biological and physical factors influencing the uptake and depuration of the PAH anthracene by this clam, it became apparent that infor- mation on the reproductive cycle of R. cuneata in the Gulf of Mexico is limited. Fairbanks (1963) observed that R. cuneata in Louisiana had two "incompletely definitive" spawning periods per year. In February, clams developed a pre-spawn- ing condition, followed by the production of ripe gametes and spawning in March through May. According to Fairbanks, there was a short period of recovery in early June, followed by a more prolonged period of gamete production beginning at the end of the same month and lasting until November. Cain (1975) investigated the gametogenic cycle of R. cuneata in the James River, Virginia, and indicated that gametogenesis started in early April and continued throughout the summer. Ripe gametes were present from May to late November, with spawning beginning in early and mid-summer, but the major spawning period occurred in the fall. 130 JOVANOVICH AND MARION: GAMETOGENIC CYCLE OF RANGIA CUNEATA 131 30 I 12.0 25 0 0- en 10.0 r-» -W 20 a: Z ;:) 8.0 ~ =i c( 15 -< 6.0 a: Cl W - Q. 8- :E 10 4.0 - W ~ I 5 2.0 JJ ASONDJ F MAMJ JASON OJ F 1983 1984 1985 Figure 1. Rangia cunea/a. Seasonal changes in water temperature and salinity at Dog River, Mobile County, Alabama between June 1983 and January 1985. The purpose of this study was to determine the gonadal development of R. cuneata under the temperature and salinity conditions prevailing in Mobile Bay, Alabama over a 16-month period. Since the gametogenic cycle of lamellibranchs can vary considerably between localities (Pfitzenmeyer, 1962; Ropes and Stickney, 1962; Shaw, 1964), knowledge of the cycle at additional locations can provide useful insights to factors controlling geographic variation in the reproduction of lamel- libranchs. MATERIALS AND METHODS Clams were collected by hand at the mouth of Dog River, Mobile County, Alabama, USA (30°34' 10"N; 88°05'OO"W) approximately every 6 weeks from September 1983 to January 1985. The salinity and temperature of the water were mcasured at every collection and, whenever possible, between collec- tions, using a YSI salinity-temperature meter. Samples for histological examination consisted of 16 mature individuals per collection. Each clam was measured by taking the total length from the anterior to the posterior of the valve. Total wet weight was recorded, after which each clam was shucked and the tissue wet weight determined. Gonads were then dissected, weighed, and fixed in Bouin's fixative for 24 h (Pantin, 1969). The tissues were embedded in paraffin, sectioned at 6 jlm thickness, stained with Mayer's Acid Hema]um as modified by Lillie (Clark, 1981) and counterstained with Eosin Y. Each slide was examined at 63 x and 250 x magnification to determine the status of gonadal deve]- opment. Since the gametogenic cycle of this clam is a nearly continuous process without sharp demarcations, five artificial stages of gonadal development, as employed by Cain (1972), were used for reproductive assessment. These were: early active, late active, ripe, partially spawned and spent. The average number of eggs in five follicles for each female collected (total number of females = 50) was also determined, and the vertical and horizontal axis of five eggs selected at random were measured, together with the diameter of the nucleus of each of these eggs. RESULTS Water Temperature and Salinity. - Water temperatures at the sampling site (Fig. 1) showed a clear seasonal cycle, with maximum values in September (29°C in 1983 and 32°C in 1984), and minimum values in February 1983 (11°C) and 132 BULLETIN Of MARINE SCIENCE, VOL. 45, NO. I, 1989 3: - m ~ I~ -f C) 10.0 ~ -•.•• 60.0 -f ~ ....I 8.0~ <C •.•• 48.0 o I •••• 6.0 -C) -, 36.0 o I Z • 4.0 ~ E E 24.0 ~ -f :J:- 2.0 ca o•••• 12.0 Z UJ ....I 1983 J A SON o J 1985 - Figure 2. Rangia cuneata. Changes in length (closed squares), total wet weight (circles), meat wet weight (open squares) and gonad wet weight (triangles) in clams between September 1983 and January 1985. N = 16 for each month, except for October 1983 (N = 13), September 1983 (N = 11), February and March 1984 (N = 12), and May and December 1984 (N = 15), December 1984 (10°C). Changes of more than 10°C were observed between April and May 1984, and again between September and October of the same year. Salinity also followed a seasonal pattern (Fig. 1) with a maximum value in November 1983 (14%0) and another peak value in October 1984 (100/00).Minimum salinity was observed in March 1984 (00/00).Salinity values reflected the discharge from the nearby rivers, being high during extended periods of low river influx and very low under flooding conditions, approaching 00/00(Schroeder and Lysinger, 1979). Seasonal Variation in Size. - The length of the shells remained nearly constant throughout the 16-month period, averaging 47.0 ± 0.6 mm (Fig. 2); however, the total weight of individual clams and the weight of their meat showed a seasonal cycle (Fig. 2). Both values were minimal between October 1983 and January 1984, while maximum weights were observed in August (58.1 ± 2.4 and 8.3 ± 0.3 g, respectively). Gonad weight also reflected this pattern (Fig. 2), reaching minimum values during October 1983 through March 1984, followed by increased weights from May 1984 through September 1984, with a maximum in August (2.7 ± 0.2 g). Histology: Female Gonadal Development. - Using the system of Cain (1972), fe- males judged to be in the early active stage had few oogonia at the periphery of the follicle (Fig. 3a, b). Oogonia appeared partially embedded in the membrane or attached by a short stalk. In the late active phase, larger oocytes (Fig. 3c), measuring between 29 and 38 !lm, with a conspicuous basophilic nucleolus present in the nucleus became prevalent. The nuclei measured between 20 and 25 !lm. Gonads of ripe females (Fig. 3d, e) contained the largest number offollicles, with densely packed oocytes in their lumina. The mature oocytes measured between Figure 3. Rangia cuneata. Sections of gonad tissue of clams from Mobile Bay, Alabama. a and b) female in early active phase of oogenesis (x 63 and x 250); c) female in late active phase (x 250); d and e) ripe female (x 63 and x 250); f and g) partially spawned ovary (x 63 and x 250); h) spent female (x 63). 134 BULLETIN OF MARINE SCIENCE, VOL. 45, NO. I, 1989 34 and 44 /lm, with a nucleus averaging 25 /lm in diameter. The number of oocytes in the follicles decreased and appeared free in the lumina when females became partially spawned (Fig. 3f, g). Oocyte diameter remained the same, averaging 44 /lm. As spawning continued, follicles were no longer well-defined, and increased numbers of leucocytes or phagocytes were observed in the lumina. The lumina of the follicles of spent females were generally void of oocytes, or a few oocytes were still scattered over the surrounding tissue (Fig.
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