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note
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Fertility (the conversion of protoplasmic contents into gametes with subsequent migration into gametangia) occurred within a 12-hour period, overnight, and can be detected, in situ, in all five genera, on the basis of color change and the appearance of gametangia [previously known for Halimeda (13)]. No evidence of spawning outside the morning observation period was ever detected.
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54
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15644373261
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This pattern of fertility and death, known as holocarpy, is diagnostic of the order (10). All traces of Caulerpa, Penicillus, and Rhipocephalus were generally gone from the reef within 24 hours. Remnants of the collapsed thalli of Halimeda and Udotea caribeae persisted 24 to 48 hours after spawn, and the blade U. flabellum often persisted for weeks.
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55
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15644375632
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-4; n = 37). This trend was present in all genera (analysis of covariance for 10 spp. with >5 observations; F(1,126) = 7.98; P < 0.005). Increasing light levels apparently initiate the process of gamete release (14) and dark mornings caused by heavy overcast, rain, or turbid water typically delayed gamete release on the reef by 5 to 15 min.
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(1996)
J. Phycol.
, vol.32
, pp. 21
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Hillis, L.1
Engman, J.2
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56
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15644369536
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I observed gamete behavior on uncovered microscope slides at 100x and 400x, using gametes released into aquaria (10 gallons) or plastic buckets (5 gallons). 24. Eyespots are a diagnostic feature of many bryopsidalian macrogametes (13). Although the expression of phototaxis within the group is not well studied, Halimeda macrogametes were observed to swim rapidly toward a light source in both the laboratory and the field.
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57
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0001217794
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Species of Halimeda (11), Penicillus (12), Udotea (12), and Rhipocephalus (confirmed, this study) are dioecious; Caulerpa is monoecious [M. Goldstein and S. Morall, Ann. N.Y. Acad. Sci. 175, 660 (1970)].
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(1970)
Ann. N.Y. Acad. Sci.
, vol.175
, pp. 660
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Goldstein, M.1
Morall, S.2
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58
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15644376442
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For dioecious species, the light green color of fertile "males" is easily distinguished, in situ, from the brownish, dark green [Halimeda (13)], or gray (Udotea, Rhipocephalus, Penicillus) colors of females. These color differences before gamete release allowed the detection of sex-specific patterns of spawning.
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59
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15644378028
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Gamete release by male algae began 1 to 19 min before female algae began releasing gametes during all 56 pairwise comparisons between sexes. Mean difference (min) ± 1 SE and sample size, by species: Halimeda incrassata (6.2 ± 1.0, 9); H. monile (2.8 ± 0.5, 5); H. opuntia (4.2 ± 0.4, 5); H. simulans (7.0 ± 0.7, 5); H. tuna (3.5 ± 0.9, 4); Penicillus capitatus (4.0 ± 0.7, 9); P. dumetosus (19.5 ± 0.5, 2); P. lamourouxii (4.0 ± 0.6, 3); P. pyriformes (4.3 ± 2.9, 3); Rhipocephalus phoenix (2.7 ± 0.9, 4); Udotea flabellum (5.0 ± 1.4, 5); U. caribeae (6.0 ± 0.3, 7).
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60
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0002859919
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Of the spawns observed between 5 June 1995 and 3 July 1996, 83% took place between March and July (48% of the sampling period). In the previous year, between 24 October 1994 and 1 May 1995, 90% of the 39 observed spawns took place after 1 March 1995 (50% of the sampling period). Panama's dry season (January through April) is characterized by strong trade winds and greater levels of solar irradiance [J. D. Cubit et al., Coral Reefs 8, 59 (1989)]. Overall levels of algal reproduction were higher in 1996 than in 1995 (for example, compare the occurrence of spawning during June 1995 and June 1996, Fig. 1A). The eight species observed spawning before 5 June 1995 (18) spawned at a rate of 0.3 spawn per day between 26 October 1994 and 3 July 1995 (172 days of observation). The same eight species spawned at more than twice that rate (0.7 spawn per day) between the same dates a year later (251 days of observation).
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(1989)
Coral Reefs
, vol.8
, pp. 59
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Cubit, J.D.1
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61
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15644380520
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The number of thalli that released gametes on a given morning was estimated qualitatively. Within a species, only 3 to 5% of the population typically became simultaneously fertile (several hundred plants), although larger spawns (15 to 20%) did occur. Levels of fertility in one species appeared unrelated to the activity of other species. The abundance of more actively spawning species (for example, Caulerpa racemosa, 39 spawns during the 125-day period between 1 March and 3 July 1996) declined noticeably during the study but was not quantified.
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62
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0021604309
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P. L. Harrison et al., Science 223, 1186 (1984).
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(1984)
Science
, vol.223
, pp. 1186
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Harrison, P.L.1
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63
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I thank the people of Kuna Yala and the Republic of Panama for permitting this work; L. Clifton, V. Francesco, P. Nachtrieb, M. Rodruigez, and M. Schlaepfer for their predawn efforts; E. Clifton, P. Raimondi, and D. Reed for their comments; and the Smithsonian Tropical Research Institute (STRI) Marine Environmental Sensing Program for water-temperature data. Supported by STRI, Smithsonian's Scholarly Studies program (SS1234-530A), and the University of California Toxics Program.
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