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The timing and spatial variation in spawning in the green sea urchin Strongylocentrotus droebachiensis (Müller) was investigated at three moderately protected sites in each of three geographic regions along the coast of Maine before the commencement of significant commercial harvesting. Urchins were sampled monthly (1987 to 1988) from subtidal hard bottoms, and test diameter (TD), height, total wet weight, and gonad wet weight were measured. To interpret reproductive and spawning patterns additional data were taken on habitat type, water temperature, salinity, urchin density, and diets. Over a range of TD (34.1–89.4 mm), 1,594 urchins were sampled. Gonad index (GI) increased as an allometric function of TD, and for urchins from the northeast and southwest regions, GI was independent of TD for animals ≥64 mm. In the central region, the size at independencewas ≥55 mm. Analysis of variance with a priori, planned contrasts was used to quantify temporal changes in GI and spawning at two spatial scales (within and between regions). This information serves as a preharvest baseline for green urchin dynamics, analysis of reproductive cycles and spawning, and for current and future ocean changes. Gonad index and spawning varied seasonally, spatially and interannually. Gonad index increased during fall and early winter, and peaked in midwinter before a major spawning event in April at seven of nine sites. Gonad index ranged from 10%to 20% from December to April. Spawning [measured as a steep decline in GI (48%–78%) between successive sampling dates) occurred between early April and mid-May, except at one site in the central (Lamoine: March to April) and one in the northeast (Jonesport: May to June) regions. Gonad index patterns during spawning corresponded inversely to increasing seawater temperatures in the range of 2.5–5 °C. Salinity, urchin density, and test size did not explain a significant proportion of the variability in mean GI through time. Diets consisted primarily of diatoms and microalgae on ledge, sediment, and coralline barrens and showed no regional trends. Sex ratio explained a significant portion of the variability in mean GI at only one site. Seawater temperature, however, explained 55%–77% of the variability in mean GI through time. Predicting when spawning occurs in natural populations is central to the sea urchin fishery by refining estimates of what are termed harvest windows (HW). The HW represents a segment of time during the general spawning season when GI are at, or above, a specified percent, for example, 10%. A review of the literature uncovered 19 different techniques to determine GI and assess spawning. Of 167 papers published between 1922 and 2013 in which methods of spawning in wild populations of sea urchins were described, 84 and 134 used histology and GI, respectively. This study contributes to the questions of dependence of GI on test size, first illuminated by Gonor (1972), and the general practice of interpreting minor declines in GI as fractional spawning events, rather than simply sampling noise. The use of statistical tests is encouraged to define aspects of the reproductive cycle in sea urchins.

Spatial and Temporal Variability of Spawning in the Green Sea UrchinStrongylocentrotus droebachiensisalong the Coast of Maine