The length of time from fertilization until larvae achieve competency remains unknown for most species of octocorals. Once larvae become competent to metamorphose they must locate a suitable environment in which to settle. Successful recruitment depends upon encounter and selection of appropriate habitat, thus an ability to delay metamorphosis in the absence of a suitable environment may enhance the probability of survival by increasing the time available to locate more favorable conditions. The energetic resources available to developing larvae have been shown to influence both competency period and metamorphosis (references in Ben-David-Zaslow and Benayahu 1998). Richmond (1989) proposed that planula with zooxanthellae may benefit from nutritive algal metabolites, allowing them to extend their competency period. Contrary to Richmond ’s prediction, Ben-David-Zaslow and Benayahu (1998) found no significant difference in competency period or larval longevity between several species of zooxanthellate and azooxanthellate soft corals from the Red Sea . This suggests that for lecithotrophic octocoral larvae, nutrient sources within the oocyte are a key important factor controlling larval competence and longevity, and ultimately dispersal capabilities.
The dispersal abilities of octocoral larvae vary widely, especially among alcyonacean-type octocorals. In brooding species with crawling larvae, settlement occurs in the immediate vicinity of the parent colony (e.g. Alcyonium siderium, Sebens 1983; Capnella gaboensis, Farrant 1986) while brooders with swimming, planktonic larvae may disperse more widely (e.g. Anthomastis ritteri, Cordes et al. 2001; Dendronephthya hemprichi, Dahan and Benayahu 1998). Despite their locomotory abilities, most planular larvae of brooding, shallow-water octocorals appear to settle shortly after release (Brazeau and Lasker 1990; Coma et al. 1995; Excoffen et al. 2004). Observations of newly released larvae have shown that they are negatively buoyant and sink rapidly (Benayahu and Loya 1983; Brazeau and Lasker 1990; Gutierrez-Rodriguez and Lasker 2004). Benayahu and Loya (1987) proposed that short-range dispersal of planula is a common trait among coral reef alcyonaceans and may enhance localized recruitment and contribute to the patchy distribution of species. This strategy may be advantageous in space-limited environments (such as coral reefs) because larvae are immediately presented with appropriate habitat type in which to settle (Sebens 1983 in Benayahu and Loya 1987). While most larvae of brooding octocorals exhibit rapid settlement behavior, the planula of the Caribbean gorgonian-type octocoral, Pseudopterogorgia elisabethae, appear to be an exception. Gutierrez-Rodriguez and Lasker (2004) noted from field observations that while settlement of some planula occurred in close proximity to the parent colony (<5 m), most planula remained in the water column (presumably due to turbulence) and were even transported to the water surface, increasing the potential for advection from the local environment.
It is generally accepted that the larvae of broadcast spawning octocorals (i.e. Pennatulaceans and some alcyonaceans) have greater dispersal capabilities than brooded larvae, however data on long-distance transport of octocoral planula is lacking. Dahan and Benayahu (1998) reported that the planular larvae of the broadcast spawning alcyonacean, Dendronephytha hemprichi, swim actively and have a relatively long competency period (65 days). Such larval life features “undoubtedly promote a considerable larval transport in the field.” (Dahan and Benayahu 1998). The cosmopolitan distribution of some deep-water Pennatulids (e.g. Kophobelemnon stelliferum, Rice et al. 1992; U. lindahli, Tyler et al. 1995) also suggests the potential for widespread dispersal of larvae from broadcast spawning. In addition to deep-water Pennatulaceans, many species of deep-water alcyonaceans display widespread distribution, often covering entire ocean basins despite the intermittent occurrence of suitable hard substrate habitats. However, at the present time very little is known about reproductive processes of deep-water octocorals. Thus, factors contributing to the widespread distribution of many species are unresolved.
Numerous environmental features including substrate, light/dark cues, and water motion may influence coral settlement and successful recruitment. Many species of shallow-water octocoral larvae preferentially settle in shaded microhabitats, such as the underside of settlement plates (Alino and Coll 1989; Zeevi Ben-Yosef and Benayahu 1999; Benayahu and Loya 1987; Dahan and Benayahu 1997 and others). This settlement behavior may be an avoidance response to conditions that exist on the upper surface of a plate, such as high light intensity, low tides, competition from filamentous algae, grazing pressure, and sedimentation (Rogers et al. 1984 in Benayahu and Loya 1987; Benayahu and Loya 1984b). Despite potential unfavorable conditions that may exist on exposed surfaces, Alino and Coll (1989) observed reduced survivorship of larvae among the tropical alcyonaceans Lobophytum crassum and Sinularia conferta resident on the undersides of settlement plates compared to the light-exposed surfaces. The general trend toward settlement in shaded microhabitats appears to decrease with increasing depth (Benayahu and Loya 1987), presumably due to both reduced competition with algae and lowered grazing pressure. Thus, it stands to reason that, in deeper waters, substrate and water motion are key factors influencing octocoral planula settlement. In addition to settlement in shaded microhabitats, alcyonacean larvae exhibited a preference for substrates with turf or crustose coralline algae, rough surfaces, and pits in the substrata (Benayahu and Loya 1984b). Water flow may also be an important factor influencing larval settlement. Benayahu and Loya (1987) reported that settlement in Red Sea alcyonacean coral, X. macrospiculata, occurred predominantly along the edges of deployed settlement plates, a pattern which may be related to regions of low flow associated with turbulent eddies created as water travels over the plates (references in Benayahu and Loya 1987).
Once an octocoral planula locates an appropriate location to settle it undergoes metamorphosis to a feeding polyp. This process involves secretion of mucus for temporary attachment to the substrate during settlement (Benayahu and Loya 1983; and others). There are few reports of larval metamorphosis in octocorals. In the alcyonacean soft coral, P. f. fulvum, attachment to the substrate is followed by development into a cone-shaped polyp with 8 tentacular buds. Within a week to ten days, the tentacles elongate and septa develop inside the polyp. In successive weeks tentacles development pinnules and within the next month additional polyps are added and sclerites are present in the polyps (Benayahu and Loya 1983). Similar processes settlement and metamorphosis have been observed in the alcyonacean soft corals Capnella gaboensis (Farrant 1986) and X. macrospiculata (Benayahu and Loya 1984).
A limited number of studies indicate that successful settlement and recruitment into a population occurs at a low rate, at least among shallow-water alcyonacean octocorals (Farrant 1987; Grigg 1977; Lasker et al. 1998). Farrant (1987) reported that for the temperate soft coral, Capnella gaboensis, the first year survival rate of newly settled colonies was 0.26%. Similarly, for the broadcast spawning alcyonacean, Plexaura kuna, Lasker et al. (1998) estimated annual survival of new colonies at 10-6. Using a Poisson probability distribution, Lasker et al. (1998) calculated that in order to produce a successful a recruitment event 95% of the time, that settlement of 700,000 individuals per year would be required. Their calculated mortality rate for P. kuna settlers of 3% per day yields an extremely low probably for successful recruitment, suggesting that this species may rely on certain environmental circumstances that enhance successful recruitment (e.g. favorable substrata, Gotelli 1988; reduced grazing, Yoshioka 1996). Extremely high post-settlement mortality of new recruits implies that successful settlement may be more closely tied to water column and post-settlement survival than to gamete production and fertilization rates (Lasker et al. 1998). Coffroth and Lasker (1998b) proposed that the life history strategy of P. kuna relies on “large and long-lived genets” which may only achieve successful sexual reproduction very infrequently over the course of the colonies’ multi-decadal lifespan. There is evidence that many species, especially those in deep and temperate waters, are long-lived (Andrews et al. 2002; Risk et al. 2002) and therefore may possess similar reproductive life history characteristics.
Table 1
FAMILY |
SPECIES |
SEX |
REPRODUCTIVE STRATEGY |
Citation |
Coenothecalia |
Heliopora coerulea |
separate |
surface brooding (6-8 days) below inflated tentacles forming brood-like chamber |
Babcock 1990 |
SOFT CORALS |
|
|
|
|
Alcyoniidae |
Alcyonium acaule |
- |
brooding |
Hartnoll 1975 |
Alcyoniidae |
Alcyonium aspiculatum |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Alcyonium digitatum |
separate |
broadcast spawning |
Hartnoll 1975 |
Alcyoniidae |
Alcyonium hibernicum |
separate |
brooding |
Hartnoll 1975 |
Alcyoniidae |
Alcyonium molle |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Alcyonium palmatum |
- |
brooding |
Hartnoll 1975 |
Alcyoniidae |
Alcyonium siderium |
separate |
brooding |
Feldman 1970 |
Alcyoniidae |
Anthomastis ritteri |
separate |
brooding (in siphonozooids) |
Cordes et al. 2001 |
Alcyoniidae |
Capnella gaboensis |
separate |
external surface brooding |
Farrant 1986 |
Alcyoniidae |
Cladiella pachyclados |
separate |
broadcast spawning |
Shinkarenko 1981 |
Alcyoniidae |
Lobophytum compactum |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Lobophytum crassum |
separate |
broadcast spawning |
Yamazato et al. 1983 & Uehara et al. 1987,Coll et al. 1995 |
Alcyoniidae |
Lobophytum hirsutum |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Lobophytum microlobatum |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Lobophytum pauciflorum |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Lobophytum planum |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Lobophytum sarcophoides |
separate |
broadcast spawning |
Dai 1989 |
Alcyoniidae |
Minabea robusta |
- |
- |
Utinomi and Imahara 1976 |
Alcyoniidae |
Paraerythropodium fulvum fulvum |
separate |
external surface brooders (entagle eggs in surface mucus) |
Benayahu and Loya 1983 |
Alcyoniidae |
Sarcophyton crassocaule |
separate |
broadcast spawning |
Dai 1989 |
Alcyoniidae |
Sarcophyton cf. ehrenbergi |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Sarcophyton glaucum |
separate |
broadcast spawning |
Benayahu and Loya 1986 |
Alcyoniidae |
Sarcophyton trocheliophorum |
separate |
broadcast spawning |
Shinkarenko 1981 |
Alcyoniidae |
Sinularia conferta |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Sinularia cruciata |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Sinularia deformis |
separate |
broadcast spawning |
Alino and Coll 1989 |
|
Sinularia dura |
|
|
Pratt 1903 |
|
Sinularia humesi |
separate |
broadcast spawning |
Benayahu et al. 1990 |
Alcyoniidae |
Sinularia liptoclados |
separate |
broadcast spawning |
Benayahu et al. 1990 |
Alcyoniidae |
Sinularia lochmodes |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Sinularia mayi |
separate |
broadcast spawning |
Benayahu et al. 1990 |
Alcyoniidae |
Sinularia polydactyla |
separate |
broadcast spawning |
Alino and Coll 1989 |
Alcyoniidae |
Sinularia rigida |
separate |
broadcast spawning |
Benayahu |
Clavulariidae |
Clavularia crassa |
|
surface brooding |
Weinbery 1986 |
Clavulariidae |
Clavularia hamra |
separate |
external surface brooders (entagle eggs in surface mucus) |
Benayahu 1989 |
Clavulariidae |
Clavularia inflata |
- |
surface brooding |
Alino and Coll 1989 |
Clavulariidae |
Pachyclavularia violacea |
- |
surface brooding |
Alino and Coll 1989 |
Cornulariidae |
Cornularia komaii |
- |
external surface brooding (spawned eggs retained in space made by closed tentacles) |
Suzuki 1971 |
Cornulariidae |
Cornularia sagamiensis |
- |
external surface brooding (spawned eggs retained in space made by closed tentacles) |
Suzuki 1971 |
Neptheidae |
Dendronepthya hemprichi |
separate |
broadcast spawning |
Dahan & Benayahu 1997 |
Neptheidae |
Dendronepthya sinaiensis |
- |
- |
Benayahu 1997 |
Neptheidae |
Litophyton arboreum |
separate |
brooding |
Benayahu et al. 1990 |
Xeniidae |
Anthelia formosa |
|
brooding |
Utinomi 1950 |
Xeniidae |
Anthelia glauca |
separate |
external & internal brooding |
Kruger et al. 1998 |
Xeniidae |
Efflatounaria sp. |
separate |
external surface brooders |
Dinesen 1985 |
Xeniidae |
Heteroxenia coheni |
hermaphroditic |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Heteroxenia elizabethae |
hermaphroditic & gonochoristic, dimorphic |
brooding |
Gohar 1940; Shinkarenko 1981 |
Xeniidae |
Heteroxenia fuscescens |
hermaphrotic, dimorphic |
external & internal brooding |
Benayahu et al. 1989 |
Xeniidae |
Heteroxenia ghardaqensis |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Sympodium caeruleum |
separate |
internal brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia biseriata |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia blumi |
separate |
brooding |
Gohar 1940 |
Xeniidae |
Xenia farauensis |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia garciae |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia hicksoni |
separate |
brooding |
Gohar 1940 |
Xeniidae |
Xenia impulsatilla |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia macrospiculata |
separate |
external brooding |
Achituv et al. 1992 & Benayahu & Loya 1984 |
Xeniidae |
Xenia membranacea |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia obscuronata |
separate |
brooding |
Benayahu et al. 1989 |
Xeniidae |
Xenia umbellata |
separate |
internal brooding |
Benayahu et al. 1989 |
GORGONIANS |
|
|
|
|
Briareidae |
Briarium asbestinum |
separate |
surface brooding (3-5 days) entrapped in mucus sheet |
Brazeau & Lasker 1990 |
Briareidae |
Briarium stechei |
separate |
surface brooding |
Alino and Coll 1989 |
Corallidae |
Corallium rubrum |
separate |
brooder |
Vighi 1970 |
Gorgoniidae |
Eunicella singularis |
- |
brooder |
Weinberg and Weinberg 1979 |
Gorgoniidae |
Eunicella stricta |
separate |
brooder |
Theodor 1967 |
Gorgoniidae |
Leptogorgia virgulata |
separate |
broadcast spawning |
Gotelli 1988 |
Gorgoniidae |
Pseudopterogorgia bipinnata |
separate |
brooder |
Kinzie 1970 |
Gorgoniidae |
Pseudopterogorgia elisabethae |
separate |
external brooding |
Gutierrez-Rodriguez and Lasker 2004 |
Isidiidae |
Acanella arbuscula |
separate |
brooder? |
Lawson 1990 |
Plexauridae |
Eunicea clavigera |
separate |
brooder |
Kinzie 1970 |
Plexauridae |
Muricea californica |
separate |
brooder |
Grigg 1979 |
Plexauridae |
Muricea fructicosa |
separate |
brooder |
Grigg 1979 |
Plexauridae |
Paramuricea clavata |
separate |
surface brooding |
Coma et al. 1995 |
Plexauridae |
Plexaura flexuosa |
separate |
broadcast spawning |
Beiring and Lasker 2000 |
Plexauridae |
Plexaura homomalla |
separate |
broadcast spawning |
Martin 1982 |
Plexauridae |
Plexaura kuna |
separate |
broadcast spawning |
Lasker |
Plexauridae |
Pseudoplexaura porosa |
separate |
broadcast spawning |
Kapela and Lasker 1989 |
Primnoidae |
Ainigmaptilon antarcticum |
separate |
? May be brooder (lrg egg size) |
Orejas et al. 2002 |
Primnoidae |
Fannyella rossii |
- |
brooder |
Orejas et al. 2002 |
Primnoidae |
Thouarella variabilis |
separate |
internal brooding |
Brito et al. 1995 |
PENNATULACEANS |
|
|
|
|
Kophobelemnidae |
Kophobelemnon stelliferum |
separate |
probably broadcast spawning |
Rice et al. 1992 |
Pennatulidae |
Ptilosarcus guerneyi |
separate |
broadcast spawning |
Chia and Crawford 1973 |
Pennatulidae |
Pennatula aculeata |
separate |
broadcast spawning |
Eckelbarger et al. 1998 |
Virgularidae |
Virgularia juncea |
- |
- |
Soong 2005 |
Veretillidae |
Cavernularia obesa |
- |
- |
Mori and Tanase 1971 |
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