An Update on Modes and Timing of Gamete and Planula Release in Hawaiian Scleractinian Corals with Implications for Conservation and Managementl
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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by ScholarSpace at University of Hawai'i at Manoa An Update on Modes and Timing of Gamete and Planula Release in Hawaiian Scleractinian Corals with Implications for Conservation and Managementl Steven P. Kolinski and Evelyn F. Cox 2 Abstract: Reproductive data for 24 of the 50 plus species of scleractinian corals in Hawai'i are available. A majority ofspecies (75 %) are broadcast spawners, just over half (58%) of which are hermaphrodites. Peak reproduction of Hawaiian corals occurs during summer months, although reproduction continues year round for some brooders. Timing, duration, mode, and location ofreproductive processes have implications for disturbance management, assessment, and con servation of reef corals. THE MINIMIZATION OF human impacts on (Harrison and Wallace 1990, Levitan 1995, coral reef dynamics is critical to maintaining Morgan 1995, Coma et al. 1998). Individual coral reef diversity and reef ecosystem func stages may differ in their exposure, vulnera tions. Human impacts can be direct and/or bility, and susceptibility to various impacts indirect, affecting reef organisms during one (Richmond 1993, 1997). or more of a variety of life history stages Understanding where and when repro (Brown and Howard 1985, Richmond 1993, ductive processes take place allows for a pre 1997, Raimondi and Reed 1996). Sexual re cautionary approach to minimizing human production in scleractinian corals involves a interference with coral reproductive dynam number of discrete stages that occur in dif ics. In this paper we summarize available data ferent zones of the marine environment. for reproduction in Hawaiian corals in an ef These fundamental stages include gamete fort to provide information pertinent to miti development (internal to the organism), ga gating human impacts on coral reproduction mete release (internal coordination leading and population dynamics. to external release), fertilization (internal, or external in the water column, at the water MATERIALS AND METHODS surface, or on the benthos), embryonic and larval development (internal, or external in the Data summarized in this report resulted from water column or at the water surface), dis both direct and indirect observations of ga persal (external in the water column and/or at mete formation and release in laboratory and/ the water surface), settlement, and metamor or field settings. Information for the majority phosis (external on a substrate) (Harrison and of species came from studies conducted at Wallace 1990, Richmond 1993, 1997). Syn the Hawai'i Institute ofMarine Biology and a chronization of various reproductive stages review of the literature (including reviews by within populations is fundamental to increas Fadlallah 1983, Richmond and Hunter 1990, ing the likelihood of reproductive success and Richmond 1997). Although Hawaiian coral taxonomy is currently in a state of flux, Maragos (1995) was used as a guide (see 1 This is contribution no. 1128 of the Hawai'i Insti Maragos 1977 for reported synonyms). tute of Marine Biology. Manuscript accepted 22 March 2002. RES.ULTS. '2 Hawiii'i rnsnriite-ofMarineBiology,-P.O~ Box 1H6;-' Coconut Island, Kane'ohe, Hawai'i 96744. Information on modes and timing of gamete and planula release for 24 of roughly 50 Ha Pacific Science (2003), vol. 57, no. 1:17-27 waiian scleractinian coral species (Maragos © 2003 by University ofHawai'i Press 1995) is provided in Table 1. The data cover All rights reserved the majority of the most common inshore 17 TABLE 1 Reproductive Characteristics of Hawaiian Scleractinian Coral Species Listed by Maragos (1995) Spawning Location of Species Sexo Modeb Seasonc Moon Phase Times Fertilization Reference I ACROPORIDAE Acropora 0't~erea (Dana, 1846) H *S *Lt. Sp-Sr *June-Aug. Kenyon (1992, 1994) A. humilis (Dana, 1846) H *S Lt. Sp *lst qtr *June *Surface Kenyon (1992, 1994) A. paniculatJ Verrill, 1902 A. valida (qana, 1846) H *S Sr *New-4th qtr? *July-Aug. *Surface Kenyon (1992, 1994) Montipora c4pitata (Dana, 1846) H S Lt. Sp-Sr New-1st qtr May-Sept., Surface Heyward (1986), 20:45-22:30 Hodgson (1988), Hunter (1988b), Cox (1991) M. dilatata Studer, 1901 HS Sr New, July-Aug., Surface Heyward (1986), Full-3rd qtr 20:30-21:45 Hodgson (1988), Hunter (1988b) M. flabellatal Studer, 1901 HS Sr-*F July-*Sept., Surface Heyward (1986), this 21:05-21:50 study M. patula Verrill, 1864 HS Sr New-1st qtr, July-Sept., Surface Hodgson (1988), this I Full-3rd qtr 22:05-23:10 study M. studeri Viaughan, 1907 H S Sr New-1st qtr, July-Sept., Surface Heyward (1986), Full-3rd qtr 22:23-23:00 Mate (1998) M. tuberculofa (Lamarck, 1816) M. verrilli Viaughan, 1907 H *S Sr Full-3rd qtr July *Surface Heyward (1986) AGARICIIDJ.\E Gardineroserys planulata (Dana, 1846) Leptoseris hawaiiensis Vaughan, 1907 L. incrustanslcQuelch, 1886) L. mycetosero~des Wells, 1954 L. papyracea I(Dana, 1846) L. scabra Val1ghan, 1907 L. tubulifera IVaughan, 1907 Pavona duerqeni Vaughan, 1907 GS *Water column G. Hodgson, unpubL data P. varians Vhrill, 1864 GS Lt. Sp Full-3rd qtr June, *Water column/ G. Hodgson, unpubL 19:05-20:15 surface data; Mate (1998); this study P. maldivensis (Gardiner, 1905) BALANOPH¥LLIDAE Balanophyllia Isp. cf. affinis (Semper, 1872) B. hawaiiensJ Vaughan, 1907 DENDROPIiYLLIDAE Tubastrea cocqinea Lesson, 1831 B Lt. Sp-W All June-Jan., Polyps Edmondson (1929, diurnal and 1946), Jokie! et a!. nocturnal (1985); A. M. Tarrant and S.P.K., unpub!. data FAVIIDAE Cyphastrea oc~llina (Dana, 1846) HB Yr All Polyps Edmondson (1929, 1946), Stimson (1978), Jokiel et al. (1985), Wright (1986) Leptastrea bo~ae (Milne-Edwards & GS 'Water column G. Hodgson, unpub!. Haime, 1~50) data L. purpurea Ipana, 1846 GS 'Water column G. Hodgson, unpub!. data FUNGIIDAE Cycloseris ten~is (Dana, 1846) C. vaughani (Boschma, 1923) Diaseris distoi,ta (Michelin, 1843) Fungia scutatia Lamarck, 1801 G S Sr-F Full-3rd qtr June-Nov., Water column Krupp (1983), 17:00-19:00 Schwarz et a!. (1999) POCILLOPORIDAE Pocillopora da'rzicornis (Linnaeus, 1758) HB Yr All Diurnal and Polyps Edmondson (1946), nocturnal Reed (1971), Harrigan (1972), Stimson (1978), Richmond and Jokie! (1984), Jokiel (1985) P. eydouxi Milne-Edwards & Haime, 1860 P. ligulata D~a, 1846 P. meandrinal Dana, 1846 H S Sp Full-3rd qtr Apr.-May, Water column Stimson (1978), 07:20-08:15 Fiene-Severns (1998); S.P.K., A. M. Tarrant, and E.F.C., unpub!. data P. molokensisVaughan, 1907 Reproduction in Hawaiian Scleractinian Corals . Kolinski and Cox 21 I2a Spawners 14 • Brooders 12 CJ) (l) '(5 10 (l) a. (J) 8 0+- 0 L.. (l) 6 .0 E ::J 4 Z 2 o Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month FIGURE 1. Known number of scleractinian coral species planulating and spawning each month in Hawaiian waters. shallow-water species but are far from com The classification of Psammacora stellata plete. Six (25%) of the 24 species brood as a brooder in this study is tentative and in larvae, 10 (42%) are broadcast-spawning her need of verification. It is based on the tank maphrodites, and eight (33 %) are gonochoric collection of swimming coral larvae 45 min broadcast spawners. Seventeen species (71 %) after previous examination of secluded display peak reproductive development and colonies. We made direct observations on gamete or planula release during summer, at mode and timing of reproductive release of least five species (21 %) extend release into Montipora patula colonies over multiple re fall, three species (13 %) continue release productive seasons, as well as observations into winter, and nine species (38%) begin or confirming the reproductive activities of M. continue reproductive release in the spring capitata, Pavona varians, Tubastrea coccinea, (Figure 1). Fertilization in the Hawaiian Cyphastrea ocellina, Fungia scutaria, Pocillopora Acroporidae is overwhelmingly a surface damicornis, P. meandrina, Porites compressa, and phenomenon. In brooding species syngamy is P. lobata. Observations of Montipora fiabellata presumed to take place within polyps. Zygote spawning were limited, and species identifi formation in gonochoric broadcast spawners cation of the morphological variant used in is likely to occur within the water column, this study is in need of verification and con but surface fertilization may also take place. gruence of multiple coral taxonomists. Although specific timing of gamete release . ·app-e~rs-discrete·~arrd-predictable, spawning DISCUSSION for most species occurs over a number ofdays and months, indicating variability and possi Any assessment of potential impacts to ble plasticity in seasonal synchrony of spawn Hawaiian reef corals must take into consider ing events. ation local population structure and timing 22 PACIFIC SCIENCE· January 2003 and location ofall stages ofcoral reproductive Raimondi and Morse 2000). Appropriate cues cycles, including recruitment and growth. for coral settlement include physical stimuli Reproductive stages for each species are sus and, in many cases, specific biochemical in ceptible to different impacts at particular ducer molecules (Morse et al. 1996, Heyward times of the year, corresponding to spatial and Negri 1999, Raimondi and Morse 2000). and temporal variation of reproductive activ Factors such as high sediment levels have ities. Because of the open nature of the been shown to inhibit coral larval settlement marine system, the ultimate effect of any lo and persistence (Hodgson 1990, Babcock and calized impact is also likely to be incurred by Davies 1991, Hunte and Wittenberg 1992, one or more sink reef areas within the region Te 1992, Gilmour 1999). Herbicides, pesti of the metapopulation, even those remote cides' heavy metals, oils and dispersants, from the reproductive source area (Keough human drug by-products, and other derived and Black 1996, Raimondi and Reed 1996). chemical compounds may act as waterborne Negative impacts to one or multiple stages toxicants that have inhibitory effects on coral of the reproductive process are likely to have recruitment as a result of reactions with a cascading effect on the success of the re larvae, settlement inducer molecules, and/or maining stages.