Gametogenesis and Inter-Annual Variability in the Spawning Pattern of Acropora Hyacinthus in Northwestern Philippines Emmeline A
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Zoological Studies 57: 46 (2018) doi:10.6620/ZS.2018.57-46 Open Access Gametogenesis and Inter-annual Variability in the Spawning Pattern of Acropora hyacinthus in Northwestern Philippines Emmeline A. Jamodiong1,*, Elizaldy A. Maboloc1,2, Ronald D. Villanueva1, and Patrick C. Cabaitan1 1The Marine Science Institute, College of Science, University of the Philippines, Diliman, 1101 Quezon City, Philippines. E-mail: [email protected] (Maboloc); [email protected] (Villanueva); [email protected] (Cabaitan) 2Present address: Department of Ocean Science, Hong Kong University of Science and Technology, Clear Water Bay, Kowloon, Hong Kong, SAR, China (Received 29 March 2018; Accepted 16 August 2018; Published 21 September 2018; Communicated by Yoko Nozawa) Citation: Jamodiong EA, Maboloc EA, Villanueva RD, Cabaitan PC. 2018. Gametogenesis and inter-annual variability in the spawning pattern of Acropora hyacinthus in northwestern Philippines. Zool Stud 57:46. doi:10.6620/ZS.2018.57-46. Emmeline A. Jamodiong, Elizaldy A. Maboloc, Ronald D. Villanueva, and Patrick C. Cabaitan (2018) Acropora hyacinthus is a fast-growing tabular coral that dominates the shallow water coral assemblage in the Magsaysay reef at the Bolinao-Anda Reef Complex (BARC), northwestern Philippines. The timing of gamete development was investigated for A. hyacinthus through dissection and histological analyses of coral fragments that were collected monthly from February 2014 to April 2015 from the 22 tagged colonies. The spawning time was identified by the presence of oocytes in the sampled A. hyacinthus colonies through rapid sampling from January to March 2014, 2015 and 2016. Results show that A. hyacinthus is a hermaphroditic broadcast spawning coral with an annual gametogenic cycle. Acropora hyacinthus exhibits an extended release of gametes across 2 to 3 months, from February to April. Major release of gametes occurred in March. Two types of extended spawning patterns that are unique in this region were observed in this species (i.e., asynchronous spawning amongst colonies and split spawning of individual colonies). This study contributes to the increasing knowledge on the coral reproductive strategies in northwestern Philippines and provides information on availability of coral materials for coral reef restoration efforts and management. Key words: Corals, Acropora, Reproduction, Spawning pattern, Philippines. BACKGROUND 1998; Mendes and Woodley 2002; Guest et al. 2005; Lueg et al. 2012; Nozawa 2012). However, Coral reefs are among the most diverse with the changing environments brought about and economically important marine ecosystems by global climate change, corals are exposed to that support various biological components and temperature anomalies that sub-lethally impact provide resources and livelihood to people (Chou coral reproduction (Ward et al. 2000). Thermal et al. 2002). The persistence and maintenance stress lessens reproductive output of reef corals of coral communities are dependent on the (Jones and Berkelmans 2011). With the prediction presence of sexually reproducing coral individuals that thermal stress events will frequently reoccur, (Kaniewska et al. 2015). Sexual reproduction in it is imminent to understand how corals maintain corals has been demonstrated to be significantly their fecundity (Victor et al. 2009). Corals may have influenced by annual temperature changes (Dai developed different spawning strategies that will et al. 1993; Van Veghel 1994; Hagman et al. allow propagules to persist and survive. *Correspondence: E-mail: [email protected] © 2018 Academia Sinica, Taiwan 1 Zoological Studies 57: 46 (2018) page 2 of 10 Reproductive strategies of Acropora species This study aimed to elucidate the reproductive have been studied in different geographical areas biology of A. hyacinthus, specifically to determine (Babcock et al. 1986; Baird et al. 2002; Guest gametogenesis, spawning synchrony among et al. 2005; Carroll et al. 2006; Mangubhai and colonies, and spawning time variability of A. Harrison 2008; Kenyon 2008; Hanafy et al. 2010; hyacinthus in the Magsaysay reef in northwestern Bouwmeester et al. 2015; Sola et al. 2016). Philippines. Insights on coral spawning patterns These acroporid species usually participate in and knowledge on the availability of populations a synchronous mass spawning event (Willis et that may settle are essential in understanding al. 1985; Hanafy et al. 2010; Bouwmeester and how coral communities may be maintained Berumen 2015; Chelliah et al. 2015; Jamodiong (Bouwmeester et al. 2015). et al. 2017), despite extended spawning characteristics (Baird et al. 2009). Extended spawning occurs when coral colonies spawn over a MATERIALS AND METHODS series of consecutive lunar cycles or months (Baird et al. 2009). In northwestern Philippines, broadcast Rapid sampling and sample collection spawning of corals, represented by species from the families Faviidae and Acroporidae, occur Fragments of Acropora hyacinthus were March to May (Bermas et al. 1992; Vicentuan sampled from 22 tagged colonies (147 ± 48.81 cm et al. 2008). A detailed study on Favites species in diameter) located at 2 to 3 m depth on the reproduction showed brief synchronous spawning Magsaysay reef in the Bolinao-Anda Reef Complex (Maboloc et al. 2016), which supports the (BARC), northwestern Philippines (16°18'38.3"N, hypothesis that extended spawning may be a 120°01'46.8"E) (Fig. 1). The tagged colonies were characteristic of Acropora species. However, sampled monthly from February 2014 to April 2015 comprehensive studies on the reproductive to determine the gametogenesis and fecundity. timing of Acropora corals in the Philippines are However, because of logistical constraints, no very scarce. Depending on geographic location, samples were collected in September 2014. A the length of spawning durations varies between more intensive sampling, approximately every 1 genus and species (Baird et al. 2009). to 2 weeks, were conducted during the expected Acropora hyacinthus (Dana 1846) is a spawning season to determine approximate fast-growing tabular coral that is widespread in the spawning time, from February to April in 2014 Indo-Pacific region (Veron 2000). They dominate and 2015 and from January to March in 2016. the Magsaysay reef (c. 60% of the coral cover, During each sampling, two to three fragments (2 to Jamodiong personal observation) in the Bolinao- 3 cm in length) per colony were collected and the Anda Reef Complex (BARC) of the Philippines. fragments were carefully checked for the presence This area is regarded as an important source of or absence of oocytes underwater (after Baird et coral larvae for the neighboring reefs because al. 2002). Samples were collected between the of its high coral coverage (coral cover 80%, center and marginal area of the coral colonies. Cabaitan unpublished data) and Acropora corals’ When oocytes were present, the color of oocytes synchronous mass spawning event (Jamodiong et was recorded. An HOBO Water Temperature Pro al. 2017). However, despite the dominance of A. v2 Data Logger (Onset Computer Corp., Bourne, hyacinthus, no detailed study on its reproduction MA) was deployed in the sampling site from has been done in the Philippines. Studying the November 2014 to November 2016 to record reproductive behavior of this particular coral seawater temperature. species, which dominates the reef assemblage, will provide knowledge on coral spawning along Histological examinations this area and implications on the reproductive phenology of corals. Furthermore, the majority of The collected coral fragments were fixed in the studies previously conducted were from the Zenkers solution for 24 hours, rinsed with running randomly sampled colonies per species (Harrison seawater for another 24 hours and decalcified by et al. 1984; Hanafy et al. 2010; Bouwmeester soaking the fragments to 10% buffered HCl for et al. 2015; Chelliah et al. 2015; Sola et al. 2 to 3 days depending on the dissolution of the 2016). Repeated sampling of the same colonies coral skeleton (Maboloc et al. 2016). Some of the provides an opportunity to see the changes in decalcified tissue samples (n = 10) were used the reproductive timing throughout the months. to determine the polyp fecundity and some were © 2018 Academia Sinica, Taiwan Zoological Studies 57: 46 (2018) page 3 of 10 used to examine the gametogenesis (n = 10). For (from 100 x to 400 x magnifications). Oocyte fecundity, decalcified fragments were dissected development was recorded in different stages: to determine the mean oocyte count per polyp for stage I, when there was enlargement of the each individual colony. The tissue sample was interstitial cells with large nuclei in the mesoglea placed on a clean petri dish and ten polyps per of the mesentery; stage II, when cytoplasm started colony were haphazardly selected for examination. to accumulate around the nuclei; stage III when Each polyp was isolated from the tissue sample cytoplasm was larger such that the nucleus was and dissected using dissecting needles under centrally located; and stage IV for the mature level a stereomicroscope. In each polyp, the number and when the oocyte achieved a full size with of oocytes was counted and the minimum and proliferation of yolk granules and indented nucleus. maximum diameters of 10 haphazardly selected These stages were based on the criteria used by oocytes were photographed and measured using Szmant-Froelich et al. (1985). The developmental the MOTIC Images plus 2.0