Brachyura, Grapsidae) That Inhabit the Oyster Reefs of Western Taiwan
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Crustaceana 90 (14) 1699-1714 POPULATION STRUCTURE AND FECUNDITY OF TWO SPECIES OF GRAPSID CRABS (BRACHYURA, GRAPSIDAE) THAT INHABIT THE OYSTER REEFS OF WESTERN TAIWAN BY KUN-NENG CHEN1), JUNG-TING HSU2) and YIH-TSONG UENG3,4) 1) Department of Electrical Engineering, Kun-Shan University, Tainan City, Taiwan 2) Tainan Hydraulics Laboratory, National Cheng Kung University, Tainan City, Taiwan 3) Department of Environmental Engineering, Kun-Shan University, 195 Kunda Road, Yongkang District, Tainan City, 71070, Taiwan ABSTRACT This study on carcinological aspects of oyster farming was carried out at the coast of Yunlin County, Taiwan, for a period of 25 months from July 2012 to July 2014. The space formed by the clustering of the oyster shells provides a habitat for at least eight species of crabs. Nanosesarma minutum was the dominant species (73.7% or n = 18 753) with an average of 59.5 ± 30.6 individuals/string/month, while the average monthly rate of ovigerous females was 12.6 ± 13.0%, and the average number of eggs per berried female was 1510.8 ± 1018.7 eggs/individual. Hemigrapsus penicillatus was the second dominant species (23.9% or n = 6074) with an average of 17.4 ± 17.1 individuals/string/month; here, the average monthly rate of ovigerous female crabs was 4.6 ± 9.0, and the average number of eggs of a berried female was 3073.4 ± 2167.5 eggs/individual. RÉSUMÉ Cette étude sur les aspects carcinologiques de la culture d’huître a été effectuée sur la côte du conté de Yunlin, Taiwan, pendant une période de 25 mois de juillet 2012 à juillet 2014. L’espace formé par l’amoncellement des coquilles d’huître a fourni un habitat pour au moins huit espèces de crabe. Nanosesarma minutum a été l’espèce dominante (73,7% ou n = 18 753) avec une moyenne de 59,5 ± 30,6 individus/corde/mois, tandis que le taux moyen mensuel de femelles ovigères était de 12,6 ± 13,0%, et la moyenne d’œufs par femelle était de 1510,8 ± 1018,7 d’œufs/individu. Hemigrapsus penicillatus a été la deuxième espèce dominante (23,9% ou n = 6074) avec une moyenne de 17,4 ± 17,1 individus/corde/mois ; une moyenne mensuelle de femelles ovigères de 4,6 ± 9,0%, et une moyenne de nombre d’œufs par femelle ovigère de 3073,4 ± 2167,5 œufs/individu. 4) Corresponding author; e-mail: [email protected] © Koninklijke Brill NV, Leiden, 2017Downloaded DOI 10.1163/15685403-00003734 from Brill.com10/02/2021 03:13:19AM via free access 1700 KUN-NENG CHEN, JUNG-TING HSU & YIH-TSONG UENG INTRODUCTION Oyster reef communities can provide habitats and change the food webs of aquatic animals in lagoons, estuaries, and intertidal zones. These communities demonstrate the enormous biodiversity of coastal wetlands (Dame et al., 1984, 2001; Newell, 1988; Grabowski & Powers, 2004). Oyster reefs also provide numerous ecosystem services including the production of oysters, maintenance of estuarine water alkalinity, shoreline stabilization, and provision of habitats for fishes and crustaceans (Meyer et al., 1997; Piazza et al., 2005; Tolley & Volety, 2005; Waldbusser & Salisbury, 2014). Crustacea are the most abundant aquatic organisms and are represented by numerous species, exhibiting diverse behaviours and habitats. This diversity in the various groups of crustaceans is a result of their life patterns and reproductive strategies (Sastry, 1983; El-Serehy et al., 2015). Some studies have investigated their size composition, growth, sexual maturity, and life history (Kwei, 1978; Fukui, 1988; Carsen et al., 1996; Doi et al., 2007). Gehrels et al. (2016) reported upon the interaction in predator-prey relationships between mud crabs, Dyspanopeus sayi (Smith, 1869), and green crabs, Carcinus maenas (Linnaeus, 1758). Brousseau & McSweeney (2016) investigated the fecundity and maturation rates of Asian shore crabs, Hemigrapsus sanguineus (De Haan, 1835) and Atlantic mud crabs, Panopeus herbstii H. Milne Edwards, 1834. Although oyster reefs constitute the most threatened coastal habitat because of the regular harvesting of oysters, numerous estuarine and tidal species still provide essential habitats and food resources (Yeager & Layman, 2011). Fecundity is closely related to two biological factors in most crustaceans; it is inversely proportional to egg mass weight and directly proportional to female size (Reid & Corey, 1991; García-Guerrero, 2004; Leme, 2004; Terossi et al., 2010). Fecundity is a key factor in the persistence of a fishery stock and is used to evaluate the status of a population, because it directly affects the recruitment of species in estuarine and marine environments (Begg & Waldman, 1999; Rodrigues et al., 2011). Oyster farming in Taiwan takes approximately 6 months to 1 year from seed to harvest. To illustrate this practice, a common way to farm oysters in Taiwan is by moving the oyster larvae to suitable areas located at coasts and estuaries, using floating frames. The advantage of this method is, that the oysters thus are able to take in nutrients during the whole day, as they are constantly suspended in sea water: they do not fall dry during low tides. Obviously, in this way the oysters can grow faster than when using other farming methods. However, this cultivation method requires harvesting the oysters before the invasion of typhoons, which limits the growth period to around six months a year. By contrast, another prevalent method is to suspend the oyster larvae on the lines on a fixed frame in a Downloaded from Brill.com10/02/2021 03:13:19AM via free access POPULATION STRUCTURE AND FECUNDITY OF TWO GRAPSID CRABS 1701 lagoon. Due to the cycles of the tides, the oysters will be out of the seawater and be exposed to air twice a day, which results in a slower growth rate compared to the previous method. On the other hand, without the threats of typhoons, the oyster farms in lagoons can extend the cultivation period to one year. During the annual oyster harvest, fishermen remove larger crabs. This man- agement strategy directly changes the life cycle factors of those crabs and indi- rectly affects their populations. To learn the sum of those effects, investigating body size distribution and age composition have provided information, including a forecast of the growth and additional strength of young organisms, as well as pos- sible seasonal variations (Pianka, 1974; Hartnoll & Bryant, 1990; Hsueh, 1991; Czerniejewski & Wawrzyniak, 2006). After the release of eggs, a series of changes in environmental factors can result in a deviation from the normal sex ratio of 1 : 1 (Fisher, 1930; Wilson & Pianka, 1963; Leigh, 1970; Josileen, 2013). Statistical studies have revealed that, of the various relationships, carapace width and fecundity are better indices for the estimation of the reproductive potential of crabs than is body weight (Josileen, 2013). The west coast of Taiwan is an important area for oyster farms, which supply oyster larvae. Oyster farming is a vital economic activity there, and the main species in the oyster farms is Crassostrea angulata (Lamarck, 1819) (cf. Wang et al., 2010; Vaschenko et al., 2013; Hsiao et al., 2016). Oyster reefs are one of the most depleted and degraded marine habitats worldwide (Geraldi et al., 2013), while the income from fisheries is often crucial for rural communities, and the decline and depletion of oyster reefs thus can result in loss of income. This means that sufficient knowledge of the ecosystems built by oyster reefs and farms is important for a sustainable farming strategy, and the crabs that live among the oysters constitute a major component in those ecosystems. Habitat loss is another critical factor contributing to the reduction of fishery re- sources. The Formosa Plastics Corporation Sixth Naphtha Cracker was established in 1993 at the Jhuoshuei estuary. Before its construction, the area of oyster farms south of the construction area with a water depth < 2 m at low tide was 1212 ha. By 2008, this area was reduced to 1084 ha, with an average annual decrease of 10.7 ha. In addition to reducing the oyster farming area, the habitat for fishery resources was diminished (Chen et al., 2016). However, few studies have analysed the popu- lations and biological patterns of crab species inhabiting oyster farms (Ijeomah et al., 2013). In this study, we investigated the biodiversity, community, and fecundity of crab populations in an oyster culture zone of western Taiwan. Downloaded from Brill.com10/02/2021 03:13:19AM via free access 1702 KUN-NENG CHEN, JUNG-TING HSU & YIH-TSONG UENG MATERIAL AND METHODS The study site of our research was located in the intertidal zone (23°4248N 120°0925E) of Taisi Township, Yunlin County, Taiwan. We set two oyster racks in the study site to simulate oyster farming habitats, and the distance between them was approximately 1 km. We hung ten batches of oyster-shell-strings on the oyster racks with the help of local fishermen once every two weeks, from 2 March 2012 to 12 July 2014. Each oyster-shell-string was composed of a 5-m long plastic rope, tied with 18 empty oyster shells that were spaced evenly along the rope. At high tide, the strings were immersed in the water to a depth of approximately 1-2 m; during low tide, they were exposed for approximately 2 hours/day. The settlement of oyster larvae was possible for 16-20 hours/day. The oyster larvae started to grow after they settled in the empty shells. Hundreds of new live oysters developed on each oyster-shell-string, thereby forming microhabitats for small crabs, barnacles, and mussels. Sample collection From July 2012 to July 2014, we procured the crabs that were gathered in oyster reef through collecting the oyster-shell-strings. These oyster-shell-strings had been farmed over a period of 6-12 months, except for the first 3 times.