Characteristics of Scylla Spp. (Decapoda: Portunidae) and Their Mangrove Forest Habitat in Ngaremeduu Bay, Republic of Palau1
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Characteristics of Scylla spp. (Decapoda: Portunidae) and Their Mangrove Forest Habitat in Ngaremeduu Bay, Republic of Palau1 Katherine C. Ewel,2,3 Stacy Rowe,4 Blake McNaughton,5 and Kimberly M. Bonine6 Abstract: Three species of mangrove crabs (Scylla spp.) were captured in live traps in Ngaremeduu Bay on the island of Babeldaob, Republic of Palau. Most were S. serrata, but one individual each of S. olivacea and S. paramamosain was also trapped, establishing existence of a biogeographic gradient in mangrove crab species diversity across the Micronesian archipelago. Species composition of mangrove trees along transects around the bay and along the three major tributaries was similar to that of other Micronesian islands, although trees are smaller in Palau. For 17 months in 1999–2000, crabs were trapped in the bay and captured by hand along the transects; they were trapped again for 1 month in 2004. Characteristics of the crabs and of burrows encountered along the transects suggested that only S. serrata was captured in 1999–2000 and that pop ulation density of this species was 40 crabs ha-1. Carapace widths for the 159 crabs captured during the entire study did not differ significantly over the 4-yr span, and averaged 153 mm for males and 137 mm for females. However, aver age carapace widths for the largest quartile of crabs declined significantly from 174 mm to 171 mm across the study period. Catch per unit effort was 0.28 crab per trap night in 1999–2000 and 0.45 in 2004. Although large crabs are still available in Ngaremeduu Bay, current regulations may not be sufficient to keep populations from decreasing gradually in size, especially in the face of increasing harvest pressure on the island of Babeldaob. Mangrove crabs (Scylla spp. [also called the westernmost country in the Pacific island mud crabs]) are an important component of group of Micronesia, mangrove crabs are local economies and mangrove forest food regularly trapped in Ngaremeduu Bay, a webs throughout the Indo-Pacific. They large, shallow, mangrove-lined estuary on often exceed 200 mm in carapace width and the island of Babeldaob, at 333 km2 the sec are commonly harvested as a source of food ond largest island in Micronesia (Figure 1). and income throughout their range. In the In spite of the popularity of these crabs as Republic of Palau (7° 29 0 N, 134° 31 0 E), food items, little is known about them in 1 Participation of K.M.B. was made possible by a School of Forest Resources and Conservation, University grant from the John D. and Catherine T. MacArthur of Florida, Gainesville, Florida 32611 (e-mail: kewel@ Foundation (no. 97-49885A-WER; B. Singer, R. Naylor, ufl.edu). K. Ewel, and S. Abraham, principal investigators). The 4 Ngatpang State Government, Republic of Palau work performed in this study complied with the current 96940. Current address: National Oceanic and Atmo laws of the Republic of Palau. Manuscript accepted 28 spheric Administration, Northeast Fisheries Science November 2007. Center, Woods Hole Laboratory, Woods Hole, Massa 2 Corresponding author. chusetts 02543. 3 U.S. Department of Agriculture Forest Service, Pa 5 Palau Department of Marine Resources, Republic cific Southwest Research Station, Institute of Pacific Is of Palau 96940. Current address: Tropical Conservation lands Forestry, Hilo, Hawai‘i 96720. Current address: Biology and Environmental Science, University of Ha wai‘i at Hilo, Hilo, Hawai‘i 96720. 6 Institute for International Studies, Center for Envi ronmental Science and Policy, Stanford University, Pacific Science (2009), vol. 63, no. 1:15–26 Stanford, California 94305-6055. Current address: Con : 2009 by University of Hawai‘i Press servation Strategy Fund, 1160 G Street, Arcata, Califor All rights reserved nia 95521. 15 Babeldaob Republic Island of Palau Ngaremeduu Bay l 5km 130' E 140' E 150' E 160' E 170' E 20' N Wake,; r,.../) ) I\_ + / Philippines l ,,,'• ~ .... '-.,.~ Guam • · \)•'""•~ '.I ~~{~ l--,';;J'~ ~ •. - (~ ?~"1',- --+---Ya_p_ , J·--•-------t------+-----..,, -, -I-M-ar-sh-a-11 10' N- ,,_,..,_., l ,;, ~ ""' Pohnpei Islands v-~,..... ('.} J ) ' z ~.,#!:J ~a -o-- ..... Q Cl C: •••• -~ p I , ° Chuuk ' v i • a au '• Kosrae Figure 1. Locations of Ngaremeduu Bay and its three main tributaries on the island of Babeldaob and of the Republic of Palau in the Micronesian archipelago. Mangrove Crabs in Palau . Ewel et al. 17 Palau and in many other parts of their range. able for overlapping populations of S. para This island nation is located in a regional mamosain and S. olivacea in Vietnam (Walton ‘‘hotspot’’ of biodiversity (sensu Atherton et al. 2006b, Le Vay et al. 2007) and Thailand 2007), and, like the rest of the Micronesian (Overton and Macintosh 2002) and for S. islands, has an economy that depends heavily tranquebarica, S. olivacea, and S. serrata in the on its renewable natural resources. Under Philippines (Walton et al. 2006a). standing more about ecological relationships This study began as an effort to estimate involving mangrove crabs may be useful not the population size and individual size charac only for Palauan resource managers but for teristics of mangrove crabs at a time when we understanding how this important organism believed S. serrata to be the only species. We varies in its population characteristics and also wanted to characterize mangrove forest habitat preferences throughout its range. The habitat, particularly where the crabs construct purpose of this paper is to summarize major burrows. Because of the opening of Palau’s characteristics of mangrove crabs and their new capitol complex on Babeldaob and the habitat in Palau. imminent completion of a road around the is Adult mangrove crabs spend most of their land, it was clear that the human population time in sheltered estuaries, bays, and the low and general traffic on that island would soon er reaches of rivers and tidal creeks; they increase, subjecting Ngaremeduu Bay to more often dig burrows, which they may use as ref environmental disturbance and its mangrove uges or for mating (Brown 1993). They are crabs to greater harvest pressure. Federal reg generalist feeders and seldom move more ulations in force at the end of the twentieth than 1–2 km from a given site (Hyland et al. century already prohibited the harvest of any 1984, Benstead et al. 2006, Le Vay et al. 2007, mangrove crabs with a carapace width less Bonine et al. 2008; D. Perrine, 1978, unpubl. than 6 inches (approximately 152 mm) and data on Scylla serrata on Ponape). of all egg-bearing females, as well as the ex Shortly before our study began, a taxo port of any crabs. Nevertheless, experienced nomic revision subdivided Scylla serrata, a fishers around the bay remarked at that time designation that for many years included all that it was increasingly difficult to trap man mangrove crabs (Stephenson and Campbell grove crabs of any size. Understanding the 1960), into four species: S. serrata, S. para status of the crab population and major char mamosain, S. olivacea, and S. tranquebarica acteristics of areas where crab burrows occur (Keenan et al. 1998). These four species over could assist future resource managers in pro lap throughout their range and are likely to tecting key areas, evaluating environmental be subjected to similar harvest pressures. impacts, and designing appropriate manage Nevertheless, the taxonomic confusion that ment and conservation protocols. Our study prevailed for many years reduces what is de was later extended to determine which species finitively known about the characteristics of of mangrove crabs were present in the bay any one species of crab, of communities of and the average sizes of crabs in these species. two or more species, and of the habitats In addition to providing information useful of different species. Studies from sites where to resource managers on Palau, the results of only one species occurs or, more recently, this study also expand our understanding of where coexisting species could be discrimi the distribution of different species of man nated, have shed some light on differences in grove crabs throughout the Indo-Pacific and population characteristics and habitat prefer provide a basis for interpreting the status of ences among the four species. For instance, mangrove crab populations in other coun studies in South Africa (Robertson and Piper tries, particularly elsewhere in Micronesia. 1991) and Kosrae, Federated States of Mi cronesia (Benstead et al. 2006, Bonine et al. materials and methods 2008), provide information on S. serrata where it is known to be the only species. Dif We first trapped mangrove crabs from Janu ferences in growth rates, habitat preferences, ary 1999 through May 2000 in Ngaremeduu and responses to exploitation are now avail Bay. At the same time, we obtained baseline 18 PACIFIC SCIENCE . January 2009 information about habitat characteristics. We from the bay, starting azimuth (0–360), and trapped again in May 2004. initial distance from shore (0–20 m). At the end of each transect, a new random azimuth was chosen along with a randomly selected Study Site distance (0–20 m) to the beginning of the Ngaremeduu Bay is shallow, with extensive next transect. intertidal and subtidal mudflats, and it is lined Within each belt transect, soil depth (cm) by mangrove forests. At 1,450 ha, it is the was measured by inserting a meter stick as largest estuary in Micronesia. Three rivers deep as possible. Soil type was classified as flow into the bay: Ngermeskang, Ngatpang silt with organic matter or sandy by visually (also called Ngetpang and Ngchelotel), and inspecting it and rubbing it between the fin Nekkeng (also called Tabecheding) (Figure gers. Tree height at 0, 10, 20, and 30 m along 1). Names and spellings of place names are the transect was estimated as less than 5 m, notably variable in Palau, in part a result of 5 m to 10 m, or greater than 10 m.