SEASONAL 1RENDS OF EPIBENTIIIC MACROFAUNAL
ASSEMBLAGES IN THE NEARSHORE WA1ERS OF THE WES1ERN
YELLOW SEA, QINGDAO, PEOPLE'S REPUBLIC OF CHINA
A University Thesis Presented to the Faculty
of
California State University, Hayward
In Partial Fulfillment
of the Requirements for the Degree
Master of Science in Marine Science
By
Kevin L. Rhodes
June, 1995 Abstract
The Yellow Sea demersal fishery has become one of the world's most extensively overfished marine resources since the introduction of the bottom trawl in the early 20th century. Since the 1960's, a continuous decline has been observed in the total catch and size composition of target species, with concurrent changes in community composition and the relative abundance and distributions of individual taxa, characteristic of ecosystem overflshing. To document the impact of overfishing and to describe seasonal trends in macrofauna! assemblages in the nearshore waters along the eastern shore of Shandong
Province, People's Republic of China, a total of 82 tows were made at three 16 Ian2 sites at bimonthly intervals from March through November 1993 with a 4.9 m otter trawl.
Invertebrates, particularly decapod shrimp, dominated the catch throughout the survey, comprising 90.4% of the total abundance and 76.4% of the total biomass, with fishes contributing ouly 9.6% of the total abundance and 23.6% of the total biomass. All individuals of the most abundant fish, the commercially important white croaker
Argyrosomus argentatus, were entirely composed of young-of-the-year (1 year of age), with a mean standard length of ouly 5.0 ± 3.2 em. Nonparametric analysis of variance showed several significant differences between sites for overall monthly mean abundance, biomass, and number of taxa per tow, with consistently higher values at the north site, perhaps due to differences in sediment type and composition, or influences from nearby
Jiaozhou Bay. Several confounding political, social and economic factors within each of the neighboring countries limit implementation of fisheries regulations within the Yellow
Sea. Establishment of an independent monitoring agency is drastically needed to document existing stocks, monitor biological and physical changes within the Yellow Sea and initiate enforcement of existing regulations.
ii Acknowledgements
I would particularly like to thank Xia Qing and the Chinese Research Academy of
Environmental Sciences (CRAES), and the government of the People's Republic of China for their kind assistance and for allowing me to conduct this study. Without his help, this project would not have been possible. I sincerely hope that this information will be useful in resolving the problems associated with the Yellow Sea fisheries and look forward to cooperating with him on future projects.
I would also like to thank Professor Zhang Zhinan, Department of Marine Biology,
Qingdao Ocean University, for providing me with laboratory space and equipment, which enabled me to complete this study. His kindness and generosity are greatly appreciated.
I am greatly indebted to Zhao Jianzhong, Department of Physical Oceanography,
Qingdao Ocean University and Captain Zhang Ruiquan and his crew, who provided enthusiastic support and assistance during the fieldwork. I would like to extend a special thanks to Mr. Xu, whose talents in the galley made each trip an especially delightful experience.
For their generous financial assistance, a special thanks to Paul and Kathy Buck, who wholly supported the research by providing funds for purchases of equipment, housing, field assistance and tuition both during and after my stay in China. Their help and kindness will not be forgotten.
Most notably, thanks to my friend, Ji Rubao, whose help, advice and friendship during the course of this survey was indispensable, whose discussions greatly increased my understanding of the Yellow Sea environment, and whose personality reflected everything that is great about the Chinese people.
IV Also, thanks to my committee members, Drs. Gregor M. Cailliet, James P. Barry and James Nybakken, whose advice, enthusiasm and support during all phases of the project helped it reach fruition.
Finally, my most heartfelt thanks to Kimberley Warren, who believed, more than anyone, in my ability to see this project through to the end. Her companionship and encouragement during all phases of the project, and her insights about the culture and personalities of the Chinese people, allowed for "smooth sailing." Her many trips into the field over the years, often in inhospitable weather, are greatly appreciated.
v Table of Contents
Page
Abstract ...... ii Acknowledgements ...... , ...... v List of figures ...... ix List of tables ...... xiii Introduction ...... 1 Materials and methods ...... 5 Site Description ...... 5 Pilot study ...... 6
Physical Measurements ...... ?
Biological Sampling ...... ? Analysis ...... 9 Results ...... 12 Pilot Study ...... 12 Physical Measurements ...... l2 Fishes ...... 13 Invertebrates ...... 19 Community Parameter ...... 23 Discussion ...... 25 Bibliography ...... , .... 3 7 Figures ...... 43 Tables ...... 63
vi Page
Appendix A: Regulations on the Protection of Aquatic Resources ...... 91
Appendix B: List of fishes captured during the March through November 1993 survey
...... 92
Appendix C: Table of values from the Kruskal-Wallis analysis of variance for comparison
of site-specific monthly fish abundance, biomass and number of taxa per tow ...... 94
Appendix D: List of invertebrates captured during the March through November 1993
survey ...... 95
Appendix E: Table of values from the Kruskal-W allis analysis of variance for comparison
of site-specific monthly invertebrate abundance, biomass and number of taxa per
tow ...... 99
vii List of Figures
Figure Page
1. Map of the Yellow Sea and inserts of the biological and sediment sampling sites
...... 43
2. Scatter plot of fish and invertebrate taxa trawled during the pilot study made in
February and March 1993, with mean and standard deviation of number of taxa
per tow from the main study ...... 44
3. Monthly cumulative species curves for fish and invertebrate taxa, by site, for samples of
combined fish and invertebrate taxa taken March-November 1993 ...... 45
4. Mean temperature and salinity measurements by month ...... 46
5. Relative contributions of the major families of fish collected in samples taken in the
March-November 1993 survey ...... 47
6. Percent contribution to total abundance for fishes collected in samples from March
through November 1993 ...... 48
7. Percent contribution to total biomass for fishes collected in samples from March
through November 1993 ...... 49
viii Figure Page
8. Monthly mean fish abundance at each of the three survey sites for samples taken March-
November 1993. Error bars represent standard error...... 50
9. Monthly mean fish biomass at each of the three survey sites for samples taken March-
November 1993. Error bars represent standard error...... 51
10. Size frequency histogram for Argyrosomus argentatus captured March- November
1993 ...... 52
11. Size frequency histogram for Cynoglossus joyneri captured March- November 1993 ...... 53
12. Rank percent abundance and biomass of the dominant flsh and invertebrate taxa...... 54
13. Percent contribution to total abundance for invertebrate taxonomic groups collected in
samples from March through November 1993 ...... 55
14. Percent contribution to total abundance for dendrobranchiate and caridean shrimps and
prawns collected in samples from March through November 1993 ...... 56
15. Percent contribution for invertebrates taxa contributing ~1% to total biomass collected
in samples from March through November 1993 ...... 57
ix Figure Page
16. Monthly mean invertebrate abundance at each of the three survey sites for samples
taken March-November 1993. Error bars represent standard error...... 58
17. Monthly mean invertebrate biomass at each of the three survey sites for samples taken March-November 1993. Error bars represent standard error...... 59
18. Monthly mean diversity values for combined fishes and invertebrates collected at each
of the three survey sites for samples taken March- November 1993. Error bars
represent standard error...... 60
19. Monthly mean evenness values for combined fishes and invertebrates collected at each
of the three survey sites for samples taken March- November 1993. Error bars
represent standard deviation ...... 61
20. Mean monthly number of combined fish and invertebrate taxa per tow. Error bars
represent standard error ...... 62
X List of Tables
Table Page
1. List of the number of tows per month by site ...... 63
2. Rank percent abundance and biomass of dominant fish and invertebrate taxa..... 64
3. Taxon-specific abundance, biomass and frequency of occurrence data for fishes for all
sites combined. N=total number collected, FO=frequency of occurrence,
SE=standard error...... 65
4. Taxon-specific fish abundance, biomass and frequency of occurrence data for north
site. N=total number collected, FO=frequency of occurrence, SE=standard
error ...... 66
5. Taxon-specific fish abundance, biomass and freqency of occurrence data for central
site. N=total number collected, FO=frequency of occurrence, SE=standard
error ...... 67
6. Taxon-specific fish abundance, biomass and frequency of occurrence data for south
site. N=total number collected, FO=frequency of occurrence, SE=standard
error ...... 68
7. Monthly trends of relative abundance and species richness for fishes ...... 69
xi Table Page
8. Mean percent similarity comparisons of fish and invertebrates ...... 70
9. Overall mean percent similarity comparison for all taxa combined ...... 71
10. Taxon-specific invertebrate abundance, biomass and frequency of occurrence data for
all sites combined. N=total number collected, FO=frequency of occurrence,
SE=standard error...... 7 2
11. Taxon-specific invertebrate abundance, biomass and frequency of occurrence data for north site. N=total number collected, FO=frequency of occurrence, SE=standard
error ...... 75
12. Taxon-specific invertebrate abundance, biomass and frequency of occurrence data for
central site. N=total number collected, FO=frequency of occurrence, SE=standard
error ...... 78
13. Taxon-specific invertebrate abundance, biomass and frequency of occurrence data for
south site. N=total number collected, FO=frequency of occurrence, SE=standard
error ...... 81
14. Monthly trends of relative abundance and species richness for invertebrates ..... 84
xii Table Page I Sa. Comparisons table of findings of fish and invertebrate abundance of the nearshore
Yellow Sea survey to results of selected coastal and deep-sea surveys ...... 87
15b. Comparisons table of fmdings of fish and invertebrate biomass of the nearshore
Yellow Sea survey to results of selected coastal and deep-sea surveys ...... 89
xiii 1
Introduction
Commercial fishing in the Yellow Sea has a history that dates back several
centuries. It is multinational in scope with over one hundred commercially harvested
species of fishes and invertebrates (Zhang, et al, 1988; Tang, 1988). Although not
extensively fished before the introduction of the bottom trawl in the early 1900's, the
Yellow Sea is now one of the most intensively fished areas in the world. From the early
1960's to the early 1980's, fishing effort tripled while resource biomass declined over
40%, relative to other Northwest Pacific continental shelf areas (Tang, 1988). During the
same period, total catch and size composition of most target species declined continuously
and changes in community composition and the relative abundance and distributions of
K: individual taxa were common (Hwang, 1977; Lee & Park, 1978; Mangan, 1982; Mio,
1984; Valencia, 1988; Tang, 1988; Yu, 1991). This was particularly noteworthy for
commercially important demersal fishes and invertebrates, which account for 60 to 90% of
the annual total catch, with semi-demersal and pelagic stocks contributing the remaining
proportion (Zhang, et al, 1988; Tang, 1988; Iverson, 1993). Total catch rates in the
Yellow Sea in 1988 were 2.3 tons·km-2yrl, or approximately 9.2 X lOS tons of catch,
which is well above maximum sustainable yield estimates (Tang, 1988).
The proportion of catch for the five most commercially important fishes in the
Yellow Sea ( Sciaenidae: small yellow croaker, Pseudosciaena polyactis, and large yellow
croaker, Pseudosciaena crocea; Trichiuridae: hairtail, Trichiurus haemula; Gadidae: Pacific
cod, Gadus macrocephalus; Pleuronectidae: flatfish, Cleisthenes herzensteinz) has declined
from 35% of the total catch in the 1950's to less than 10% in recent years (Zhang, et al,
1988; Tang, 1988). The catch of small yellow croaker, once a major commercial species,
was reduced from 330,000 tons in 1957 to only 63,000 tons in 1983 (Mio, 1984; Tang, 1988). This same trend has been shown for the hairtail, whose catch reflected a 20 year low in 1988 that was an 18.8% decline from the previous year alone (Yu, 1991).
To compound the problem of overfishing, countries that have jurisdictional claim to this area (Japan, China and North and South Korea) have attempted to boost yield by targeting nursery areas in coastal zones and overwintering grounds of spawning stocks
(Zhang, et al, 1988; Tang, 1988). This has resulted in a reduction in abundance and distribution of several commercially important species and a replacement of these species by less desirable fishes, providing additional evidence that the Yellow Sea has reached a state of ecosystem overfishing (Pauly, 1979, 1988a, 1988b). Higher trophic level species have been replaced by small-bodied, lower trophic level species and at least one Korean survey reports that in 1985, 70% of the catch biomass of all commercial species of fishes was composed of individuals with mean body length of only 12 em (Zhang, et al, 1988).
Further evidence is provided by catch statistics of the small yellow croaker, which show that: (1) from the late 1970's to the present, individuals in the 0-1 year age class accounted for 80% of the catch, and, (2) that an overall increase in growth rate, early maturation and a decrease in the mean age and body length of spawning stocks occurred (Hwang, 1977; Lee
& Park, 1978; Zhang, et al, 1988). Additional findings from catches of the white croaker
Argyrosomus argentatus in the Yellow Sea and East China Sea from 1970-75 showed that
87% of the fish caught were 0-1 year age class (Lee and Park, 1978).
In China, politics, economics and a growing population have all played crucial roles in fisheries overexploitation (Yu, 1991). During Mao's Cultural Revolution (1966-76), conservation measures were nonexistent and fishermen throughout the country were advised to catch any and all fishes, without regard to size or species (Mangan, 1982).
Recently, management and conservation issues have become more relevant, given the obvious crisis in China's seas, so that seasonal fishery closures, mesh size restrictions and taxon-specific size regulations have been proposed.
2 In 1979, a comprehensive doctrine on national fisheries resource conservation, the
"Regulations on the the Protection of Aquatic Product Resources," was enacted by China's
State Ministry of Agriculture to curtail overfishing through fines, confiscation of gear and catch, and compensation from violators (National People's Congress of the People's
Republic of China, 1982) (Appendix A). However, cost of enforcement, corruption within and conflicts between regulatory agencies, and a powerful and politically-effective local fishing lobby have all prevented effective enforcement of existing laws and have contributed significantly to the continuation of exploitative fishing practices (Yu, 1991).
Adding to the dilemma is the recent shift toward a free market economy. This shift resulted in changes in economic policy which allows fishermen within cooperative fishing units to sell much of their catch to individual buyers for a much higher price than that paid previously by the state (Mangan, 1982). This has resulted in intensified efforts to increase catches to boost personal profits. When combined with the increased availability of hard currency, that enables fishermen to buy from foreign sources, and the import of more previously unavailable, technologically-advanced equipment, remaining stocks can now be easily identified and targeted, fueling further decline of the fisheries. Additionally, China currently has few options for retraining or repositioning the growing labor force which mans 190,000 vessels currently fishing in Chinese waters. Moreover, the "one child" policy that was implemented to stabilize China's population at 1.5 billion in the year 2000 has fallen short as revealed by the most recent census that shows the population to have exceeded that cap in 1994 (Edmonds, 1994). As the current population continues to grow, so will the demand for food resources.
Under Deng Xiaoping, scientific research has again been looked upon as progressive and is touted as one of the "Four Modernizations" that are essential to China's transformation into an economic superpower. But the country is still developing and money for scientific research is scarce, with projects generally short-term and focusing on
3 only commercially important species (Gu, 1975). In addition, techniques and equipment are largely outdated and communication among agencies is virtually nonexistent, causing a duplication of data or an incomplete understanding of the impacts to the fisheries (Milliman,
1981). The focus of this project was to descrioe the nearshore epibenthic macrofauna! assemblage of the waters on the western edge of the Yellow Sea near Qingdao, Shandong
Province, People's Republic of China, and to document the seasonal changes in this assemblage. The paucity of information on China's marine biological communities provided incentive to complete this survey, the results of which will hopefully give biologists a reference point, or standard for comparison, to use for further study within this area. It will also provide information on the marine biological commuuity for the western
Yellow Sea that can perhaps be used to compare to other marine communities with similar physical attributes within and outside of the Yellow Sea. Finally it is intended to support previous evidence that the Yellow Sea is in a state of ecosystem overfishing and, if so, suggest more drastic and immediate changes in China's fisheries policies.
4 5
Materials & Methods
Site description
The Yell ow Sea is part of a semi-enclosed, shallow water system bordered by
China to the west and the Korean Peninsula to the east (Figure 1). It is one of the largest shallow water areas of continental shelf in the world wi'th about 400,000 kJn2 of total area.
(fang, 1988). Mean depth is 44 m and its bottom is primarily sandy mud with occasional rocky outcrops in the nearshore areas (Mio, 1984). Isobaths typically run in a north-south direction (Guan, 1984). The northernmost limits are at approximately north latitude
N39°00.00, at the Bohai Sea boundary, which runs from Shandong Province to the
Liaodong Peninsula. To the south it is bounded by the East China Sea at approximately
N32°.00.00, by a line running from the Yangtze River mouth to Cheju Island, located just south of the Korean peninsula. Average rates of primary productivity are estimated to be
60 gC·m-2·yrl with a benthic biomass of 20-41g·m-2 (2.0-4.1 X 1Q5 g ha-l) and an annual maximum sustainable yield of about 9.6 X 1Q5 tons (fang, 1988; Yang, 1989).
The Bohai Sea and Yellow Sea receive oceanic water from an extension of the
Kuroshio Current, which diverges near the western part of Kyushu Island and flows northward at less than 0.5 knots along the western portion of the Korean Peninsula
(Zhang, et al, 1988; Guan, 1994). Together with the less saline Huanghai (Yellow Sea)
Cold Water Mass, which forms in winter and flows southward from the Bohai Sea in summer, a basin-wide cyclonic current is formed and constitutes the overall pattern observed for this area (Guan, 1984; Su and Weng, 1994). Along the eastern coastal border of the Yellow Sea, bottom currents flow predominantly southward along isobaths year round, except during summer monsoon when localized and temporary anticyclonic eddies form along the coast Within the sampling area, persistent semidiurnal tidal currents flow through nearby
Jiaozhou Bay, varying both in speed and direction, and generally run in a northwest southeasterly direction at a speed of up to 2 knots (Qingdao Ocean University, 1990).
Temperature of bottom water within the sampling area varies seasonally with highest values in late summer, the lowest in late winter, and an annual average of 22.8° C. Salinity remains relatively stable year round, ranging from 29.9 to 31.1°/00, and averaging
30.80/00. Wind direction is predominantly from the southeast at 5.3 mls (Qingdao Ocean University, 1990). Annual primary productivity estimates for this region range from 110-
183 gC·m-2-yrl with peak production in summer for phytoplankton and peak biomass in summer and winter for zooplankton (Guo, 1994). Primary productivity measurements within nearby Jiaozhou Bay average 154 gC·m-2·yrl, with lowest values around 9 gC·m-
2.yrl in winter and high value of 321 gC·m-2·yrl in summer (Guo, 1994).
Pilot Study
An estimate of optimal towing time was determined from a pilot study conducted in
February 1993, during which two tows of 20, 30, and 40 min duration each were made.
An additional 1 hr tow was made in March 1993. Tows for the pilot study were made at a haphazardly-chosen site in close proximity to the main study area. Sampling protocol for the pilot study was identical to the main study, with a boat speed of 2.5 knotslhr and a 4:1 scope. Samples were sorted in the field at the time of capture, preserved in 10% formalin, and returned to the lab for identification to lowest taxonomic leveL Towing time for the main study was based on results of the pilot study and a desire to maximize the number of tows allowed during normal daylight hours.
6 Physical Measurements
Physical measurements made during most surveys included temperature and salinity, recorded with a conductivity, temperature, depth (CID) recorder. The CID was not available for use during the March sampling period and salinity and temperature values for that month were derived from the literature. Logistical constraints prevented current measurements from being taken during the course of the survey. Maximum tidal velocity for the sampling area has been estimated to be 0.8 knots, with tidal current flow running along the coast in a predominantly northeast-southwest direction (Qingdao Ocean
University, 1990).
From observations during sampling and analysis, obvious differences in sediment type and grain size were noted and an a posteriori survey of the sediment was conducted in random fashion in September 1994 at each of the three sites (Figure 1). Five replicate samples were taken using a 4 em diameter corer and/or a Mcintyre grab sampler. Samples were returned to the lab, oven dried at 100 OC, sorted by grain size and analyzed for differences in fractional percentages and means of grain size.
Biological Sampling ·
Sampling was conducted at three sites, located 8 to 12 km east-southeast of the mouth of Jiaozhou Bay, Shandong Province, between N 35054.00 toN 36010.00 and
E 120°18.00 toE 120027.00 (Figure 1). Sites were chosen along depth gradients that range from 14 to 28 m and roughly parallel the coast Each site covered 16 km2, and was separated from other sites by at least 1 km but less than 4 km. The three sites were selected randomly from a pool of seven potential sites that paralleled the coast for 32 km.
Prior to the initiation of the main study, each 16 km2 site was subdivided into 1 km2 'blocks'. Six 'blocks' were randomly chosen from each site prior to each bimonthly sampling and trawling was initiated within those 'blocks'. Direction of trawling was
7 haphazardly determined at the time of trawling, based on sea conditions, primarily swell direction and force.
Bimonthly trawling was conducted from March through November 1993 using a standard commercial otter trawl with a 4.9 m headrope, a nylon net with a bag length of 6.1 m of 3 em mesh and a 3 mm stretch-mesh cod end. Effective trawl path width was estimated to be 3.4 m and faunal densities were calculated from this estimate (Research Net
Shop, Bothell, WA). The net was spread with 12 by 24 inch wooden doors and towed at an average speed of2.5 knots with a 4:1 scope. Depth of trawling was between 14 and 28 m. Eighty-six tows were taken during the survey. Six tows were taken at each site bimonthly, except in November, when equipment loss on the last scheduled sampling day prevented a complete set of tows from being taken at the north site (Table 1). Four tows were removed from the analyses because of questionable bottom contact, leaving a total of
82 tows for analysis of benthic faunal distribution. Tows were timed from the point when all line was paid out to the initiation of gear retrieval. All positions were recorded using a portable Koden KP-109A Global Positioning Satellite (GPS) receiver.
At the time of captore, specimens were preserved in 10% formalin, returned to the lab for sorting and identification, and later transferred to 70% ethanol. Voucher specimens for each taxon were retained, and whenever possible, photographs were taken in the laboratory or in the field at the time of capture. For all fishes, the standard length (SL) of specimens was measured to the nearest 1.0 mm, and weight taken to the nearest 0.1 g.
Similarly, invertebrates were measured to the nearest 1.0 mm and weighed to the nearest
0.1 g. Weights for specimens weighing less than 0.1 g were not recorded.
Voucher specimens of fishes were donated to the California Academy of Sciences,
San Francisco, California (CAS), Natural History Museum of Los Angeles County, Los
8 Angeles, California (LACM), or to the Smithsonian Institution National Museum of
Natural History, Washington, D.C. (USNM). All invertebrates and some fish specimens were donated to the USNM, except octopods, which are housed at the Santa Barbara
Museum of Natural History, Santa Barbara, California (SBMNH).
Fishes were identified using both general and specialized references (Tchang, 1954;
Lindberg and Legeza, 1959, 1965; Lindberg and Krasyukova, 1969, 1989; Menon, 1977).
Invertebrates were identified and later verified using several general and specialized taxonomic references (D'yakonov, 1950; Ushakov, 1955; Shen and Dai, 1964; Qi, eta!,
1989). Unidentifiable taxa were sent to taxonomists at either CAS or USNM .
The identification of some of the specimens is still pending. For purposes of analysis, all unidentified taxa were minor components within the community, yet were treated as individual species, with the exception of the cyclopterid genus Liparis. Liparis is a dominant fish group and a major contributor to the fish community. It is possible that this genus comprises 3 separate species. However, for analytical purposes at this stage,
Liparis were treated as a single taxon.
Analysis
With few exceptions, data on fishes and invertebrates were analyzed separately.
For community parameters, such as diversity (Shannon-Weaver index H') (Shannon and
Weaver, 1963), evenness (J') (Pielou, 1969), and species richness (S), catch data for fishes and invertebrates were analyzed separately, and in combination, for comparative purposes. Reported mean values of of H', J', and S were derived from an analysis of individual tows for each of these parameters. An overall value for these parameters (H', J, and S) was derived from the combination of data from all tows and is presented for comparison. For combined indices, a Spearman's rank correlation coefficients test (Sokal and Rohlf, 1981) was used to justify combining sites for these forms of analyses using
9 ranked percent abundances for individual fish and invertebrate taxa from each of the respective sites.
Similarity of the fish and invertebrate samples among months and sites were evaluated using a percent similarity index (PSI), derived from combined PSis on paired tow comparisons (Whittaker, 1952). For the site/month comparisons, mean PSI values were determined by summing individual PSI values from each possible individual tow-tow comparison, for all of possible combinations. For comparisons of sites, a grand mean PSI was calculated by summing means from PSI measurements, with corresponding values representing an overall site-to-site comparison. The level of significance for PSI values was set at 80% (Silver, 1975).
Frequency of occurrence (FO), percentages of total catch (%N), and means and standard errors (SE) for both biomass and abundance for all individual taxa were calculated and are shown for both the combined sampling area and for individual sites.
Presence/absence trends, relative monthly abundance, and historical and observed data regarding resident/transient status, were also established. Monthly trends for mean abundance and biomass are given for fishes and invertebrates by site to show temporal changes within the community. For the dominant fish species Argyrosomus argentatus and
Cynoglossus joyneri, size frequency histograms were plotted to determine the size distribution of commercially and noncommercially harvested fishes within the sampling area, and to see how these findings compare to recently reported trends in sizes of these two taxa in commercial catches from the Yellow Sea. For some dominant taxa, a graphical analysis of percent biomass, abundance, and frequency of occurrence, by site, was made to demonstrate outstanding differences among taxa. Again, a Spearman's rank correlation coefficient comparison was used prior to combining data from individual sites.
A nonparametric analysis of variance (Kruskal-Wallis, a=0.05) was used to compare salinity and temperature between areas. Nonparametric analysis of variance was
10 also used to determine (1) if there were differences in monthly mean values for abundance and biomass of fish or invertebrates among sites, (2) if there were differences in the monthly mean number of taxa per tow among the three sites, and, (3) whether differences existed between sites regarding sediments composition and mean grain size.
Finally, for comparative purposes, resUlts of several surveys of fishes and macroinvertebrates from coastal and deep-sea sites in the U.S. were selected from the literature and compared to fmdings from this survey. Most surveys used in the comparison were from latitudinal gradients similar to those of Yellow Sea survey, although a few were chosen from the southern coast of California because of similarities in depth and sampling regime.
11 12
Results
Pilot study
A technique that included three tows of 30 min each at each site was selected as the principal sampling method as a compromise between logistic constraints and variability among tows. The pilot study conducted in February 1993 indicated that the number of taxa and the variability between tows was still rising at tow times up to 40 min (Figure 2). An additional tow of 60 min, made in March 1993, captured a lower number of taxa than the
February 1993 tows of 40 min, suggesting that variability would also potentially increase with tow times greater than 40 min. Therefore, for logistical reasons and to increase sample size, while trying to decrease variability among tows, a tow time of 30 min was selected and used in the main study.
Cumulative species curves calculated for each month and site show that the number of taxa levelled after -3 tows, demonstrating that both the duration and number oftows made at each of the sites satisfactorily characterized the epibenthic faunal community
(Figure 3). This trend is also shown for fish and invertebrates when treated separately, but the results of that analysis are not presented here.
Physical measurements
Salinity remained relatively constant throughout the sampling period with temperature fluctuating seasonally (Figure 4). Bottom temperatures were highest in July and lowest in March (22.7 and 4.2 OC, respectively). Analysis of variance (Kruskal-
W allis, a.=O.OS) showed that there was a significant difference in salinities between sites in
September, although the maximum range of difference was less than 0.2 0/00 (F= 19.40,
P= 0.39, n= 17). Significant differences in temperature were found in July and September, with a maximum mean difference of 0.8° C between north and south sites in September and o.4o C difference in July (F= 12.39, p= 0.02, n= 18, and F= 20.61, p < 0.001, n=
17, respectively). In both cases, the site with the highest mean temperature also had the highest abundance. . Sediment mean grain size and fractional percentages differed significantly among the north and the central and south sites. Sediment samples from the north site were dominated by medium to fine sand fraction with a mean grain size of 0.2 ± 0.0 mm and a mean fractional percentage of silt/clay of only 23.8 ± 16.5%. By comparison, the central and south sites had much higher mean percentages of silt/clay, at 76.1 ± 1.5% and 87.2 ±
1.2%, respectively. Mean grain size for both central and south sites was 0.06 ± 0.00 mm.
Nonparametric analysis of variance (Kruskal-Wallis, Fishes A total of 1840 individuals with representatives from 15 families and at least 26 species of fishes was collected in trawls made between March and November 1993 (Appendix B). Fishes were a relatively minor component within the community, in both abundance and biomass, contributing only 9.6% of the overall abundance and 23.6% of the biomass for the survey. Combined data for fishes show that the ten dominant taxa account for 94.9% of the total abundance and 83.2% of the total biomass. 'Rare' taxa, defmed as those which contributed less than 1% to the total abundance of fishes, accounted for 66.7% of the total number of fishes collected. In terms of rank abundance for fishes and invertebrates combined, only three fishes (the white croaker, Argyrosomus argentatus, the cyclopterid taxa, Liparis sp., and the cynoglossid, Cynoglossus joyneri) ranked in the top ten, and contributed only 7.3% to the 13 combined total abundance for the survey (Table 2). These same fishes accounted for 17.3% of the total biomass. Families represented by the greatest number of individual taxa were Sciaenidae and Gobiidae, each with six representatives. Sciaenids included, in order of highest to lowest abundance, A. argentatus, Johnius belengerli, the small yellow croaker, Pseudosciaena polyactis, Collichthys lucidus, Nibea albiflora, and Collichthys niveatus. Gobiid representatives were, in order of rank abundance, Gobius pflaumi, Chaeturichthys hexane rna, Cryptocentrus filifer, Synechogobius hasta, Tridentiger nudiventris and Ctenopauchen chinensis. In terms of percent composition of total abundance for fishes at all sites, Sciaenidae was the roost dominant family, followed by the Cynoglossidae, with the single taxon Cynoglossus joyneri, Cyclopteridae, represented by Liparis sp., and Gobiidae (Figure 5). The most dominant fish taxon for the survey, in terms of both biomass and abundance was A. argentatus, with an overall density of 11.2 indivlha and a biomass of 71.8 g!ha. The flatfish Cynoglossus joyneri was the most frequently caught fish at all locations, occurring in 69.0% of the samples, and second only to A. argentatus in biomass, at 51.7 g/ha. Other taxa which comprised at least 1.0% of the total catch for fishes, included Liparis sp., Gobius pjlaumi, Apogon lineatus, Callionymus punctatus, Ammodytes sp., Chaeturichthys hexanerna and Pholis fangi (Table 3). The relative contributions of these taxa varied among sites, with the greatest variation shown for the three most abundant taxa (Argyrosomus argentatus, Liparis sp., and Cynoglossus joyneri) (Figure 6). Abundance contributions also varied substantially among sites for some minor taxa. At the north site, A. argentatus dominated catches overall and contributed 41.0% to the total abundance and 38.8% of the total fish biomass (Table 4). Liparis sp. accounted for 31.2% of the total abundance, but only 11.0% of the total fish biomass. By 14 comparison, C. joyneri made up 20.5% of the total biomass and only 7.8% of the total fish abundance (Figure 7). Other major contributors to the abundance totals at the north site were Gobi us pflaumi, Apogon lineatus, Callionymus punctatus and Ammodytes sp. The gobiid Cryptocentrus jilifer was the only fish sampled exclusively at the north site site, but contributed less than 1.0% to the total abundance. Overall abundance for fishes at the north site were estimated to be 60.5 indivlha with a total biomass of 576.5 glha. In comparison, at the central site, both C.joyneri and A. lineatus constitoted a much higher proportion of the total abundance and biomass (Table 5). At this site, C. joyneri contributed 24.8% and 38.4%, and A. lineatus 10.9% and 7.8% of the respective totals for percent abundance and biomass. The contribution of Liparis sp. was considerably lower than at the north site, with only 11.2 % of the total abundance. The roost notable differences between the north and central sites were the differences in the total abundance and biomass per hectare. For both measurements, a nearly fivefold difference is shown between sites, with the north site clearly having a higher biomass and abundance of fishes. Estimates of total fish abundance at the central site were 14.0 indivlha and a total biomass of 128.1 glha. Abundance and biomass measurements for dominant fishes at the south site were similar to those of the central site and also differed greatly from estimates from the north site (Table 6). A. argentatus and C. joyneri were again major contributors to abundance and biomass totals, with a combined 56.4% of the total abundance and 69.8% of the total biomass. In contrast to the central site was the large contribution of G. pflaumi, which accounted for 17.9% of the total abundance, whileA.lineatus made up only 1.7% of the total. Biomass percentage totals were not greatly affected by the increased abundance of G. Pflaumi because of its relatively small body size. Density estimates for fish abundance at the south site was 16.8 indiv/ha, with biomass estimates of 134.7 glha. 15 Temporal changes in abundance were largely inconsistent between sites with highest overall abundance at the north site during most of the sampling period except in November when catches were highest at the south site (Figure 8). Major abundance peaks at the north site occurred during March and September and coincided with an influx of two dominant taxa (Iiparis sp. and A. argentatus) into the area Variability among tows during March and September was high in comparison to other months, largely due to these two taxa. The minor increase in abundance at the south site during November was attributed to a relatively higher number of G. pflaumi and C. joyneri. Abundance peaks in March and September were primarily due to a few transient taxa entering the area during these months (Table 7). Specifically, young and juveniles of Liparis sp. and A. argentatus were present in relatively large numbers during these months in comparison to the number of other taxa, including adults of Liparis sp. and A. argentatus. Except for the increase of C. joyneri during July at the north and central sites, resident taxa maintained a relatively low abundance throughout the sampling period. By comparison, catches of most transients varied greatly, both temporally and spatially, but in general were caught in relatively small numbers. It should be noted that because of the low overall abundance of fishes, minor intrusions by one or a few taxa, particularly transient taxa such as A. argentatus, often dramatically altered the level of abundance within a sampling area. Overall abundance of fishes for the survey was estimated to be ouly 29.3 indivlha. Changes in biomass generally followed abundance trends, except at the north site. Major biomass peaks occurred in samples from July, September and November at the north site, in July at the central site, and November at the south site (Figure 9). Peaks in biomass during July at both the north and central site were attributed to an increase of C. joyneri and A. argentatus, with a relatively minor contribution by A. lineatus. The most notable 16 difference in biomass measurements between the two sites in July was related to the larger number of A. argentatus adults at the north site. The sustained biomass level at the north site in September and the minor increase in the south site reflected the influx of young and juvenile A. argentatus in the area. In November, the increase at the south site was related to the introduction of the 'rare' species of the synodontid Saurida elongata, the pleuronectid Pleuronichthys co mutus and the gobiid Synechogobius hasta. Although these taxa occurred in small numbers, they contributed greatly to the overall biomass due to their large body size. The cap tore of one very large adult liparis resulted in a dramatic increase in biomass in November. As with abundance trends, large-scale changes in overall biomass often resulted from the introduction of a small number of individuals of large body size. Overall biomass for fishes during the survey was estimated to be 189.1 glha. Size frequency histograms for the 531 individuals of A. argentatus captored during the survey showed a predominance of both young and juveniles, with adults making a relatively minor contribution (Figure 10). A bimodal distribution occurred with peaks at 3.1-4.0 em and 11.1-12.0 em SL, or 0-1 year age class, and an overall mean ± SD of 4.98 ± 3.23 em SL. For C. joyneri, a non-commercial fish, five possible size modes were observed from the size frequency distribution (Figure 11). The overall mean for all specimens was 11.43 ± 3.71 em SL. Gravid females were captured in July and September ranged in size from 11.7 em to 19.5 em SL (mean= 15.61 ± 1.43 em SL). Non parametric analysis of variance showed consistent significant differences between sites for May, July and September comparisons in the mean number of taxa per tow, density, and biomass of fishes (Kruskal-Wallis, a=0.05, Appendix C). For November comparisons, no differences were detectable between sites for any comparison, possibly related to the small sample size used for these comparisons. Multiple comparisons 17 teSting (fukey test) showed that south and the central sites grouped together in 92% of the comparisons. In contrast, groupings between north and south sites occurred in only 2 of 30 comparisons. Comparisons of fish abundance between sites using a PSI showed few significantly similarities among sites at the arbitrarily-set 80% significance level (fable 8). This was particularly true for comparisons between different months, probably due to seasonal movements of fishes through the sampling area (fable 7). In the majority of cases, only within-month comparisons between sites showed PSI values of>70%, reflecting general agreement between sites in terms of seasonal catch and relative abundances of taxa. PSI values were lowest in May, ranging from 6% for central-south comparisons to 52% for north-central comparisons, concurrent with lows for that month in terms of abundance, biomass and taxa per tow. Differences during May were largely due to the dominance of two taxa, Liparis sp. and Pholis fangi at the north site, which occurred in low proportions at the central and south sites during this period. Highest PSI values were in March, with 82% similarity for all site comparisons, when Liparis sp. dominated fish catch at all sites. Overall grand mean PSI values between sites were surprisingly similar and low, ranging from 31 to 33%, with the greatest similarity occurring between the central and south sites and the lowest values for the north: south comparison (fable 9). These values reinforce fmdings from Tukey' s test analysis in which central and south sites grouped most often in abundance comparisons. Invertebrates Invertebrates dominated catches throughout the survey, contributing 76.4% to overall biomass and 90.4% to total abundance. A total of 17,409 individuals from at least 35 families and 46 taxa was sampled (Appendix D). Invertebrates also occupied the top 18 five positions of rank biomass and abundance within the community and generally had catch frequencies above 70% (Figure 12). Eleven taxa accounted for 91.0% of the total abundance and 79.0% of the total biomass for invertebrates (fable 10). Crustaceans were the largest invertebrate group sampled, with representatives from at least 23 taxa, including the seven most abundant taxa for the survey: T. curvirostris, Metapenaeopsis dalei, Latreutes laminirostris, Squilla aratoria, Leptochela gracilis, Crangon ajfinis and Acetes chinensis. Among invertebrates, the percent contribution of dendrobranchiate and caridean shrimp and prawns to overall abundance was high compared to other taxonomic groups, with combined levels of total abundance~ 80% (Figure 13). With the exception of stomatopods (Squilla aratoria), the contribution by all other groups to overall abundance, including other arthropods, molluscs and echinoderms, was relatively minor. The asteroid Asterias rollestoni was the only major contributor not belonging to the arthropods, and dominated overall catch biomass for fishes and invertebrates combined, with 24.8% of the total, while the species contributed only 1.2% to overall abundance. For catches of both fishes and invertebrates, the most dominant taxon was the commercially important southern rough shrimp, Trachypenaeus curvirostris, whose overall mean abundance was 89.3 indiv/ha and biomass was 102.2 glha (Table 10). A. rollestoni, which was captured in 89.0% of the samples, contributed greatest to overall biomass, with 32.5% to the total among invertebrates. The mantis shrimp Squilla aratoria, was similar in biomass contribution, with 25.0% overall, but only 4.8% total abundance. As with fishes, the relative abundances of many invertebrate taxa, particularly dominants, often varied dramatically among sites (Figure 14). At the north site, T. curvirostris accounted for 47.5% of the total abundance and L. laminirostris contributed 28.7% (fable 11). T. curvirostris, S. aratoria and A. rollestoni had a combined total of 62.4% of the total biomass, with relatively minor contributions from each of the other 50 19 taxa found at this site. Abundance totals for all invertebrates at the north site were estimated to be 450.9 indiv/ha and 639.3 glha. Abundance percentages among the four major taxa at the central site were more evenly distributed than at the north site (Table 12). The proportions of M dalei and S. aratoria rose to 12.5% and 12.1 %, respectively, while those ofT. curvirostris dropped to 29.1% and L. laminirostris to 17.2%. Again, A. rollestoni and S. aratoria contributed most to overall biomass, with a combined total of 75.7% (Figure 15). Except forT. curvirostris and the urchin Temnopleurus toreumaticus, whose biomass values were 9.0% and 5.9%, respectively, biomass contributions from each of the remaining taxa were generally less than 1.0% of the total. Only 39 taxa occurred in samples at the central site, and several taxa which were taken at the north site were not caught during the survey, particularly the nudibranchs Sakuraeolis enosimensis, Dendronotus arborescens and the cephalaspidean Philine kinglipini. Abundance and biomass estimates for the central site were 155.5 indiv/ha and 710.0 gfha, respectively. In contrast to other sites, catches of invertebrates at the south site were dominated by the Kishi velvet shrimp M. dalei, which contributed 64.7% to total abundance, but only 13.5% to total biomass (Table 13). T. curvirostris, L laminirostris, S. aratoria, L. gracilis and C. affinis, which made up a relatively large proportion of the total abundance at the other sites, had a combined total at the south site of only 26.6%. However, T. curvirostris, whose contribution to abundance percentages was only 8.9%, was responsible for 16.4% of the total biomass. Other major biomass contributors were S. aratoria and A. rollestoni (Figure 15). As with the central site, only 39 taxa were sampled during the survey. Abundance and biomass totals for the south site were 243.9 indiv/ha and 464.7 glha, respectively. Patterns of seasonal abundance for invertebrates roughly paralleled those for fishes, with major abundance peaks in March, September and November, coincident with the 20 highest variability (Figure 16). Compared to other sites, the north site generally maintained a relatively higher abundance throughout the survey. Abundance peaks in March at the north site corresponded to increased numbers of the resident shrimp Latreutes laminirostris, whose abundance dropped thereafter (Table 14). Increases at all sites occurred in July and were related to an influx of several resident taxa, which included Crangon ajjinis and S. aratoria, and the transient T. curvirostris, which continued to be sampled in large numbers for the remainder of the survey. Other notable changes during July were the substantial increase in abundance of the crinoid Compsometra serrata, and a minor jump in numbers of the brachyuran crabs Dorippe granulata and Carcinoplax vestitus. The 42 taxa sampled during July were the highest for the survey. The largest increases in abundance among all taxa were for T. curvirostris in September at the north site, and in November forM. dalei at the south site. Overall abundance for invertebrates was estimated to be 277.3 indivlha. Invertebrate biomass trends generally followed abundance patterns, except at the central site during July and November (Figure 17). The July increase at the central site was due to a relatively large catch of A. rollestoni in several tows, in addition to a single tow that included over 1100 grams of S. aratoria. These same taxa caused a subsequent peak in biomass at the south site during November, particularly A. rollestoni, which contributed over 1000 grams to a single tow. As with fiShes, the inclusion of a relatively small number of large bodied animals, such as A. rollestoni, dramatically affected the overall biomass trends within a month. For invertebrates, monthly comparisons among sites for overall abundance, biomass and number of taxa per tow showed fewer differences than did those for fishes (Appendix E). Except for the November comparison, significant differences were consistently found between sites for the number of taxa per tow. For this measure, the north site consistently had the highest mean values, averaging 12.1 ± 5.8 taxa per tow, compared to 9.4 ± 4.2 and 8.1 ± 4.6 taxa per tow for the central and south sites, 21 respectively. Again, differences were greatest between the north and south site, and grouping (Tukey' s test) occurred in only 1 of 9 cases. Differences between sites in density and biomass measurements of invertebrates were found in March and September. Differences during March are attributed to the disproportionately high number of Latreutes laminirostris which occurred at the north site, and the large number of Asterias rollestoni at the central site. It should be noted that A. rollestoni often had a major effect on the overall biomass for a site, given both the large mean weight of these organisms (mean= 55.2 ± 34.5 g/indiv), the frequency of capture (mean= 2.7 indiv/tow) and the rather low overall biomass per tow (mean= 468.2 ± 514.3 g/tow). Differences in biomass and abundance during September are related primarily to the presence of large numbers of Trachypenaeus curvirostris at the north site. Percent similarity comparisons reflect the disproportionality in seasonal abundance and community composition of invertebrates among sites (Table 8). Several within-month comparisons show values well below the significance value of 80%, although most have values above 50%. As with fishes, lowest PSI values are shown for May comparisons, with highest values in March. The largest differences consistently occurred between sites in July when one or more dominant taxa, which included T. curvirostris, C. affinis, Squilla aratoria and M. dalei, were present in disproportionately large numbers within a particnlar site. Overall mean PSI values for all months combined were similar to those for fish, ranging in value from 29.0 to 32.0%, with the greatest similarity occurring between the south and central sites (Table 9). These values once again reinforce findings from group analysis (Tukey' s test) that the central and south sites are most similar in biological makeup. PSI fmdings also show that although there is low similarity among sites overall, there is general agreement in terms of biological makeup among sites even when temporal differences are included. 22 Community parameters Spearman's rank correlation coefficient comparison showed no significant differences for rank abundances of individual taxa among sites. Therefore, estimates of overall diversity and evenness measurements were calculated from combined site data. In general, patterns for overall diversity and evenness followed those for individual sites (Figures 18). Seasonal changes in diversity remained relatively consistent between sites, even though catch composition and relative abundance often differed dramatically. Overall diversity estimates were highest in July (H0 '=3.98) with similarly high values estimated for both the north (Hn'=3.45) and central sites CHc'=3.16) when the highest species richness and abundances were recorded. Highest values at the south site were recorded in May (H5'=2.34) when the lowest values of abundance and species richness for the survey occurred. Low overall values of diversity were observed in March, September and November, which differed substantially from individual values measured at the north and central sites. Evenness values at all sites were highest in May (Figure 19), possibly related to the low numbers of transient taxa entering the area, which affected catch composition in September and November. The domination of September and November catches by decapod shrimps and prawns is reflected in the low overall evenness values for those months CHsept'=0.310, HNov'=0.309). Low March values (HMar'=0.295) reflect the relatively large catch of both Liparis sp. and Latreutes laminirostris. Repressed evenness values in September resulted from the immigration ofT. curvirostris into the area, particularly at the north and central sites. The greatest disparity between sites occurred in November at the north site, with a dramatic drop in values at the south site, reflective of the disproportionately high catches of M. dalei there. 23 The mean number of taxa per tow for the survey was 12.2 ± 7.0 taxa, with a invertebrates to fish ratio of 2.3:1 taxa per tow. Changes in the mean number of taxa per tow occurred both spatially and temporally, with highest values generally found at the north site (Figure 20). Highest overall values were in July when a combined total of 57 fish and invertebrate taxa were present in the sampling area, suggesting a seasonal response to abiotic and biotic changes, and coincident with increases in temperatures and possibly availability of prey . Lowest species richness values were seen in May, when only 27 taxa were captured. Surprisingly, the ratio of fish to invertebrates was virtually even during May, the only month in which invertebrates did not dominate in terms of species richness. 24 25 Discussion The low abundance and biomass of fishes and invertebrates captured in the nearshore waters of the western Yellow Sea adjacent to Jiaozhou Bay support previous evidence that the area has been impacted severely by overfishing. Analysis of the size frequency of theA. argentatus population in the survey area and casual observations on reproductive status show that this commercially important fish is demonstrating characteristic signs of growth overfishing. According to the size:age relationship reported for A. argentatus by Lee and Park (1978), all fishes sampled for this survey and those sampled from a recent survey of fishes from Jiaozhou Bay belong to the 0-1 year age class (Wu and Wu, 1992). These results are consistent with earlier fmdings by Korean trawlers from 1970-75 in the Yellow Sea, which showed that all commercial catches of A. argentatus consisted primarily of fish under 4 years of age, with a mode of 0-1 year (Lee and Park, 1978). For larger specimens of A. argentatus examined for identification purposes during this study, well developed pre-spawn stage ovaries and testes were apparent, suggesting that these fish may be reproductively maturing within the first year. These data are strikingly similar to those reported for overfished stocks of the small yellow croaker (Pseudosciaena polyactis) and the large yellow croaker (P. crocea) in the Yellow Sea (Hwang, 1977; Mio, 1984; Tang, 1988; Zhang, et al, 1988; Yang, 1989; Yu, 1991). Findings are also consistent with those from several other areas known to be under intense fishing pressure (Regier and Loftus, 1972; Spangler, et al, 1977; Munro, 1983; Chan and Liew, 1986; Koslow, et al, 1988; Pauly, 1988a). The most obvious cause of early maturation, and the reduction of population and body size, is the targeting of spawning grounds for several species, including the large yellow croaker Pseudosciaena crocea and hairtail Trichiurus lepturus, which have now shown dramatic declines in catch biomass, mean body length and age, and relative proportions within the fish community (Tang, 1988; Zhang, et al, 1988; Yang, 1989). Although regulations have been imposed, and many of these fisheries have been closed or have become non-targets, there has been no definite sign that any of these stocks is recovering, suggesting a need for stricter regulations and enforcement Size frequency histograms for C. joyneri suggest that the effects of fishing pressure may not only be limited to commercially important fishes. Although information for age and growth of C.joyneri are not readily available, both mean (11.4 em) and maximum standard length (19.1 em) of specimens trawled for this survey were below the reported maximum of 30 em. Size distribution of this taxon from a recent survey of fishes from Jiaozhou Bay conforms to findings from nearshore waters C:Wu and Wu, 1992). However, several gravid females collected during the survey approached the upper limit of the known size distribution, indicating that this taxon may not be as heavily impacted as A. argentatus and Pseudosciaena sp. For the commercially important penaeid shrimps and prawns, the case for overfishing is not as clear. Size frequencies of the southern rough shrimp T. curvirostris and the Kishi velvet shrimp M. dalei indicate that a large percentage of the catch were adults, and most were large enough to be reproductively active (Zhang, 1990). Since catch statistics for these taxa were unavailable from this area, and recruitment among shrimps is often highly variable, it would be difficult from this limited survey to make a strong case for overfishing of either of these stocks. Additionally, since shrimp generally can maintain healthy stock levels from a relatively small spawning stock, and are sexually mature at a relatively young age (2 years oldforT. curvirostris), these taxa are likely to be less susceptible to changes brought about by intense fishing pressure than fishes (Zhang, 1990). It is noteworthy, however, that the main target of the Chinese shrimp fishery in this area, the fleshy prawn Penaeus orienta/is (=chinensis), was captured on only three 26 occasions for the entire survey with a total of only four adults taken (Table 10). Adults of P. orienta/is traditionally migrate in large nwnbers into the sampling area and the more northern Bohai Sea in April to spawn and disperse to the overwintering grounds in November. Thus, P. orienta/is would have been expected to appear in trawls in much higher numbers (Cheng, 1984). Although seasonal restrictions have also been placed on the P. orienta/is fishery, this survey suggests that stocks of this genus may be in trouble. A more detailed survey of the stocks of each of these taxa is necessary to determine their current fishery status. Recent acoustic surveys of pelagic ftshery stocks within the Yellow Sea suggests that the abundance of lower trophic level species, such as the anchovy Engraulis japonica, may be increasing and are now large enough to warrant expansion of the fishery (Iverson, et al, 1993). Until recently, these fisheries have been virtually unexploited and a recent estimate has put the potential annual yield of anchovy at 0.5 X 1()6 tons. Catch levels for major pelagic species was reported in 1988 to be between 3.0-33.0 X lo4 tons per year (Tang, 1988). If a shift towaril targeting pelagic stocks is made, pressure on demersal stocks may be somewhat reduced, which could improve the longevity of the Yellow Sea demersal fisheries. Comparison of results from this survey to trawling surveys from similar latitudes show that the epifaunal biomass and abundance densities in the nearshore waters of Shandong Province are relatively low. Interestingly, abundance levels are more similar to those of the deep sea than to the coastal habitats surveyed, and biomass levels for fishes are strikingly lower than any of the habitats compared (Table 15). Similar findings are shown for invertebrates, with lower levels of abundance at most sites compared, except from south of San Francisco, where abundance levels from the Yellow Sea were higher, although biomass was lower. Although each of these habitats are biologically and physically unique, and only a few studies were compared, abundances of macrofauna in 27 the Yellow Sea seem quite depressed. One reason may be the lower average primary productivity values characteristic of the survey area, which are estimated at 110-183 gC·m· 2.yrl. When compared to levels from coastal upwelling zones like the Califoruia coast, that can be up to 1000 gC·m-2-yrl, the lower values of benthic biomass from the Yellow Sea seem somewhat more reasonable. However, given the close proximity of the sites to nearby Jiaozhou Bay, with an annual average primary productivity of 154 gC·m-2yrl, one should expect a benthic biomass much higher than 0.8 kg/ha. For example, the Scotian shelf, with a primary productivity of 102 gC·m-2-yrl has a benthic biomass of 8.0 g·m-2 (80 kglha); benthic biomass from the Andaman Sea was estimated at 7.3 g·m·2 (73 kglha) and from the west coast of India at 5.3 g·m-2 (53 kglha), with productivity values estimated at 286 gC·m-2·yrl and 435 gC·m-2-yrl, respectively (Dwivedi, 1993). Indeed, by correlating global primary productivity values from Koblentz-Mishke (1970) to benthic biomass estimates from Zenkevitch, et al (1971) one can determine that benthic macrofaunallevels should range from at least 1-10 kg/ha. Certainly, comparisons of this type are problematic for a number of reasons, not the least of which are differences in sampling gear, environmental makeup and commuuity composition, and the inclusion/exclusion of a variety of different faunal groups within each of the respective analyses. However, by including several different surveys from various locations, this comparison should help to reinforce the fact that overfishing in the Yellow Sea is substantial, particularly when compared to areas where fishing pressure is less intense. Gear size may have contributed largely to both the low overall estimates of macrofauna! abundance and the high variability, particularly for fishes, within the area. One recent study within the deep sea using camera sled, in combination with otter and beam trawls, has shown that the use of trawls can severely underestimate fish abundance and biomass (Cailliet, et al, 1992). Certainly the small path width (3.4 m) of the trawl used for this survey could have allowed a relatively high degree of escapement, herding and/or 28 avoidance in comparison to larger nets (Ramm and Xiao, in press). Escapement may be particularly problematic for smaller taxa, such as certain gobiids, which could easily slip through the net mesh. Net avoidance could also have resulted in an underestimation of more mobile organisms, such as schooling fishes, given the low turbidity and shallow nature of the survey sites. In at least one study for net escapement, it was shown that extrusion of A. argentatus is minimal and occurs mostly in the net ceiling for smaller specimens and in the cod end for large specimens (Chow, et al, 1990). Unfortunately, no information was given in that report for avoidance or detection by this or any other taxon. Without comparative estimates using different net sizes, and evaluation of the response of individual taxon to approaching gear, it is impossible to gauge the accuracy of the results from this survey. However, a comparison of results of scientific surveys in nearby Jiaozhou Bay and casual observations of catches from local fishermen working in nearby areas confirms that the composition of the epifaunal assemblage was sampled fully and that density estimates should be viewed as at least the very minimum. Since the effects of herding and escapement, particularly as it applies to individual taxa, could not be determined, estimates assumed 100% effectiveness of the net in capturing fishes and invertebrates that were encountered. For shrimps and prawns, it has been shown that herding is less problematic and, as a result, estimates for this group may more accurately reflect real densities (Andrew, et al, 1991). It is suggested that because of the low areal abundances of macrofauna in this area and the high variability between tows, future surveys should incorporate a larger sampling unit, or at least compare effectiveness of various gears, to decrease variability and reduce potential avoidance by more mobile epibenthic organisms. Night trawling, which was deemed unsafe during the course of this survey, may also have helped to verify the exact nature of the epifaunal community composition and abundance, since die! catch rates have been shown to be substantially 29 different in some areas (Livingston, 1976; Ross, et al, 1987; Blaber, et al, 1990; Albert and Bergstad, 1993). Although there are several obvious problems associated with the use of small trawls in benthic surveys, estimates of benthic biomass from this survey (791.2 glha) should not be looked upon as altogether inaccurate. They are, however. dramatically lower than the 20-41 g!m2 (20-41 X lOS glha) benthic biomass value proposed by Tang (1988) for the South Yellow Sea. The rank importance of the faunal groups given by Tang (molluscs, echinoderms, polychaetes and crustaceans) is also much different than that found in nearshore waters during this study (crustaceans, fishes, echinoderms, molluscs and cnidarians), but since Tang included infauna in his estimates, a meaningful comparison of values is highly problematic. However, Tang's estimates do conform to those from a recent study in nearby Jiaozhou Bay that included infaunal organisms, such as polychaetes and molluscs, which may at least partially explain the discrepancy (Liu, et al, 1992). Given the small spatial scale of this study, i.e., total area of 48 km2, and the observation that large variability in biomass occurs within the area, larger-scaled surveys should be made before support is given to either estimate. Large variation in the abundance and biomass of epibenthic macrofauna! among sites suggests that some physical or biological characteristics are present in the north site that are responsible for the increase. Certainly, the close proximity of the north site to nearby Jiaozhou Bay, which has a reported average armual primary productivity of 154 mgC·m-2-d-1, could result in an increase in benthic biomass from transport into that area, in comparison to the central and south sites. Additionally, data suggest that differences in the grain size and composition of the sediments in these areas may be also least partially responsible for the differences. In studies on adult penaeids in the Gulf of Mexico and Western Australia, it was demonstrated that some species show diel burrowing patterns associated with specific 30 sediment types (Penn, 1981). ·Information on the burrowing habits ofT. curvirostris and M. dalei is scarce, but given the prevalence of T. curvirostris at the north site and M. dalei at the south site, data suggest that they may prefer specific sediment types. Penaeus orienta/is ( =chinensis) has been shown to be a non-burrowing, diurnal species which prefers highly turbid waters and muddy· bottom habitats characteristic of river mouths, which may explain its absence during the survey. A recent study of C. joyneri in Asan Bay, Korea found that abundance of this taxa was higher on fine sediment than on sandy sediment (Lee, 1993), which is consistent with findings from data from my survey. A comprehensive study of this area is necessary to determine which, if any, other physical characteristics are important in controlling population densities within these areas, or if other differences, alone or in combination, such as differences in food availability, competition and/or predation, or possibly even differences in nutrient load or pollution levels from nearby Jiaozhou Bay influence the faunal densities at these sites. Several taxa sampled during this survey are known to use Jiaozhou Bay as a temporary or permanent residence and probably move through the sampled areas on different occasions (Tian and Sun, 1992). Several species of sciaenid, such as A. argentatus, Johnius belengerii, Nibea albijlora, Collichthys lucidus and C. niveatus are known to use nearby Jiaozhou Bay as a temporary residence and as a spawning and nursery ground in warmer months. Generally, these fishes reside in the bay from April through November, after which time they move to deeper water. Argyrosomus argentatus spawns in Jiaozhou Bay from May through September, Nibea albijlora from May through August, and Collichthys niveatus from May through July. Information on C. lucidus was unavailable. For the most abundant taxon, A. argyrosomus, larger specimens first appeared in nearshore samples in July followed by a large increase in the number of young and juveniles in September, indicating that older fish probably leave the bay after spawning and are followed subsequently by their progeny in September and November (Figure 7). 31 Young and juvenile sciaenids trawled in November were not readily identifiable to generic level and may belong to one or more of the other taxa which reside in the bay. Taxa which have similar terms of residency within the bay include Apogon lineatus, Saurida elongata and Astroconger myriaster. Saurida elongata is known to spawn within the bay during June and July and Apogon lineatus from July through September. Other taxa, such as Liparis sp., utilize the bay for spawning from November and December, when temperature within the bay is reduced, and migrate into deeper water during summer. The capture of juvenile and young cyclopterids in large numbers during March and May samples suggests a prolonged residency in the bay for young and juveniles of this taxon. Larger adult cyclopterids, trawled ouly in November, were undoubtedly returning to the bay to begin the next seasons spawning. One other species which is known to follow this particular pattern of residency is Pholis fangi, whose capture was similar to that of Liparis sp. during the survey. Spawning data for this taxon was unavailable. Several resident taxa from Jiaozhou Bay occurred in samples in relatively stable numbers throughout the survey. These include Cynoglossus joyneri, Gobius pflaumi, Callionymus punctatus and Chaeturichthys hexanema. Juveniles of C. joyneri were most prevalent during March and May samples, although spawning is known to occur from June through September within the bay. Juvenile callionymids (Callionymus sp.) appeared only in November trawls, and young and juvenile gobiids, thought to be those of Gobius pflaumi were captured in September. Spawning periods for these two taxa are in summer months for Callionymus sp. and June for Gobius pflaumi. Other residents of Jiaozhou Bay captured only occasionally included Synechogobius hasta, Zoarces elongatus, Hexagrammos otakii, Cryptocentrus filifer and Pleuronichthys co mutus. Several of the more mobile invertebrates sampled during the survey also showed seasonal changes in abundance among sites that can also be related to traditional spawning and/or feeding migrations, most notably that of the southern rough shrimp T. curvirostris. 32 This commercially important shrimp has a distribution ranging from the Mediterranean and Red Sea through the Indo-Pacific to the northern Bohai Sea of China It migrates annually from overwintering grounds in the deeper waters into coastal areas of the Yellow Sea in spring and summer months where it spawns. A recent study has shown that this prawn spawns at the mouth of the Chiangjiang (Yangtze) River in May and June (Wu, et al, 1991). Findings here suggest that this shrimp enters the area to spawn in early July, which coincides with period of peak primary production, and begins its overwintering migration in November. The timing of the appearance of T. curvirostris coincides with a period of warm water during the summer monsoon season (Guan, 1984). This area represents the northernmost distributional boundary for this taxon (Holthuis, 1980). The Kishi velvet shrimp M. dalei has a more limited distribution than that ofT. curvirostris, but seems to follow a similar movement pattern, although specific information is lacking on M. dalei . Several other commercially important shrimp taxa, such as Latreutes laminirostris, Leptochela gracilis and Crangon affinis have wide-ranging depth distributions and appeared in high numbers at various times during the survey that did not conform to changes in water temperature or traditional peak periods of primary productivity within the area (Guo, 1994). Additional study is needed on these taxa to determine what factors influence abundance and distribution patterns in this area Capture of Octopus ace/latus and 0. variabilis occurred between March and July at which time these taxa may have been moving to and from spawning grounds inside the bay. These two species are known to move from deeper water to more shallow environs from March to late May or early June for spawning, during which time they are targeted for catch (Qi, et al, 1989). However, the sporadic capture and low sample size does not allow for any definite conclusions for these species. 33 Several factors affect the enforcement of fisheries regulations and protection of stocks in China. From observations and inquiries made during the course of this study, it seems that the majority of decisions related to enforcement occurs at the local level. In Qingdao, as is probably the case in most coastal fishing areas, fisheries cooperatives (which are much like local unions) wield a great deal of power and influence over regulatory agencies and the amount of enforcement that is enacted. Although the 1982 "Regulations of State Aquatic Resources" (National People's Congress, 1982) set several provisions for the protection of stocks by prohibiting catches of sexually immature animals, violations of these regulations were observed frequently during the course of this survey. Discard of any portion of the catch is virtually nonexistent and sexually immature individuals of many commercial species were opeuly displayed and sold both at the docks and markets. From an international perspective, attention has recently focused on increasing cooperation between China, Japan, and North and South Korea for coordination of fishing and conservation efforts within the Yellow Sea. Because at least ten of the major commercial fishes move annually across jurisdictional boundaries, a joint cooperation agreement and a multinational sharing of information among nations on fisheries-related issues is necessary for restoration and maintenance of the commercial stocks. Although these nations have historic and cultural similarities, discrepancies in economic conditions, internal political structure, and external economic and political alignments have prevented them from achieving meaningful cooperation on a variety of issues (Valencia, 1988; Yang, 1990; Yu, 1991). Several bilateral fishing agreements are currently in effect, including those between China and Japan, China and North Korea and Japan and South Korea, but recognition of these agreements by the nonaligned nations generally does not exist, and the agreements do not always cover all of the Yellow Sea resources. Moreover, China and South Korea, although cooperating in some regional economic ventures, still have no formal dialogue between governments. The imposition of economic exclusion zones (EEZs), which has proven useful in many instances globally, is impossible within the Yellow Sea due to its limited size and the close proximity of bordering nations. In spite of these problems, North Korea has already recently established an EEZ along its western coastline. In addition, in 1989, South Korea announced plans to extend its fishing activities, possibly into areas protected under a previous Sino-Japanese agreement, which will inevitably increase the potential for future conflict (Yang, 1989). And Japan, although it has no borders with the Yellow Sea, is allowed through their 1972 agreement with China, to send at least 400 vessels into the area (Yang, 1989). Several recent suggestions have been both forwarded by scientists and international agencies regarding implementation of conservation measures and expansion of cooperation between nations in the areas of fishing and sharing of scientific data. One suggestion is the establishment of an independent agency and/or research body, that would implement meaningful, large-scale scientific investigations of Yellow Sea fisheries in order to establish conservation and fishing guidelines that could be used by all nations to set restrictions on fishing within the Yellow Sea. This process might reduce or eliminate future conflict and may help to restore stocks to sustainable levels. Data from this survey, when combined with recent historical data on catch biomass trends and changes in community composition, provide overwhelming evidence that the Yell ow Sea is at or entering a state of ecosystem overfishing. 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Preliminary study on the population structure of Trachypenaeus curvirostris (Stimpson) in Huanghai and Bohai Seas. Acta Ecol. Sin. (2):145-149. (in Chinese) 43 Figure 1: Map of the study site. The north, central and south sites are shown, along with major cities and islands. Isobathic contours are shown for the Yellow Sea and Jiaozhou Bay. Sediment samples were taken in each of the 1 km2 'blocks' marked by an "X". Insert map shows the area of the Yellow Sea surrounding the sample site and the bordering seas and nations. 44 35r------, El Taxa per tow: pilot study OMean no. taxa per tow: maffi study 30 El 25 20 15 10 El El El 5 El Tow length (min) Figure 2: Results of the pilot study from tows made in February and March 1993. Points represent individual tows and include both fishes and invertebrates. Mean and standard deviation number of taxa per tow (20.2±10.2 taxa) for the maffi study is represented by the oval and refletts the high degree of variability which occurred between individual tows throughout the study. 45 50 40 30 20 10 March May 1993 t 1993 0 40 "' "' = 30 ·;:::-5 [!l ·o... 20 r/l=- 10 July t 1993 1993 0 ~ 1 2 3 4 5 6 40 Trawl number 30 20 ~ ~North 10 -<>-Central November t 1993 --south 0 1 2 3 4 5 6 Trawl number Figure 3: Cumulative species curves for samples collected March through November 1993 at nearshore stations, PR China. Data points represent actual numbers of taxa trawled and were randomly selected from each data set Note that the November plot has a maximum number of five tows. Data include both fish and invertebrate taxa. Individual curves in most cases begin to show levelling at approximately three tows (indicated by arrows), suggesting that both the number and duration of tows used in the survey satisfactorily characterize the epibenthic macrofauna! assemblage in the nearshore waters of Qingdao. 46 35~------, 30 c:: ·p0 25 ·~"';> 0) '1:) 20 '1:)~ § +I-"' 15 § 0) ::E 10 5 1-a- Xtemp • X sal Month Figure 4: Mean monthly salinity and temperature values from samples taken from March through November 1993. March values are derived from the literature. Temperature is listed in degrees Celsius. Salinity 0 is listed in parts per thousand ( / 00). Family Sciaenidae Cynoglossidae Gobiidae Cyclopteridae 10 30 10 20 30 40 50 Percent composition Figure 5: Relative percent contribution of fish famiies by site for all samples collected March through November 1993 at nearshore station, Yellow Sea, PR China. Sciaenidae dominated overall catches at all sites and were represented by six taxa. Other major families contributing to catches include in rank order, Cynoglossidae with one taxon, Cyclopteridae represented by at least one taxon, and Gobiidae with six taxa. Other families contributing at least 1% to the total abundance include Apogonidae and Callionymidae, each with a single taxon. :!3 Taxon South All othertaxa Pholis fangi (P) Ammodytes sp. (Am) Gobius pflaumi (G) Callionymus punctatus (Ca) Apogon lineatus (A) Liparis sp. (Cy) Cynoglossus joyneri (C) Argyrosomus argentatus (S) 40 50 Percent composition Figure 6: Percent composition of total abundance by site for fishes collected in samples from March through November 1993 in nearshore waters, Yellow Sea, PR China. Each of the individual taxon shown contributed~ 1% of the total abundance for a site. The sciaenid taxon Argyrosomus argentatus clearly dominated catches at all sites, with major contributions by the cynoglossid Cynoglossus joyneri at both the central and south sites. The cyclopterid taxon Liparis sp. was the second most dominant fish at the north site overall. Abundances of the gobiid Gobi us pflaumi were disproportionately higher at the south site, as were other minor taxa. Proportions were calculated from total catches of fishes only. .J:>. 00 Taxon Central South Cantherhines septentrionalis Pleuronichthys cornutus (G) Gobius pflaumi (G) Chaeturichthys hexanema (G) Callionyrnus punctatus Saurida elongata Astroconger myriaster Synechogobius hasta Apogon lineatus Liparis sp. Cynoglossus joyneri Argyrosomus argentatus Percent composition Figure 7: Percent composition of total biomass by site for fishes collected in samples from March though November 1993 in nearshore waters, Yellow Sea, PR China. Each taxon shown contributed;:: 1% of the total biomass for the study. At all sites the sciaenid Argyrosomus argen(atus was the major contributor. Other major biomass contributors were Cynoglossus joyneri and Liparis sp, with all other taxa accounting for a relatively minor proportion of the total biomass. Biomass contributions by all taxa were most evenly distributed at the north site where the highest species richness was found for all sites. ~ 50 200 180 North 160 140 120 100 80 ., 60 ~ 40 "'., 20 .=..., 0 !:~< 180 fll Central 'a 160 "C= 140 ·;: :a 120 100 ·~= ...0 80 ...... 60 "5 40 = 20 r..l= til 0 +I = 180 South ..= 160 ~ 140 120 100 80 60 40 20 -f m = $---I 0 .<:: >. >. ... t <1) ~ ::E= .....,-::l ,I:J ,I:Js <1) ::E ~ ;> !:~< <1) 0 Cll z Month Figure 8: Mean fish abundance for samples collected March through November 1993 at nearshore stations, Yellow Sea, PR China. Error bars represent one standard error. 51 1000 900 North 800 700 600 500 400 300 200 100 0 "'t;; 900 Central 't 800 ..="'... 700 "'~ 600 rll § 500 ;, 400 r..l 300 1:/l 200 +I § 100 0 ~ 900 800 South 700 600 500 400 300 200 100 0 ...... <:: Figure 9: Mean fish biomass for samples collected March through Novemer 1993 at nearshorestations, Yellow Sea, PR China. Error bars represent one standard error. 52 250~------~------, Mean=4.98±3.23 em SD t:::] November TotalN=531 D September ll!:l July 200 ~ ;:I 't:l 150 ·~> ·~ 't:l.s 0 ""':;; e 100 ~ 50 Standard length (em) Figure 10: Size frequency distribution for Argyrosomus argentatus collected in samples taken from March through November 1993 in the nearshore waters of the western Yellow Sea, PR China. Two size modes are represented and all individuals collected were young-of the year. 53 35,------, 121 November Mean=11.43±0.24 em SE 1111 September Total N=239 j 30 I<] July '*l 1111 May .g 25 0 March ·-;> ·-'"0 .s..... 20 0 til .c ~ 15 10 5 0 Standard length (em) Figure 11: Size frequency distribution for Cynoglossus joyneri collected in samples from March through November 1993 at nearshore stations, Yellow Sea, PR China. Arrows indicate that five, or possibly even seven, size modes were represented by individuals taken during the survey. Figure 12: Overall comparison of biomass, abundance, and frequency of occurrence values for each of the major dominant fishes and invertebrates caught during the March-November 1993 trawling survey in the nearshore waters of Shandong Province, China. Frequency of occurrence values range from 0 to 100%, and are shown as the combined width of abundance and biomass bars. Ranks fro abundance and biomass of each taxon are in parentheses. Solid bars represent biomass values; open bars represent abundance values. ~'''~~m'~-~'''''''''''~-''''''_' __ '''-"'~''' ~- _,,~~- ''''~'''~ Taxonomic group North Central South Crabs Stomato pods Caridean shrimp· Dendrobranch shrimP" 0 10 20 30 40 50 60 70 80 10 20 30 40 50 60 70 80 10 20 30 40 50 60 70 80 Percent composition Figure 13: Percent abundance contribution of major invertebrate groups collected in samples from March through November 1993 at nearshore stations, Yellow Sea, PR China. Dendrobranchiate shrimps and prawns were the most dominant crustacean group at all sites, consistently contributing over 50% of the total abundance, followed in rank abundance by the caridean shrimps. Another dominant crustacean group, the stomatopod, represented by the single taxon Sguilla oratoria, was a major abundance contributor at the central site, in particular. Molluscs, echinoderms, and annelids (not shown) were minor contributors to abundance totals at all sites. lJl lJl Taxon Parapenaeopsis tenella Crangon affmis Leptochela gracilis Acetes chinensis Latreutes laminirostris Metapenaeopsis dalei Trachypenaeus curvirostris 0 10 w w ~ ~ 00 0 10 w w ~ ~ 00 0 10 w w ~ ~ 00 w Percent composition Figure 14: Percent abundance contribution of dendrobranchiate and caridean shrimps and prawns collected in samples from March through November 1993 at nearshore stations, Yellow Sea, PR China Clear shifts in dominance among these taxa occurred among sites, with Trachypenaeus curyirostris dominanting catches at the north and central sites, but replaced by Metapenaeopsis dalei at the south site. One resident taxon, Latreutes Iaminirostris. was a major contributor at both the north and central sites, and surpassed M. .dllli:i in abundance overall at those sites. Highest evenness among these taxa was demonstrated at the central site. l1\ 0\ Taxon North Central South Rapana sp. (M) Octopus variabilis (M) Carcinoplax vestitus (C) Portunus trituberculata (C) Charybdis bimaculata(C) Charybdis japonica (C) Temnopleurus toreumaticus(E) Temnopleurus hardwickii (E) Metapenaeopsisdalei (D) Trachypenaeuscurvirostris (D) Squilla oratoria (S) Asterias rollestoni (A) 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 50 Percent composition Figure 15: Percent biomass contribution of all invertebrate taxa contributing. Ut -l 58 1400 North 1200 1000 800 600 400 ~ ...... 200 -=.. 0 ~ 1400 Central ~ .; 1200 :s "CC 1000 :a-~ 800 .•= 600 'Cl ~ 400 200 e:s 0 f;l;l= !:ll 1400 +I South Iii 1200 ~ 1000 800 600 400 200 0 ..c: ...... ~ .Q .., .., ~ ;:;s .c ;:;s ~ s s.., ~ fr > Cl) Month z Figure 16: Mean invertebrate abundance for samples collected March through November 1993 at nearshore stations, Yellow Sea, PR China. Error bars represent one standard error. 59 2000.------, 1800 North 1600 1400 1200 1000 800 600 400 200 0+---~~--~------~------~------~ 1800 Central 1600 1400 1200 1000 800 600 400 200 0+------~ 1800 South 1600 1400 1200 1000 800 600 400 200 0~--~----~------~------~ ..c >. ..,...... ,.... u ~ ,.0 ~ :;a -~ s El.., :;a B :> .., 0 CZl""' z Month Figure 17: Mean invertebrate biomass for samples collected March through November 1993 at nearshore stations, Yellow Sea, PR China. Error bars represent one standard error. 60 5 North 4 3 2 1 -Q-Overall H' --a--Mean± SD H' 0 =0 Central ..::= ·;:"' 4 ~ "0 3 til "0 ! 2 +I"' ; 1 ~"' 0 South 4 3 2 1 0 ...... <:: :>.. Figure 18: Monthly mean diversity by site for samples collected from March through November 1993 at nearshore stations, Yellow Sea, PR China. Overall diversity (H'), calculated from all samples combined, is included for reference and reflects the trends shown for most sites in a given month. Error bars represent one standard deviation. T l I 61 I I 1.0 ,------=------:=-----, ' 0.9 North 0.8 0.7 0.6 0.5 0.4 0.3 0.2 -a--Overall J' 0.1 -o-Mean± SD J' 0+---~~--~--~--~------~~--~--~ 0.9 Central 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0+---~~--~------~------~--~ 0.9 South 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0~--~~--~------~------~--~...... <:: :>. Figure 19: Monthly mean eveness by site for samples collected from March through November 1993 at nearshore stations, Yellow Sea, PR China. Overall evenness (J') calculated from all samples combined, is included for reference and reflects trends for most sites in a given month. Error bars represent one standard deviation. 62 50 45 illil North D Central 40 .~~! South 35 ....""t;j 'S.. 30 1"' 25 z= 20 15 10 5 0 ~ >. :2 '3,_, Month Figure 20: Monthly mean number of taxa caught per tow in samples taken March through November 1993 at nearshore stations, Yellow Sea, PR China. Values for the north site in November were calculated for two tows only, possibly resulting. in low mean values for that month. Error bars represent one standard deviation. Table 1: Distribution of tows taken between March and November 1993 at nearshore stations in the western Yellow Sea Note that a maximum of five tows for samples taken in November and a total of only two tows for the north site during that month due to equipment loss. 64 Species/taxon Percent Rank Percent Rank abundancf biomass Trachypenaeus curvirostris (S) 29.1 1 13.3 3 Metapenaeopsis dalei (S) 22.4 2 3.8 6 Latreutes larninirostris (S) 17.0 .. 3 0.4 - Squilla oratoria (MS) 4.3 4 19.2 2 Leptochela gracilis (S) 3.7 5 0.1 - Argyrosomus argenteus (F) 3.7 6 9.5 4 Crangon affmis (S) 3.3 7 0.1 - Acetes chinensis (S) 2.7 8 0.0 - Liparis sp. (F) 2.2 9 1.9 9 Cynoglossus joyneri (F) 1.4 10 5.9 5 Asterias rollestoni (A) 1.2 11 24.8 1 Parapenaeopsis tenella (S) 0.9 12 0.1 - Compsometra serrata (CR) 0.8 13 0.0 - Charybdis bimaculata (C) 0.7 14 1.3 - Gobius pflaurni (F) 0.6 15 0.2 - Temnopleurus hardwickii (U) 0.4 - 3.5 7 Charybdis japonica (C) 0.1 - 2.7 8 Temnopleurus toreumaticus (U) 0.2 - 1.6 10 Apogon lineatus (F) 0.3 - 0.9 11 Zoarces elongatus (F) 0.0 - 0.7 12 Rapana sp. (M) 0.0 - 0.7 13 Portunus trituberculata (C) 0.0 - 0.7 14 Carcinoplax vestitus (C) 0.3 - 0.6 15 Totals 94.7 88.4 Table 2: Overall rank percent biomass and abundance for dominant taxa taken in trawls between March and November 1993 at nearshore stations, Yellow Sea, PR China. A= asteroid; C= crab; CR= crinoid; F= fish; M= mollusc; MS= mantis shrimp; S= shrimp or prawn; U= urchin. 65 Species/taxon %N Mean SE FO % Total Mean SE no./ha. no./ha biomass !!/ha <>/ha Argyrosomus argentabls (S) 38.26 11.21 4.61 0.42 37.99 71.83 19.23 Liparis sp. (Cy) 23.10 6.77 ··2.72 0.31 7.56 14.30 7.63 Cynoglossus joyneri (C) 14.24 4.17 0.75 0.69 27.33 51.66 10.41 Gobius pflaumi (G) 5.92 1.74 0.65 0.36 0.80 1.52 0.46 Apogon lineabls (A) 3.64 1.07 0.29 0.21 3.53 6.68 1.72 Callionymus punctabJS (Ca) 3.48 1.02 0.32 0.24 1.33 2.52 0.97 Arumodytes sp. (Am) 3.15 0.92 0.32 0.16 0.25 0.48 0.16 Chaeturichthys hexanema (G) 1.20 0.35 0.10 0.20 1.59 3.01 0.80 Pholis fangi (P) 1.03 0.30 0.15 0.07 0.55 1.03 0.56 Johnius belengerii (S) 0.92 0.27 0.11 0.11 2.29 4.34 1.93 Syngnathus acus (Sy) 0.87 0.25 0.09 0.11 0.22 0.41 0.19 Cryptocentrus f"tlifer (G) 0.60 0.18 0.09 0.07 0.63 1.20 0.64 Callionymidae (C) 0.54 0.16 0.10 0.04 0.00 0.00 0.00 Zoarces elongabls (Z) 0.49 0.14 0.06 0.09 2.98 5.64 2.40 Astrocongermyriaster (Co) 0.43 0.13 0.05 0.09 2.09 3.95 1.70 Pseudosciaena polyactis (S) 0.33 0.10 0.05 0.05 1.25 2.37 1.58 Synechogobius basta (G) 0.27 0.08 0.03 0.06 2.26 4.27 2.14 Tridentigernudiventris(G) 0.27 0.08 O.D3 0.06 0.01 0.01 0.01 Collichthys lucidus (S) 0.16 0.05 0.05 0.01 0.25 0.47 0.47 Hippocarnpusjaponicus (Sy) 0.16 0.05 0.04 0.03 0.00 0.00 0.00 Sauridaelongata(Sd) 0.16 0.05 0.03 0.04 1.66 3.14 2.21 Minous monodactylus (Sc) 0.11 0.03 0.02 O.D3 0.27 0.51 0.48 Ctenopauchen chinensis (G) 0.11 0.03 0.02 0.03 0.00 0.00 0.00 Gobiidae(G) 0.11 0.03 0.03 0.01 0.00 0.00 0.00 Pleuronichthys comubls (PI) 0.11 0.03 0.02 0.03 1.99 3.76 2.84 Cantherhines septentrionalis (B) 0.11 0.03 0.02 0.03 1.94 3.67 2.62 Sciaenidae(S) 0.05 0.02 0.02 0.01 0.00 0.00 0.00 Nibea albiflora (S) 0.05 0.02 0.02 0.01 0.25 0.47 0.47 Collichthys niveabls (S) 0.05 0.02 0.02 0.01 0.00 0.00 0.00 Hexaorammos otakii (H) 0.05 0.02 0.02 0.01 0.45 0.85 0.85 Totals 100.00 29.30 5.84 100.00 189.1 31.88 Table 3: Taxon-speciltc abundance, biomass and frequency of occurrence ofltshes capblred in all samples collected between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled; SE=standard error of the mean; FO=frequency of occurrence. A=Apogonidae;Am=Arumodytidae;B=Balistidae;C=Cynoglossidae;Co=Congridae;Ca= Callionymidae; Cy=Cyclopteridae; G=Gobiidae;H=Hexagrammidae; P=Pholidae; PI= Pleuronectidae;S=Sciaenidae;Se=Scorpaenidae;Sd=Synodontidae;Sy=Syngnathidae; Z=Zoarcidae. 66 Species/taxon %N Mean SE FO % Total Mean SE no./ha no.lha biomass <>/ha !!/ha Argyrosomus argentarus (S) 41.01 24.82 14.20 0.44 38.80 226.50 121.69 Liparis sp. (Cy) 31.20 18.89 8.37 0.44 10.98 67.38 38.69 Cynoglossus joyneri (C) 7.76 4.70 122 0.76 2051 121.91 62.76 Gobius pflaumi (G) 2.39 1.44 0.49 0.48 0.47 7.28 4.98 Apogon lineatus (A) 2.39 1.44 0.54 0.32 2.83 20.80 9.76 Callionymus punctarus (Ca) 3.67 2.22 0.86 0.48 2.46 18.68 8.80 Arnmodytes sp. (Am) 4.43 2.68 0.95 0.40 0.42 7.03 4.96 Chaetnrichthys hexanema (G) 0.26 0.15 0.09 0.12 0.55 7.73 5.10 Pholis fangi (P) 1.53 0.93 0.48 0.20 0.90 9.75 5.59 J ohnius belengerii (S) 0.60 0.36 0.17 0.20 2.94 21.39 10.82 Syngnathus acus (Sy) 1.02 0.62 0.27 0.24 0.38 6.78. 4.95 Cryptocentrus filifer (G) 0.94 0.57 0.28 0.24 1.21 11.54 6.10 Callionymidae (C) 0.17 0.10 0.10 0.04 0.00 4.61 4.88 Zoarces elongatns (Z) 0.43 0.26 0.17 0.12 2.16 16.94 8.90 Astrocongermyriaster(Co) 0.60 0.36 0.14 0.24 3.19 22.85 11.09 Pseudosciaena polyactis (S) 0.43 0.26 0.15 0.12 1.96 15.84 8.59 Synechogobius basta (G) 0.26 0.15 0.09 0.12 1.91 15.55 8.23 Tridentigernudiventris (G) 0.17 0.10 O.D7 0.08 0.00 4.63 4.87 Collichthys lucidus (S) 0.00 0.00 0.00 0.00 0.00 4.61 4.88 Hippocampus japonicus (Sy) 0.17 0.10 0.10 0.04 0.00 4.62 4.88 Sauridaelongata(Sd) 0.09 0.05 0.05 0.04 1.62 13.89 8.17 Minous monodactylus (Sc) 0.09 0.05 0.05 0.04 0.03 4.78 4.87 Ctenopauchen cbinensis (G) 0.09 0.05 0.05 0.04 0.00 4.62 4.88 Gobiidae(G) 0.00 0.00 0.00 0.00 0.00 4.61 4.88 Plenronicbthys cornutus (PI) 0.09 0.05 0.05 0.04 2.64 19.72 11.88 Cantherbines septentrionalis (B) 0.09 0.05 0.05 0.04 2.16 16.98 10.08 Sciaenidae(S) 0.00 0.00 0.00 0.00 0.00 4.61 4.88 Nibea albiflora (S) 0.00 0.00 0.00 0.00 0.00 4.61 4.88 Collicbthys niveatus (S) 0.09 0.05 0.05 0.04 0.00 4.61 4.88 Hexaorammosornkiinn 0.09 0.05 0.05 0.04 0.86 9.50 5.90 Totals 100.00 60.53 16.22 100.00 576.46 300.3 4 Table 4: Taxon-specific abundance, biomass and frequency ofoccurrence offtsbes caprured in samples at the north site between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled; SE=standard error of the mean; FO=frequency of occurrence. A=Apogonidae;Am=Arnmodytidae;B=Balistidae;C=Cynoglossidae;Ca=Callionymidae; Cn=Congridae; Cy=Cyclopteridae; G=Gobiidae;H=Hexagrammidae; P=Pbolidae; PI= Plenronectidae; S=Sciaenidae; Se=Scorpaenidae; Sd=Synodontidae; S y=Syngnathidae; Z=Zoon:idae. 67 Species/taxon %N Mean SE FO % Total Mean SE no./ha no./ha biomass 1!/ha 1!/ha Argyrosomus argentatus (S) 37.62 5.25 1.91 0.43 37.58 48.13 27.24 Liparis sp. (Cy) 11.22 1.57 .. 0.70 0.25 1.38 1.77 1.10 Cynoglossus joyneri (C) 24.75 3.46 1.06 0.54 38.40 49.19 14.34 Gobius pflaumi (G) 5.28 0.74 0.27 0.36 0.58 0.74 0.30 Apogon lineatus (A) 10.89 1.52 0.64 0.25 7.76 9.94 3.80 Callionymus pnnctatus (Ca) 3.30 0.46 0.37 0.11 0.09 0.11 0.09 Armnodytes sp. (Am) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Chaeturichthys hexane rna (G) 2.64 0.37 0.16 0.21 3.08 3.94 1.76 Pholis fangi (P) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Johnius belengerii (S) 0.66 0.09 0.06 O.D7 0.43 0.55 0.54 Syngnathus acus (SY) 0.99 0.14 0.10 0.07 0.06 O.D7 0.05 Cryptocentrus filifer (G) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Callionymidae (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Zoarces elongatus (Z) 0.99 0.14 0.08 0.11 6.21 7.95 5.05 Astrocongermyriaster (Co) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pseudosciaena polyactis (S) 0.33 0.05 0.05 0.04 0.96 1.23 1.23 Synechogobius basta (G) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TridentigernudivenUis (G) 0.33 0.05 0.05 0.04 0.01 0.01 0.01 Collichthys lucidus (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Hippocarnpusjaponicus (Sy) 0.33 0.05 0.05 0.04 0.00 0.00 0.00 Sauridaelongata(Sd) 0.33 0.05 0.05 0.04 0.00 0.00 0.00 Minous monodactylus (Sc) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Ctenopauchen chinensis (G) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Gobiidae(G) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pleuronichthys comutus (PI) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Cantherhines septenUionalis (B) 0.33 0.05 0.05 0.04 3.46 4.43 4.44 Sciaenidae(S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Nibea albiflora (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Collichthys niveatus (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 HexaQrammos otakii (H) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Totals 100 13.96 2.93 100.00 128.08 39.38 Table 5: Taxon-specific abnndance, biomass and frequency of occurrence of fishes captured in samples at the central site between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled; SE=standard error of the mean; FO=frequency of occurrence. A=Apogonidae;Am=Armnodytidae;B=Balistidae;C=Cynoglossidae;Ca=Callionymidae;Co= Congridae; Cy=Cyclopteridae; G=Gobiidae;H=Hexagrammidae; P=Pholidae; Pl=Pleuronectidae; S=Sciaenidae;Sc=Scorpaenidae;Sd=Synodontidae;Sy=Syngnathidae;Z=Zoarcidae 68 Species/taxon %N Mean SE FO % Total Mean SE no./ha no.lha biomass P"/ha 1!/ha Argyrosomus argentams (S) 29.95 5.02 3.08 0.36 38.19 51.45 21.47 Liparis sp. (Cy) 6.87 1.15 0.63 0.25 6.02 8.11 7.36 Cynoglossus joyneri (C) 26.37 4.42 1.58 0.79 31.55 42.51 19.08 Gobius pflaumi (G) 17.86 2.99 1.80 0.21 1.70 2.29 1.19 Apogon lineams (A) 1.65 0.28 0.20 0.07 1.37 1.85 1.06 Callionymus punctaUJs (Ca) 3.02 0.51 0.26 0.14 0.12 0.16 O.D7 Ammodytes sp. {Am) 1.65 0.28 0.17 0.11 0.12 0.17 0.14 Chaeturichthys hexanema (G) 3.02 0.51 0.21 0.25 2.50 3.37 1.20 Pholis fangi (P) 0.27 0.05 0.05 0.04 0.30 0.40 0.41 Johnius belengerii (S) 2.20 0.37 0.26 0.07 2.65 3.57 2.54 Syngnathus acus (Sy) 0.27 0.05 0.05 0.04 0.02 0.02 0.02 Cryptocentrus filifer (G) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Callionymidae (C) 2.20 0.37 0.26 0.07 0.00 0.01 0.00 Zoarces elongams (Z) 0.27 0.05 0.05 0.04 2.18 2.93 2.28 Astrocongermyriaster(Co) 0.27 0.05 0.05 0.04 1.64 2.21 2.25 Pseudosciaena polyactis (S) 0.00 0.00 0.00 0.00 0.15 0.20 0.20 S ynechogobius basta (G) 0.55 0.09 0.06 O.D7 4.93 6.64 4.72 Tridentigernudiventris (G) 0.55 0.09 0.06 O.D7 0.01 0.01 0.01 Collichthys lucidus (S) 0.82 0.14 0.14 0.04 0.97 1.31 1.33 Hippocampusjaponicus (Sy) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Sauridaelongata(Sd) 0.27 0.05 0.05 0.04 3.18 4.28 4.36 Minous monodactylus (Sc) 0.27 0.05 0.05 0.04 0.99 1.33 1.36 Ctenopauchen chinensis (G) 0.27 0.05 0.05 0.04 0.00 0.00 0.00 Gobiidae(G) 0.55 0.09 0.09 0.04 0.01 0.01 0.01 Pleuronichthys comums (PI) 0.27 0.05 0.05 0.04 2.37 3.20 3.26 Cantherhines septenlrionalis (B) 0.00 0.00 0.00 0.00 0.53 0.72 0.73 Sciaenidae(S) 0.27 0.05 0.05 0.04 0.01 0.01 0.01 Nibea albiflora (S) 0.27 0.05 0.05 0.04 0.97 1.30 1.33 Collichthys niveams (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Hexavrammos otakii fl.n 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Totals 100.00 16.77 5.28 100.00 134.74 40.21 Table6: Taxon-specific abundance, biomass and frequency ofoccurrence of fishes captured in samples at the south site between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled; SE=standard error of the mean; FO=frequency of occurrence. A=Apogonidae;Am=Ammodytidae;B=Balistidae; C=Cynoglossidae; Ca=Callionymidae; Co= Congridae; Cy=Cyclopteridae; G=Gobiidae; H=Hexagrammidae; P=Pholidae; Pl=Pleuronectidae; S=Sciaenidae;Se=Scorpaenidae;Sd=Synodontidae;Sy=Syngnathidae;Z=Zoarcidae 69 Species/taxon March May July Sept Nov R/T Argyrosomus argenteus XX xxxx XX T Liparis sp. XXX XX X T Cynoglossus joyneri XX XX XXX XX XX R Gobius pflaumi X X X X XX R Apogon lineabJs XX XX T Callionymus punctatus X X XX XX X R Arnmodytes sp. XX X X T ChaebJrichthyshexanema X X X X R Pholis fangi X XX X X R Johnius belengerii X X X R Syngnathus acus X X X X R Cryptocentrus filifer X X T Callionyrnidae XX Zoarces elongabJs X X T Astrocongermyriaster X X X T Pseudosciaenapolyactis X X T Synechogobius basta X X T Tridentigernudiventris X T Collichthys lucidus X T Hippocampus japonicus X T Sauridaelongata X T Minous monodactylus X T Ctenopaucben cbinensis X X R Gobiidae X Pleuronicbthys cornubJs X X R Cantherbines septentrionalis X T Sciaenidae X Nibea albiflora X T Collichthys nivearus X T HexagrammosotU Table 7: Monthly trends of relative ftsh abundance as determined from samples collected between March and November 1993 at nearshore stations, Yellow Sea, PR China. X= 1-9 individuals; XX= 10-99 individuals; XXX= 100-499 individuals; XXXX > 500 individuals. Species richness totals for eacb monthly sample are shown above. R=Resident, T=transient (based on catch and natural history data) ---- ~~~"""'~~~"""' ·~ -~'"'"-~·~·~~ ~·~~~~"~' '''~'''''''''""'''~~··-·--1 Fish Site North Central South Site Month 3 5 7 9 11 3 5 7 9 11 3 5 7 9 11 Month 11 O.D7 0.15 0.51 0.12 0.74 0.10 0.37 0.46 0.33 0.74 0.14 0.36 0.46 0.32 11 9 0.05 0.11 0.47 0.77 0.31 O.Dl 0.05 0.40 0.92 0.44 0.12 0.18 0.38 0.43 9 s 7 0.07 0.09 0.69 0.28 0.35 0.08 0.02 0.77 0.42 0.41 0.12 0.43 0.26 0.20 7 s 5 0.22 0.13 0.44 0.03 0.19 0.12 0.06 0.44 0.20 0.22 0.24 0.20 0.27 0.23 5 3 0.82 0.75 0.15 0.03 0.18 0.82 0.48 0.12 0.12 0.12 0.23 0.06 0.03 0.03 3 11 0.05 0.12 0.47 0.24 0.69 0.09 0.36 0.41 0.40 0.13 0.30 0.33 0.56 0.63 11 9 0.05 0.06 0.49 0.79 0.27 0.08 0.02 0.44 0.16 0.05 0.05 0.15 0.53 0.09 9 c 7 0.07 0.09 0.79 0.32 0.34 0.07 0.01 0.20 0.46 0.14 0.37 0.52 0.43 0.25 7 c 5 0.47 0.52 0.02 0.00 0.39 0.57 0.42 0.09 0.43 0.41 0.65 0.25 0.39 0.33 5 3 0.82 0.77 0.09 0.03 0.12 0.14 0.12 0.05 0.13 0.98 0.26 0.05 0.03 0.02 3 11 0.12 0.16 0.35 0.20 0.08 0.40 0.35 0.14 0.78 0.10 0.27 0.30 0.48 0.73 11 9 0.05 0.04 0.33 0.08 0.02 0.02 0.13 0.75 0.09 0.02 0.01 O.D7 0.45 0.07 9 N 7 0.10 0.12 0.10 0.31 0.14 0.32 0.58 0.18 0.37 0.33 0.26 0.42 0.32 0.18 7 N 5 0.79 0.32 0.02 0. 70 0.18 0.58 0.41 0.08 0.59 0.21 0.46 0.28 0.42 0.72 5 3 0.19 0.15 0.00 O.D7 0.83 0.34 0.14 0.01 0.07 0.83 0.16 0.06 0.01 0.03 3 Month 3 5 7 9 111 3 5 7 9 111 3 5 7 9 11 Month ! Site North I Central I South Site Invertebrates Table 8: Mean percent similarity index by month and site combinations. Numbers represent mean PSI values for all possible tow-tow combinations within each respective month and site. Fish and invertebrates are treated separately. Values abnve 50% are highlighted for reference. Significance levels are set at 80%. cl Fish Site North Central South Site North 0.324 ± 0.283 0.309 ± 0.264 North Central 0.304 ± 0.256 0.332 ± 0.263 Central South 0.289 + 0.238 0.316 + 0.239 South Site North Central South Site Invertebrates Table 9: Overall mean and standard deviation percent similarity index (PSI) values for fish and invertebrate samples collected March through November 1993 at nearshore stations, Yellow Sea, PR China. Means were calculated by summing the values from all possible tow-to-tow combinations for all sample months. Although overall mean values are not significantly similar according to the arbitrarily set 80% PSI value, numbers indicate a moderate level of agreement in composition and relative contribution of individual taxa among sites, given the large temporal variation shown in month-to-month comparisons (see Table 8). Values shown here also reinforce results of group analysis using Tukey's test, in which central and south sites consistently grouped together in terms of relative abundances of fish and invertebrate groups. --..! 72 Table 10: Taxon-specific abundance, biomass and frequency of occurrence of invertebrates captured in samples collected between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled, SE= standard error of the mean, FO= frequency of occurrence. A= asteroid; C= crab; Ce= cephalaspidean; CR= crinoid; I= isopod; M=mollusc; MS= mantis shrimp; N= nudibranch; No= notaspidean; P= polychaete; S= shrimp or prawn; U= urchin; X= pycnogonid. 73 Species/taxon %N Mean SE FO % Total Mean SE no./ha. no./ha. biomass /ha /ha Trachypenaeuscurvrrostris(S) 32.19 89.25 34.17 0.73 16.98 102.22 25.38 Metapenaeopsisdalei (S) 24.74 68.59 28.18 0.85 5.27 31.73 9.60 Latreutes laminirostris (S) 18.82 52.19 24.93 0.95 0.49 2.98 1.52 Squillaoratoria (MS) 4.81 13.33 . 2.60 0.85 24.98 150.41 30.95 Leptochela gracilis (S) 4.08 11.32 2.68 0.69 0.12 0.71 0.19 Crangon affmis (S) 3.69 10.24 3.10 0.57 0.17 1.01 0.49 Acetes chinensis (S) 2.94 8.14 2.66 0.57 0.03 0.19 0.06 Asterias rollestoni (A) 1.31 3.63 0.65 0.89 32.49 195.61 31.50 Parapenaeopsis tenella (S) 1.00 2.77 1.14 0.19 0.19 1.16 0.50 Compsometra serrata (CR) 0.87 2.40 1.13 0.14 0.01 0.04 0.03 Charybdis bimaculata (C) 0.78 2.15 0.59 0.51 1.63 9.79 2.89 Temnopleurus hardwickii (U) 0.42 1.16 0.33 0.41 4.83 29.10 7.82 Palaemon sp. (S) 0.36 0.99 0.33 0.27 0.03 0.17 0.07 porcellanidcrab (C) 0.35 0.97 0.28 0.33 0.00 0.03 0.01 limpet (Type A) (M) 0.30 0.84 0.64 0.06 0.01 0.06 0.04 Carcinoplax vestitus (C) 0.31 0.86 0.27 0.23 0.78 4.71 1.87 Pleurobranchaeanovazealandia(No) 0.27 0.75 0.26 0.25 0.52 3.11 1.00 Alpheus japonicus (S) 0.24 0.67 0.16 0.32 0.11 0.66 0.23 Alpheus distinguendus (S) 0.24 0.67 0.24 0.25 0.35 2.12 0.89 Palaemon gravieri (S) 0.20 0.56 0.32 0.14 0.12 0.71 0.36 lnquisitorflavidula (M) 0.20 0.54 0.12 0.28 0.22 1.30 0.32 Temnopleurus toreurnaticus (U) 0.20 0.54 0.16 0.27 2.06 12.42 3.50 Diogenes edwardsii (C) 0.17 0.46 0.20 0.14 0.13 0.76 0.48 anemones (A) 0.17 0.46 0.23 0.07 0.03 0.19 0.11 Charybdis japonica (C) 0.16 0.45 0.18 0.19 3.48 20.93 8.43 flabelliferan isopod (1) 0.11 0.30 0.08 0.21 0.01 0.04 0.02 Cirolanajaponica (1) 0.10 0.29 0.20 0.05 0.01 0.07 0.05 Alpheus sp. (S) 0.09 0.25 0.13 0.07 0.01 0.03 0.03 Metapenaeusjoyneri(S) 0.09 0.24 0.09 0.12 0.15 0.89 0.39 Pyrene marteusi (M) 0.08 0.22 0.16 0.06 0.00 0.01 0.00 Heptacarpus rectirostris (S) 0.07 0.19 0.14 0.05 0.01 0.06 0.04 Dorippe granulata (C) 0.07 0.21 0.08 0.12 0.01 0.85 0.36 Dendronotusarborescens(N) 0.06 0.16 0.09 0.06 0.00 0.02 0.01 hippid crab (C) 0.06 0.16 0.07 0.09 0.00 0.01 0.01 Patiria pectinifera (A) 0.05 0.13 0.05 0.11 0.73 4.39 1.95 nudibranch (Type A) (N) 0.05 0.13 0.07 0.06 0.00 0.01 0.00 Okenia plana (N) 0.04 0.11 0.06 0.06 0.01 0.03 0.02 Lambros validus (C) 0.03 0.10 0.05 0.06 0.23 1.39 0.81 majid crab (C) 0.03 0.08 0.05 0.05 0.00 0.01 0.01 Octopus ocellatus (M) 0.03 0.08 0.03 0.07 0.39 2.38 1.25 Rapana sp.. (M) O.D3 0.08 0.04 0.06 0.89 5.38 3.88 Sakuraeolis enosirnensis (N) 0.02 0.06 0.05 0.02 0.00 0.01 0.01 pycnogonid(X) 0.02 0.06 0.03 0.06 0.00 0.00 0.00 Octopus variabilis (M) 0.02 0.06 0.03 0.06 0.92 5.55 3.26 74 Species/taxon %N Mean SE FO %Total Mean SE noJha. no.lha. biomass Plha Plha Penaeus orientalis (S) 0.02 0.06 0.04 0.05 0.35 2.13 1.27 Aphrodita australis (P) 0.02 0.05 . 0.04 0.02 0.06 0.38 0.29 Sternaspis scutata (P) 0.02 0.05 0.03 0.05 0.00 0.00 0.00 Annina compta (N) 0.02 0.05 O.D3 0.05 0.00 0.01 0.01 Philine kinglipini (Ce) 0.02 0.05 0.03 0.05 0.02 0.13 0.10 Dimorpbostylis asiatica (S) 0.02 0.05 0.03 0.05 0.00 0.00 0.00 Portunus trituberculata (C) 0.01 0.03 0.03 0.01 0.89 5.37 5.34 Luidia quinaria (A) 0.01 0.03 0.02 0.02 0.11 0.63 0.53 Ceritbidea cingulata (M) 0.01 0.02 0.02 0.01 0.00 0.00 0.00 Arcania undecimsninos~ !Cl O.DI 0.02 0.02 0.01 0.02 0.15 0.1 'i Totals 100.00 277.25 50.73 99.87 602.06 73.97 75 Table 11: Taxon-specific abundance, biomass and frequency of occurrence of invertebrates captured at tbe north site in tows made between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled; SE=standard error of tbe mean; FO=frequency of occurrence. A= asteroid; C=erab; Ce= cephalaspidean; CR=erinoid; !=isopod; M=mollusc; MS=mantis shrimp; N=nudibranch; No=notaspidean; P= polychaete; S=shrimp or prawn; U=urchin; X=pycnogonid 76 Species/taxon %N Mean SE FO % Total Mean SE noJha no./ha biomass /ha /ha Trachypenaeus curvirostris (S) 47.50 214.19 104.54 0.48 27.00 172.62 70.02 Metapenaeopsis dalei (S) 5.24 23.63 11.14 0.56 2.23 14.26 5.53 Latreutes laminirostris (S) 28.70 129.41 78.16 0.84 1.19 7.59 4.78 Squilla aratoria (MS) 2.47 11.15 3.18 0.68 15.66 100.12 32.87 Leptochela gracilis (S) 5.13 23.12 6.84 0.80 0.23 1.45 0.51 Crangon affinis (S) 1.24 5.57 2.07 0.60 0.30 1.94 1.54 Acetes chinensis (S) 2.12 9.55 4.58 0.44 0.02 0.14 O.D7 Asterias rollestoni (A) 0.62 2.79 0.57 0.84 19.73 126.10 24.90 Parapenaeopsis tenella (S) 0.26 1.19 0.69 0.16 0.09 0.54 0.30 Compsometra serrata (CR) 1.60 7.22 3.49 0.28 0.02 0.12 0.10 Charybdis bimaculata (C) 0.47 2.12 1.02 0.32 1.62 10.34 5.25 Temnopleurushardwickii (U) 0.43 1.96 0.93 0.36 5.76 36.81 20.26 Palaemon sp. (S) 0.08 0.36 0.14 0.24 0.04 0.24 0.21 porcellanidcrab(C) 0.29 1.29 0.68 0.28 0.01 0.05 0.04 limpet (Type A) (M) 0.60 2.68 2.04 0.12 0.03 0.17 0.12 Carcinoplax vestitus (C) 0.13 0.57 0.42 0.12 0.38 2.45 1.87 Pleurobranchaeanovazealandia(No) 0.05 0.21 0.12 0.12 0.15 0.95 0.64 Alpheus japonicus (S) 0.17 0.77 0.28 0.32 0.10 0.64 0.22 Alpheus distinguendus (S) 0.10 0.46 0.18 0.28 0.17 1.11 0.50 Palaemon gravieri (S) 0.34 1.55 1.02 0.16 0.24 1.51 1.08 Inquisitorflavidula (M) 0.10 0.46 0.23 0.16 0.20 1.30 0.70 Temnopleurus toreumaticus (U) 0.29 1.29 0.46 0.36 4.31 27.54 10.02 Diogenes edwardsii (C) 0.27 1.24 0.62 0.24 0.17 1.12 0.93 anemones (A) 0.14 0.62 0.39 0.12 0.04 0.23 0.18 Charybdisjaponica (C) 0.26 1.19 0.55 0.28 9.26 59.20 25.52 llabelliferan isopod (I) 0.10 0.46 0.18 0.24 0.02 0.10 0.06 Cirolana japonica (I) 0.21 0.93 0.65 0.12 0.03 0.22 0.14 Alpheus sp. (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Metapenaeus joyneri (S) 0.01 0.05 0.05 0.04 0.07 0.47 0.47 Pyrene martensi (M) 0.02 0.10 0.10 0.04 0.00 0.00 0.00 Heptacarpusrectirostris (S) 0.13 0.57 0.43 0.08 0.03 0.18 0.13 Dorippe granulata (C) 0.14 0.62 0.25 0.28 0.37 2.38 1.10 Dendronotusarborescens (N) 0.09 0.41 0.30 0.08 0.01 0.07 0.05 hippid crab (C) 0.11 0.52 0.22 0.24 0.01 0.04 0.02 Patiria pectinifera (A) 0.06 0.26 0.13 0.16 1.69 10.79 5.87 nudibranch (Type A) (N) 0.08 0.36 0.20 0.12 0.00 0.02 0.01 Okenia plana (N) 0.05 0.21 0.16 0.08 0.01 0.05 0.04 Lambros validus (C) 0.05 0.21 0.14 0.08 0.45 2.90 2.20 majid crab (C) 0.05 0.21 0.16 0.08 0.00 0.03 0.03 Octopus ocellatus (M) 0.05 0.21 0.10 0.16 0.67 4.27 2.24 Rapana sp. (M) 0.05 0.21 0.12 0.12 2.70 17.23 12.33 Sakuraeolis enosimensis (N) 0.05 0.21 0.16 0.08 0.01 0.04 0.03 pycnogonid (X) 0.02 0.10 0.07 0.08 0.00 0.00 0.00 Octopus variabilis (M) 0.01 0.05 0.05 0.04 1.27 8.10 8.10 77 Species/taxon %N Mean SE FO %Total Mean SE noJha noJha biomass Plha ""'a Penaeus orientalis (S) 0.02 0.10 0.10 0.04 0.51 3.28 3.28 Aphrodita australis (P) 0.03 0.15 0.11 0.08 0.19 1.22 0.92 Sternaspis scutata (P) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Armina compta (N) 0.01 0.05 0.05 0.04 0.00 0.02 0.02 Philine kinglipiui (Ce) 0.01 0.05 0.05 0.04 0.00 0.00 0.00 Dimorphostylis asiatica (S) 0.01 0.05 0.05 0.04 0.00 0.00 0.00 Portunus trituherculata (C) 0.02 0.10 0.10 0.04 2.69 17.19 17.19 Luidia quiuaria (A) 0.01 0.05 0.05 0.04 0.26 1.67 1.67 Cerithidea cingulata (M) 0.01 0.05 0.05 0.04 0.00 0.00 0.00 Arcauia undecimsninosa fC\ 0.01 0.05 0.05 0.04 O.D7 0.47 0.47 Totals 100 450.93 122.20 100.00 639.30 264.53 1 I 78 Table 12: Taxon-specific abundance, biomass and frequency of occurrence of invertebrates captured at the central site in tows made between March and November 1993 at nearshore stations, Yellow Sea, PR China. N= total number sampled; SE=standard error of the mean; FO=frequency of occurrence. A=asteroid; C=crab; Ce=eephalaspidean; CR=erinoid; I=isopod; M=mollusc; MS=mantis shrimp; N=nudibranch; No=notaspidean; P= polychaete; S=shrimp or prawn; U=urchin; X=pycnogonid 79 Species/taxon %N Mean SE FO % Total Mean SE no./ha no./ha biomass /ha /ha Trachypenaeus curvirostris (S) 29.09 45.24 21.89 0.61 9.04 64.16 22.74 Metapenaeopsisdalei (S) 12.53 19.49 5.38 0.75 2.19 15.55 5.47 Latreutes Iaminirostris (S) 17.21 26.77 13.28 0.79 0.19 1.36 0.72 Squilla aratoria (MS) 12.14 18.89 6.37 0.79 32.36 229.78 72.71 Leptochela gracilis (S) 3.35 5.21 1.85 0.61 0.05 0.33 0.12 Crnngon affinis (S) 4.35 6.77 2.58 0.43 O.D7 0.53 0.16 Acetes chinensis (S) 7.26 11.29 6.35 0.50 0.04 0.29 0.17 Asterias rollestoni (A) 3.67 5.71 1.69 0.79 43.28 307.32 76.57 Parapenaeopsis tenella (S) 1.13 1.75 1.14 0.18 0.09 0.63 0.30 Compsometra serrata (CR) 0.33 0.51 0.39 O.D7 0.00 0.00 0.00 Charybdis bimaculata (C) 1.90 2.95 1.27 0.50 1.80 12.81 6.07 Temnopleurushardwickii (U) 0.71 1.11 0.34 0.43 5.87 41.67 12.22 Palaemon sp. (S) 0.92 1.43 0.76 0.21 0.03 0.19 0.09 porcellanid crab (C) 0.80 1.24 0.52 0.32 0.00 0.03 0.01 limpet (Type A) (M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Carcinoplax vestitus (C) 0.68 1.06 0.43 0.25 0.87 6.17 2.59 Pleurobrnnchaeanovazealandia(No) 0.62 0.97 0.62 0.18 0.54 3.84 2.03 Alpheus japonicus (S) 0.62 0.97 0.38 0.29 0.17 1.17 0.64 Alpheus distinguendus (S) 0.80 1.24 0.64 0.18 0.69 4.89 2.51 Palaemon gravieri (S) 0.09 0.14 0.08 0.11 0.05 0.36 0.25 Inquisitor flavidula (M) 0.36 0.55 0.21 0.25 0.17 1.17 0.49 Temnopleurus toreumaticus (U) 0.15 0.23 0.10 0.18 0.97 6.92 3.05 Diogeues edwardsii (C) 0.06 0.09 0.09 0.04 0.00 0.00 0.00 anemones(A) 0.18 0.28 0.28 0.04 0.00 0.00 0.00 Charybdis japonica (C) 0.09 0.14 0.08 0.11 0.26 1.88 1.20 flabellifernn isopod (I) 0.06 0.09 0.06 0.07 0.00 0.01 0.01 Cirolana japonica (I) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Alpheus sp. (S) 0.27 0.41 0.29 0.11 O.Dl 0.08 0.08 Metapenaeus joyneri (S) 0.15 0.23 0.12 0.14 0.12 0.87 0.46 Pyrene martensi (M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Heptacarpus rectirostris (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Dorippe grnnulata (C) 0.03 0.05 0.05 0.04 0.04 0.31 0.23 Dendronorusarborescens (N) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 hippid crab (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Patiria pectinifera (A) 0.03 0.05. 0.05 0.04 0.17 1.23 1.21 nudibranch (Type A) (N) 0.03 0.05 0.05 0.04 0.00 0.00 0.00 Okenia plana (N) 0.09 0.14 0.10 0.07 0.01 0.05 0.04 Lambrusvalidus(C) 0.06 0.09 0.06 O.D7 0.20 1.45 1.28 rnajid crab (C) 0.03 0.05 0.05 0.04 0.00 0.00 0.00 Octopus ocellatus (M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Rapana sp. (M) O.D3 0.05 0.05 0.04 0.00 0.00 0.00 Sakuraeolis enosiroensis (N) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 pycnogonid(X) 0.06 0.09 0.06 0.07 0.00 0.00 0.00 Octopus variabilis (M) 0.06 0.09 0.06 0.07 0.70 4.95 4.86 80 Species/taxon %N Mean SE FO %Total Mean SE noJha no.lha biomass g/ba .g/ba Penaeus orientalis (S) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Aphrodita australis (P) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Sternaspis scutata (P) 0.03 0.05 0.05 0.04 0.00 0.00 0.00 Armina compta (N) 0.03 0.05 0.05 0.04 0.00 0.01 0.01 Philine kinglipini (Ce) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Dimorphostylis asiatica (S) 0.03 0.05 0.05 0.04 0.00 0.00 0.00 Portunus tritnherculata (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Luidia quinaria (A) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Cerithidea cingulata (M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Arcania undecimspinosa (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Totals 100.00 155.54 30.00 100 710.03 218.26 81 Table 13: Taxon-specific abundance, biomass and frequency of occurrence of invertebrates captured at the south site in tows made between March and November 1993 at nearshore stations, Yellow Sea, PR China. N=total number sampled; SE= standard error of the mean; FO=frequency of occurrence. A=asteroid; C=crab; Ce= cephalaspidean; CR=crinoid; !=isopod; M=mollusc; MS=mantis shrimp; N= nudibranch; N o=notaspidean; P=polychaete; S=shrimp or prawn; U =urchin; X= pycnogonid l ! lj l 82 Species/taxon %N Mean SE FO % Total Mean SE no./ha no./ha iomass /ha /ha Trachypenaeus curvirostris (S) 8.90 21.70 9.89 0.61 16.37 76.07 28.65 Metapenaeopsis dalei (S) 64.71 157.84 78.94 0.75 13.54 62.92 25.93 Latreutes laminirostris (S) 3.55 8.66 3.29 0.68 0.09 0.42 0.15 Squilla oratoria (MS) 3.99 9.72 2.74 0.61 25.56 118.77 42.64 Leptochela gracilis (S) 2.83 6.91 3.78 0.29 0.09 0.42 0.25 Crangon affinis (S) 7.33 17.88 8.31 0.43 0.14 0.66 0.26 Acetes chinensis (S) 1.53 3.73 1.64 0.43 0.03 0.13 0.06 Asterias rollestoni (A) 0.94 2.30 0.56 0.57 32.26 149.93 39.76 Parapenaeopsis tenella (S) 2.13 5.21 3.02 0.11 0.48 2.23 1.37 Compsometra serrata (CR) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Charybdis bimaculata (C) 0.57 1.38 0.70 0.39 1.37 6.38 3.58 Temnopleurus hafdwickii (U) 0.21 0.51 0.26 0.18 2.17 10.09 5.22 Palaemon sp. (S) 0.45 1.11 0.59 0.21 0.02 0.11 0.05 porcellanidcrab(C) 0.17 0.41 0.18 0.21 0.00 0.00 0.00 limpet (Type A) (M) 0.02 0.05 0.05 0.04 0.00 0.01 0.01 Can:inoplax vestitns (C) 0.38 0.92 0.55 0.18 1.15 5.33 4.48 Pleurobranchaeanovazealandia(No) 0.42 1.01 0.43 0.29 0.93 4.32 1.98 Alpheus japonicus (S) 0.11 0.28 0.12 0.18 0.04 0.20 0.12 Alpheusdistinguendus(S) 0.11 0.28 0.15 0.14 O.D7 0.35 0.19 Palaemon gravieri (S) 0.04 0.09 0.06 0.07 O.D7 0.33 0.23 Inquisitor flavidula (M) 0.25 0.60 0.20 0.29 0.31 1.42 0.53 Temnopleurus toreurnaticus (U) 0.08 0.18 0.09 0.14 0.91 4.21 2.02 Diogenes edwardsii (C) 0.06 0.14 0.10 0.07 0.25 1.18 1.12 anemones (A) 0.21 0.51 0.51 0.04 0.07 0.33 0.29 Charybdisjaponica (C) 0.04 0.09 0.06 O.D7 1.11 5.14 3.57 flabelliferanisopod (1) 0.15 0.37 0.16 0.21 0.00 0.02 0.01 Cirolana japonica (1) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Alpheus sp. (S) 0.13 0.32 0.24 0.07 0.00 0.02 0.01 Metapenaeusjoyneri (S) 0.17 0.41 0.24 0.11 0.28 1.29 0.96 Pyrene martensi (M) 0.23 0.55 0.46 0.11 0.00 0.01 0.01 Heptacarpus rectirostris (S) 0.02 0.05 0.05 0.04 0.00 0.02 0.02 Dorippe granulata (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Dendronotnsarborescens(N) 0.04 0.09 0.06 0.07 0.00 0.00 0.00 hippid crab (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Patiria pectinifera (A) 0.04 0.09 0.06 0.07 0.37 1.72 1.26 nudibranch (Type A) (N) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Okenia plana (N) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Lambrus validus (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 majid crab (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Octopus ocellatns (M) 0.02 0.05 0.05 0.04 0.64 2.98 2.98 Rapaua sp. (M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Sakuraeolis enosimensis (N) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 pycnogonid(X) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Octopus variabilis (M) 0.02 0.05 0.05 0.04 0.83 3.86 3.86 83 Species/taxon Mean SE IFO 1":' Total Mean SE noJha noJba hnomass Wh a Wba Penaeus oneritiillsW T.U4 u.u~ u.uo I U.U/ U.M ::l.l~ :l.l~ Aphrodita australis (P) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Stemaspis scutata (P) 0.04 0.09 0.06 O.D7 0.00 0.00 0.00 Armina compta (N) 0.02 0.05 0.05 0.04 0.00 0.00 0.00 Philine kinglipini (Ce) 0.04 0.09 0.06 0.07 0.08 0.37 0.28 DimoiJJhostylis asiatica (S) 0.02 0.05 0.05 0.04 0.00 0.00 0.00 Portnnus trituberculata (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Luidia quinaria (A) 0.02 0.05 0.05 0.04 0.07 0.32 0.32 Cerithidea cingulata (M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Arcania undecimspinosa (C) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Totals 11oo.oo 243.90 87.89 1 1 too 464.69 174.36 84 Table 14: Monthly trends of relativeinvertebrate abundance as determined from samples collected between March and November 1993 at nearshore stations, Yellow Sea, PR China. X= 1-9 individuals; XX= 10-99 individuals; XXX=100-999 individuals; XXXX= 500-999 individuals; XXXXX=1000-2000 individuals; XXXXXX=2000-7000 individuals. R=Resident taxa, T=Transient taxa (based on catch and natural history information) 85 S ecies/taxon March Ma Nov R/T Trachypenaeus curvirostris XXX XXX XXX XXX T Metapenaeopsis dalei XXX XXX XXX XX XXX R Latreutes laminirostris xxxxxx XX XXX X XX R Squilla oratoria X XX XXX XXX XXX R Leptochelagracilis XXX X XXX XX XXX R Crangon affinis X XX xxxx X XX R Acetes chinensis XX X XXX XX R Asterias rollestoni XX XX XX XX XX R Parapenaeopsis tenella X XXX T Compsometraserrata X XXX X T Charybdis birnaculata XX XX XX X R Temnoplenrus hanlwickii X XX XX XX R Palaemon sp. X X XX X porcellanid crab XX XX X XX R limpet (Type A) XX X R Carcinoplax vestitns XX X X T Plenrobranchaea novazealandia X X XX X R Alpheus japonicus XX X XX X X R Alpheusdistinguendus X X XX X X R Palaemon gravieri XX X T fuquisitorflavidula X XX XX XX R Temnoplenrus torenrnaticus X X X X XX R Diogenes edwardsii XX XX R anemones XX X R Charybdisjaponica X XX X T flabelliferan isopod X X X R Cirolana japonica XX T Alpheus sp. X X R Metapenaeusjoyneri X XX T Pyrene martensi X X XX R Heptacarpus rectirostris XX T Dorippegranulata XX X T Dendronotns arborescens X X R hippidcrab X X T Patiria pectinifera X X X X R nudibranch (Type A) X R Okenia plana X R Lambros validus X X T majidcrab X X X T Octopus ocellatns X X R Rapanasp. X X R Sakuraeolisenosirnensis X R Ascorhyncusrarnipes X X R Paranymphon magnidigitum X R Octopus variahilis X X R Penaeus orienlalis X X T Aphrodita australis X R 86 Species/taxon March May July September November !Rtf Stemaspis scutata X X R Annina compta X X R Pbiline kinglipini X X R Dimorphostulis asiatica X R Portunus trituberculata X T Lnidia quinaria X T Cerithidea cingulata X R Arcania undecims inosa X T Species richness 31 14 42 31 28 87 Table 15a: Comparative summary of results of the nearshore survey of the Yellow Sea near Jiaozhou Bay, Shandong Province, PR China, to results of several recent coastal and deep-sea macrofauna! surveys. The table shows that abundance levels for fishes in the Yellow Sea survey area are only slightly higher than those for deep sea areas along the western and eastern seaboards of the United States. Abundance levels for invertebrates are similar to those from at least one study from south of San Francisco, California. This comparison is not meant tu suggest that these areas are physically or biologically eqnivalent, but is merely shown as a reference to other areas and studies. Converted estimates were made whenever possible from original estimates based on net mouth width, boat speed, and trawling times from each of the individual surveys. Results are given as numbers of individuals or biomass (in kilograms) per hectare. ·····~·~ Type Location Habitat Depth Sampling unit Orlg. abundance ~onv. abundance References estimates estimates Shandong Province, coastal !6-28 m 4.9 m otter trawl 29±6 29 ± 6 Rhodes, this study PR Chinn indivlha indiv/ha Fnrallon lslaods, deep-sea 2, 000 - 12.1 m otter trawl 14 14 Cnilliet, et nl, 1992 San Franciso, California 3,200 m 2.1 m beam trawl indivlha indivlha Farallon Islands, deep-sea 2,000- camera sled 207 207 Cnilliet, et nl, 1992 San Franciso, California 3,200 m indivlha indivlha F Narragansett Bay, semi -enclose to 40m 9.2 m balloon 117±151 289 ± 373 pviatt and Nixon, 1973 I Rhode Island bay otter trawl indiv/acre indivlha s Oxnard to San Diego, coastal 10-49 m 7.6 m otter trawl 173±33 415 ± 79 Allen and Voglin, 1976 H Southern California indiv/tow indiv/ba Porcupine Bight deep-sea 2, 200 - 8.6 m semi-balloon 5 indiv/ 50 Merrett, et a~ 1991 NW Atlantic, New England 2, 400m otter trawl 1000 sq. m indiv/ba Orange County, California coastal 18-90 m 7.5 m otter trawl 469±26 500 ± 27 Mearns, 1977 indiv/baul indiv/ha Southwest Ocean Outfall, San coastal 56-92 m 7.6 m otter trawl 246±132 590 ± 317 CH2M Hill, 1980 Francisco, California indiv/haul indivlha Shandong Province, coastal 16-28 m 4.9 m otter trawl 277±51 277 ±51 Rhodes, this study PR Chinn indivlha indiv/ha I I Santa Monica Bay Control, coastal 30-60m 7.6 m otter trawl 1018±977 2027 ± 1944 SCCWRP, 1982 N Southern California indivlhaul indiv Ibn I v Snntn Monica Bay Outfall coastal 30-60m 7.6 m otter trawl 1565±757 3114± 1506 SCCWRP, 1982 E Southern California indiv/haul indiv/ha R Palos Verdes coastal 30-60m 7.6 m otter trawl 669±316 !331 ± 629 SCCWRP, 1982 i T Southern California indiv/hnul indiv/ba s Southwest Ocean Outfall, San coastal 56-92 m 7.6 m otter trawl 72±68 173 ± 163 CH2M Hill, 1980 Francisco, California indiv/haul indiv/ha Fnrnllon Islands, deep~sea 2, 000- camera sled, beam 255 ± 772.6 2550 ± 77,260 Nybnkken, et nl, 1992 San Franciso, California 3,200 m and otter trawl indiv/100 SQ. m indiv/ha 00 00 l 1 89 Figure 15b: Comparative summary of results of nearshore survey of the Yellow Sea near Jiaozhou Bay, Shan dong Province, PR China, to results of several recent coastal and deep-sea macrofauna! surveys. The table shows that biomass levels of fishes in the western Yellow Sea are at least one order of magnitude lower than those from any of the other areas compared. Biomass levels for invertebrates are similarly low, although abundance levels were lower for invertebrates surveyed at the site south of San Francisco, California. This comparison is not meant to suggest that these areas are biologically or physically equivalent, but is merely shown as a reference to other areas and studies. Converted estimates were made whenever possible from original estimates based on net mouth width, boat speed, and trawling times from each of the individual surveys. Results are given as numbers of individuals or biomass (in kilograms) per hectare. .... ~•. ~ Type Location Habltnt Depth Sampling unit Orlg. biomass Conv. biomass References estimates estimates Shandong Province, coastal 16-28 m 4.9 m otter trawl 189 ± 32 0.2 ± 0.0 Rhodes, this study PR China g/ba kg/ha Farallon Islands, deep-sea 2, 000 - 12.1 m otter trawl N/A N/A Cnillie~ et al, 1992 San Franciso, California 3,200 m 2.1 m beam trawl Farallon Islands, deep-sea 2, 000 - camera sled N/A N/A Caillie~ et al, 1992 San Franciso, California 3,200 m F Narragansett Bay, semi-enclosed to 40m 9.2 m balloon 28.5 ± 16 31.7 ± 17.8 Oviatt and Nixon, 1973 I Rhode Island bay otter trawl lbs/acre kg/ha s Oxnard to San Diego, coastal 10-49 m 7.6 m otter trawl 7.1 ± 1.4 12.1 ± 2.4 Allen and Voglin, 1976 H Southern California kg/tow kg/ha Porcupine Bight deep-sea 2, 200 - 8.6 m semi-balloon 1.7 17 Merrett, et al, 1991 NW Atlantic, New England 2, 400 m otter trawl kg/1000 sq. m kg/ha Orange County, California coastal 18-90 m 7.5 m otter trawl N/A N/A Meams, 1977 Southwest Ocean Outfall, San coastal 56-92 m 7.6 m otter trawl 176.5 ± 18.75 kg/ 12.6 ± 1.4 CH2M Hill. 1980 Francisco, California bimonthly sample kg/ha Shandong Province, coastal 16-28 m 4.9 m otter trawl 602.0 ± 74.0 0.6 ± 0.1 Rhodes, this study PR China g/ha kg/ha I Santa Monica Bay Control, coastal 30-60m 7.6 m otter trawl N/A N/A SCCWRP, 1982 N Southern California v Santa Monica Bay Outfall coastal 30-60m 7.6 m otter trawl 19.6 ± 4.6 335 ± 7.9 SCCWRP, 1982 E Southern California kg/haul kg/ha R Palos Verdes coastal 30-60m 7.6 m otter trawl 9.7 ± 2.4 16.6 ± 4.1 SCCWRP, 1982 T Southern California kg/haul kg/ha s Southwest Ocean Outfall, Sau coastal 56-92 m 7.6 m otter trawl 36.32 ± 20.66 kg/ 2.6 ± 1.5 CH2M Hill, 1980 Francisco, California bimonthly sample kg/ha Farallon Islands, deep-sea 2, 000 - camera sled, beam NIA N/A Nybakken, et al, 1992 San Frnnciso, California 3,200 m and otter trawl ------\0 0 91 Appendix A: Text of the State Ministry of Agriculture's 'Regulations on the Protection of Aquatic Resources (National People's Congress of the People's Republic of China, 1982) - (1) Protection shall be extended to all aquatic animals and plants of economic and scientific importance, including their parent bodies, larvae, eggs and spores, and to the aquatic habitats which sustain their growth and reproduction. (2) Allowable catch shall ensure the sustainable yield. Therefore, standards for allowable catch shall be formulated in terms of length or weight limits, on a species-by-species basis, and in terms of the ratio of the catch in keeping with the standards to the catch of those smaller than the standard. Adequate numbers of parent fish must be allowed to survive. Ouly sexually mature aquatic animals can be harvested. (3) Seasonal closures, areal closures or gear restrictions shall be applied, as necessary, to certain important spawning grounds, feeding grounds, or overwintering grounds. Different size limits shall be effected in harvesting different species of fish. Any fishing methods which act to the detriment of fishery resources must be eliminated. Explosives and electrofishing are forbidden. (4) Penalties against violation include compensation, fmes and confiscation of harvests and/or gears. Administative, disciplinary, civil and criminal punishments may be enforced as appropriate. (5) Protection of aquatic product resources shall be administered by the State Aquatic Product Bureau attached to the Ministry of Agriculture, and by the provincial and the local fishery administrations. The establishment of a fishing license system and registration of fishing vessels and operations shall be actively pursued. 92 Appendix B: List of Fishes Collected from Nearshore Stations, Yellow Sea, PR China, March through November 1993 Family Congridae Astroconger myriaster (Brevoort, 1856) Family Synodontidae Saurida elongata (Temminck et Schlegel, 1846) Family Syngnathidae Syngnathus acus (Linne', 1758) Hippocampus japonicus (Kaup) Family Scomaenidae Minous monodactylus (Cuvier et Valenciennes, 1829) Family Hexagrammidae Hexagrammos otakii (Steller in Tilesius, 1809) Family Cyclopteridae Liparis sp. (Rose, 1793) Family Apogonidae Apogon lineatus (Temminck et Schlegel, 1842) Family Sciaenidae Collichthys lucidus (Richardson, 1844) Collichthys niveatus (Jordan et Starks, 1906) Nibea albijlora (Richardson, 1846) Johnius belengeri (Cuvier, 1839) Pseudosciaena polyactis (Bleeker, 1877) Argyrosomus argentatus (Houttuyn, 1782) Family Zoarcidae Zoarces elongatus (Kner, 1868) Family Pholididae Pholis fangi (Wang et Wang, 1935) Family Ammodytidae Ammodytes sp. (Linne', 1758) Family Callionymidae Callionymus punctatus (Ochiai et al, 1955 93 Family Gobiidae Tridentiger nudiventris (Tomiyama, 1934) Gobius pflaumi (Bleeker, 1853) Cryptocentrusfilifer (Valenciennes, 1837) Synechogobius hasta (Temminck et Schlegel, 1845) Chaeturichthy hexanema (Bleeker, 1853) Ctenotrypauchen chinensis (Steindachner, 1867) Family Pleuronectidae Pleuronichthys comutus (Girard, 1854) Family Cynoglossidae Cynoglossus joyneri (Gunther, 1878) ·.~~ Month N df Fa Pa Fb Pb Ft Pt March 17 2 22.66 < 0.001 1.19 ns 21.97 < 0.001 May 17 2 17.12 < 0.001 17.79 < 0.001 16.82 < 0.001 July 18 2 16.64 < 0.001 6.04 0.049 17.7 < 0.001 September 17 2 16.98 < 0.001 16.28 < 0.001 20.84 < 0.001 November 6 2 1.14 ns 2.57 ns 1.54 ns Appendix C: Summary table of results from Kruskal-Wallis non parametric analysis of variance testing for differences in fish abundance and biomass densities, and taxa per tow among sites for the sample months. Tests were run at an alpha level of 0.05. (Fa, Pa =abundance, Fb, Pb =biomass, and Ft, Pt =taxa per tow, where F = F-ratio and P =probability; ns = not significant) \0 -!:>- 95 Appendix D: List of invertebrates collected from nearshore stations in the western edge of the Yellow Sea near Qingdao, People's Republic of China Phylum Cnidaria Class Anthozoa Order Actinaria unknown anemone Phylum Annelida Class Polychaeta Family Aphroditidae Aphrodita australis (Baird) Family Stemaspididae Sternaspis scutata (Ranzani) Phylum Mollusca Class Gastropoda Subclass Prosobranchia Order Patellogastropoda Family Patellidae limpet (Type A) Order Mesogastropoda Family Potamididae Cerithidea cingulata (Gmelin) Order Neogastropoda Family Muricidae Rapana sp. Family Turridae Inquisitor jlavidula (Lamark) Subclass Opisthobranchia Order Nudibranchia Family Okeniidae Okenia plana (Baba) Family Facelinidae Sakuraeolis enosimiensis (Baba) Family Arminidae Armina compta (Bergh) Family Dendronotidae Dendronotus arborescens (Miiller) 96 Phylum Mollusca, cont Subclass Opisthobranchia Order Notaspidea Family Pleurobranchidae Pleurobranchaea novaezealandia (Cheeseman) Order Cephalaspidea Family Philinidae Philine kinglipini (Tchang) Class Cephalopoda Subclass Coleoidea Order Octopoda Family Octopodidae Octopus ocellatus (Gray) Octopus variabilis (Sasaki) Phylum Arthropoda Subphylum Chelicerata Class Pycnogonida Ascorhynchus ramipes (Bohn) Paranymphon magnidigitum (Hong & Kim) Subphylum Crustacea Class Malacostraca Subclass Eumalacostraca Superorder Hoplocarida Order Stomatopoda Family Sqnillidae Squilla aratoria (de Haan) Superorder Peracarida Order Js6poda Suborder Flabellifera Family Spaeronidae flabelliferan isopod (Type A) Family Cirolanidae Cirolana japonica Superorder Eucarida Order Decapoda Suborder Dendrobranchiata Family Sergestidae Acetes chinensis (Hansen) Family Pasiphaeidae Leptochela gracilis (Stimpson) 'i 97 Phylum Arthropoda, cont Subphylum Crustacea Class Malacostraca Subclass Eumalacostraca Superorder Eucarida Order Decapoda Suborder Dendrobranchiata Family Penaeidae Penaeus orienta/is (Osbeck) Metapenaeopsis dalei (Rathbun) Trachypenaeus curvirostris (Stimpson) Metapenaeopsis joyneri (Miers) Parapenaeopsis tenella (Bate) Suborder Pleocyemata Infraorder Caridea Family Crangonidae Crangon affinis (Say) Family Hippolytidae Latreutes laminirostris (Ortmann) Heptacarpus rectirostris (Stimpson) Family Palaemonidae Palaemon gravieri (Yu) Family Alphaeidae Alpheusjaponicus (Miers) Alpheus distinguendus Infraorder Anomura Family Paguridae Diogenes edwardsii (de Haan) Family Hippiidae hippid crab (fype A) Family Porcellanidae porcellain crab (fype A) Infraorder Brachyura Family Dorripidae Dorippe granulata (de Haan) Family Leucosiidae Arcania undecimspinosa (de Haan) Family Parthenopidae Lamhrus validus (de Haan) 98 Phylum Arthropoda, cont Subphylum Crustacea Class Malacostraca Subclass Eumalacostraca Superorder Eucarida Order Decapoda Infraorder Brachyura Family Goneplacidae Carcinoplax vestitus (de Haan) Family Portunidae Ponunus trituberculata (Miers) Charybdisjaponica (A Milne-Edwards) Charybdis bimaculata (Miers) Phylum Echinodermata Subphylum Crinozoa Class Crinoidea Subclass Antedonacea Family Antedonidae Compsometra serrata Subphylum Echinozoa Class Echinoidea Subclass Euechinoidea Family Temnopleuridae Temnopleurus toreumaticus (Leske) ·Temnopleurus hardwickii (Doderlein) Subphylum Asterozoa Class Asteroidea Order Platyasterida Family Lnidiidae Luidia quinaria (von Martens) Order Spinulosida Family Asteriidae Patiria pectinifera (Muller et Troschel) Order Forcipulatida Asterias rollestoni (Bell) j Month N df Fa Pa Fb Pb Ft Pt March 17 2 8.01 0.018 14.28 0.001 20.71 < 0.001 May 17 2 4.06 ns 15.84 < 0.001 13.9 0.001 July 18 2 3.61 ns 3.52 ns 7.72 0.021 September 17 2 18.5 < 0.001 16.14 < 0.001 16.44 < 0.001 November 6 2 2.00 ns 1.14 ns 0.52 ns Appendix E: Summary table of results from Kruskal-Wallis non parametric analysis of variance testing for differences in invertebrate abundance and biomass densities, and taxa per tow among sites for the sample months. Tests were ron at an alpha level of 0.05. (Fa, Pa =abundance, Fb, Pb =biomass, and Ft, Pt =taxa per tow, where F represents F-ratio and P =probability; ns = not significant) 1.0 1.0