Diversity of Marine Snakes on Trawling Grounds in the Straits of Malacca and the South China Sea

2017·3.3 MB·

by  Voris, Harold K

Tropical Natural History 17(2): 65–87, October 2017
2017 by Chulalongkorn University
Diversity of Marine Snakes on Trawling Grounds in the Straits of
Malacca and the South China Sea

HAROLD K. VORIS

Science and Education, Field Museum of Natural History, 1400 South Lake Shore Drive,
Chicago, Illinois 60605 USA
* Corresponding author. Harold K. Voris ([email protected])
Received: 5 June 2017; Accepted: 28 August 2017

ABSTRACT.– Prior to World War II, traditional hand-operated fishing methods prevailed throughout Southeast
Asia. However, by 1950, mechanized diesel-powered trawlers were being introduced and the modern boom-and-
bust of fisheries was set into motion. Besides targeted fish, squid, and prawns, otter trawls on the bottom brought
up a vast diversity of demersal by-catch including marine snakes. This paper reports on the diversity of the marine
snake by-catch obtained from otter trawlers operating at two locations in the Straits of Malacca (Sungai Buloh and
Parit Botak) and one location on the eastern coast of the Malay Peninsula (Endau) in the mid-1970s. At Sungai
Buloh 11 species of snakes were observed and a single species, Hydrophis curtus, strongly dominated the
assemblage, comprising 82% of the by-catch. Eleven species were also observed in the by-catch from Parit Botak
but there, three species (Hydrophis fasciatus, H. curtus, and Aipysurus eydouxii) shared dominance with each
making up more than 22% of the assemblage. At Endau, 13 species appeared in the by-catch and two species (H.
curtus and Hydrophis viperinus) dominated, comprising 33% and 32% of the catch respectively. Besides
differences between locations, some small differences in species diversity were detected between collection
periods at Sungai Buloh. In addition, a review is undertaken of published trawl surveys of marine snakes in
Southeast Asia in the context of the steady depletion of fisheries in the region that took place in the second half of
the twentieth century. This review emphasizes that marine snake species diversity needs to be understood on a
relatively fine spatial scale and in the context of the health of the fishery as a whole.

KEY WORDS: Elapidae, Hydrophiinae, Acrochordidae, sea snake, otter trawl, demersal by-catch, species
richness, species evenness, Southeast Asian commercial fisheries

INTRODUCTION that “A full list of all the specimens
examined has not been given in every case,”
The first major survey of marine snakes (Smith, 1920, p.4) thus making it impossible
was published by Malcolm Smith in 1920. to know the relative abundance of species
Smith reported on marine snakes collected collected.
from the coastal waters of the Gulf of Two decades later, Bergman (1938,
Thailand and the Malay Peninsula between 1943) began to report on another large
1915 and 1918. Most of the sea snakes were collection of marine snakes from coastal
obtained from local fishermen using a areas near Sourabaya (Surabaya, Java).
variety of traditional hand-operated fishing Made by local fishermen between 1936 and
methods including stake nets and baskets 1942, this collection consisted of six
placed in river mouths. Some also were identified species represented by 984
obtained by hand nets and trawling along specimens, and four rare species represented
coasts. In all, 17 species were collected. The by fewer than four snakes each that were
number of specimens collected per species disregarded in his report (Bergman, 1943).
ranged from two to 98, but the author notes Although the purpose of the collection was
66 TROPICAL NATURAL HISTORY 17(2), October 2017
descriptive anatomical studies (e.g. 10"E; Parit Botak, 1°41'45"N, l03°6'15"E)
Bergman, 1949; 1954; 1956) it represents and one was sampled in the South China
the first major collection of marine snakes in Sea (Endau, 1°41'45"N, l03°6'15"E). Sungai
which all specimens from a single coastal Buloh was sampled in November 1971,
area were collected, identified, and counted, December 1974, and between January and
thus providing nearly complete data on August 1975. Parit Botak was sampled
species richness and relative abundance. between January and April 1975, while
Unfortunately, the methods of collection Endau was sampled only once in April
were not defined but most likely traditional 1975.
fishing gear accounted for most of the snake Snakes were obtained as by-catch from
catch. small diesel-powered otter board trawlers
By 1950, fishing methods in Southeast (~12–16m lg) at Sungai Buloh, Parit Botak
Asia were becoming mechanized and the and Endau (See Appendix). The trawlers
subsequent systematic depletion of fisheries normally remained at sea for 12–24 hours
was underway (Morgan and Staples, 2006). and ranged no more than 20–40 km from
The focus of this paper is on documented port. Generally they were limited to 3 to 4
collections of marine snakes obtained by hauls of 2 to 3 hours each before returning
diesel-powered trawlers operating in to port. Trawlers usually would leave port
Southeast Asia following World War II. In on high tides in either the early morning or
particular, new results on species richness late afternoon. Throughout the year trawling
and relative abundance are presented for was limited to fair weather, taking special
two locations in the Straits of Malacca, and care to avoid high winds and waves. The
one location on the eastern coast of the number of trawlers contracted to save
Malay Peninsula, and then these findings are marine snake by-catch on a given day varied
compared and contrasted to other from 1 to 6.
comparable studies conducted in the region. Environmental monitoring.– On some
Finally, all the studies of the by-catch of sea days the author accompanied one of the
snakes from trawls carried out in Southeast boats to the trawling grounds to collect
Asia are placed into the context of the environmental data. Data were collected at
simultaneous overexploitation of the the beginning and end of each haul, and the
fisheries, largely resulting from the speeds and depths of trawls were monitored
expansion of mechanized diesel-powered at regular intervals. Salinity was measured
bottom trawlers. using a LaMotte salinity kit (LaMotte Co.,
Chestertown, MD, USA) based on the
titration of silver nitrate (Boyle’s method).
MATERIALS AND METHODS
Turbidity was measured using both a Secchi

disk (plain white circular disk 50.8 cm
Trawlers and trawling grounds.– In
diameter) providing Secchi depths (m) and
November 1971 and between 18 December
the Jackson turbidity method (LaMotte
1974 and 27 August 1975, 1,575 marine
turbidity kit, LaMotte Co., Chestertown,
snakes were collected from three sites along
MD, USA) providing readings in Jackson
the coast of the Malay Peninsula (Fig. 1).
Turbidity Units (JTUs). The speed of the
Two localities were sampled in the Straits of
trawlers was measured using a mechanical
Malacca (Sungai Buloh, 3°14'45"N, 101°18'
propeller flowmeter with a standard rotor
VORIS — MARINE SNAKE DIVERSITY IN SOUTHEAST ASIA 67

FIGURE 1. Map showing the location of the three trawling grounds at Sungai Buloh, Parit Botak, and
Endau
(General Oceanics, Miami, FL, USA; Peninsula (Ministry of Agriculture, 1967)
Threshold approx. 10 cm/s: Range 10 cm/s provided a robust context for the
– 790 cm/s). Depth was measured using a measurements associated with the trawl
portable sonar depth finder, Model LFP-300 days of this study.
(Lowrance Co., USA). Water-temperatures Processing the by-catch and identification
were recorded at a depth of 1 m using a of snakes.– After snakes were separated
mercury thermometer. Substrate samples from the haul, living snakes were placed in
were collected using a galvanized steel Styrofoam boxes and kept in the shade
cylinder (15.2 cm diam, 45.7 cm lg) with the while drowned snakes were placed in
top 30.5 cm perforated by 0.31 cm holes. Styrofoam boxes with crushed ice for the
Retrieved samples were processed in duration of the trip. All snakes were brought
sequence through coarse mesh (~8 squares back to the laboratory, where living ones
per 2.54 cm), medium mesh (~18 squares were euthanized with an injection of sodium
per 2.54 cm), and fine mesh (~40 squares pentobarbital. Prior to preservation each
per 2.54 cm) sieves. Extensive hydrographic relaxed snake was tagged and measured
surveys both in the Straits of Malacca (snout-vent and tail length, and girth at the
(Hydrographer of the Navy, 1958; Ministry neck and mid-body). Stomach contents were
of Agriculture and Fisheries, 1972; Chong, removed, preserved in 12% formalin and
1974) and along the east coast of the Malay later transferred to 70% ethanol for
68 TROPICAL NATURAL HISTORY 17(2), October 2017

TABLE 1. Numbers of marine snakes collected as by-catch from trawlers operating on three Malaysian trawling
grounds in 1971 and 1974-1975. The species names in the first column are the names applied at the time of the
field work. The names used throughout this paper appear in column two and largely follow Figueroa et al., 2016.

Sungai Buloh Parit Botak Endau
Name (1975) Name (2016) Number % Number % Number %
Aipysurus eydouxi A. eydouxii* 0 0 120 22.7 16 8.2
Acrochordus granulatus A. granulatus* 59 6.9 21 4.0 0 0.0
Hydrophis brookii H. brookii 2 0.2 1 0.2 0 0.0
Hydrophis caerulescens H. caerulescens 48 5.6 51 9.7 0 0.0
Lapemis curtus H. curtus 702 82.4 147 27.8 65 33.3
Hydrophis cyanocinctus H. cyanocinctus 11 1.3 13 2.5 9 4.6
Hydrophis fasciatus H. fasciatus 22 2.6 164 31.1 5 2.6
Microcephalophis gracilis H. gracilis 1 0.1 1 0.2 3 1.5
Hydrophis inornatus H. inornatus 0 0.0 0 0.0 2 1.0
Hydrophis klossi H. klossi 0 0.0 0 0.0 1 0.5
Hydrophis ornatus H. ornatus 3 0.4 1 0.2 11 5.6
Acalyptophis peronii H. peronii 0 0.0 0 0.0 13 6.7
Enhydrina schistosa H. schistosus 2 0.2 7 1.3 5 2.6
Hydrophis spiralis H. spiralis 1 0.1 0 0.0 0 0.0
Hydrophis torquatus H. torquatus 0 0.0 2 0.4 0 0.0
Thalassophina viperina H. viperinus 0 0.0 0 0.0 63 32.3
H. sp. H. sp. 1 0.1 0 0.0 1 0.5
H. sp. H. sp. 0 0.0 0 0.0 1 0.5
Total 852 528 195
* The generic names from 1975 remain unchanged.
permanent storage. Other particulars on the largely consistent with Figueroa et al.
data collection can be found in Voris and (2016). Table 1 provides the list of species
Glodek (1980), Lemen and Voris (1981) and collected from the three trawling grounds.
Voris and Voris (1983). The first column gives the names applied in
The species collected represent two 1975 which largely followed Smith (1926)
families of marine snakes, Acrochordidae and the second column gives the names
(Acrochordus granulatus) and Elapidae used in the present publication that largely
(Hydrophiinae, true sea snakes). follow Figueroa et al. (2016).
Identifications were based on the key in Diversity measures and statistics.– Several
Smith (1926), data and descriptions in estimates of species diversity were applied
Smith (1920; 1926), and confirmed by in the analysis of the snake assemblages
comparisons to preserved specimens sampled from the three trawling grounds.
collected by Malcolm Smith and deposited All the measures assess information both on
at the Field Museum of Natural History numerical species richness and species
(FMNH) and the British Museum of Natural evenness, and each has been used widely
History (BMNH). The nomenclature has over the past half century. Simpson’s index,
been updated where appropriate to be D (Simpson, 1949; Magurran, 1988) and the
VORIS — MARINE SNAKE DIVERSITY IN SOUTHEAST ASIA 69
complement of Simpson’s D, the probability within a trawling area on a given day. Local
of interspecific encounter, PIE (Hurlbert, weather conditions were usually fair but
1971), were used for some comparisons. afternoon squalls were not uncommon.
The Shannon index, H’ (Shannon and Because of the duration of hauls and the
Weaver, 1963; Magurran, 1988), the distances covered over three hours, snake
Brillouin index, HB (Brillouin, 1960; Lloyd, by-catch could not be attributed to any
1968; Pielou, 1966; Magurran, 1988), and specific condition or feature observed within
the related evenness index, E (Pielou, 1966; the trawling grounds.
Hurlbert, 1971; Magurran, 1988) were Published data document seasonal shifts
applied in other analyses. in salinity, temperature, precipitation, river
Several of the above measures of discharge, and plankton productivity in the
diversity have statistics associated with Straits of Malacca. For example, at Sungai
them and those may be found in Magurran Buloh, salinity varied from 29.0 p.p.t. in
(1988) or the other citations provided for the November to 30.5 p.p.t. in June
measures. In addition, the non-parametric (Soeriaatmadja, 1956). Sea surface
Chi-square test (Sokal and Rohlf, 1995) was temperature ranged from 29.5°C in
used to compare observed species September-March to 30.5°C in April-June
abundances with those that would be (Hydrographer of the Navy, 1958). Also,
expected on the basis of chance alone. there were seasonal variations in rainfall
from about 10 cm of rain in June and July to
about 23 cm in November and December
RESULTS
(Dale, 1959). The river discharge rates on

the Selangor River near Sungai Buloh vary
Environmental conditions.– Trawler
seasonally with lows in June and July of
captains normally did not attempt to trawl in
0.022 m3/sec/km2 to highs in November of
an exact location. The environmental data
about 0.092 m3/sec/km2 (taken from five-
within each trawling ground indicate
year averages given by the Ministry of
considerable variation in many physical
Agriculture and Fisheries, 1972).
conditions within individual three-hour
Trawling grounds: Size and Patchiness.–
hauls (continuous trawl pull), between
Table 1 provides the numbers and relative
hauls, and between days. For example,
abundance (%) of marine snakes collected
during one three-hour haul, a trawler
as by-catch from otter trawls operating on
traveled in one general direction for a
the three trawling grounds: Sungai Buloh,
distance of about 14 km. The course of the
Parit Botak, and Endau (Fig. 1). The
track was subjectively determined by the
trawling grounds were large in comparison
captain and intersected depths ranging from
to the area fished on a given day. For
6 to 18 m. Substrate varied from rather
example, the trawling ground at Sungai
homogeneous fine sand and mud to
Buloh was roughly oval in shape and was
particulate samples containing 5 to 10
normally set off from the coast by a 3 km
percent broken shells and coarse sand.
no-trawl zone. The oval extended about 20
Generally, measurements of Secchi disk
km parallel to the coast with a width of
turbidity ranged from 0.8 to 1.6 meters.
about 6 km. Thus, the area was
Salinity (28–31 p.p.t.) and surface water
approximately 377 sq km. A single trawler
temperature (28–30°C) were fairly constant
traveling at 3 km/h with a net width at the
70 TROPICAL NATURAL HISTORY 17(2), October 2017

FIGURE 2. Bar graph showing the species composition and relative abundance for 852 marine snakes
collected as by-catch from otter trawlers operating on the trawling grounds at Sungai Buloh (See Fig. 1). The
snakes were collected in 1971 and 1974-75 and represented 10 species of true sea snake (Elapidae) and one
species of file snake (Acrochordidae). The assemblage is strongly dominated by Hydrophis curtus (see inset
in figure, FMNH 202179) that made up 82% of the snake by-catch. The three most common species,
Hydrophis curtus, Acrochordus granulatus, and Hydrophis caerulescens, made up 95% of the sample.
mouth of 18 meters would cover only a tiny (Hydrophiinae, true sea snakes). This
proportion of the trawling ground in one assemblage is strongly dominated by the
pull of 3 hours (roughly only 0.16 sq km). short sea snake, Hydrophis curtus (Fig. 2).
The catch of fish and prawns along with the The overall diversity of the assemblage as
snake by-catch varied greatly, documenting measured by Simpson’s index (D), the
patchy distributions of fish, prawns, and probability of interspecific encounter (PIE),
snakes over the grounds. For example, 18 the Shannon index H’, Evenness (E), and
individual three-hour trawl pulls at Sungai the Brillouin index (HB) were as follows: D
Buloh taken over three months resulted in = 0.688, PIE = 0.313, H’ = 0.729, E =
snake by-catches that ranged between 0 and 0.304, and HB = 0.976.
27 snakes while both the median and modal Sungai Buloh: Species diversity
number of snakes per trawl pull was just 2 accumulation curves.– To examine species
snakes. diversity (species richness and evenness)
Sungai Buloh: Species richness and over time, the Brillouin diversity index (HB)
evenness.– The 852 marine snakes caught was calculated for all snakes (adults and
as by-catch on the Sungai Buloh trawl juveniles) and adult snakes only captured in
ground were distributed among 11 species December 1974 and in 1975 (Fig. 3). The
in two families of snakes, Acrochordidae final HB values for all snakes captured
(Acrochordus granulatus), and Elapidae (adults and juveniles, n = 768) and adult
VORIS — MARINE SNAKE DIVERSITY IN SOUTHEAST ASIA 71

FIGURE 3. A comparison of plots of the values of the Brillouin diversity index (HB) for all snakes (adults
and juveniles) and for adult snakes only caught on the Sungai Buloh trawling grounds in 1974-1975. A
trawler day is the catch from one boat during a one-day trip from port. The values of the HB diversity at the
end of the curves (far right) were 1.579 for adult snakes only and 0.976 for all snakes (adults plus juveniles).
snakes only (n = 363) were 0.976 and 1.579 January (combined) were compared to
respectively. The plots in Figure 3 document catches from February and from March.
three phenomena: 1) The diversity values Collections from April and August were not
for adult snakes attained the highest levels at compared due to the presence of many
the very beginning of the sampling in the neonates and juveniles. The single most
first few trawler days; 2) The two curves are noticeable difference among the samples is
identical until trawl day 12 (23 February the high proportion (15.9 %) of A.
1975) when the first neonate appeared in a granulatus in the trawls made in December
trawl pull; 3) The curve based on adult and early January. A Chi-square test
snakes only remains remarkably flat through comparing the distributions of the three
41 trawler days while the capture of sampling periods revealed that the observed
numerous neonate and juvenile snakes distributions differed significantly from the
(mainly H. curtus) in March, April, and expected (χ² = 17.57, df = 8, p < 0.05). A
August steadily reduced species diversity. pairwise Chi-square test comparing the
Sungai Buloh: Diversity through time.– distributions from December-January with
Table 2 gives the total number of marine February also proved significantly different
snakes caught and relative abundance for (χ² = 13.99, df = 4, p < 0.01) while the
species caught in November 1971, and comparisons of December-January with
between December 1974 and August 1975. March, and February with March showed no
To determine if the assemblage of snakes in significant differences (χ² = 4.50, df = 4, p >
the by-catch varied between time periods, 0.05; χ² = 9.10, df = 4, p > 0.05). The PIE
the catches from late December and early value of 0.055 (Table 2) also shows that the
72 TROPICAL NATURAL HISTORY 17(2), October 2017
TABLE 2. Number of marine snakes collected and their relative abundance (%) for 11 species collected on the
trawling grounds at Sungai Buloh during six fishing periods. Samples from November 1971 and from December
1974 to February 1975 included adults only while samples from March to August 1975 included neonates and
juveniles (See Fig. 3). Comparisons of the monthly distributions using Chi-square tests were limited to the five
most common species to avoid the numerous zero cells present in the last six rows.

November Dec-Jan February March April August
1971 1974-75 1975 1975 1975 1975
N % N % N % N % N % N %
H. curtus 67 79.8 96 63.6 73 79.3 52 70.3 191 93.6 223 90.3
A. granulatus 1 1.2 24 15.9 8 8.7 4 5.4 5 2.5 17 6.9
H. caerulescens 1 1.2 23 15.2 4 4.3 11 14.9 6 2.9 3 1.2
H. fasciatus 9 10.7 3 2.0 6 6.5 3 4.1 0 0.0 1 0.4
H. cyanocinctus 3 3.6 4 2.6 1 1.1 2 2.7 1 0.5 0 0.0
H. ornatus 3 3.6 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0
H. brookii 0 0.0 0 0.0 0 0.0 1 1.4 0 0.0 1 0.4
H. schistosus 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 0.8
H. gracilis 0 0.0 0 0.0 0 0.0 1 1.4 0 0.0 0 0.0
H. spiralis 0 0.0 0 0.0 0 0.0 0 0.0 1 0.5 0 0.0
H. sp. 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 0 0.0
Total 84 151 92 74 204 247
PIE 0.354 0.550 0.361 0.485 0.123 0.181
1 STD 0.064 0.039 0.061 0.065 0.031 0.032

relatively large numbers of A. granulatus the Brillouin index (HB) were as follows: D
and H. caerulescens in the December – = 0.237, PIE = 0.764, H’ = 1.615, E =
January sample contribute to a significantly 0.674, and HB = 2.290.
more diverse sample. Parit Botak: Species diversity
Parit Botak: Species richness and accumulation curves.– Figure 5 shows two
evenness.– The by-catch of marine snakes plots of the Brillouin diversity index
from the Parit Botak trawling grounds (Fig. accumulation over time. The main plot of
1) included 528 snakes distributed among diversity values for all snakes shows HB
11 species in two families, Acrochordidae increasing sharply at first as species were
(Acrochordus granulatus), and Elapidae added to the collection but then the curve
(Hydrophiinae, true sea snakes). This levels off after just four or five trawler days.
assemblage was dominated by three species, The second graph (inset in Fig. 5) shows
H. fasciatus, H. curtus, and Aipysurus two similar curves starting with trawler day
eydouxii (Fig. 4). Four other species made 26 on March 9 when neonates first appeared
up from 2 to 9 percent of the total catch. The in the catch. The neonates and juveniles (24
overall diversity of the assemblage as H. curtus and 1 H. schistosus) appeared
measured by Simpson’s index (D), the across the catches in March and April and
probability of interspecific encounter (PIE), because of their modest numbers the HB
the Shannon index H’, Evenness (E), and accumulation curves of adults only and all
VORIS — MARINE SNAKE DIVERSITY IN SOUTHEAST ASIA 73

FIGURE 4. Bar graph showing the species composition and relative abundance for 528 marine snakes
collected as by-catch from otter trawlers operating on the Parit Botak trawling grounds (See Fig. 1). The
snakes were collected in 1975 and represented 10 species of true sea snake (Elapidae) and 1 species of file
snake (Acrochordidae). The assemblage is dominated by Hydrophis fasciatus (see inset in figure, FMNH
198841), Hydrophis curtus, and Aipysurus eydouxii that combined made up 81.6% of the snake by-catch.
Table 3 provides the monthly results as well as the totals.
snakes remained nearly identical. The final square tests were used to compare the
HB values for all snakes captured (adults distributions from the three sampling
and juveniles, n = 528) and adult snakes periods with each other. The comparisons
only (n = 503) were 2.280 and 2.290 included only the seven most common
respectively. species in Table 3 to eliminate the numerous
Parit Botak: Diversity through time.– The zero cells among the four least common
total number of sea snakes caught and species. None of the three tests (Jan. vs.
relative abundances for species caught Mar.; Jan. vs. Apr.; Mar. vs. Apr.) showed a
during three periods, January (plus February significant difference at the p < 0.05 level.
1-4), March, and April are given in Table 3. Endau: Species richness and evenness.–
The table shows that the samples from the The 195 marine snakes caught as by-catch
three periods are remarkably similar in that on the Endau trawling grounds over just two
the same four species are the most common days were distributed among 13 species of
species in each of the periods. The order of true sea snakes (Elapidae, Hydrophiinae).
relative abundance among the four species This assemblage was sharply dominated by
is also nearly identical, with H. fasciatus two species, H. curtus and H. viperinus
most common in January and March, and H. (Fig. 6). The overall diversity of the
curtus most common in April. Pairwise Chi- assemblage as measured by Simpson’s
74 TROPICAL NATURAL HISTORY 17(2), October 2017

FIGURE 5. Plots of Brillouin diversity index (HB) values for all snakes (adults and juveniles) caught on the
Parit Botak trawling grounds in 1975. A trawler day is the catch from one boat during a one-day trip from
port. The HB diversity value at the end of the curve (far right) was 2.290 for all 528 snakes (adults plus
juveniles). The inset graph shows that the effect of the 25 juveniles (24 Hydrophis curtus and 1 Hydrophis
schistosus) appearing in the samples after day 26 on the species diversity was very small (< 0.01 HB units on
any day). This was the case because the young were distributed across the last 25 of the 50 trawler days.
index (D), the probability of interspecific periods was remarkably constant within
encounter (PIE), the Shannon index H’, both trawling grounds in the Straits of
Evenness (E), and the Brillouin index (HB) Malacca.
were as follows: D = 0.234, PIE = 0.770, H’ The relative abundance of species over
= 1.801, E = 0.702, and HB = 2.435. time at Sungai Buloh did show one
significant difference between periods,
namely, among adult snakes collected in
DISCUSSION AND CONCLUSIONS
Dec.-Jan. and February (Table 2). This
could be due to any number of factors. For
Comparisons within trawl grounds.– example, trawlers may have fished in unique
Overall the species composition on trawling areas in December and January resulting in
grounds remained very similar over the time their obtaining the larger numbers of A.
periods observed. At Sungai Buloh, the granulatus and H. caerulescens, or these
three most common species, H. curtus, A. species may have moved into the trawling
granulatus, and H. caerulescens (Fig. 2), grounds in larger numbers. The relative
were observed in all six sampling periods abundance of species taken during three
(Table 2). At Parit Botak, the seven most periods on the Parit Botak trawling grounds
common species (Fig. 4, Table 3) were (Table 3) did not differ significantly from
present in all three sampling periods. Thus, each other.
the species composition over the sampling