Few studies have compared water quality and plankton along the eutrophication gradient from Kingston Harbour to oceanic waters around Jamaica. To compare the planktonic community along the expected nutrient gradient, we sampled every two weeks at four stations, from eutrophic Kingston Harbour to oceanic California Bank. Phytoplankton was assessed from whole water Niskin ]]>
-1 at the Harbour Shoal Beacon to 0.2 μg L-1 at California Bank). Characteristic estuarine phytoplankton genera -such as Ceratium, Gonyaulax, Gyrodinium and Rhizosolenia- dominated harbour samples while ]]>
Asterionelliopsis, Navicula, Nitzschia,Rhizosolenia and Thalassionema- dominated California Bank. Highest phytoplankton densities (mean values of 34 174 cells L-1) were found at the harbor mouth. Mean zooplankton abundances ]]>
Lucifer faxoni and Penilia avirostris were indicative of harbor waters and
Microsetella sp. and Farranula carinata of offshore waters. Some zooplankton taxa, like L. faxoni, Paracalanus parvus and copepod nauplii, despite showing gradual decline with distance from Beacon to the Edge, increased in abundance at the furthest station, California. California Bank clearly experiences enrichment which at times can be as high as near-shore areas, but the planktonic community is unique and similar only to those at other offshore station. To be reliable, characterization of water ]]>
Key words: Plankton, Water quality, Kingston Harbour, South-east coast, Jamaica.
Resumen
Pocos estudios han comparado la calidad de agua y plancton a lo largo de un gradiente de eutrofización en el Puerto Kingston con aguas oceánicas en Jamaica. Para comparar la comunidad de plancton a lo largo del gradiente de nutrientes esperado, muestreamos cada dos semanas en cuatro estaciones, desde el eutrófico Puerto Kingston hasta el Banco oceánico California. Evaluamos el fitoplancton de agua ]]>
L-1 en el Puerto Shoal Beacon a 0.2µg L-1 en el Banco California). Géneros característicos estuarinos como Ceratium, Gonyaulax, Gyrodinium y Rhizosoleniadominaron en las muestras del Puerto Kingston mientras que géneros característicos de costa afuera como Asterionelliopsis, Navicula, Nitzschia,Rhizosolenia y Thalassionema dominaron en el Banco California. Las densidades más altas de fitoplancton (valores promedio de 34 174 células L-1) se encontraron en la boca del ]]>
Lucifer faxoni y Penilia avirostris fueron indicativos de las aguas del puerto y
Microsetella sp. y Farranula carinata fueron identificados como indicadores de aguas costa afuera. Taxones seleccionados de de zooplancton, como L. faxoni
, Paracalanus parvus y nauplios de copépodos, a pesar de mostrar un decline gradual con la distancia desde el puerto de Kingston a la otra estación costa afuera, aumentan su abundancia en la estación más lejana, Banco California. Banco California es claramente una zona que experimenta un enriquecimiento que a veces puede ser tan alto como las áreas cerca de la costa, pero la comunidad planctónica es única y similar únicamente a las estaciones costa afuera. Para ser confiable, la caracterización de masas de agua debe implicar una serie de ]]>
Palabras clave: Plancton, costa sur-este, calidad de agua, Puerto Kingston, Jamaica.
There is now a great deal of information concerning the planktonic communities of the Kingston Harbour and the nearby coastal shelf areas (Moore & Sander, 1979; Webber & Roff, 1996; Webber & Webber, 1998; Dunbar & Webber, 2003; Campbell, Manning, Webber & Webber, 2008). Some of these studies have been quite extensive covering distances from Kingston Harbour to as far as 5km to the shelf edge (Moore & Sander, 1979) or parallel to the coast for 15km (Lindo, 1991). Other studies have covered extensive areas of the Port Royal cays (Webber et ]]>
Few studies have adequately compared water quality and plankton along this eutrophication gradient from Kingston Harbour to oceanic waters around Jamaica. Indeed, the only previous study was conducted in 1964 and compared the zooplankton community and water quality between Kingston Harbour, the Port Royal Cays and an offshore station located 5 km away from the coastline of Jamaica (Moore & Sander, 1979). Moore and Sander (1979) indicated a ]]>
The area beyond the southern ]]>
The paucity of studies comparing oceanic, shelf and harbor waters on the south coast is due primarily to the extent of the south coast shelf (28km at its widest point) and the ]]>
The primary aim of this investigation was therefore to compare water quality and plankton along a gradient from expectedly pristine areas well beyond the shelf with areas close to shore and ultimately high nutrient areas like Kingston Harbour. The study was intended to:
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1. Identify and confirm a eutrophication gradient between Kingston Harbour, the Port Royal Cays and offshore areas,
2. Compare the planktonic communities along the gradient and
3. Identify planktonic indicator species or communities characteristic of the areas along the gradient.
Materials and Methods
Study sites: Four sites were selected (Fig. 1) to represent four contrasting marine environments with increasing linear distance from Kingston Harbour (Table 1). The linear distance represented a nutrient gradient from the eutrophic Kingston Harbour to oligotrophic California Bank (furthest away). The first site (Harbour Shoal Beacon - HSB) was located at the mouth of Kingston Harbour which has been characterized as eutrophic since the 1970’s (Goodbody, 2003; Wade, Antonio & Mahon, 1972). Kingston Harbour receives agricultural runoff and partially treated sewage effluent which serve as a source of nutrient enrichment. The second site (South East Cay - SEC) was located 5 km from HSB. This location represents a coral cay and is one of a number of cays on the south coast shelf of Jamaica referred to as the Port Royal Cays. The Port Royal Cays are important for near-shore fishery and recreational sites for locals and visitors (Steele, 1998). The third location (Windward Edge - WE) was located 7km from the SEC, on the south-coast shelf edge and as such is exposed to oceanic influences. The coral reefs of this location are deep, patchy and often senescent (Munro, 1983). The fourth sampling location (California Bank - CB) was the furthest being ten 10km from the third site (WE) or 22km from Kingston Harbour. California Bank represents a small offshore bank, 8.6km2 in area with immediate surrounding waters of approximately 700km depth. The water depth on the bank ranges between 35-45m. Munro (1983) also describes the reefs of this location to be sparse in coral cover and with sills that are not well developed. ]]>
Sampling programme: Sampling was carried out every two weeks from April to December 2004. Table 1 has the G.P.S coordinates of the stations that were occupied for sampling. On each occasion, sampling commenced from the station furthest offshore (California Bank) and progressed towards the more eutrophic station ]]>
Water quality and phytoplankton assessment: Physical and chemical parameters including: temperature, salinity, oxidative redox potential, turbidity and light were collected in situ on each sampling occasion using a hydro-lab multi-parameter ]]>
Whole water samples, collected ]]>
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A 250mL portion of each sample was immediately preserved (poured directly from the Niskin bottle into 250mL plastic bottle) containing 2.5mL of acidized Lugol’s solution for counting and identification while the remainder of the sample was filtered through Nalgene size fractionating tower into three size classes. Separation was done using the appropriate size filters as follows: Nitex screening - 20µm; Whatman glassfibre filters, GDF - 2.7µm and Whatman glassfibre filters, GDF - 0.7µm.
Chlorophyll a content was determined using 6mL of 90% acetone to extract the chlorophyll (Arar & Collins, 1997). The quantity of Chlorophyll a in the extract was determined using a Turner Design Fluorometer TD700. Lugol’s preserved phytoplankton samples were gently homogenized and 100mL portions used to fill settling chambers which were allowed to stand for 24 hours before examination using a Carl Zeiss inverted microscope with phase ]]>
Nitrates and phosphate were ]]>
Zooplankton assessment: SCOR, WP2 pattern plankton nets (UNESCO, 1968) of three different mesh sizes: 600µm, 200µm and 64µm were used to collect zooplankton samples in replicate (n=2) vertical hauls. The hoop diameter of the 600µm net was 1 m while the diameter of the 200µm and 64µm net was 0.5. Samples were collected from a depth of 20m from California Bank, Windward Edge and South East Cay but from a 5m depth at Harbour Shoal Beacon. The net was hauled at a speed of approximately 0.5ms-1 using a manual winch. Plankton nets were washed between stations to prevent artificial transfer of species.
Zooplankton samples were fixed upon collection by pouring the contents of the cod end into 1 l containers containing 10mL of full strength formalin. All samples were later preserved in 10% formalin. Zooplankton species identification and enumeration were conducted on a sub-sample obtained using a modification of the beaker split method (Van Guelphen, Markle & Duggan, 1982). The sub-sample was transferred into a Bogorov tray and ]]>
Results
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A total of 10 physicochemical, five phytoplankton and 13 zooplankton parameters were collected and analyzed each as 105-108 discrete samples during this investigation. All data were assessed for normal distribution using STATISTICA 7 and Log (x +1) transformations applied where necessary. Temporal differences were not significant (MANOVA, p>0.05) but spatial variability was significant for eight of the physicochemical, all of the phytoplankton (size fractionated biomass (pico, nano and netplankton), abundance and ]]>
Physicochemical parameters: Mean temperature values showed a slight decrease with increasing distance from Harbour Shoal beacon (HSB). Values ranged from 29.53oC at HSB to ]]>
salinity values (MANOVA, P < 0.001) increased sharply from Harbour Shoal Beacon (35.76o/oo) to South East Cay (36.19o/oo) and remained almost constant as distance increased from South East Cay (SEC) to offshore stations (Fig. 2). Significant differences in turbidity values were not observed (MANOVA, P=0.165) and there was no trend in the ]]>
Fig. 3).
Oxidative redox potential (ORP) values (MANOVA, p = 0.011) increased as distance offshore increased (Fig. 3) but Phosphate (MANOVA, P = 0.027) showed similar values at the Harbour Shoal Beacon as at California Bank. Phosphate values ranged from a low of 0.055 µg at L-1 at Windward Edge to a high of 0.099µg at L-1 at Harbour Shoal Beacon with California Bank being a close second with high values of 0.098µg at L-1 (Fig. 4). However, nitrate values showed no significant differences between stations (MANOVA, P = 0.079).
Phytoplankton: A severe decline in mean chlorophyll a total biomass (MANOVA p = < 0.001) was seen from Harbour Shoal Beacon (1.25µg L-1) to South East Cay (0.25µg L-1). A more gradual decline was observed towards Windward Edge (0.2µg L-1) then to California Bank (0.125µg L-1). Also as depth increased, chlorophyll a values showed small increases at all stations and were found to not be significant (Fig. 5).
Examination of the proportion contributed by each phytoplankton size fraction (Fig. 6) showed that the picoplankton (0.7µm) contributed the dominant portion of the biomass at California Bank, Windward Edge and South East Cay while the netplankton (20µm) dominated only at Harbour Shoal Beacon. Generally the picoplankton fraction increased and netplankton decreased with distance from Kingston Harbour. The nanoplankton fraction (2.7µm) showed the least pattern across all stations (Fig. 6). Chlorophyll a values for all size classes of phytoplankton showed significant spatial variation (MANOVA P <0.001) after log transformation.
Stepwise, variable selection multiple regression tests showed that 56% of the variation in netplankton was related to variation in distance, temperature, pH and dissolved oxygen (R2=55.75%, P<0.005), while 72% of the variation in nanoplankton was related to variation in distance, temperature, specific conductivity and phosphates (R2=71.38%, P<0.005). Variation in distance, dissolved oxygen and salinity were attributable for 54% of the variation in picoplankton (R2=53.29, P<0.005).
The number of phytoplankton taxa (richness) increased from Harbour Shoal Beacon (mean of 12) to South East cay with a mean of 19 (MANOVA, P < 0.001). However, as distance increased further offshore a general decline was then observed with Windward Edge having a richness of 15 genera, the same as California Bank (Fig. 7). Stepwise variable selection multiple regression showed that variations of 13% in the number of phytoplankton genera could be directly attributable to variation in nitrates and dissolved oxygen (R2=12.58%, P<0.05), with both having a negative relationship to richness.
Phytoplankton abundance (# of cells l-1) followed a similar spatial distribution pattern (MANOVA, P<0.001) where a severe decrease was seen from Harbour Shoal Beacon (34 173.51 cells l-1) to South East Cay (507.67 cells l-1). A gradual decrease was then obtained from South East Cay as distance offshore increased. Windward Edge had mean abundance values of 320.30 cells l-1 while California Bank had 137.71 cells l-1 (Fig. 6). Stepwise variable selection multiple regression showed that 61% of the variation in number of cells was attributable to distance and total dissolved solids (R2=60.79%, P<0.05), both having negative relationships to the number of cells l-1.
Jaccard community coefficient (JCC) analysis indicated that the communities at Harbour Shoal Beacon and South East cay were 45% similar, Harbour Shoal Beacon and Windward Edge were 46% similar while Harbour Shoal Beacon and California Bank were 38% similar. South East Cay and Windward Edge were 63% similar and South East Cay and California Bank 49% similar. Finally, phytoplankton communities at Windward Edge and California Bank were 53% similar (Table 2).
Zooplankton species richness and total abundances: The overall number of zooplankton species encountered during the sampling period (total richness) ranged from 81 species at Harbour Shoal Beacon (HSB) to 115 species at Windward Edge (MANOVA, P<0.001). HSB also consistently had the lowest mean richness (Fig. 8). Contrastingly, HSB had the greatest mean zooplankton abundance (Nos. m-3) when compared to the other three stations (MANOVA, P<0.001). Mean zooplankton abundance at HSB was 5 963 individuals m-3 while South East Cay (SEC), Windward Edge (WE) and California Bank (CB) had abundance values of approximately 2 000 individuals m3 (Fig. 9). ]]>
When all net collections were combined, larvae (invertebrate and vertebrate) were the most abundant group being the greatest proportion of the collections from all four stations. Larvae accounted for more than half of the total abundance at CB (Fig. 9). Of the larval group, copepod nauplii were numerically the most dominant taxon. These occurred universally across all stations and sampling occasions with highest numbers observed at HSB followed by CB. The other two stations (SEC and WE) showed similar low numbers. Adult copepods were the second most abundant taxonomic group (after larvae). Cnidarians made the smallest contribution at each station, but were still most abundant at Harbour Shoal Beacon.
Potential zooplankton indicator species were examined in terms of their relative abundance across the four stations. Species like Penilia avirostris and Lucifer faxonihad been identified as Kingston harbor indicators (Lindo, 1991; Webber, Roff, Chisholm & Clarke, 1996). P. avirostris showed greatest abundance at Harbour Shoal Beacon (Table 4) and then declined exponentially to the stations further offshore (MANOVA P > 0.001). L. faxoni was similar with maximum abundances at HSB, followed by a severe decline at SEC and WE. However, abundances showed a slight increase at CB, above values recorded at SEC and WE.
Microsetella sp. occurred at all four stations investigated and abundances gradually increased with distance (MANOVA, P<0.001) from the Harbour Shoal Beacon, where mean abundances were less than 5 individuals m-3. However it was present as 30 and 37 individuals m-3 at Windward Edge and California Bank, respectively. F. carinata (MANOVA, P= 0.001) seemed to have similar distribution (increase in the numbers of this species as distance offshore increased) but it was only observed at South East Cay, Windward Edge and California Bank. It was not observed at Harbour Shoal Beacon over all the sampling occasions.
Using the Jaccard Community Coefficient (JCC), the zooplankton community at the stations increased in similarity from Harbour Shoal Beacon (HSB) towards California Bank (CB). HSB was the least similar compared to the other stations with similarities of 38% and less (Table 3). The highest community similarities were observed between CB and WE (66% similarity). Communities at South East Cay (SEC) and Windward Edge (WE) showed 63% similarity while SEC and CB were 46% similar (Table 4). This pattern was almost identical to the phytoplankton community.
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Discussion
Comparison of water quality and plankton across areas spanning coastal bays or harbors and offshore, oceanic banks are rare as intensive plankton and water quality sampling requires relatively short intervals between collecting and processing events. Alternatives include availability of large enough vessels for ]]>
Thus it was expected that the physicochemical parameters would show severe changes from the eutrophic ]]>
L. faxoni, Paracalanus parvus and copepod nauplii. Values at California bank for these parameters indicated unexpected enrichment of the area. Offshore banks and shelf areas may receive enrichment from upwelling which is aided by horizontal circulation patterns like gyres that concentrate nutrients and retain planktonic communities (Genin, 2004; Rakhesh, Raman & Sudarsan, 2006). Such circulation is said to lead to increased numbers in especially larval forms, phytoplankton and ]]>
While it was unexpected for the offshore bank to have nitrate values as high as at the mouth of Kingston Harbour, upwelling has been shown to promote significant enrichment of surface waters (Caldeira et al., 2002). Nevertheless, it is important to note that nutrient values between the eutrophic Kingston harbor and relatively pristine offshore sites were not as ]]>
The identification of indicator species and groups from the plankton was more reliable at characterizing different water masses (McGehee, Derner & Warren, 2004; Fehling et al., 2012; Tseng, Dahms, Chen & Hwang, 2013). Netplankton (>20µm) is known to be dominant in high nutrient ]]>
The taxonomic richness (number of zooplankton species and phytoplankton genera) did not follow the expected gradient as lowest number of species occurred at both Harbour Shoal Beacon and California Bank. Furthermore, while total abundances (zooplankton Nos. m-3 and Chlorophyll a L-1) followed the expected gradient, individual species and groups such as copepod nauplii showed secondary increases at the station located furthest offshore due to enrichment associated with the changes in circulation associated with the submerged bank.
While abundances may be unpredictable, community composition has been shown to be better when identifying distinct water masses (Campbell et al., 2008). Thus greater ]]>
Gonyaulax, Gyrodiniumand Perdinium being dominant while oceanic stations having diatoms such as: Navicula,
Nitzschia, Rhixosloenia and Thallassionema being dominant; irrespective of nutrient levels.
Throughout the study, zooplankton groups were dominated by copepods and larvae which accounted for more ]]>
Within each group/taxa a few species displayed high abundances and occurrence at the stations and therefore have the potential for use as indicators of different water masses/conditions. L. faxoni has been identified as an indicator of ]]>
L. faxoni, clearly displayed highest numbers at the Harbour station but did not have lowest mean values at the furthest offshore station. The distribution of the species appears to be less affected by salinity levels and more by nutrients.
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The Cladoceran, P. avirostris has often been paired with L. faxoni. As shown in the present investigation, it also tends to have highest numbers at the harbor station and low numbers at offshore areas. P. avirostris has been previously reported in high abundances in the Kingston Harbour and at ]]>
P. avirostris’ distribution, thereby limiting its presence to areas with shallow low-salinity waters. ]]>
The copepod Microsetella sp. was the dominant harpacticoid at the offshore stations and Moore and Sander (1979) reported this species as only occurring at stations outside of the Harbour and while Webber et al. (1996) did not report it from their study of the Port Royal Cays, it was found offshore Discovery Bay, north coast of Jamaica (Webber & Roff 1995). The cyclopoid,
F. carinata was important at the offshore stations, while being absent from the Harbour Shoal Beacon. This species has also been found to be important in oceanic waters off the north coast at Discovery Bay, Jamaica by Webber & Roff (1995). Microsetella sp. and F. carinata therefore could be indicators of offshore waters.
Larvae followed the general trend of highest numbers at the Harbour mouth; however the numbers at station California Bank were second highest. Moore and Sander (1979) had reported relatively low mean numbers (135m-3) at their offshore station, a magnitude lower than reported by the present study. In using a single mesh (~200µm) for their study they would have underestimated copepodites by a factor of three to six, and also ]]>
In general there was a change in ]]>
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1. Department of Life Sciences, University of the West Indies, Mona, Kingston 7, Jamaica; hugh.small@uwimona.edu.jm, mona.webber@uwimona.edu.jm. dale.webber@uwimona.edu.jm, krislue@gmail.com.
Received 19-IX-2013 Corrected 31-III-2014 Accepted 01-IV-2014
