Decapod Shrimps in the Antarctica

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Chapter 3 Diversity and Distribution of deep-sea shrimps in the Ross Sea Region of Antarctica

Abstract

Although decapod crustaceans are widespread in the oceans, only Natantia (shrimps) are common in the Antarctic. Because remoteness, depth and ice cover restrict sampling in the South Ocean, species distribution modelling is a useful tool for evaluating distributions. We used physical specimen and towed camera data to describe the diversity and distribution of shrimps in the Ross Sea region of Antarctica. Eight shrimp species were recorded: Chorismus antarcticus; Notocrangon antarcticus; Nematocarcinus lanceopes; Dendrobranchiata; Pasiphaea scotiae; Pasiphaea cf. ledoyeri; Petalidium sp., and a new species of Lebbeus. For the two most common species, N. antarcticus and N. lanceopes, maximum entropy modelling was used, based on records of 60 specimens and over 1130 observations across 24 sites in depths from 269 m to 3433 m, to predict distributions in relation to environmental variables. Two independent sets of environmental data layers at 0.05° and 0.5° resolution respectively, showed how spatial resolution affected the model. Chorismus antarcticus and N. antarcticus were found only on the continental shelf and upper slopes, while N. lanceopes, Lebbeus n. sp., Dendrobranchiata, Petalidium sp., Pasiphaea cf. ledoyeri, and Pasiphaea scotiae were found on the slopes, seamounts and abyssal plain. The environmental variables that contributed most to models for N. antarcticus were depth, chlorophyll-a concentration, temperature, and salinity, and for N. lanceopes were depth, ice concentration, seabed slope/rugosity, and temperature. The relative ranking, but not the composition of these variables changed in models using different spatial resolutions, and the predicted extent of suitable habitat was smaller in models using the finer-scale environmental layers. Our modelling indicated that shrimps were widespread throughout the Ross Sea region and were thus likely to play important functional role in the ecosystem, and that the spatial resolution of data needs to be considered both in the use of species distribution models.

Introduction

Natant decapod crustacea (shrimp and prawns) are ubiquitous in the world’s oceans and shallow seas, including the Antarctic, where other decapod taxa are largely absent (Clarke, 1990; Briggs, 1995; Arntz et al., 1999; Gorny, 1999; Thatje & Arntz, 2004; Griffiths, 2010). As they are predominantly benthic particulate feeders and predators, they can be important in processing of material at the seabed (Coull & Bell, 1983; Field, 1983; Cartes et al., 2007). Studies by Arntz & Gorny (1991) and Gutt et al. (1991) using underwater photography, have described species composition, distributions, and habitats of three shrimp species in the Weddell Sea but no similar studies have been conducted for the Ross Sea. The benthic fauna of the Ross Sea continental shelf has been relatively well-studied, particularly in coastal regions, and shares many taxa with other sectors of the Antarctic (Berkman et al., 2005; Thrush et al., 2006; Smith et al., 2007; Cummings et al., 2010), but deeper benthic habitats of the shelf edge, slope and abyssal depths remain little-sampled (Arntz et al., 1994; Clarke & Johnston, 2003; Brandt et al., 2007). Brandt et al. (2007) have highlighted the high rate of discovery for new species from the deep Southern Ocean, where up to 86% of isopod crustacean species were new to science, and argued that priority should be given to identifying the spatial distribution and abundance of key species in each trophic group across the region. Decapods are key species in the functioning of marine ecosystems, world-wide, as predators, scavengers, detritivores, and prey (Welsh, 1975; Coleman & Williams, 2002). In the Ross Sea region, information on the distributions and population densities of shrimps is necessary for producing ecosystem models which will improve understanding of trophic interactions and inform environmental management (Pinkerton et al., 2010).
Because sampling in the Antarctic is restricted by remoteness, intense seasonality, and sea-ice, species distribution models (SDM) may provide a useful tool for estimating species’ occurrences from limited field sample data. The basic assumption of SDM is that the fundamental niche of a species, defined by physiological and ecological tolerances, is the primary driver of its realized distribution (Soberón & Peterson, 2005; Soberón & Nakamura, 2009). Few SDM studies have focused on marine invertebrates, yet these groups have several attributes that make them well suited to species distribution modelling. Restricted availability of marine data (Kaschner et al., 2006), and a limited number of high quality species occurrence records were considered as obstacles behind the application of SDM in the ocean (Elith & Leathwick, 2009; Tyberghein et al., 2012). In recent years, as more sophisticated modelling algorithms have become available, the potential to model species’ distributions across un-sampled marine regions is now realistic. In this study, we used MaxEnt, a machine-learning algorithm based on the principle of maximum entropy (Jaynes, 1982), which has been shown to have superior performance among presence-only algorithms (Elith et al., 2006) for species distribution modelling (Phillips et al., 2004; Phillips et al., 2006).
Marine environmental datasets available for use in SDM have varying spatial resolutions and are frequently provided in different file formats, making the data assembly a time-consuming aspect of SDM studies (Tyberghein et al., 2012). Studies in the terrestrial domain have found that coarser spatial resolution resulted in reduced accuracy of predicted area although overall in SDM performance was not affected (Tobalske, 2002; Guisan et al., 2007; Seo et al., 2009; Song et al., 2013). However, the effect of spatial resolution on the relative influence of environmental variable on species distributions has not been assessed. The availability of several environmental datasets for the present study area, each with different spatial resolution provided an opportunity to investigate the effect of spatial resolution on the influence of environmental variables and the accuracy of the predicted area in the marine environment for the first time. In this study we used sample data on shrimp distribution in the Ross Sea, Antarctica, with historical records of occurrence, and two sets of environmental variables to (a) explore the diversity and distribution of shrimps in the Ross Sea region, (b) model distributions of suitable habitat for two common species, and (c) investigate the effect of using datasets with differing spatial resolutions on model predictions in the marine environment.

Methods

Study area

Our study area was bounded by 65° S, 150° E, 140° W, and the Ross ice shelf in the south. It included the entire Ross Sea continental shelf area, the Balleny Islands, and Scott and Admiralty seamounts (Figure 3.1). The mean depth of the Ross Sea continental shelf is about 500 m, although depth varies widely between deep troughs and shallow banks, and the area free of glaciers and permanent ice shelves is ca. 433,061 km2 (delineated by 800m isobath and the Ross Ice Shelf).

Sampling

Samples were collected from 12 February to 11 March 2008 during New Zealand’s 2008 International Polar Year – Census of Antarctic Marine Life voyage (IPY-CAML, RV Tangaroa, TAN0802) at 10 sites on the Ross Sea continental shelf, 10 sites on the northern continental slope, 3 sites on the abyssal plain (>3000 m depth), and 5 seamounts to the north (Figure 3.1). At each site, at least one, and up to 7, 1 hour deployments of a towed camera system with high definition digital video and still image cameras were made (Hill, 2009). The camera array (NIWA’s Deep Towed Imaging System, DTIS) was held ca. 2.5 m above the seabed and towed at 0.25–0.5 ms-1. In total, 55 camera transects were run. The seabed position was recorded in real time using an ultra short baseline (USBL) acoustic transponder system (Simrad HPR 410). Camera transects at each site were followed by physical sampling gear including a beam trawl (4 m width, 25 mm mesh), a large demersal fish trawl (25 m wing spread, 40 mm mesh), and two types of epibenthic sled; a fine mesh (1 m width, 0.5 mm mesh) sled used on flat, smooth seabeds (Brenke, 2005), and a coarse mesh (1 m width, 25 mm mesh) sled used on seamounts. In addition, a fine-mesh midwater trawl, was used following acoustic surveys. It had a circular mouth opening of about 12 m diameter and a cod end mesh of 10 mm and was generally towed for 20-30 min at 3-4 knots. All shrimp specimens collected by trawls and sleds were preserved (except in large hauls where representative 2-5 specimens/station were preserved) in 99% ethanol and were identified to species level.
Post-voyage analyses of video transects were run using Ocean Floor Observation Protocol (OFOP; www.ofop-by-sams.eu) software. Raw USBL transponder positions were first smoothed using a running mean and splined with associated metadata (e.g., time, depth, heading, vehicle altitude) to yield corrected seabed tracks with position coordinates and metadata values at 1 -s intervals. The digital video files were then synchronised with the corrected position data to enable re-running of transects in the laboratory with full video playback control and precise spatial and temporal logging of events. All shrimps on all transects were recorded and identified as close to species level as possible, using the high-resolution still images to confirm identities.

Environmental variables

We compiled environmental variables from two different sources (referred to hereafter as SET 1 and SET 2), each with different spatial resolution (Table 3.1, 3.2 and Figure 3.2). We selected variables that were likely to be ecologically relevant to benthic distributions: depth; seabed slope or rugosity; bottom temperature; ice concentration (proportion of the year with >85% ice cover in SET 1, annual mean in SET 2); chlorophyll-a concentration (mean summer in SET 1, mean annual concentration in SET 2) and for SET 1 only, bottom current speed.
SET 1 had a spatial resolution of 0.05˚ longitude and 0.05˚ latitude (Grant et al., 2006; Rickard et al., 2010), representing approximately 5.5 km by 2 km at areas between 67˚S and 68˚S , and consisted of 7 variables derived from satellite observations and modelled climatologies. SET 2 had a spatial resolution of 0.5˚ and consisted of 6 variables obtained from AquaMaps (Kaschner et al., 2008) (Table 3.1).
All datasets were received in raw csv format, and interpolated to raster layers at the respective spatial resolutions using the “Spatial Analyst” extension in ArcGIS 10. Inverse distance weighted (IDW) multivariate interpolation (Shepard, 1968; Daly, 2006) was used in the ArcGIS Spatial Analyst extension with default setting and smoothing (p=2) option to assign the final interpolated cell value in the generated raster layers. Chlorophyll-a was transformed to natural log to improve normality in SET 1 (Table 3.2). Raster layers were converted to ASCII grid with WGS84 Antarctic Polar Stereographic projection. The finer resolution dataset (SET 1) had almost 45 times more grid cells across the study region than the coarser resolution dataset (ca. 450,000 vs 10,000).
There were missing values in some layers in SET 2 but not in SET 1. During raster interpolation, these ‘no data’ pixels were assigned average values of 12 surrounding (ocean) cells using ArcGIS raster calculator. The “Band Collection Statistics” multivariate toolset function (Snedecor & Cochran, 1968) of Spatial Analyst was used to calculate Pearson’s correlation coefficient between the variables in each dataset (Appendix I). Correlation coefficients over ± 0.7 were considered significant (Cohen, 1988; Dormann et al., 2013) and are known to affect model prediction capability (Jiménez-Valverde et al., 2009; Veloz, 2009).

Modelling fitting procedure

Using all occurrence data from TAN0802 physical and photographic samples, we modelled the two most commonly-occurring shrimp species, Notocrangon antarcticus and Nematocarcinus lanceopes, using MaxEnt version 3.3.3e (http://www.cs.princeton.edu/~schapire/maxent/), with each of the two sets of environmental variables as predictors in consecutive runs for each species (Table 3.1 and Figure 3.2). Our occurrence records were distributed over 160,000 km2, which is sufficiently spatially segregated to reduce the probability of spatial-correlation between observations (Phillips et al., 2004; Mateo et al., 2010). MaxEnt is flexible with respect to the types of variables used and the form of their relationship to a species’ presence (e.g. linear, nonparametric, etc.). A review comparing 16 models of >200 taxa found that machine-learning methods including MaxEnt consistently outperformed traditional linear methods (Elith et al., 2006) and that presence-only models were preferable because limited sampling may mean that apparent absences may not be true. We selected the ‘Auto features’ function for model fit in MaxEnt, which automatically applies the feature or features estimated to be appropriate for the particular sample size of occurrence records (Phillips & Dudik, 2008). As the number of records varied depending the resolution of the datasets in this study, only linear, quadratic and hinge features (See Elith et al., 2011 for definitions) were utilized for model fitting.
MaxEnt models were generated using 100 bootstrap replicates run with the ‘random seed’ option turned on. The ‘Remove duplicate presence records’ feature was enabled to exclude duplicate records that fell within individual pixels of background environment layers on each dataset and the occurrence records were split into 75% for training and 25% for testing for bootstrap replications. The Maximum number of background points (randomly selected in each replication) was increased to 100,000 instead of the default value of 10,000 because of our large-scale mapping objective. Maximum iterations were also increased to 1000 allowing enough time for model convergence. As suggested by Phillips & Dudik (2008) the default regularisation value was used because it results in better performance of evaluation data for presence-only datasets. We also used the settings ‘fade by clamping’ option to minimize unreliable extrapolation into areas with environmental conditions that were not encountered during model training. The 51 relative contributions of variables were calculated in the MaxEnt models in training steps where the algorithm keeps track of how much each environmental variable contributes to fitting the model and adjusts the overall gain to calculate contributions of individual variables.

Model evaluation

Various test statistics are available to test the ability of models to discriminate suitable versus unsuitable habitat (Fielding & Bell, 1997; Ferrier et al., 2007). Several studies have highlighted issues with using only one statistic to evaluate model performance (Elith & Graham, 2009; Merow et al., 2013). Options for model validation include: (1) internal validation, or cross-validation in which the data are partitioned randomly into ‘training’ and ‘test’ sets, thus creating quasi-independent data for model evaluation (Fielding & Bell, 1997; Guisan et al., 2007) using the Area Under the receiver operating Curve (AUC; Phillips & Dudik, 2008; Elith et al., 2011; Dambach et al., 2012) criterion; (2) omission rates (Anderson et al., 2003; Shcheglovitova & Anderson, 2013); (3) low presence threshold (LPT; Pearson et al., 2007), and (4) completely independent datasets (Fleishman et al., 2003; Elith et al., 2006; Williams et al., 2009; Newbold et al., 2010; Wilting et al., 2010). We validated our models using all four of these methods.
AUC measures the quality of a ranking of sites (Elith & Burgman, 2002). Use of AUC analysis with presence-only evaluation datasets has been justified for the presence versus random classification problem (Wiley et al., 2003). AUC is measured on a scale of 0–1, where 1 indicates no errors of omission or commission, 0.5 indicates no better than random selection, and 0.9 indicates that there is a 90% chance that predicted habitat suitability for a randomly drawn species presence will be higher than that of randomly drawn absence (Wiley et al., 2003; Fawcett, 2006; Williams et al., 2009). MaxEnt provides AUC values based on the evaluation localities used in each model run. In this study, mean AUC values calculated from 100 bootstrap models were used to measure model performance. MaxEnt’s built-in Jack-knife validation method was also used as an independent estimate of each variable’s contribution to overall model performance allowing comparison with AUC values for each variable.
The threshold-dependent intrinsic (based on training data) or extrinsic (based on test data) omission rate, is the fraction of the known presence localities that fall into pixels not predicted as suitable for the species. A low omission rate is indicative of a good model (Anderson et al., 2003). High-quality models should show zero or low omission of evaluation localities, or at least predict evaluation localities statistically better than random.
LPT sets the lowest threshold value of the prediction for any of the presence localities in the training dataset (measured on a scale of 0-1) (Pearson et al., 2007). This yields a binary prediction that includes all pixels that are at least as suitable (according to the model) as those where the species was known to be present (in the training dataset). These threshold values generally vary by model. We also checked the models using a fixed threshold value of 10 out of 100 for the cumulative output. MaxEnt provides a convenient interpretation for the output of cumulative probabilities, where the expected omission rate for localities of the species is equal to the threshold employed. For example, an ideal model and a threshold of 10 would be expected to yield approximately 10% omission in an independent, unbiased sample of localities of the species. Hence, use of the fixed threshold of 10 is expected to lead to omission levels of approximately 10%.
Using an independent dataset is the optimal method for evaluating model performance (Phillips & Dudik, 2008; Kumar & Stohlgren, 2009). We used 6 N. lanceopes and 58 N. antarcticus occurrence records in the Ross Sea area extracted from the Ocean Biogeographic Information System (OBIS, www.iobis.org), the SCAR-Marine Biodiversity Information Network (SCAR-MarBIN, www.scarmarbin.be), and the published literature (Appendix II). Records were filtered to remove duplicates (i.e. same co-ordinates or same records from different sources) and apparent geographic errors (i.e. co-ordinates plotting on land or in different regions) before combining them into a single data set for model verification using GIS. Probability of occurrence values, which ranged from 0 to 1, where 0 meant no probability of presence and 1 meant highest probability of presence at that particular location, were extracted from the average of all bootstrap models on each data set using the “Extract Values to Point” function of Spatial Analyst in ArcGIS. We evaluated model accuracy with the independent dataset by seeing how successfully the model predicted the species’ potential distribution outside its sampled distribution using six model evaluation metrics (each measured on a scale of 0-1), namely: Percent Correct Classification (PCC, overall accuracy); Sensitivity (the proportion of actual presences that are accurately predicted); Specificity (the proportion of actual absences that are accurately predicted); False Positive Rate; False Negative Rate, and True Skill Statistics (TSS, correct classification rate in relation to false positive rate) (Franklin & Miller, 2009).

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Results

Sampled diversity and distribution

In total, 921 shrimp specimens (91 preserved) were collected and 1249 individuals observed in video transects across 24 different sites (Table 3.3). Eight species were identified; Chorismus antarcticus (Pfeffer, 1887); Notocrangon antarcticus (Pfeffer, 1887); Nematocarcinus lanceopes (Bate, 1888); Pasiphaea scotiae (Stebbing, 1914); Pasiphaea cf. ledoyeri (Hayashi, 2006); Petalidium sp.; an unidentified damaged specimen of the suborder Dendrobranchiata; and a new species of Lebbeus (S. Ahyong, unpublished data). Chorismus antarcticus and Notocrangon antarcticus were found only on the continental shelf in depths shallower than 1000 m. Chorismus antarcticus was largely restricted to depths shallower than 700 m, whereas N. antarcticus was found down to ca 1000 m at sites out to the edge of the continental slope. None of the other species were found on the continental shelf or at depths shallower than 450 m. Nematocarcinus lanceopes, Petalidium sp., Pasiphaea cf. ledoyeri and Pasiphaea scotiae were found on the continental slope and northern seamounts, but only N. lanceopes and P. scotiae were found at abyssal depths. Dendrobranchiata was found only at one site on the continental slope, and Lebbeus n. sp. was found only on the northern seamounts (Table 3.3 & Figure 3.3). Notocrangon antarcticus was the most frequently recorded species on the continental shelf (440 individuals, depth range 269–930 m) and N. lanceopes was the most frequently recorded species elsewhere (1554 individuals, depth range 570–3433 m). Larvae of N. lanceopes were also recorded from MOCNESS (Wiebe et al., 1976) samples on the slope and seamounts (4 individuals, 110-800 m). The distributions of these two species overlapped at one site (D27) on the northern continental slope (Table 3.3 &Figure 3.3).

Modelled distributions

A total of 281 N. antarcticus and 909 N. lanceopes occurrence records were available from the TAN0802 cruise, including both physical specimens and records from video transects. When duplicate presence records within each grid cell were excluded there were 22 and 41 presence records at the fine spatial resolution (SET 1) for N. antarcticus and N. lanceopes, respectively, and 12 and 17 records for the two species, respectively, at the coarser resolution (SET 2) (Table 3.4). For both N. antarcticus and N. lanceopes, the extent of predicted suitable habitat was greater in the coarser spatial resolution model (SET 2) and less in the finer model (SET 1). There were also differences in the locations of highest probability of occurrence values between SET 1 and SET 2 models. This was particularly noticeable for N. antarcticus, for which the coarser resolution SET 2 models show wider distribution of suitable habitat across northern and western areas of the continental shelf than do the finer resolution SET 1 models (Figure 3.5). Both of the modelled distributions indicated geographic separation of the two species at the shelf break (Figure 3.5). The predicted distribution for N. antarcticus was restricted to the Ross Sea continental shelf, whereas suitable habitat for N. lanceopes was predicted to occur on the continental slope, Scott and Admiralty seamounts, and around the Balleny Islands, with lower probability of occurrence on the abyssal plain near these features (Figure 3.5).

Model evaluation

AUC values for both models were high (>0.9) and significantly different from a random prediction (Wilcoxon rank-sum test, p <0.01) (Table 3.4). High test gain (all values > 1), indicated that <0.1% of the withheld test presences were misclassified. Intrinsic omission rates for all models were zero and extrinsic omission rates were <0.1, indicating acceptable model performance (Phillips et al., 2006).
LPT and 10th percentile presence threshold values were lowest at the finer spatial scale of SET 1 for both species; 0.168 and 0.291 for N. antarcticus, and 0.031 and 0.141 for N. lanceopes, respectively. Corresponding values using the coarser spatial scale in SET 2 were higher (0.431 and 0.431 for N. antarcticus, and 0.432 and 0.438 for N. lanceopes, Table 3.4). Because LPT is considered more suitable than the 10th percentile in cases where presence records have been collected in a short period of time and with high spatial accuracy (Rebelo & Jones, 2010) as in the present study, we used the LPT values as the suitability cut-off value for model validation using independent records.
For both species, mean probability of independent location records plotting within the predicted habitat suitability area was highest at the coarse spatial scale (SET 2, mean ± SD; 46.7 ± 0.19% and 80.9 ± 0.21% for N. antarcticus and N. lanceopes, respectively) and somewhat lower at the finer spatial scale (SET 1, 24.03 ± 0.11% and 19.6 ± 0.08%, respectively) (Table 3.4). Models of N. lanceopes had the highest accuracy based on the independent record evaluation metrics; in particular, PCC scores of 0.86 and 0.93 and TSS scores of 0.54 and 0.92 for SET 1 and SET 2 models, respectively. Corresponding values for N. antarcticus models were lower, at 0.62 and 0.60 for PCC and 0.27 and 0.17 for TSS, respectively.

Environmental variables

Temperature and depth were correlated with each other in SET 1 (r = 0.75), and salinity and slope were correlated with each other in SET 2 (r = 0.88) (Appendix I). MaxEnt has robust mechanisms integrated in the algorithm to deal with interactions of correlated variables (Phillips et al., 2009; Elith et al., 2011) so we did not exclude any variables from our variable pool.
MaxEnt model response curves show how the logistic prediction changed across the sampled range of each environmental variable, while keeping other variables at their average value (Figure 3.6). Each of these response curves represents a separate MaxEnt model created using only the named variable. The principal differences in environmental envelopes between the models of each species were in temperature range, chlorophyll-a, and ice concentration (Figure 3.6). The response curves indicated that N. antarcticus was likely to be found in lower seabed rugosity and slope areas that had colder waters with higher chlorophyll-a concentrations and lower ice concentrations than N. lanceopes.
In fine scale models of N. antarcticus using SET 1 variables, temperature, chlorophyll-a concentration, and depth had the highest contributions to the models, whereas using the coarse-scale SET 2 variables highest contributions were from depth, salinity, and chlorophyll-a concentration (Table 3.5). In the fine-scale SET 1 N. lanceopes models, highest variable contributions were from ice concentration, seabed rugosity, and depth, whereas in the coarse-scale SET 2 model, highest variable contributions were from depth, ice concentration, and temperature (Table 3.5). The maximum contribution of an individual variable to any model was 46.67 % (depth, SET 2, for N. antarcticus). Jack-knife analyses of model gains, and test AUC scores for models generated with a single variable indicated that the same variables listed above were the top predictors regardless of covariation.

Discussion

Diversity and distribution

The NZ IPY-CAML survey has extended the number of known sites with species-level records of deep-sea shrimps in the Ross Sea and provided the first such records from seamounts and abyssal regions in the north of the region (Appendix II, Figure 3.3 and Figure 3.4). These new observations have enabled us to re-evaluate known shrimp diversity and distribution in the Ross Sea region. Shrimps occurred throughout the region, with N. antarcticus being the most abundant species on the continental shelf, and N. lanceopes on the continental slope and seamounts to the north. Notocrangon antarcticus and Chorismus antarcticus occurred only on the shelf, whereas the five other species were only recorded off-shelf. These distributions reaffirm previous findings (Kirkwood, 1984; Childress et al., 1990; Arntz & Gorny, 1991; Gutt et al., 1991; Gorny et al., 1992 and citations therein; Gorny et al., 1993; Gorny & George, 1997; Gorny, 1999; Guzmán & Quiroga, 2005; Dambach et al., 2012). However, previous surveys (Bullivant, 1959, 1967; Gambi & Bussotti, 1999) did not find N. lanceopes, Petalidium sp. and Dendrobranchiata in the Ross Sea region, although a 2004 survey (NIWA unpublished data, Pinkerton et al., 2010) found N. lanceopes in six locations at north western Ross Sea around slopes near Cape Adare and Balleny Islands (Appendix II). Our results also show distinct depth zonation of C. antarcticus, N. antarcticus, and N. lanceopes, with a broad overlap between C. antarcticus and N. antarcticus in shelf regions, and between N. antarcticus and N. lanceopes on the upper slope; N. lanceopes being widely distributed in depths greater than 1000 m but less frequent in depth shallower than this (Figure 3.4). The new records of N. lanceopes and Pasiphaea spp. on seamounts north of the Ross Sea show that their distributions are more widespread than previously reported.

Modelled distributions

Although the present data increase the number of records of the shrimps in the Ross Sea region considerably, the available data remain insufficient to map their distributions with confidence. Therefore, we used species distribution models to predict the geographic distribution of the two most common shrimps, N. antarcticus and N. lanceopes, based on their occurrences at 23 different locations in the Ross Sea region. This study is also the first in the marine environment to assess of the effect on species distribution model performance of using different environmental datasets at different spatial resolutions.
For all MaxEnt models of the predicted habitat suitability for both N. antarcticus and N. lanceopes, independent validation records plotted into areas with predicted maximum probability of presence between 59-86 %, and all models had high AUC scores supported by high training gain and low omission rates, regardless of environmental dataset resolution. The AUC value tends to increase when the selected background area is larger than the species observed presence area (Phillips & Dudik, 2008; Merow et al., 2013). Thus, inclusion of other validation metrics is required for a thorough evaluation of model performance, particularly when our modelled species are known to have restricted distribution ranges (N. antarcticus in the shelf and N. lanceopes off-shelf) in a large geographic area. These results suggest that any of the modelled predictions are likely to be useful indications of distributions for these species, regardless of the spatial resolution of the underlying environmental data (Phillips et al., 2006; Anderson & Gonzalez Jr, 2011; Davies & Guinotte, 2011). However, there was appreciable variation between outputs of the different models (Figure 3.5, Table 3.4) and it is important both to understand which environmental variables are influencing the models and to consider factors that might underlie the differences between the models.
A recent study that modelled the distributions of N. antarcticus and N. lanceopes over the entire Southern Ocean using MaxEnt showed depth, ice concentration and salinity to have the highest explanatory power for models of N. antarcticus, while N. lanceopes distribution was better explained by depth, ice concentration and temperature (Dambach et al., 2012). In our study, at the scale of the Ross Sea region, depth, temperature, chlorophyll-a concentration, and salinity had highest explanatory power for N. antarcticus, whereas for N. lanceopes, ice concentration, depth, seabed rugosity, and temperature contributed most to the models. Given the spatial separation of these two species between the extreme high-Antarctic environment of N. antarcticus on the Ross Sea shelf and the more moderate oceanic environment of N. lanceopes beyond the shelf-break front, it is perhaps not surprising that these variables should contribute most to the models. Depth and seawater temperature are obvious distinctions between the two environments, shelf habitats being characterised by temperatures <0°C and depths <1000 m whereas beyond the shelf break temperatures are always >0°C and depths, other than on the seamounts, are >1000 m. However, the influence of the Ross Sea polynya also causes strong distinctions in ice concentration, salinity, and chlorophyll-a concentration between the environments of the two species (Figure 3.2). Seabed slope and rugosity are also important influences on benthic faunal distributions in the deep sea, e.g. by influencing food supply via current flow amplification (Mohn & Beckmann, 2002). Their influence in models, however, is likely to be strongly influenced by the spatial scales at which they are calculated. In our regional-scale models, the continental shelf break and slope, and the northern seamounts, are areas with high computed values for both slope and rugosity which contrast strongly with the comparatively uniform morphology of continental shelf and abyssal environments.
Because the steepest gradients in several potentially important variables coincide at the Ross Sea shelf break (depth, temperature, slope/rugosity, ice concentration, Chlorophyll-a concentration), determining which of these variables are most ecologically important to the realised distributions of the two species is problematic. Adaptation to cold has been postulated as the primary reason why Antarctic shrimps are capable of living at the extremely low temperatures of the continental shelf where other decapod taxa are absent (Wittmann et al., 2010), and as an explanation of why they were able to re-colonize high southern latitudes after past glaciation cycles (Gutt et al., 1991; Frederich et al., 2001; Wittmann et al., 2010). That only two of the eight species identified here have distributions on the continental shelf, and that there is strong demarcation between species’ ranges at the shelf break, suggests either that such adaptation is species-specific or that factors other than physiological adaptation to low temperature per se have a stronger influence on realised distributions.
Physiological studies have suggested that many Antarctic benthic invertebrates on the continental shelf are highly stenothermal, and thus have limited capacity to withstand future environmental warming (Peck, 2004). If the shrimp species studied here were currently range-limited by temperature, predicted warming might be expected to result in southward range shifts of those species currently found only in warmer waters north of the shelf break front (e.g. N. lanceopes, Figure 3.3). Conversely, for the two species with shelf-only distributions (C. antarcticus and N. antarcticus) at present, the only potential range shift would be southward into the region currently covered by the Ross Ice Shelf.
In addition to the suite of environmental variables used in species distribution modelling, three other factors were likely to affect the final outputs of the models and how well individual models rated in evaluation metrics. First, the number of independent records used to validate models can influence the test statistics (Newbold et al., 2010). In the present study, only 6 independent presence records were available to validate the N. lanceopes models, compared to 58 records for N. antarcticus, and it is likely that this will have had some effect on their respective validation metrics. Second, validation using independent records assumes geographic accuracy of the independent records; i.e., that the position data associated with these records are both accurate and precise. The accuracy of records derived from biodiversity databases can be uncertain, however (Feeley & Silman, 2010; Mesibov, 2013), and in the present study none of the records used for independent validation had spatial accuracy information associated with them. Therefore, it is possible that some of the independent records that plotted outside predicted areas of suitable habitat here might be as a consequence of such inaccuracies. Finally, the spatial resolution of the environmental datasets used in the models clearly influenced the predictions of the resulting models; this is discussed in more detail below.

Table of Contents
Abstract
Acknowledgements
Chapter 1 Thesis Overview
1.1 Natant Decapods
1.2 The Southern Ocean
1.3 The Ross Sea
1.4 Species Distribution Modelling to predict species potential distribution
1.5 Thesis objectives
1.6 Chapters overview and aims
1.7 References
Chapter 2 Decapod Shrimps in the Antarctica
2.1 Introduction
2.2 Biodiversity
2.3 Geographic distribution
2.4 Depth distribution
2.5 Ecology
2.6 References
Chapter 3 Diversity and Distribution of deep-sea shrimps in the Ross Sea Region of Antarctica
3.1 Introduction
3.2 Methods
3.3 Results
3.4 Discussion
3.5 References
Chapter 4 Dispersion and Faunal association pattern of common Antarctic shrimps
4.1 Introduction
4.2 Methods
4.3 Results
4.4 Discussion
4.5 References
Chapter 5 Global Marine Environment Datasets for environment visualisation and species distribution modelling
5.1 Introduction
5.2 Methods
5.3 Results
5.4 Discussion
5.5 Conclusion
5.6 References
Chapter 6 Past, Present and Future distribution of deep-sea shrimp in the Antarctica
6.1 Introduction
6.2 Methods
6.3 Results
6.4 Discussion
6.5 Conclusion
6.6 References
Chapter 7 General Discussion, Conclusion and Future direction
7.1 General Discussion
7.2 General conclusions of the thesis
7.3 Future direction
7.4 Reference
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