Assessment ofASU sampling package
CHAPTER FIVE
Assessment of Artificial Substrate Unit sampling package
5.1 Introduction
In Australia, comprehensive inventory data for most marine invertebrate taxa for a region is the exception rather than the rule (Ponder et al. 2002; Smith, 2005). For this reason, rapid assessment methods are increasingly being regarded as an important tool for addressing the current demand for biodiversity information by government and management agencies (Arvanitidis el a/., 2005; Smith, 2005). There are many different approaches to assess, monitor and conserve biodiversity and combinations of methods are proving useful (Giangrande el al., 2005). Challenges facing the cataloguing ofterrestrial invertebrates are similar to those experienced by marine invertebrate surveys. Developing rapid assessment methods to encompass such a diverse assemblage has led to novel approaches, many ofwhich include the used ofsampling packages. For example, to efficiently assess arthropod assemblages Kitching el al., (200 I) developed a 'sampling package' approach, using a number ofcollecting methods to sample regional fauna. As different collection techniques were more successful for certain taxa, a greater range of fauna could be sampled and a better overall community assessment was obtained.
In freshwater habitats, sampling packages based on Artificial Substrate Units (ASUs), and, combining a number ofstructures, are widely implemented across the United States,
United Kingdom, France and Belgium as a standardised monitoring method (De Pauw el aI., 1986; Costello & Thrush, 1991). Research has gone into developing effective and efficient ASU sampling protocols with experimentation into ASU composition (material, volume, particle size), deployment length, placement in the environment, representativeness of local fauna, and pollution sensitivity ofcolonizing fauna (De Pauw et al., 1986). The use ofa standardised method of monitoring freshwater habitats, which
153 Assessment ofASU sampling package
is repeatable across time and location, is valuable for long-term monitoring ofchanges and is a sound means ofcomparing biodiversity (De Pauw et al., 1986).
The development ofrapid assessment sampling packages for the marine environment is also beginning to progress. Zimmerman & Martin (2004) outline the construction of 2 ARMs - artificial reef matrixes, a 0.5 m , mainly cement structure with added features such as crevices, PVC pipe, coral rubble tray, filter matting and kitchen scourers. This structure targeted the macroinvertebrate fauna such as echinoderms, sponges and molluscs, and the attachment of small complex ASUs in the form ofthe kitchen scourers sampled the small invertebrate fraction ofthe reef community. Constructing and deploying these structures may be time consuming but is important as part of a non destructive method ofcataloguing invertebrate biodiversity in fragile and/or protected coral reef areas (Zimmerman & Martin, 2004).
Once a method is determined to be useful, it is important to test and re-test outcomes and look at its wider application. Re-testing is part ofthe application of logical progression in scientific research, thus refining models and further elaborating hypotheses (Underwood, 1990; 1991). Continuing research on successful techniques has demonstrated that some methods that are appropriate for one habitat/region may not be as functional or accurate elsewhere (Arvanitidis et al., 2005).
For studies investigating ASUs for monitoring and assessment, one ofthe most pressing questions being tested is how representative the assemblages recruiting to ASUs are of natural habitats and/or the local species pool (Myers & Southgate, 1980; Smith & Rule, 2002; Mirto & Danovaro, 2004). For this project, the aim was to determine if ASUs could recruit assemblages that were representative ofbiodiversity, defined by species richness and taxonomic distinctness measures. Previous studies have found that ASU assemblages were representative of local fauna in red coralline algae, kelp forests and other macroalgal habitats (Myers & Southgate, 1980; Edgar, 1991 a; Norderhaug et al., 2002). However, in a local study of fauna recruiting to scourer ASUs, Smith & Rule (2002) found ASU assemblages to be largely unrepresentative ofthose occurring in
154 Assessment ofASU sampling package
adjacent natural habitats; in particular, the ASUs under-sampled amphipod fauna. Smith & Rule (2002), however, deployed ASUs for a period of five months, a long deployment period, which may explain the low diversity and abundance ofamphipod fauna (see section 4.6).
The objective ofthis study is to formulate and test an ASU sampling package for the rapid assessment ofamphipod biodiversity. Therefore, the aim ofthe first section is to evaluate the most efficient ASU sampling package for collecting amphipod fauna. To determine the most efficient ASU sampling package, comparisons ofspecies richness of combinations ofASU types are made. As previous the section demonstrated that amphipod assemblages on different ASU types are variable across locations (section 4.4), combinations of ASU types were evaluated for each location, then, drawing from these results, a final ASU sampling package is formulated.
This ASU sampling package is then rigorously tested by assessing its ability to adequately sample local and regional biodiversity, using measures oftaxonomic distinctness. Finally, the ASU sampling package is trialed at an additional location, Bare Island, Botany Bay, as a further test ofthe ability ofthis package to effectively sample amphipod biodiversity. This trial was conducted at Bare Island (- 3 degrees south ofthe Solitary Islands region) because the subtidal habitat is different from that found at the study locations in the Solitary Islands, and the fauna at this site is particularly well catalogued (J. K. Lowry, pers. comm.).
155 Assessment of ASU sampling package
5.2 Methods
5.2.1 Additional study site - Bare Island, Botany Bay
Bare Island (33 0 59' 56"E 151 0 13' 88"S) is a small, 60-m wide, landmass 20 m from the shore in the protected waters of Botany Bay, Sydney, New South Wales (Figure 43; Figure 44). The island is a surrounded by a hard, sandstone reef which is dominated by the kelp Ecklonia radiata over the depth range from 0 - 6 m. Under the macroalgal canopy is a layer ofalgae consisting ofa sparse mix ofSargassum sp., dictyotaleans, Zonaria sp. and a turfoffilamentous coralline algae. The latter was heavily silted with a ~3 cm layer oforganic detritus at the time ofsampling. The topography below 6 m comprises walls and ledges covered in encrusting coralline algae. These "barrens" habitats are maintained by the grazing ofthe urchin Centrostephanus rodgersii and the limpets Patel/oida alticostata and Cel/ana tramoserica (Underwood et al., 1991). The reefterminates at a depth ofapproximately 10m and grades into a coarse, shelly, sedimentary habitat.
Bare Island was chosen as the site for this experiment as it is an area with a well known local amphipod fauna (J. K. Lowry, pers. comm.) and also represents a very different benthic habitat to those at locations previously sampled in the Solitary Islands region. In addition, Bare Island is situated in a cool temperate bioregion, as opposed to the subtropical bioregion ofthe Solitary Islands.
156 Assessment ofASU sampling package
~o KIn
Figure 43: Map ofthe Bare Island region, New South Wales, Australia. Modified from Kennelly & Underwood (1992).
• Study site
\ \ (J
{ I
l....----.J 10m
Figure 44: Map ofBare Island, Botany Bay, Sydney, New South Wales. Blue box indicates study site.
157 Assessment ofASU sampling package
5.2.2 Field and laboratory method
5.2.2.1 Collection of natural habitats at the Solitary Islands
During the dive to retrieve the final sixteen-week ASU deployment, a general collection was made all natural habitats within a 10m radius ofthe deployed racks. An airlift was used to sample fixed habitats such as corals, corallimorphs, and bare rock. All other habitats were collected using the "butcher's bag" method, (see section 2.1.3 for full description of habitats at each location). These samples, along with previous collections of natural habitats, were used to establish the source ofrecruiting amphipod fauna and to establish a master list ofamphipod fauna for the location. Natural habitats were not sampled at the beginning ofthe experiment, as this would have removed a potential source ofrecruits and so natural habitats had the same recent recruitment history as the ASUs prior to sampling.
5.2.2.2 Deployment method at Bare Island
Experimental racks were deployed on a flat, barren, sandstone ledge at 8 m at the edge of the kelp forest. The barrens area was chosen over a site within the thick kelp canopy due to the logistics ofclearing a space to house the racks with a buffer margin to reduce interference from kelp fronds. Two racks (described in section 2.3.5) containing two replicates ofeach ASU type were deployed at the study site giving a total offour replicates of each ASU type. Holes (6 cm deep) were drilled into the sandstone substrate using a pneumatic drill supplied by a SCUBA tank. Racks were secured in place by dynabolts (screws with expanding collars), which had an eyelet at the top, with four dynabolts used per rack. Malleable galvanized wire was looped through the dynabolt eyelet and the comer grid ofthe rack and the two ends ofthe wire were twisted together several times to secure the rack in place.
158 Assessment ofASU sampling package
Samples were deployed in February 2004 and retrieved four weeks later in March 2004. Samples were removed from the racks using the "butcher's bag" method (see section 2.3.1). Following collection, samples were processed using the method described in section 2.3.2.
5.2.3 Statistical methods
5.2.3.1 Construction ofsampling package
Following on from the colonization section, which investigated variation in assemblage with deployment time, this section used the dataset from the optimal deployment time (four weeks) as the best estimator ofamphipod species richness. To determine the optimal combination ofASU types to provide the most efficient sampling package, comparisons were made for every combination ofthe six different ASUs, using the cumulative species richness ofeach combination. ASU packages were tested as combinations oftwo, three, four and five ASU types with each type represented by four replicates. Species richness results ofmultiple combinations were compared within each location. The ASU combination which was considered the most effective was the grouping which had a species richness value equivalent to the total ASU species richness for the location and used the least number ofASU types. Based on the most efficient ASU combination from each location, a final ASU 'sampling package' was determined using one replicate ofeach ofthe ASU types identified.
Note, comparisons using a single ASU type for a sampling package were not made as there was insufficient replication within the experimental design to allow this. This option was not pursued in the design ofthe experiment due to logistical constraints, namely time taken to retrieve an additional 144 samples in the field was limited by sea and diving hours, and further by changing weather conditions in the area. It was therefore decided to focus on a range ofASU types rather than testing each ASU type for potential use as a single, universal sampler.
159 Assessment ofASU sampling package
5.2.3.2 Local and regional assessment of the ASU sampling package
The dataset used in this set ofanalyses was from the four-week ASU collections from each ofthe four Solitary Islands locations (North Solitary, North West Solitary, Muttonbird Island and Korffs Islet). The data for Holdfast ASUs were removed leaving the most effective ASU combination ofOnion, Poofie, Scourer, Rope and Astro ASUs (see section 4.3).
To assess how representative ASU sampling packages were ofamphipod biodiversity at each location, and also for the Solitary Islands region, measures oftaxonomic distinctness (average taxonomic distinctness - Delta+, and variation in taxonomic distinctness Lambda+) were applied (Clarke & WaIwick, 1998). Average taxonomic distinctness is a measure ofthe average distance ofa taxonomic path-length between all ofthe species present in an assemblage, and variation in taxonomic distinctness is a measure ofthe variance associated with this mean value and, therefore, reflects the evenness ofthe distribution ofspecies across the taxonomic tree for the assemblage (Clarke & Warwick, 1998). The taxonomic levels used in the analysis were species, genus and family. Further, hierarchical structure was not used as the phylogenetic treatment ofgroups within the Amphipoda is uneven and higher classification for amphipods is largely unresolved (Martin & Davis, 2001).
Master lists ofspecies for a location and the region were compiled from: i) the ASU collections; ii) collections made from natural habitat at a range of locations over the previous three years; iii) collections held by the Australian Museum; collections made by S.D.A. Smith from holdfasts (Smith, 1993); and collections made by MJ. Rule from Kitchen scourer ASUs (Rule, 2004). As the ASU sampling package developed here was designed to target epifaunal amphipod assemblages, species master lists against which the sampling packages were tested included only epifaunal species (Appendix 2.1). Therefore, scavenging, obligate commensal and fossorial amphipods were excluded (and
160 Assessment ofASU sampling package
further, were unlikely to recruit to ASU structures) (Appendix 2.1). Species lists from ASU sampling packages were compared to master lists for the relevant location as well for the region using the TAXDTEST function in PRIMER (2001) (Clarke & Warwick, 1998).
Results of local TAXDTESTs were generated as histograms for comparison ofASU packages at each location. In these plots, average, and variation in, taxonomic distinctness values for the ASU sampling package were plotted against an expected distribution generated by using 1000 random permutations from the master list (Warwick & Clarke, 1998).
For regional comparisons, sampling packages for each location, and for mixed locations, were tested. Mixed sampling packages were formulated to test ifa random set ofASU replicates sampled from the four islands are more or less representative than a sampling package deployed at a single location. Mixed location sampling packages consisted of a single replicate ofeach ASU type from each location. Combinations of replicates from locations were randomly selected and a total of four mixed sampling packages were formulated from the available pool ofreplicates. Results oftaxonomic distinctness measures for regional ASU sampling packages were plotted as points on a confidence funnel which defines the 95% probability limits ofthe null distribution for random re samples from the regional master list (Warwick & Clarke, 1998).
5.2.3.3 Assessment of Bare Island, Botany Bay deployment
Data were analysed using univariate and multivariate techniques in the Minitab (1996) and PRIMER (2001) software packages. One-way ANOVA was used to compare differences in species richness and average taxonomic distinctness between ASU types. Variation in the structure of amphipod assemblages on ASU types was investigat.ed using a one-way ANOSIM.
161 Assessment of ASU sampling package
To assess how representative ASU sampling packages were ofthe taxonomic composition ofthe local amphipod assemblage, measures oftaxonomic distinctness were applied. Average taxonomic distinctness and variation in taxonomic distinctness ofASU assemblages were compared to values drawn from a master list ofamphipod fauna for Bare Island using the TAXDTEST function in PRIMER (2001). The master list were complied from Australian Museum collection data and records provided by Dr. A.G.B. Poore (University ofNew South Wales), who has worked extensively on epifaunal species in the area.
162 Assessment ofASU sampling package
5.3 Results
5.3.1 Construction ofASU sampling package
At North Solitary, the minimum combination of ASU types required to achieve the highest species richness consisted ofonly three ASU types (Table 34). A combination of Onion, Poofie, and Astro ASUs formed the most efficient package sampling 23 species. The shared species richness between two ofthe three ASU types was also high, with Onion and Poofie ASUs each contributing only one unique species to the cumulative species total.
At North West Solitary, 28 species were collected on ASUs and five ASU types were required for an optimum combination. Poofie, Scourer, Astro, and Holdfast formed the most efficient package with either Onion or Rope ASUs as interchangeable units. Shared fauna between each ofthe ASU assemblages was high with Scourer contributing two unique species and all other units with only one unique contribution.
Muttonbird Island had the highest species richness ofall the islands (34 spp.). As seen for North West Solitary Island, a combination offive ASU types was required with two ASU types, Onion and Rope, as interchangeable units. There was less shared fauna between ASU types at Muttonbird Island; Astro ASUs collected four unique species, Holdfast two and Onion and Poofie one unique species each. At Korffs Islet 29 amphipod species were recorded from ASUs. Only four ASU types, Onion, Poofie, Scourer and Astro were required to form the most efficient combination. There was only one unique species on each of Rope and Astro and two on Onion ASUs.
Different ASU types were more successful at some locations than others. Based on the results ofthe most efficient ASU combination for each location, a final sampling package of five ASU types Onion, Poofie, Scourer, Rope and Astro was determined. Holdfast
163 Assessment ofASU sampling package
ASU were omitted from the sampling package because: i) they consistently failed to add extra species to the cumulative total; and ii) the durability ofthe units was poor.
Table 34: Listing ofthe most species-rich ASU combination for multiple comparisons ofASU type using combinations ofone to five ASU types, and total species richness (S). The species richness from adjacent natural habitats (NH) is also shown..r indicates that ASU types are interchangeable for an ASU combination. '*' indicates that three ofthe four ASU types listed in brackets are interchangeable to form the optimal ASU combination. 0 = Onion bag, P = Poofie, S = Scourer, R = Rope, A = Astra and H = Holdfast ASUs.
ASU NH Location Single Pair Triplet Quartet Quintet S S
North 18 22 23 23 7 A P,A O,P,A
North West 17 25 26 27 28 28 27 S P,S P,S,RlH S, (P/RIA/H)* P, S, A, H, RIO
Muttonbird 23 27 30 33 34 34 25 A P,A P/R, A, S/H P,S,A, H P, S, A, H, RIO
Korffs 23 28 28 29 29 22 A O,A 0, A, P/S/H 0, P, S,A
To assess how efficient ASUs were as a biodiversity sampling technique, comparisons were made between the species richness ofASU sampling packages (total ASU species richness) and the species richness ofsurrounding natural habitats for each location. At each location, the ASU sampling packages recruited more species than the extensive collections from the natural habitats (Table 34). At North Solitary, adjacent natural habitats were particularly depauperate with ASU packages recruiting 16 species that were not found in natural habitats, At North West Solitary, comparison between ASUs and natural habitats were more similar, with 29 species recruiting to ASUs and 28 species sampled from natural habitats. At Muttonbird Island ASUs once again collected a much
164 Assessment of ASU sampling package
higher species richness with 34 species recorded from ASUs and 25 from natural habitats. Similarly, ASUs at Korffs Islet collected a greater number ofspecies richness (29) than found in adjacent natural habitats (22).
5.3.2 Representativeness of local biodiversity
Results ofthe average and variation in average taxonomic distinctness ofsampling packages with location were mixed. Not all packages were reflective ofthe local species pool (Fig. 45).
At North Solitary and Muttonbird Island, both average and variation in taxonomic distinctness were within the 95% confidence intervals. At North West Solitary both average and variation in taxonomic distinctness were outside the 95% confidence intervals ofthe distributions generated from the master lists (Fig. 45). Values for these locations were at the higher (for average taxonomic distinctness) and lower (for variation in taxonomic distinctness) ofvalues. This indicated that the assemblages recruiting to ASUs at these locations were closely related and some species were over represented in the sampling package. At Korffs Islet average taxonomic distinctness but not variance in taxonomic distinctness was with in 95% confidence intervals ofthe local distribution of species, with the assemblage recruiting to ASU sampling packages over represented some groups.
Table 35: Summary ofvalues for average taxonomic distinctness (Delta+) and variation in taxonomic distinctness (Lambda+) ofASU sampling packages for location, as tested against respective local master lists.
Source ofvariation Delta+ P Lambda+ P
North 96.179 0.432 200.597 0.230 North West 95.862 0.030 197.437 0.012 Muttonbird 96.296 0.304 183.146 0.286 Korffs 95.627 0.060 222.217 0.044
165 Assessment ofASU sampling package
a) ! >-. 0 c: Q) ::J 1 c:r ~ u.. -. - I b)
...... ;..~- c)
1. .J
d)
-~ .1oI'" -_-..-..
Average taxonomic distinctness Variation in taxonomic distinctness Figure 45: Histogram ofaverage taxonomic distinctness (Delta+) and variation in taxonomic distinctness (Lambda+) ofASU sampling packages plotted against the distribution generated from the master list for: a) North Solitary; b) North West Solitary; c) Muttonbird Island; and d) Korffs Islet. Blue arrows indicate values for the AU sampling packages. (see also Table 35 for values).
166 Assessment ofASU sampling package
5.3.3 Representativeness of regional biodiversity
For the regional analysis, the results for average taxonomic distinctness indicated that not all ASU sampling packages were within 95% confidence intervals ofthe distribution drawn from the regional master list. The sampling package from Korffs Islet was below the confidence intervals while the Muttonbird Island package was on the lower borderline (Figure 46). This indicated that species recruiting to these packages were drawn from a narrower taxonomic range (i.e. were more closely related) than the "average" sample from the master list. The remaining two single location packages, from North and North West Solitary, were within the 95% confidence intervals (Fig. 46). The four mixed location packages, which represented a combination ofASUs from all locations, were all within the 95% confidence intervals and supported relatively high species richness (Figure 46).
For variation in taxonomic distinctness, three ofthe eight sampling packages, North Solitary and two mixed packages, were within the 95% confidence intervals (Figure 46). The remaining five sampling packages were either on the upper borderline or showed a higher variance in taxonomic distinctness, indicating that assemblages on these packages were over-representing certain family or generic groups.
To investigate which ofthe amphipod epifauna were not sufficiently sampled by the sampling packages, and to potentially improve the method, the master list for the region was scrutinized to determine which species were absent from ASUs.
Ofthe total of72 species that were considered potential ASU recruits, 48 species were recorded on at least one sampling package. Ofthe 24 species not recorded on ASUs, 11 species have been collected as less than 20 individuals, and a further 8 species as less than 40 individuals, in total, for all collections from the region. Several ofthese 'rare' species have not been reported since their original collections (many over 13 years ago from Australian Museum collections), despite considerable sampling within and around their original collection localities.
167 Assessment ofASU sampling package
a) b)
75 - 200
74 150 KI A N + A + 73 A B ro MB ~ A c "'0 NW NW A~ .0 Q) 100 N A 0 A A E 72 K1 ro A i t .....J A A A 50 71
70 0- 10 15 20 25 30 35 40 10 15 20 25 30 35 40 Number of species Number of species
Figure 46: Probability funnels (95% confidence intervals) for a) average taxonomic distinctness (Delta+) and b) variation in taxonomic distinctness (Lambda+) forregional analysis. N = North Solitary, NW = North West Solitary, MB = Muttonbird Island, KI = Korffs Islet and A - 0 = mixed location packages.
To confirm that lower values ofaverage taxonomic distinctness were due to this group of 'rare' fauna, ASU sampling packages were re-tested against a restricted regional master list from which the above mentioned 19 species had been removed. Results ofthe restricted regional analysis confirmed that the lack ofthese rare species contributed strongly to the high taxonomic variance seen in original analysis (Figure 47). Results for both average, and variation in, taxonomic distinctness show more representative results, with all sampling packages falling within the 950/0 confidence intervals (Figure 47).
The Muttonbird Island sampling package recruited the highest species richness and three mixed sampling packages also recruited a high species richness (Figure 47). One mixed location sampling package (package A), did not perform as well as other packages. Based on these results, mixed sampling packages are recommended as the best approach for sampling regional amphipod fauna.
168 Assessment of ASU sampling package
a) b)
76 250
74 200
N A + + 72· ... ro 150· KI .. "t:l ~ .0 .. MB Q) E N NW 0 ro 100 - .. A .. 70 ....J .. n"
68 50
66 0 10 15 20 25 30 35 40 10 15 20 25 30 35 40 Number of species Number of species
Figure 47: Probability funnels (95% confidence intervals) for a) average taxonomic distinctness (Delta+) and b) variation in taxonomic distinctness (Lambda+) for analysis ofrestricted regional list.
5.3.4 Results from Bare Island
5.3.4.1 Univariate analysis ofASU assemblages at Bare Island
A total of 1,544 individuals representing 38 species from 15 families were collected using the ASU sampling packages at Bare Island. Onion ASU assemblages recruited the highest species richness, closely followed by Rope and Poofie ASUs (Figure 48). The one-way ANOVA for species richness and taxonomic distinctness showed significant differences between ASU types (Table 36) and Tukey's pairwise comparisons ofASU types for species richness revealed a significant difference for two ofthe ten pairwise comparisons (Table 36). Onion ASU assemblages were significantly different from Scourer and Astro ASUs, and this was a result ofa higher species richness on the Onion ASUs (Table 36; Figure 48). For taxonomic distinctness there was a significant difference between Astro and Scourer ASU assemblages, these being the extreme ends of the range ofvalues recorded (Figure 49).
169 Assessment ofASU sampling package
III,. III ~14• U -;: 12 III c:i• ,. •Q. III 8
•DI l! • ~.•
Onion Poofie Scourer Rope Astro
Figure 48: Mean species richness (±SE) ofamphipods by ASU type at Bare Island.
:: "'+------=----I CIl .su 82+------+---- :E ~ 80+------+------u eo .. c o ~ .. ~
•01"' l! Ql > 82 ~
Onion Poofie Scourer Rope Astro
Figure 49: Average taxonomic distinctness (±SE) ofamphipods by ASU type at Bare Island.
From observations ofthe dataset, several species appeared to show a significant association with particular ASU types and these relationships were tested using a one way ANOVA for each species and subsequent pairwise analysis, were appropriate, were used to explore this pattern. Seven species were shown to have a significantly higher abundance on a specific ASU type (Table 37).
Aora maculata showed significant pairwise contrast for comparisons between Onion and Astro ASUs, with a greater abundance on later ASU type (Table 37). Ampithoe caddi has a high abundance on Rope ASUs, compared to Poofie, Scourer and Astro ASUs.
170 Assessment of ASU sampling package
Amaryllis sp., Cyproidea ornata and Ericthonius forbesii had a greater abundance on Pootie, Rope and Onion ASUs respectively, in comparison to all other ASU types. Two species, Paradexamine moorhousei and Telsosynopia trifidilla has a greater abundance on Astra ASUs. The former in comparison to Pootie and Rope ASUs, the later with all other ASU types (Table 37).
Table 36: Summary ofone-way ANOVA and pairwise comparisons between ASU types for species richness and taxonomic distinctness ofamphi pod fauna at Bare Island. *** - significant P
Source of Species Taxonomic variation richness distinctness
MS FP MS FP One-way ANOVA ASU type 29.26 4.32 0.011 33.56 5.30 0.004
Pairwise comparisons Onion vs Poofie ns ns
Onion vs Scourer *** ns
Onion vs Rope ns ns
Onion vs Astra *** ns
Poofie vs Scourer ns ns
Poofie vs Rope ns ns
Poofie vs Astra ns ns
Scourer vs Rope ns ns
Scourer vs Astra ns ***
Rope vs Astra ns ns
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Table 37: Summary ofone-way ANOYA results and pairwise comparisons of listed species with ASU types. Result for pairwise comparison is given as number ofcomparisons significant for the respective ASU type.._. indicates that there were no significant comparisons.
One-way ANOVA Pairwise comparisons
Source ofvariation F P Onion Poofie Scourer Rope Astro
Aora maculata F= 3.26 0.033 Ampithoe caddi F= 8.59 <0.001 3 Amaryllis sp. F= 12.13 <0.001 4 Cyproidea ornata F= 6.28 0.002 4 Ericthonius forbesii F= 5.76 0.003 4 Paradexamine moorhousei F= 4.93 0.006 2 Telsosynopia trifidilla F= 6.46 0.002 4
5.3.4.2 Multivariate analysis ofASU assemblage at Bare Island
A 3D nMDs ofASU types at Bare Island showed some ASUs were better at consistently sampling a similar suite ofspecies than others (Figure 50). The Onion and Rope ASUs were tightly cluster together just above the centre ofthe ordination (IMD = 0.69 and 0.72, respectively). Poofie ASUs were also fairly closely grouped at the top ofthe ordination (IMD = 0.73). Scourer ASUs were loosely spread at the upper centre ofthe ordination (IMD = 1.27), whereas Astro ASUS were highly dispersed (IMD = 1.60) and spread across the bottom left ofthe ordination (Figure 50). ANOSIM analysis showed no significant difference between ASU types (global R = 0.986, P = 0.063), primarily due to high levels ofvariation in the assemblage structure within some ofthe ASU types.
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Figure 50: nMDS ordination ofamphipod assemblage structure by ASU type at Bare Island. Light blue triangles = Onion bags, purple diamonds = Poofies, red circles = courers, yellow inverted triangles = Rope, green squares = Astra. Kruskal's stress level = 0.12.
5.3.4.3 Representativeness of ASU samplipg package at Bare Island
For both average taxonomic distinctness and variation in taxonomic distinctness, the ASU sampling packages at Bare Island recruited amphipod assemblages that were within the 95% confidence intervals ofthe distribution derived from the master list. This indicated that the amphipod assemblages collected by the sampling package were highly representative ofthe epifaunal amphipods at Bare Island (Figure 51).
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a) b)
75 200
74 150 73 + + BI ro 19 ... "'0 BI 72 .0 100 Q) E ... 0 ro .....J 71 50 70
69 0 10 15 20 25 30 35 40 10 15 20 25 30 35 40 Number of species Number of species
Figure 51: Probability funnels (95% confidence intervals) for a) average taxonomic distinctness (Delta+) and b) variation in taxonomic distinctness (Lambda+) for Bare Island.
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5.4 Discussion
In this section, different ASU types were more successful at some location than others (Table 34). As seen in the previous chapter assemblages recruiting to different ASU types showed different responses across locations (see section 4.4). Therefore it is not surprising that a final ASU sampling package included most ASU types tested. At three locations, North West Solitary, Muttonbird Island and Korffs Islet, optimal combinations ofASUs could be achieved by several different configurations with ASU ~ypes interchangeable in some instances (North West Quartet combination). This serves to reiterate that there was a high degree ofshared fauna between ASU types, as discussed in section 4.5.
ASUs versus natural habitats ASUs were a more effective sampling method than the collection of natural habitats, yielding a higher species richness than natural habitat samples (despite a much greater volume of natural habitats being collected overall). Many ofthe amphipod species recorded on ASUs were not collected from natural habitats within a 10m radius ofthe deployment rack. This indicates that amphipods were recruiting from a greater distance, and this further supports the earlier hypothesis that amphipods, particularly epiphytic species, are highly mobile and therefore rapid colonizers (Gunnill, 1982; Edgar, 1992; Edgar & Robertson, 1992; Taylor, 1998; Christie & Kraufvelin, 2004; Poore, 2005).
ASUs may have sampled a greater species richness than natural habitats as a product of being a new habitat. The short colonization history or 'age' ofASUs would favour rapidly-colonizing, mobile fauna. Natural habitats would be expected to have a more mature assemblage, having been exposed over a greater period oftime, increasing their potential to be colonized by later, dispersal recruiters, whose recruitment can be largely seasonally dependent (Smith & Rule, 2002; Rule, 2004; Watson & Barnes, 2004). In addition, niches in these mature assemblages are more likely to be filled allowing little scope for the addition of new species. Such a mechanism would potentially explain the
175 Assessment of ASU sampling package
presence ofspecies solely on ASUs, as motile species moving through the area could readily recruit to ASUs but would find it more difficult to recruit to natural habitats.
The density ofspecies in adjacent habitats can largely influence recruiting fauna (Tanaka & Leite, 2003). Studies offaunal movement in terms ofdispersal, have shown that distances of 8 - 15 m can be readily covered by amphipods (Virstein & Curran, 1986; Tanaka & Leite, 2004). Indeed, Virstein & Curran (1986) found amphipods living on seagrass covered distances of 0 - 15 m in one day. However, Gunnill (1982) found that ASUs isolated by a distance of30 m from other habitats were not colonized by amphipod fauna after a period of 18 days.
Habitat fragmentation, including the absence ofa target host plant, or the presence of bare sand patches has, however, been demonstrated as a non-limiting factors to amphipod dispersal (Taylor, 1998; Roberts & Poore, 2005). Poore (2005), investigating movement of herbivorous amphipods, demonstrated that "structural fragmentation of habitats does not necessarily imply a functional fragmentation to dispersing amphipod fauna."
Given the lack ofsome species in adjacent natural habitats, amphipods recruiting to ASUs in this study included fauna from source habitats greater than 10m from the deployment racks. It must be considered, however, that proximate natural habitats assessed in this experiment were only collected at the end ofthe experiment (so as not to remove a potential source ofcolonists). Thus, the assemblages on these natural habitats at the time of sampling may not represent the assemblages that were present over the duration ofthe associated ASU deployment period. Within a local species pool, many amphipod species are considered rare or transient and may only be present temporarily in an assemblage (Costello & Myers, 1996). The contribution ofthese species to amphipod assemblages from the Solitary Islands region is discussed in the next chapter.
176 Assessment of ASU sampling package
ASU sampling packages as representative ofamphipod biodiversity The ASU sampling package was found to recruit assemblages that were representative of regional biodiversity but were less so of local suites ofspecies for the four Solitary Island locations assessed (Fig. 45, 47). At a regional level, the single location and mixed location ASU sampling packages were further found to be representative ofamphipod assemblages once rare fauna had been removed from the master list (Figure 47). Given that some locations within a region may naturally have a lower species richness~ as seen at North Solitary for this study (and also found in Rule's (2004) work), a regional deployment strategy which considers several locations is recommended ifthe objective is to maximise the number ofspecies sampled.
Naturally low species richness values at certain locations is not unusual within different regions. Arvanitidis et al. (2005) found two of 14 lagoons surveyed supported lower species richness than other locations and indicate that this is an important reason to test a number of locations for assessing biodiversity. This also broadly reiterates the importance ofconsidering appropriate scales of spatial variation in studies (section 4.5; Rule, 2004; Rule & Smith, 2005; Fraschetti et al., 2005). For example, Edgar & Klumpp (2003) found disparity in species richness and host specificity ofamphipod assemblages between three tropical locations greater than 1 km apart and highlighted how different conclusions would have been drawn about assemblages ifall locations had not been investigated.
A key issue identified through the regional analyses in the Solitary Islands., was the large number of"rare" species. Approximately 26% of regional epifaunal amphipods were known from only a limited number ofrecords (Appendix 2.2). Costello & Myers (1996) similarly documented that between 22 - 44% ofspecies were transient, that is, with few individuals that occurred over time. The phenomenon ofrare species may be a result of many factors including diet specificity, competitive interactions, predation and high grazing levels, to name a few (Costello & Myers, 1996; Chapman, 1999). Further species-specific studies are needed to attempt to understand the trends in rarity observed here. The high number ofrare species, many ofwhich were not collected either in ASUs
177 Assessment ofASU sampling package
or in general collections made at the same time, has some important implications for biodiversity assessment; this is discussed in. the following chapter.
At Bare Island, ASU sampling packages were, again, successful in recruiting a representative sample ofthe known biodiversity for the area (Figure 51). fvtost interestingly, the previously observed trend for associations between some species and specific ASU types were repeated (Table 30,37). One Ischyrocerid, Ericthoniusforbesii, showed a consistent higher abundance on Onion ASUs, at both the Solitary Islands and Bare Island. This observation has obvious application if such fauna were ofspecific interest (for example, ecotoxicology studies), as they could be targeted using particular ASU types.
The sampling package ofASUs, (Onion bags, Shower poofie, Kitchen scourers, Rope Fibre and Astro turt), when formulated randomly or by location was effective at recruiting assemblages representative of regional amphipod biodiversity (Figure 47). At the local scale of islands, the sampling package was less representative (Fig. 45). At Bare Island, assemblages recruiting to the sampling package (five ASU types wiith four replicates, deployed for four weeks) were representative ofthe local fauna. The re-testing ofthis method further supported the viability ofthe ASU sampling package as a tool for sampling, cataloguing and monitoring amphipod fauna in a different benthic habitat to the one where the method was developed. Further this method ofamphipod collection is more effective than the sampling ofnatural habitats, making ASU an efficient rapid assessment method for targeting amphipod biodiversity.
178 Amphipod Taxonomy
CHAPTER SIX
Amphipod taxonomy
6.1 Introduction
6.1.1 Marine amphipod taxonomy in Australian
Studies into Australian marine amphipod taxonomy began with W.A. Haswell who published from 1879 to 1885. As with some early works his description and drawing of species were sparing. Several later authors have redescribed Haswell's material (Sheard, 1939; J.L. Barnard, 1972, 1974; Lowry & Springthorpe, 2005), however, for instances where type material is lost, some species may remain unidentifiable as a result ofthese limited original descriptions (Springthorpe & Lowry, 1994; Lowry & Stoddart, 2003).
C. Chilton published on Australian and New Zealand marine amphipod fauna from 1885 - 1923 and, most notably, described amphipods collected from South Australia and Tasmania by the Fisheries Investigation Steamer 'Endeavour' (Chilton, 1921). K.S. Sheard studied the South Australian amphipod fauna during the period 1936-1939, publishing on a wide range offamilies, including benthic and pelagic groups.
The famous 'Challenger' expedition, the first scientific voyage to circumnavigate the globe, sailed though Australian waters and T.R.R. Stebbing (1888) described the amphipod fauna collected therein. Stebbing (1910) also published the results of another cruise, the'Thetis, ' which sampled offthe coast of Sydney and, from 1891 to 1914, wrote several other papers that included Australian amphipod fauna. In total, these early workers contributed approximately 190 Australian amphipod species descriptions to the fauna.
The next wave of research on Australian amphipods began with J.L. Barnard in 1972. J.L. Barnard's publications on the marine amphipod fauna of Australia commenced with
179 Amphipod Taxonomy
a five part series suitably titled 'Gammaridean Amphipoda of Australia' (1972- I982), co authoring parts III, IV and V with M.M. Drummond. J.L. Barnard & Drummond continued to publish together (1976-1992), focusing on the fossorial amphipod families Phoxocephalidae, Oedicerotidae and Urohaustoridae.
The second major contemporary contributor to Australian amphipod fauna has been J.K. Lowry (1981- present), publishing with a number ofco-authors. Research projects include: Lowry & Stoddart (1983 - present) on the Lysianassoid fauna ofthe Australasian region; Lowry & Berents (1989; 1996; 2005), on the genus Cerapus; Lowry & S.D.A. Smith (2003; in prep.) on the Scavengers of Eastern Australian Seas; Lowry & Springthorpe (2005a) on the Melitidae, and most recently, two works by Coleman & Lowry (in press) on Iphimedioid groups. Other relevant publications of note, include: Freewater & Lowry (1994); Lowry & G.C.B. Poore (1985; 1989); A.G.B. Poore & Lowry (1997); Watson, Lowry & Steinberg (2004); and Lowry & Springthorpe (2005b).
P.G. Moore (1981-1992) also made a significant contribution to Australian amphipod fauna. Working mainly with collections from Western Australia and Tasmania, Moore published on a large range ofamphipod families mainly associated with macroalgal and seagrass habitats, including: Ampeliscidae; Amphilochidae; Aoridae; Colomastigidae; Cyproidiidae; Iphimediidae; Melitidae; Leucothoidae; Lysianassidae; Photidae; and Sebidae (Moore, 1981; 1982; 1987a,b; 1988a,b; 1989). A collaboration by Moore and Myers dealt with the genus Aora and several other corophioid groups (Myers & Moore, 1983; Moore & Myers, 1988). The hyperiidean amphipods ofAustralia have been studied by W. Ziedler, 1978-1999 with over 120 species recorded in Australia. J. Just (1983 - present) has made a considerable contribution, working on the ampithoid, siphonoecetine and iphimedioid amphipod groups. P. Berents's (1983) work on the family Melitidae from the Lizard Island region is one ofthe few published tropical amphipod studies for Australia.
Several visiting researchers have also made a significant contribution to Australian amphipod taxonomy, many as specialists on particular family groups: Myers (1988) on
180 Amphipod Taxonomy
the Aoridae; Berge (2001) and Berge & Vader (2000; 2003) on the Stegocephalidae; Guerra-Garcia (2003; 2004) and Guerra-Garcia & Takeuchi (2004), on the Caprellidea. Vader (1996) and Vader & Myers (1996) have described species of Liljeborgiidae and Ischyroceridae commensal on hermit crabs (Paguroidea).
Recently, doctoral research projects by J.D. Taylor (2002) and R.A. Peart (2002) have studied the families Phoxocephalidae and Ampithoidae, respectively. Current publications from these works include Taylor & G.C.B. Poore, 2001; Taylor, 2002 and Peart, 2004.
Since J.L. Barnard (1972), modern amphipod taxonomy in Australia, has recorded over 750 species from Australia waters, equating to approximately 80% ofthe known Australian marine amphipod fauna. Lowry & Stoddart (2003) published a catalogue of the Australian amphipod fauna as part ofthe Zoological Catalogue ofAustralia, which includes family and generic diagnoses, synonymies, distribution and ecological information.
Despite the above works, most ofthe amphipod research in Australia has focused on the southern halfofthe continent, with the majority ofamphipod fauna described from the southern region from Coffs Harbour to Perth in Western Australia. An exception to this is K.H. Barnard (1931) "Amphipoda. Great Barrier Reef Expedition 1928-29," this work was predominately on hyperiidean amphipods, with only 14 species ofgammaridean amphipods reported. As a result, Australia's tropical amphipod fauna is at present largely unknown. However, a recent expedition to Lizard Island (Great Barrier Reef, North Queensland) by a team of 15 international amphipod researchers will begin to address this faunal gap. The Lizard Island project collected 176 species ofamphipod, ofwhich approximately 154 are new species (J.K. Lowry, pers. comm.).
Australian amphipod taxonomy is now entering the new age ofweb-based taxonomic databases, through the 'The Australian Amphipod Project' established in 2000 and led by J.K. Lowry. This project will provide interactive identification keys and monographs to
181 Amphipod Taxonomy
Australian (and world) amphipod families. The project forms part ofa larger network to provide taxonomic information to all crustacean groups on-line at www.crustacea.net. Interactive information is already available for several Australian amphipod families: Ampeliscidae; Aoridae; Eusiridae; Hyalidae; Leucothoidae and Melitidae (Lowry & Springthorpe, 200 I-onwards).
6.1.2 Amphipod taxonomy in the Solitary Islands region
Published information on amphipod fauna for the Solitary Islands region is predominantly from ecological studies, including: Smith & Simpson (1991); Smith et al. (1992) and Smith (1996; 2000) investigating kelp holdfast macrofauna. More recently, Smith & Rule (2002) and Rule & Smith (2005) published on epifaunal recruitment to artificial substrates. However, these studies have not always kept up with changes to taxonomic nomenclature, and identification at species level has been coarse, with a tendency to lump specimens (pers. obs.).
The Australian Museum and visiting researchers have undertaken several collection events in the Solitary Islands. These have been both project specific: e.g. Scavengers of Eastern Australian Seas (SEAS project) - J.K. Lowry; Ampithoidae ofAustralia - R. Peart; Caprellids ofNew South Wales - I. Takeuchi (Ehime University, Japan); The Systematics and Ecology of Marine Interstitial Amphipods of Eastern Australia and New Caledonia - M. Yerman; and general collection by Australian Museum parties, in 1992 and 2005. Aside from Lowry & Smith (2003), publications from the majority ofspecific projects are in preparation, while work by M. Yerman is still in progress. Material collected by Australian Museum parties is sorted and identified to various levels (family, genus, species) and catalogued within the Australian Museum collection.
Work by Lowry & Springthorpe (2005a; 2005b) within the Eusiridae and Melitidae families, has recently described two new species (Regalia juliana, Mal/acaata malua)
182 Amphipod Taxonomy
from the Solitary Islands and several other species with range distributions that include the Solitary Islands region.
The taxonomic work from this thesis has focused on epifaunal amphipod species between 0-15 m, one ofthe most diverse ecological environments for amphipod fauna (Thomas, 1993). Taxonomic information has been gathered as part ofwider ecological experimentation into the development ofa method to rapidly assess amphipod biodiversity. This rapid assessment approach to collecting information is hoped to encourage the assessment ofamphipod fauna in other regions. This experimental approach to collecting information is an efficient and objective technique which will allow the comparisons of information from different habitats and locations (Gee & Warwick, 1996).
6.1.3 New species from the Solitary Islands region
As a result ofthe collections made during the course ofthis project, 93 species are reported from the Solitary Islands representing 27 family and 55 genera. Among these species a number ofcommonly occurring taxa were recognised as new to science and one species, Protohyale pusilla (Chevreux 1907), a new record for Australian waters. Eight new species from six families are described here and the subgenus Telsosynopia Karaman, (1986) in the family Synopiidae is elevated to generic level. The species Protohyale solitaire n. sp. (Hyalidae); Ericthonius rodneyi n. sp. and Ericthoniusforbesii n. sp. (Ischyroceridae); Liljeborgia polonius n. sp. (Liljeborgiidae); Elasmopus arrawarra n. sp. and Hoho cornishi n. sp. (Melitidae); Gammaropsis legoliath n. sp. (Photidae); and Telsosynopia trifidilla n. sp. (Synopiidae) are described here.
As this is the first time Protohyale pusilia (Chevreux ,1907) has been reported since the original description from French Polynesia, its occurrence from the Solitary Islands region is supported with illustrations ofthe new material (Figure 52 and Figure 53). A
183 Amphipod Taxonomy
catalogue ofspecies currently known for the Solitary Islands between 0-15 m, with emphasis on epifaunal amphipods (excluding the Caprellidae) is given (Table 38).
6.2 Methods
Material described here comes from collections made during the 2002-2004 fieldwork component ofthis project and additional material examined from the Australian Museum's collection. A list ofamphipod species found between 0-15 m from the Solitary Islands region, with focus on epifauna (excluding the Caprelloids) (Table 38) was tabulated from both material collected during this project and Australian Museum records. Some undescribed species collected during this project belonged to groups that require considerable taxonomic attention, which was beyond the time and scope ofthis project. For these species, identification remained at the generic level and a species number was assigned. For instances where a species was thought to belong to a larger species complex, the binomial species name is given to infer the relationship and cf.. used to denote its tentative allocation.
184 Table 20 continued...
Table 38: A list ofamphipod species found between 0-15 m from the Solitary Islands region, with focus on epifauna (excluding the Caprelloidea).!... indicates the dominant habitat from which species were collected. Habitat and location abbreviations: N-North Solitary; NW-North West Solitary: SW-Southwest Solitary: SP-Split Solitary; S-South solitary: MB-Muttonbird Island: K-Korffs Islet W- WoolgooIga. Museum indicates specimens from the Australian Museum collections.
Family Species Location N NW SW SP S MB K W Museum
Amaryllididae Amaryllis sp. x Amaryllis kamata x x x x
Ampeliscidae Ampelisca euroa x x x Amphilochidae Amphi/ocus marionus x x Amphi/ochid sp. ] x x x
Amphi/ochid sp. 2 x Amphi/ochid sp. 4 x x Ampithoidae Ampithoe caddi x x x x x Ampithoe kava x x x x x x Ampithoe sp.] x x x x x x
Ampithoe sp. 2 x x x x Ampithoe sp. 3 x Ampithoe sp. 4 x x x x Cymadusa sp. x Peramphithoe sp. x x x P/umithoe quadrimanus x x x x x x x Aoridae Aora hebes x x x x x x x x Aora typica x x x x x x x x Bem/os australis x x x Bem/os ephippium x x x x x x Bem/osfasciata x x x x x Bem/os trudis x x Lembos sp. x Chevaliidae Chevalia cr. avicu/ae x x Colomastigidae C%mastix cf. brazieri x x
185 Table 20 continued...
Family Species Location N NW SW SP S MB K W Museum
Corophiidae Monocorophium acherusicum x Paracorophium excavatum x Cyproideidae Cyproidea ornata x x Narapheonoides mul/aya x x Dexaminidae Paradexamine moorhousei x x x x x x x Paradexamine thada/ee x x x x Po/ycheria cf. antarctica x x x x x Eusiridae Eusiroides monocu/oides x x x x x x x
Tethygeneia mega/optha/ma x x x x x x x x Hyalidae Protohya/e pusil/a x Protohya/e rubra x x x x x x Protohya/e solitaire x x x x x Protohya/e yake x x x x Iphimediidae Iphimedia sp_ x x x x x Ischyroceridae Ambicho/estes magel/ani x x Cerapus murrayae x x Ericthonius forbesii x x x x x x x Ericthonius rodneyi x x x x x x Ischyrocerus /ongimanus x Ischyrocerid sp.1 x x x x x Ischyrocerid sp.2 x x x x Jassa s/atteryi Ventojassa he/enae x x Ventojassa zebra Leucothoidae Leucothoe assimi/is x Leucothoe boo/pooli x x x x x x x x Leucothoe commensa/is x x x x x Leucothoe gooweera x x x x Leucothoel/a gracilis x Leucothoe sp_ x x x x
186 Table 20 continued...
Family Species Location N NW SW SP S MB K W Museum
Liljeborgiidae Li/jeborgia aequabilis x Li/jeborgia polonius x x x x Lysianassidae Parawaldeckia yamba x x x x x x Shoemakerel/a sp. x x x x x x x Waldeckia sp. x x x x x Melitidae Ceradocus ramsayi Ceradocus serratus x x Dulichiel/a australis x x x x x x x Elasmopus arrawarra x x x x x x x x Elasmopus warra x Hoho cornishi x x Linguimaera hamigera x x x x x x Linguimaera octodens x Mallacoota euroka x x x x x Mal/acoota kameruka x x x x x Mallacoota malua x x x x x x x Melita matilda x x
Quadrimaera reishi x x x x x x Pardaliscidae Parda/isca sp. x x Phliantidae lphiplateia whiteleggei x x Pereonotus thomsoni x x x Photidae Gammaropsis cf. atlantica x x x x Gammaropsis dentifer x x x x x x Gammaropsis haswelli x x x x x Gammaropsis legoliath x x x x x x x Photis cf. do/ichommata x x x x x Phoxocephalidae Brolgus sp. x x x x Tipimegus sp. x x x x x
Phoxocepha/id sp. x x x x
187 Table 20 continued...
Family Species Location N NW SW SP S MB K W Museum Platyischnopidae Platyischnopus mirabilis x Tittakunara katoa x
Tomituka doowi x x
Pleustidae Tepidolpleustes coffsia x x x x x x x
Podoceridae Podocerus sp. x x x x x x x x
Stenothoidae Stenothoe c[ valida x x Stenothoe miersi x
Stenothoe sp. x x x x Synopiidae Telsosynopia trifidilla x x x x
188 Amphipod Taxonomy
6.2.1 Method ofspecies description
The taxonomic species descriptions were generated from several DELTA (Dallwitz et al., 1993; Dallwitz et aI., 1998) databases. Characters in bold type represent diagnostic descriptions. Descriptions of hyalid species were generated from a Delta database to world hyalid genera and species. This database was an existing work developed by J.K. Lowry, R.T. Springthorpe, (Australian taxa); A.A. Myers, R.A. King, (Northeast Atlantic and Mediterranean taxa) & C.S. Serejo (Brazilian taxa). This database was expanded to include East and West Pacific hyalid fauna. Additional characters were also entered into the database to develop the description and identification of fauna. The new species of Protohyale were diagnosed against all other species in the genus.
Descriptions ofEricthonius species were generated from an ischyrocerid database to Australian, northeast Atlantic and Mediterranean species developed by J.K. Lowry, R.T. Springthorpe & R.A. King. Additional species and characters were entered into the database to develop the description and identification of fauna. The new species were diagnosed against world Ericthonius species.
A database was established for the Liljeborgiidae genus Liljeborgia which included 15 species ofLiljeborgia. These particular species form a monophyletic group on the basis ofdorsal serration pattern (of 3-3-0-1-1 for the pleonite/urosomite), a character considered to be synapomorphic. Characters and character states were based on relevant literature for genera and species. The new species ofLiljeborgia was diagnosed against the 15 species included in the database.
Descriptions ofElasmopus and Hoho species were generated from the world Melitidae database established by J.K. Lowry and R.T. Springthorpe. The new taxa and additional characters were entered into the database to develop the description and identification of fauna. The new species ofElasmopus was diagnosed against other Australian Elasmopus and the new species ofHoho was diagnosed against all known species ofHoho.
189 Amphipod Taxonomy
The description ofthe new species Gammaropsis legoliath was generated from a partial database to 32 world species ofGammaropsis developed by A.A. Myers which was integrated with a second Gammaropsis database by J.K. Lowry. Characters and character states were added to these works to develop the description offauna.
A database was developed for the world Synopiidae, which includes all 16 genera in the family. The database also includes species in the genera Synopia and Telsosynopia. Characters and character states were based on relevant literature ofgenera and species. The new species of Telsosynopia was diagnosed against all known species of Te lsosynopia.
Material described in this study is lodged at the Australian Museum, Sydney (AM). The following abbreviations are used on the plates: A, antenna; Ep, epimeron; G, gnathopod; Hd, head; L, labium; Mn, mandible; Mxp, maxilliped; Mx, maxilla; P, pereopod; T, telson; U, uropod; Ur, urosomite; I, left; r, right.
190 Protohyale pusilla (Chevreux, 1907)
6.3 Description of new species and new records
6.3.1 Family Hyalidae Bulycheva 1957
Serejo (2004) revised the higher classification ofthe talitridan amphipods based on cladistic analysis. The Talitroidea Rafinesque, 1815 is elevated to infraorder (Talitrida) and the Hyalidae is considered to be a sister taxon ofthe Dogielinotidae Gurjanova, 1953. The subfamilies ofthe Dogielinotidae include Dogielinotinae, Gurjanova, 1953, Hyalellinae Bulycheva, 1957 and Najniinae, J.L. Barnard, 1972 (new status). The Hyalidae is divided into two subfamilies, Hyacheliinae Bousfield & Hendrycks, 2002 and
Hyalinae Bulycheva, 1957. This classification differs from Bousfield & Hendrycks (2002) who also included the Kuriinae as a subfamily of Hyalidae, which has now been elevated to family (Kuriidae) and superfamily (Kurioidea) status based on Serejo (2004). Bousfield & Hendrycks (2002) revised the genera ofthe Hyalinae (of Serejo 20(4) and established five new genera, Apohyale, Protohyale, Ptilohyale, Ruffohyale and Serejohyale. In Australia, the genera Apohyale, Parhyale and Ptilohyale are each represented by single species, Apohyale media (Dana, 1853), Parhyale longicornis (Haswell, 1879) and Ptilohyale crassicornis (Haswell, 1879). The genus Neobule Haswell, 1879, described from Australia, remains unidentifiable owing to the poor descriptive state ofthis taxon (Bousfield & Hendrycks 2002; Lowry & Stoddart 2003; Serejo 2004).
The remaining Australian hyalids fall into the genus Protohyale, which is divided into four subgenera Boreohyale Bousfield & Hendrycks, 2002, Diplohyale Bousfield & Hendrycks, 2002, Leptohyale Bousfield & Hendrycks, 2002, and Protohyale Bousfield & Hendrycks, 2002. This subgeneric classification was only partially applied to the Australian species, with only six ofthe then known eleven Australian "Hyales" species placed in subgeneric groups. The omitted species and the two species described here do
191 Protohyale pusilia (Chevreux, 1907)
not conform to the proposed Protohyale subgeneric classification. Therefore, the subgeneric classification is not applied to the Australian Protohyale species.
Protohyale Bousfield & Hendrycks 2002
Protohyale pusilla (Chevreux, 1907) new combination
Figure 52 - Figure 53
Hyale pusilla Chevreux, 1907: 415.-Chevreux, 1908: 506-510, fig. 23-25.
Type material. Holotype male 2.5 mm, MNHN-Am5773, Lagoon, Mangareva Island, Gambier Archipelago, French Polynesia, on coralline algae and Udotea sp., 5 m.
Additional material examined. I male 4.6 mm, AM P71873; 3 specimens. AM P71874, North Solitary Island, Solitary Islands, 29°55'2T'S 153°23'18E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 17 February 2003, 1 specimen, AM P71875, North West Solitary Island, Solitary Islands, 300 1'7"S 153°18'11"E, 60 g plastic artificial substrate, 10m, L.E. Hughes, 30 January, 2003.
Description. Based on male 4.6 mm, AM P71873. Head. Eyes round. Antenna 2 less than or equal to half body length, not densely setose. Maxilla 1 palp present, long, extending beyond end ofouter plate. Gnathopod 1 subchelate; coxa about as long as broad; propodus similar length to carpus (less than 1.5 as long as carpus), broad, length less than 2.5 x width, expanded distally, palm acute, with v-shaped distal sinus. Gnathopod 2 subchelate, carpus not lobate, not projecting between merus and propodus, without setae on the posterior margin; propodus enlarged relative to carpus, palm acute, sculptured, with two proximal humps, with robust setae, with fine simple setae, mid-palm with bowl-shaped hollow, with deep dactylar socket, corner of palm with large robust setae. Pereopods 3-
192 Protohyale pusilia (Chevreux, 1907)
4 with large robust striated setae on propodus. Pereopods 3-7 propodus distal robust seta not fusiform; dactylus with very thin seta on posterior margin. Pereopods 5-7 propodus with large robust striated setae, robust setae subequal in length. Pereopod 7 basis about as broad as long. Epimeron 3 posteroventral corner produced. Uropod I peduncle with enlarged distolateral robust seta. Uropod 3 uniramous, peduncle sides parallel, with distal robust setae, ramus subequal or shorter than peduncle. Telson longer than or subequal to uropod 3 peduncle, cleft, subtriangular, apically rounded.
Habitat. Living on the green alga Chlorodesmis sp.; the brown algae Sargassum sp. and the holdfasts ofEcklonia radiata; and the red algae Amphiroa anceps, Delisea pulchra, Haliptilon roseum, Jania verrucosa; 0-15 m.
Distribution. Gambier Archipelago, French Polynesia; North Solitary Island, South Solitary Island, Korffs Islet, Solitary Islands, Australia.
Remarks. This is the first record ofProtohyale pusilla since its initial description by Chevreux (1907) and additional description in Chevreux (1908). The Solitary Islands material represents a large range extension from the type locality, Gambier Archipelago, French Polynesia, so the holotype was obtained for comparison. The largest Solitary Islands specimen is larger (male 4.6 mm) than those ofChevreux (1908) (male 2.6 mm). In the Solitary Islands specimens, the distal locking setae ofpereopods 3-7 are subequal. The distal locking setae of pereopod 5-7 in the type material is originally described with distal setae shorter than proximal, but given the small difference in relative size ofthe robust setae observed on the holotype, this character is now considered subequal for the type material. The character state 'distal setae shorter than proximal' is reserved for species such as Apohyale media (Dana, 1853) where the difference in robust setal size is marked. The posterodistal lobe of pereopods 5-7 merus ofthe original material is more strongly developed than in the Solitary Islands specimens. Neither ofthese two pereopod
193 Protohyale pusilia (Chevreux, 1907)
characters, along with the larger size ofthe Solitary Islands specimens, are presently considered significant enough to separate the material here as a new species.
Both Protohyale pusilia and P. wilari (J. L. Barnard, 1974) have a distinctive bowl shaped hollow on the mid-palm ofgnathopod 2. This structure is also weakly present in P. grenfelli (Chilton, 1916). In P. pusilla the gnathopod 2 palm proximal margin has two humps bearing setae, while in P. wilari the palm is smooth. In P. pusilla gnathopod 1 propodus palm uniquely has a V-shaped incision at the distal end.
194 Protohyale solitaire n. sp.
,\ Ep \3(:' .. / f 1\ l .// /._-_Y.1 ' j
Figure 54: Protohyale solitaire n. sp., Holotype, male, 9.4 mm, AM P7I88I, North Solitary Island, cons Harbour. Scales for Hd and Ep represent 0.5 mm, remainder represent 0.2 mm.
199 Protohyale solitaire n. sp.
Figure 55: Protohyale solitaire n. sp., Holotype, male, 9.4 mm, AM P71881, North Solitary Island, Coifs Harbour. Scales represent 0.2 mm.
200 Ericthonius forbesii n. sp.
6.3.2 Family Ischyroceridae Stebbing 1899
Siphonoecetini Myers & Lowry 2003
Ericthonius Milne Edwards 1830
The genus Ericthonius has undergone much revision (see Myers & McGrath 1984). Lowry & Berents (1996) considered Ericthonius and Pseudoericthonius to be paraphyletic taxa in a cladistic analysis ofthe 'Ericthonius' group, which established the Cerapus and Siphonoecetes clades as sister taxa. Myers & Lowry (2003) analysed Ericthonius as part ofa much larger corophiidean phylogeny and placed the 'Ericthonius' group in the Photoidea: Ischyroceriidae: Ischyroceriinae: Siphonoecetini. In a review ofthe North-East Atlantic Ericthonius species Myers & McGrath (1984) divided the genus into two groups. The two new Australian species are part of 'group one' in which the gnathopod 2 coxa has stridulating ridges, coxa 2 is widely separated from coxa 1 and 3, and gnathopod 2 carpus has two teeth or one in some hyperadults. The six Australian species ofEricthonius can be divided based on the gnathopod 2 basis, broad in E. brevicarpus Vader & Myers, 1996, E. coxacanthus Moore, 1988 and E. tacticus Moore, 1988 and thin and elongate in E. forbesii n. sp., E. pugnax Dana, 1852 and E. rodneyi n. sp.
Ericthoniusforbesii n. sp.
Figure 56 - Figure 58
Type material. Holotype male, 6.0 mm, AM P71886, Muttonbird Island, Solitary Islands, 30° ITS 153° 1OlE, on the brown alga Ecklonia radiata holdfasts, 8 m, L.E. Hughes, 23 May 2002. Paratypes: 1 female, 6.0 mm, AM P71887; 1 male, 4.8 mm, AM P71888; 1 male, 4.8 mm, AM P71889; 1 male, 5.5 mm, AM P71890; 23 specimens, AM71891, type locality.
201 Ericthoniusforbesii n. sp.
Additional material examined. New South Wales: 8 males, 8 females, AM P71892, North Solitary Island, Solitary Islands, 29°55'27"S 153°23'18"E, 60 g plastic artificial substrate, 10m, L.E. Hughes, 17 February 2003.2 males, 7 females, AM P71893, Korffs Islet, Solitary Islands, 30°19'8"S 153°9'12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003. Many specimens, AM P71894, eastern side of Bare Island, Botany Bay, Sydney, 33°59'58"S 151°13'50"E, 60 g plastic artificial substrate, 6 m, L.E. Hughes, 12 March 2004. 2 specimens, AM P70425, NE Botany Bay, 33°58'11 "s 151 °12'39"E, sand, 3 m, State Pollution Control Commission, 9 December 1976, stn 32. 3 specimens, AM P70427, NW Botany Bay, 33°58'S 151°10'E, sand, 3.4 m, N.S.W. State Pollution Control Commission, 22 November 1976, stn 12.42 specimens, AM P25548, west of La Perouse, Botany Bay, 33°59'30"S 151 ° 13'E, sand, 7 m, New South Wales State Pollution Control Commission, 20 December 1976, stn 43. 1 specimen, AM P49051, Weeney Bay, Botany Bay, 34°01'18"S 151°09'42"E, mud, 1 m, A.R. Jones, A. Roach, 30 March 1995. 1 specimen, AM P70423, east of Brighton-Ie-Sands, Botany Bay, 33°58'S 151 °1 O'E, mud, 7 m, State Pollution Control Commission, 18 November 1976, stn 11. I specimen, AM P70428, east of Brighton-Ie-Sands, BotanyBay, 33°58'20"S 151°10'14"E, muddy sand, 7 m, N.S.W. State Pollution Control Commission, 3 December 1976, stn 26. 1 specimen, AM P70424, SW Botany Bay, off Ramsgate, 33°59'S 151 ° 1O'E, mud, 4.9 m, New South Wales State Pollution Control Commission, 23 November 1976, stn 20. 1 specimen, AM P70426, south of Banksmeadow, Botany Bay, 33°58'47"S 151 °12' 16"E, mud, 8 m, State Pollution Control Commission, 13 December 1976, stn 37. Many specimens, AM P23111, 1 km east of Burwood Beach, 32°57'31 "s 151 °44'43"E, bottom fine sand, 26 m, 18 December 1975.3 specimens, Al\Il P23112, 1.5 km east of Burwood Beach, 32°57'31 "s 151 °44'43"E, bottom mud and gravel, 28 m, 18 December 1975.
202 Ericthonius forbesii n. sp.
Description of holotype. Head. Eyes large (1/4 or more of head length). Maxilla 1 inner plate setae along medial margin. Mandibular palp article 3 distinctly shorter than article 2. Gnathopod 1 basis posterior margin without knob-like process; carpus distinctly longer than propodus; propodus very wide (less than 1.5 times as long as wide). Gnathopod 2 carpochelate; coxa 2 widely separated from coxa I and 3, rounded, distinctly broader than deep, ventral margin with stridulating nodules or ridges; carpus with distinct free posterior expansion, margin extended into a single carpal tooth or with two carpal teeth; propodus subequal to or shorter than carpus, rectangular; dactylus with cluster ofdistal setae. Pereopods 3-4 basis expanded, flask shaped. Pereopods 5-7 basis expanded, dactyli with accessory spines on anterior margins. Pereopod 5 merus without posterior distal lobe. Uropod 1 peduncle without distoventral corona ofcuticular spines; outer ramus with more than 2 apical robust setae. Uropod 2 peduncle with several dorsal robust setae. Uropod 3 peduncle short (length 2 x or less breadth), expanded proximally.
Female (sexually dimorphic characters). Based on paratype female 6.0 mm, AM P71887. Gnathopod 2 subchelate; carpus margin with a blunt extension; propodus very long (more than 2.5 times as long as carpus), palm acute, with distinct corner or spine on posterior margin, posterodistal margin with robust setae; dactylus without cluster of distal setae.
Etymology. Named for Alistair Roderick Forbes, who supported the technical aspects ofthis project.
Habitat. Living on the green alga Halimeda sp.; the brown algae Lobophora sp., Sargassum sp. and the holdfasts ofEcklonia radiata; the red algae Amphiroa anceps, Haliptilon roseum; and in association with the tunicate Herdmania momus; and on sand, mud and gravel bottoms; 0-15m.
203 Ericthoniusforbesii n. sp.
Distribution. New South Wales: North Solitary, North West Solitary, South Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Bare Island.
Remarks. See remarks for Ericthonius rodneyi n. sp.
204 Ericthoniusforbesii n. sp.
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Figure 56: Ericthonius forbesii n. sp., Holotype, male, 6.0 mm, AM P71886, Muttonbird Island, Coils Harbour. Scale represents 0.2 mm.
205 Ericthonius forbesii n. sp.
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Figure 57: Ericthoniusforbesii n. sp., Holotype, male, 6.0 mm, AM P71886; Gl & G2, female, 6.0 mm, AM P71887, Muttonbird Island, Coffs Harbour. Scale represents 0.2 mm.
206 Ericthonius forbesii n. sp.
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Figure 58: Ericthoniusforbesii n. sp., Paratypes, G2 "a", hyperadult male, 5.5 mm, AM P71890; G2 "b", male, 4.8 mm, AM P71889; G2 "c", juvenile male, 4.8 mm, AM P71888, Muttonbird Island, Cotls Harbour. Scale represents 0.2 mm.
207 Ericthonius rodneyi n. sp.
ErictllOnius rodneyi n. sp.
Figure 59 - Figure 60
Type material. Holotype male, 4.5 mm, AM P71895, Muttonbird Island, Solitary Islands, New South Wales, 300 1TS 153°10'E, on the brown alga Ecklonia radiata holdfasts, 8 m, L.E. Hughes, 23 May 2002. Paratypes: 1 female, 4.5 mm, AM P71896~ 3 males, AMP71897, type locality.
Additional material examined. New South Wales: 1 male, 6 females, AM P71898, Southwest Solitary Island, Solitary Islands, 30°9'48"S 153° 13'48"E, on the brown alga Ecklonia radiata, 10m, L.E. Hughes, 11 September 2003. 1 male, 9 females, AM P71899, Korffs Islet, Solitary Islands, 30° 19'8"S 153°9'12"E, on the brown alga Ecklonia radiata holdfast, 8 m, L.E. Hughes, 5 August 2003.
Description of holotype. Head. Eyes small (less than 1/4 head length). Maxilla 1 inner plate with 2-3 setae. Mandibular palp article 3 medially broad, distinctly shorter than article 2. Gnathopod 1 basis posterior margin without knob-like process; carpus distinctly longer than propodus~ propodus very wide (less than 1.5 times as long as wide). Gnathopod 2 carpochelate~ coxa 2 widely separated from coxa 1 and 3, about as deep as broad, rounded, ventral margin with stridulating nodules or ridges~ carpus with distinct free posterior expansion, margin with two carpal teeth~ propodus subequal to or shorter than carpus, short, rectangular; dactylus without cluster ofdistal setae. Pereopods 3-4 basis expanded, flask shaped. Pereopods 5-7 basis expanded; dactyli with accessory spines on anterior margins. Peraeopod 5 basis without posterodistal lobe. Uropod 1 peduncle without distoventral corona ofcuticular spines~ outer ramus with 1-2 apical robust setae. Uropod 2 peduncle without dorsal robust setae. Uropod 3 peduncle short (length 2 x or less breadth), expanded proximally.
208 Ericthonius rodneyi n. sp.
Female (sexually dimorphic characters). Based on paratype female 4.5 mm~ AM P71896. Gnathopod 2 subchelate; propodus long (less than 2.5 times as long as carpus), palm acute, with distinct comer or spine on posterior margin, posterodistal margin without robust setae.
Etymology. Named for Professor Rod Simpson, the inaugural director ofthe National Marine Science Centre.
Habitat. Living on the green algae Halimeda sp.; the brown algae Lobophora sp., Sargassum sp. and the holdfasts ofEcklonia radiata; the red algae Amphiroa anceps; and in association with the tunicate Herdmania momus, 0-15 m.
Distribution. New South Wales: North West Solitary; Southwest Solitary; Split Solitary; Muttonbird Island; Korffs Islet; Solitary Islands.
Remarks. Ericthonius forbesii and E. rodneyi have no anteroproximal projection on the gnathopod 2 coxa. This separates them from E. brevicarpus and E. coxacanthus which have subquadrate and subtriangular proximal projections respectively on coxa 2. E ricthonius tacticus (and also E. coxacanthus) have a row of4 robust setae along the posterior margin ofthe gnathopod 2 carpus not present in other species ofAustralian E ricthonius.
Both E.forbesii and E. pugnax (males) have long setae at the distal end ofthe dactylus of gnathopod 2 that separates them from E. rodneyi. In E. forbesii and E. pugnax the gnathopod 2 carpal projection of male hyperadults is developed as a single carpal spur, but this is as yet unknown for E. rodneyi. Ericthonius forbesii may be distinguished from E. pugnax by the posterior distal lobe on the basis ofpereopod 5 in E. pugnax. Ericthonius forbesii and E. rodneyi appear to be most similar to each other in the male gnathopod 2 coxa shape, basis width and carpal lobe. However, both species, can be readily distinguished by eye size; E.forbesii has a large orange eye (more than 1/4 ofthe
209 Ericthonius rodneyi n. sp.
head length) and E. rodneyi has a small brownish eye. In Ericthonius forbesii, maxilla 1 has many setae on the inner plate, the apical margins ofthe uropod 1 rami and the peduncle of uropod 2 have more robust setae than E. rodneyi. Erichtonius forbesii also has the long distal setae on the gnathopod 2 dactylus in males, and adult specimens are slightly larger (6.0 mm) than E. rodneyi (4.5 mm). Gravid females were observed in E. rodneyi which helped distinguish it from juvenile E. forbesii. In both species, stridulations are present on the male and female gnathopod 2 coxa, unlike E. coxacanthus and E. tacticus, in which stridulations are only present in the male.
210 Ericthonius rodneyi n. sp.
Mx2,:
Figure 59: Ericthonius rodneyi n. sp., Holotype, male, 4.5 mm, AM P71895, Muttonbird Island. Corts Harbour. Scale represents 0.2 mm.
211 Ericthonius rodneyi n. sp.
Figure 60: Ericthonius rodneyi n. sp., Holotype, male, 4.5 mm, AM P71895; G 1 & G2, Paratype, female, 4.5 mm, AM P71896, Muttonbird Island, Coffs Harbour. Scale represents 0.2 mm.
212 Liljeborgia polonius n. sp.
6.3.3 Family Liljeborgiidae Stebbing 1899
Liljeborgia Bate 1862
Liljeborgia polonius n. sp.
Figure 61 - Figure 62
Type material. Holotype male, 7.2 mm, AM P71900, Muttonbird Island, Solitary Islands, New South Wales, 30°ITS 153° IO'E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 25 May 2003. Paratypes: I female, 6.5 mm, AM P7I90I; I specimen, AM P7I902, type locality
Additional material examined. New South Wales: 3 specimens, AM P7I903, North West Solitary Island, Solitary Islands, 30° I'7"S 153°18' 11 "E, 60 g plastic artificial substrate, 10m, L.E. Hughes, 23 May 2003. 3 specimens, AM P58595, Bulk Liquids Berth, Botany Bay, 33°58'30"S 151 °12'36"E, piling scraping, 7 m, NSW Fisheries, 19 October 1998. 1 specimen, AM P70353, NW side ofTollgate Islands, Bateman's Bay, 35°44'45"S 150° 15'26"E, Ecklonia radiata holdfast & attached rock, 12 m, G.D. Wilson, AJ. Millar, 29 October 2002, NSW stn 2011. 3 specimens, AM P70431, 50 m east of Sullivan's Reef, Ulladulla, 35°21 '20"S 1500 29'22"E, Ecklonia radiata holdfasts between boulders, 23 m, A. Murray, 5 May 1997, NSW stn 1328. 1 specimen, AM P70432, Golf Course Bommie, 500 m NE of Ulladulla Head, 35°20'29"S 150°29'12"E, Cluster of sponges under ledge, 15 m, K. Attwood, 2 May 1997, NSW stn 1295.2 specimens, AM P70433, GolfCourse Bommie, 500 m NE ofUlladulla Head, 35°20'29"S 1500 29'12"E, grey & orange sponges on side of large boulder, 15 m, K. Attwood, 2 May 1997, NSW stn 1297.
Description of holotype. Head. Eye very large (> 1/3 of head length). Lateral cepahlic lobe apically truncate. Mandibular palp article 3 much shorter than article 2.
213 Liljeborgia polonius n. sp.
Gnathopod 1 basis posterior margin setose; dactylus inner margin with 9 teeth. Gnathopod 2 basis posterior margin with rows of thick setae; propodus palm evenly convex; dactylus inner margin 7-10 teeth. Pereopods 3-4 propodus with row of 8-9 small robust setae, with distal seta similar to proximal setae. Pereopod 5 basis posterior margin serrate. Pereopods 5-6 basis subrectangular; dactylus apically subacute, dactylus short. Pereopod 7 propodus with long fine setae. Pleonite/urosomite dorsal serration formula (3-3-0-1-1). Epimeron 3 with single postereoventral tooth, posteriodistal corner with notch. Uropod 1 inner ramus with 4-5 robust setae; outer ramus with 4-5 robust setae. Uropod 2 inner ramus with 4-5 robust setae; outer ramus 3-4 robust setae. Uropod 3 rami longer than peduncle; inner ramus with 4-6 robust setae; outer ramus 3-4 robust setae. Telson apical margin equal, apical setae well developed, single pair of robust apical setae.
Female (sexually dimorphic characters). Based on paratype female 6.5 mm, AM P7190 1. Pereon. Pleonite/urosomite dorsal serration formula (0-0-0-0-0).
Etymology. Named after the Shakespearian character Polonius (from Hamlet), as a play on his words "Neither a borrower (lumper) nor a lender (splitter) be", reflecting the difficulty in making decisions on species taxonomy within the Liljeborgia. Used as a noun in apposition.
Habitat. Living on the brown algae Sargassum sp. and the holdfasts ofEcklonia radiata; and in association with sponges and the tunicate Herdmania momus; 0-15 m.
Distribution. New South Wales: North West Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Bateman's Bay.
Remarks. Two species ofLiljeborgia have been described from Australian waters, L. dubia (Haswell, 1879) and L. aequabilis (Stebbing, 1888). Liljeborgia dubia is very
214 Liljeborgia polonius n. sp.
distinctive with numerous serrate body features, including uni-, tri- and quindentate dorsal serrations on the pleon and urosome and a heavily serrate posterior margin on the basis of pereopods 5-7. This new species most closely resembles 1. aequabilis, which is aligned with a group of 16 other liljeborgids based on dorsal serration pattern.
A 3-3-0-1-1 dorsal serration formula is present in 17 species ofLiljeborgia: 1. aequabilis; 1. akaroica (subspecies 1. a. akaroica and1. a. maria) Hurley, 1954~ 1. dellavallei Stebbing, 1906 (= 1. mixta Schellenberg, 1925); 1. enigmatica Ledoyer, 1986; 1. eurycradus Thurston, 1974; 1. geminata J. L. Barnard, 1969b; 1. japonica Nagata, 1965; 1. kinahani Bate, 1862; 1. longicornis Schellenberg, 1931; 1. macrodon Schellenberg, 1931,' 1. marcinobrio J.L. Barnard, 1969b; 1. octodentata Schellenberg, 1931; 1. petrae Lyons & Myers, 1991; 1. polonius n. sp., 1. psaltrica Krapp-Schickel, 1975,' 1. serrata Nagata, 1965 and 1. serratoides Tzvetkova, 1967.
Liljeborgia polonius is similar to 1. akaroica, 1. geminata and 1. petrae, which all have 4 or 5 serrate teeth on the dactylus ofgnathopod I, and telsonic lobes with even apices (telson unknown for 1. pe/rae). Liljeborgia polonius has a single tooth and small sinus on the posterior distal comer ofthe epimeron 3 as does 1. pe/rae, while in 1. geminata and 1. akaroica, the epimeron 3 corner is developed with one and two teeth respectively, and a large sinus.
Liljeborgia polonius is most similar to 1. petrae from the GulfofAqaba, Red Sea. Liljeborgia polonius differs by the 9 robust setae on the anterior margin ofthe propodus ofpereopods 3 and 4, while 1. petrae has only 4 robust setae. Liljeborgia polonius also has more robust setae on the peduncle ofthe uropods 1-3.
Based on the geographic isolation of1. polonius from 1. petrae and the above morphological differences, 1. polonius is considered as a new species. Nagata (1965) remarked on a similar situation with 1. serrata (from Seto Inland Sea, Japan) and 1.
215 Liljeborgia polonius n. sp.
macrodon (from southern South America), where species were morphologically very similar but widely separated geographically.
Having grouped a set ofLiljeborgia species on the basis ofdorsal serrations, it is worth noting this character is somewhat unstable (Lyons & Myers 1991). The number of dorsal serrations in an individual is known to develop with age, and the dorsal serration pattern ofa species may vary geographically. J.L. Barnard (1962b) and Ledoyer (1986) provide tables ofdorsal serration formulae for several species but both emphasize that these should be used cautiously in initial identifications. In the species at hand the 3-3-0-1-1 pattern is stable for adult males examined. Strikingly, dorsal serrations are absent in females and juveniles examined. This sexually dimorphic character easily separates female L. polonius from female L. petrae and other closely affiliated species.
216 Liljeborgia polonius n. sp.
I I ! I 1 --1~l
1
Figure 61: Liljeborgia polonius n. sp., Holotype, male, 7.2 mm, AM P71900, Muttonbird Island, Corts Harbour. Scales for U and T represents 0.5 mm, Hd represents 0.2 mm, all remaining represent 0.1 mm.
217 Liljeborgia polonius n. sp.
I P3 l
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'/ I / ' " / / (;t.~ ------2::~ , \ ~\ \ \ 'xJ // /
Figure 62: Liljeborgia polonius n. sp., Holotype, male, 7.2 mm, AM P71900, Muttonbird Island, Corts Harbour. Scales represents 0.5 mm.
218 Elasmopus arrawarra n. sp.
6.3.4 Family Melitidae Bousfield 1973
Elasmopus Costa 1853 Elasmopus arrawarra n. sp.
Figure 63 - Figure 64
Type material. Holotype male, 6.8 mm, AM P71904, Muttonbird Island, Solitary Islands, New South Wales, 300 17'S 153°1 O'E, Amphiroa anceps, 8 m, L.E. Hughes, 25 May 2003. Paratypes: female, 5.4 mm, AMP71905; 18 specimens, AM P71906, type locality.
Additional material examined. New South Wales: 9 males, 7 females, AM P71907, North West Solitary Island, Solitary Islands, 30° I'7"S 153°18'11 "E, on Chlorodesmis sp., 10m, L.E. Hughes, 23 May 2003. 18 specimens, AM P71908, Korffs Islet, Solitary Islands, 30°19'8"S 153°9'12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003. 1 specimen, AM P54991, South Ledge, Cook Island, 28° II '39"S 153 °34'3 8"E, yellow green sponge and crinoid, 12 m, A.R. Parker, 9 June 1993, NSW stn 853. 3 specimens, AM P54990, Cook Island, northeast of Mary's Rock, 28° 11 '25"S 153°34'47"E, orange frilly bryozoan, 19 m, R.T. Springthorpe, 8 June 1993, NSW stn 816.24 specimens, AM P57117, Solitary Islands Jetty, Solitary Islands, 30° 18'24"S 153°08'30"E, worm tubes encrusted with sponge on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 725. I specimen, AM P57257, Solitary Islands Jetty, Solitary Islands, 30°18'24"S 153°08'30"E, Diopatra tubes at base ofjetty pilings, 8.5 m, SJ. Keable, 9 March 1992, NSW stn 727. Many specimens, AM P56657, Solitary Islands Jetty, Solitary Islands, 30° 18'24"S I53°08'30"E, Pyura praeputialis on jetty pilings, 8 m, P.B. Berents SJ. Keable, 9 March 1992, NSW stn 733. Many specimens, AM P56656, Solitary Islands Jetty, Solitary Islands, 30° 18'24"S 153°08'30"E, arborescent sponge on jetty pilings, 7 m, SJ. Keable, 9 March 1992, NSW stn 735. Several specimens, AM P57258, Solitary Islands Jetty, Solitary Islands, 300 18'24"S 153°08'30"E, coral scrapings
219 Elasmopus arrawarra n. sp.
on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 726. Many specimens, AM P56658, Solitary Islands Jetty, Solitary Islands, 30°18'24"s 153°08'30"E, orange sponge on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 729. 3 specimens, AM P57259, Solitary Islands Jetty, Solitary Islands, 30°18'24"S 153°08'30"E, finger sponge on jetty pilings, 4 m, R.T. Springthorpe, 9 March 1992, NSW stn 734. 19 specimens, AM P57118, Solitary Islands Jetty, Solitary Islands, 30° 18'24"S 153°08'30"E, coral scrapings on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 738. 1 specimen, AM P56659, Boambee Creek, Sawtell, 300 20'24"S 153°05'30"E, exposed flat ofsandy mud, 0 m, Australian Museum party, 8 March 1992, NSW stn 723.
Description of holotype. Head. Lateral cephalic lobe broad, truncated, anteroventral corner rounded. Antenna 1 longer than antenna 2; peduncular article 1 longer than article 2, with 1 robust seta on posterior margin. Antenna 2 peduncular article 2 cone gland reaching at least to end of peduncular article 3; article 4 subequal to article 5. Mandible palp article 1 about twice as long as broad, not produced distally; article 2 subequal to article 3.
Gnathopod 1 coxa anteroventral corner produced, rounded; propodus palm acute, convex. Gnathopod 2 left and right gnathopods subequal in size; merus with rounded distoventral corner; propodus setose, without medial depression, with well developed mid-palmar process, palm extremely acute, convex, with groups ofanterodistal fine setae, without posteroventral corner, posterodistal robust setae absent, distal tooth apically subacute; dactylus closing along palm, not reaching end ofpalm. Pereopod 5 carpus and propodus with few (or no) long slender setae along anterior margin. Pereopod 6 basis posterior margin convex, castelloserrate. Pereopod 7 basis posteroventral corner narrowly rounded or subquadrate, with posterior margin castelloserrate.
Epimera 1-2 margin without spines above posteroventral comer. Epimeron 1 posteroventral corner with small acute spine. Urosomite 1 without dorsal carina. Uropod 3 rami distally truncated; inner ramus shorter than outer ramus; outer ramus longer than
220 Elasmopus arrawarra n. sp.
peduncle, I-articulate. Telson apical conical extension absent, each lobe with 3 or more apical/subapical robust setae.
Female (sexually dimorphic characters). Based on paratype female 5.4 mm, AM P71905. Gnathopod 2 propodus palm acute, with sparse robust setae, palm defined by posterodistal robust setae.
Etymology. Named from the field station where part ofthis study was based.
Habitat. Living on the green algae Chlorodesmis sp., Halimeda sp.; the red algae Delisea pulchra, Haliptilon roseum, Hypnea sp., Pterocladia sp.; the brown algae Dictyota sp., Dilophus marginatus, Lobophora sp., Sargassum sp. and holdfasts of Ecklonia radiata,; barnacle rubble; sponges; coral; sandy mud; bryozoans; and in association with the tunicates Herdmania momus and Pyura praeputialis,· and in tubes of the polychaete Diopatra sp.; 0-15m.
Distribution. New South Wales: Cook Island, Tweed Heads; North West Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Sawtell.
Remarks. Krapp-Schickel & Ruffo (1990) recognised a group ofElasmopus defined by a dense fringe of setae along the palm ofgnathopod 2 and a castelloserrate margin on the basis ofpereopods 6 and/or 7: Elasmopus canarius Krapp-Schickel & Ruffo, 1990; E. crenulatus Berents, 1983; E. laufolii Myers, 1986; E. pectinicrus J.L. Barnard, 1955; E. serricatus J.L. Barnard, 1969a; E. spinibasus Sivaprakasam, 1970 and E. yunde Barnard, 1974. Elasmopus arrawarra n. sp., along with E. lapu Myers, 1985, E. nanshaensis Ren, 1998 and E. steelei Appadoo & Myers, 2003, also belong in this group. Elasmopus arrawarra, E. canarius E. nanshaensis, E. serricatus can be further separated by the mid palmar process on the propodus ofgnathopod 2. In E. lapu, and E. yunde the mid-palmar process is present as a long low ridge. The distal palmar tooth on the propodus of gnathopod 2 in E . canarius and E. nanshaensis is truncate while in E. arrawarra and E.
221 Elasmopus arrawarra n. sp.
serricatus the tooth is subacute. Elasmopus arrawarra is distinguished from all other species by the combination ofthe produced anterior corner ofthe gnathopod 1 coxa, the gentle parabolic shape ofthe mid-palmar process on gnathopod 2, the convex posterior margin ofthe basis ofpereopod 6, the castelloserrate posterior margin ofthe basis of pereopods 6 - 7, the weak sharp tooth on the posteriodistal corner ofepimera 1 - 3 and the inner ramus of uropod 3 without medial setae.
In comparison to other Australian species ofElasmopus, E. arrawarra is superficially similar to E. warra Kelaher & Lowry, 2002 in the form ofthe gnathopod 2 with the acute distal palmar tooth, however the pereopods 6 - 7 basis in E. warra is not castelloserrate, and the setae along pereopods 6 - 7 are much longer than in E. arrawarra. Elasmopus crenulatus has a crenulate posterior margin on the basis ofthe pereopods 6 - 7 and a well developed distal tooth on the propodus ofgnathopod 2, but does not have a mid-palmar process on the propodus ofgnathopod 2. The arrangement of robust setae on the acute distal tooth on the palm ofgnathopod 2 is also different from that ofE. arrawarra.
222 Elasmopus arrawarra n. sp.
Figure 63: Elasmopus arrawan"a n. sp., Holotype, male, 6.8 mm, AM P71904, Muttonbird Island, cons Harbour. Scale represents 0.2 mm.
223 Elasmopus arrawarra n. sp.
/
Figure 64: Elasmopus arrawarra n. sp., Holotype, male, 6.8 mm, AM P71904; G 1 & G2, Paratype, female, 5.4 mm, AM P71905, Muttonbird Island, Coffs Harbour. Scale represents 0.2 mm.
224 Hoho cornishi n. sp.
Hollo Lowry & Fenwick 1983 Hollo cornislli n. sp.
Figure 65 - Figure 66
Type material. Holotype male, 12.5 mm, AM P71909, Muttonbird Island, Solitary Islands, New South Wales, 300 1TS 153°1 O'E, Sargassum sp., 8 m, L.E. Hughes, 21 May 2002. Paratypes: 6 males, AM P71910, type locality.
Additional material examined. New South Wales: Many specimens, AM P71911, eastern side of Bare Island, Botany Bay, Sydney, 33°59'58"S 151°13'84"E, 60 g plastic artificial substrate, 6 m, L.E. Hughes, 12 March 2004. 1 specimen, AM P57245, NE of Mary's Rock, Cook Island, 28° 11 '25"S 153°34'47"E, Zonaria sp., 17 m, R.T. Springthorpe, 8 June 1993, NSW stn 814.1 specimen, AM P57246, NE of Mary's Rock, Cook Island, 28° 11 '25"S 153°34'47"E, orange frilly bryozoan, 19 m, R.T. Springthorpe, 8 June 1993, NSW stn 816. 2 specimens, AM P57247, North Ledge, Cook Island, 28°11'26"S 153°34'40"E, Zonaria sp., 10 m, L. Albertson, 8 June 1993, NSW stn 817.1 specimen, AM P57248, South Ledge, Cook Island, 28° 11 '39"S 153°34'38"E, rock, 15 m, K.B. Attwood, 9 June 1993, NSW stn 851. 7 specimens, AM P57249, 100 m NW of Split Solitary Island, 300 14'S 153°10'48"E, red algae, 17 m, SJ. Keable, 7 March 1992, NSW stn 682. 8 specimens, AM P57250, 100 m NW of Split Solitary Island, 30°14'S 153°10'48"E, mixed red algae, 17 m, SJ. Keable, 7 March 1992, NSW stn 693.2 specimens, AM P57242, 100 m NW of Split Solitary Island, 30° 14'S 153° 10'48"E, Halimeda sp., 17 m, SJ. Keable, 7 March 1992, NSW stn 689. 12 specimens, AM P57243, 100m NW of Split Solitary Island, 300 14'S 153°10'48"E, brown algae, 17 m, SJ. Keable, 7 March 1992, NSW stn 694. 1 specimen, AM P42820, Moe's Rock, south of Jervis Bay, 35°09'S 1500 45'E, foliose bryozoan, 18 m, J.K. Lowry & R.T. Springthorpe, 29 June 1981, NSW stn 55. 1 specimen, AM P52794, Moe's Rock, south ofJervis Bay, 35°09'S 150 0 45'E, foliose bryozoan, 18 m, J.K. Lowry, R.T. Springthorpe, 29 June 1981, NSW stn 55. 1 specimen, AM P42816, off Snapper Point, Kioloa, 35°34'30"S 150 0 22'E,
225 Hoho cornishi n. sp.
algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15. 1 specimen, AM P42817, off Snapper Point, Kioloa, 35°34'30"S 1500 22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15. 1 specimen, AM P42818, off Snapper Point, Kioloa, 35°34'30"S 150 0 22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15.1 specimen, AM P42819, off Snapper Point, Kioloa, 35°34'30"S 1500 22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15. 1 specimen, AM P42815, north side ofO'Hara Head, Kioloa, 35°34'S 1500 23'E, coralline algae, 2 m, P.B. Berents, J.K. Lowry, R.T. Springthorpe, P.M. Berents, 25 April 1981, NSW stn 12. 1 specimen, AM P70342, NE of Point Upright, SW side ofGrasshopper Island, 35°38'01 "S 150°19'51 "E, mixed algae, 11 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 10 February 2003, NSW stn 2037. Many specimens, AM P70343, west side of Wasp Island, north of Bateman's Bay, 35°40'02"S 150°18'29"E, Me/anthalia po/ydactylis (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 10 February 2003, NSW stn 2045. Many specimens, AM P70334, north of Burrewarra Point, East Wall, 35°50'01 "S 1500 14'10"E, from macroalga Peyssonelia nova-hollandiae, 25 m, G.D. Wilson, AJ. Millar, N. Yee, 25 October 2002, NSW stn 1985. Many specimens, AM P70333, north of Broulee Island, 35°51 '20"S 150° 11 '38"E, from macroalga Phace/ocarpus apodus, 16 m, G.D. Wilson, AJ. Millar, N. Yee, 30 October 2002, NSW stn 2017. 2 specimens, AM P7034I, south ofBroulee Island, 35°50'50"S 1500 11'05"E, from macroalga Ha/opteris p/atycena, 8 m, AJ. Millar, N. Yee, 25 October 2002, NSW stn 1990. 5 specimens, AM P70337, south of Broulee Island, 35°50'49"S 150° 11 '05"E, from corallines Amphiora anceps & Jania nata/ensis, 8 m, G.D. Wilson, AJ. Millar, N. Yee, 25 October 2002, NSW stn 1989. 1 specimen, AM P70336, NW side of Tollgate Islands, Bateman's Bay, 35°44'46"S 150° 15'28"E, from macroalga Sporochnus radiciformis, 12 m, G.D. Wilson, AJ. Millar, N. Yee, 28 October 2002, NSW stn 2006. 2 specimens, AM P70335, NW side ofTollgate Islands, Bateman's Bay, 35°44'46"S 1500 15'28"E, from macroalga Ga/axaura marginata, 12 m, G.D. Wilson, AJ. Millar, N. Yee, 28 October 2002, NSW stn 2005.5 specimens, AM P70338, NW side ofTollgate
226 Hoho cornishi n. sp.
Islands, Bateman's Bay, 35°44'50"S 150° 15'32"E, from macroalga Caulerpaflexilis, 7.9 m, G.D. Wilson, A.J. Millar, 28 October 2002, NSW stn 2001. 2 specimens, AM P70339, NW side ofTollgate Islands, Bateman's Bay, 35°44'46"S 150°15'28"£, from macroalga Scinaia tsinglanensis, 12 m, G.D. Wilson, A.J. Millar, 28 October 2002, NSW stn 2004. 1 specimen, AM P70340, NW side of Tollgate Islands, Bateman's Bay, 35°44'45"S 150°15'26"E, from macroalga Ecklonia radiata holdfast and rock, 12 m, G.D. Wilson, A.J. Millar, 29 October 2002, NSW stn 2011. Many specimens, AM P70345, west side of North Tollgate Island, Bateman's Bay, 35°44'55"S 150°15'27"£, Chondria succulenta (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 11 October 2003, NSW stn 2053.2 specimens, AM P70344, west side ofNorth Tollgate Island, 35°44'55"S 1500 15'28"E, Caulerpa cactoides (alga), 9 m, P.B. Berents, J. Eu. AJ. Millar, G.D. Wilson, 11 February 2003, NSW stn 2052. 2 specimens, AM P70332, north of Jimmy's Island, 35°48'56"S 150°14'06"E, from macroalga Dictyota dichotoma, 16 m, G.D. Wilson, AJ. Millar, N. Vee, 29 October 2002, NSW stn 2010.
Description of holotype. Head lateral cephalic lobe broad, truncated, anteroventral comer rounded. Antenna 1 peduncular article I longer than article 2, with 4 or more robust setae along posterior margin. Antenna 2 peduncular article 4 longer than article 5. Mandibular palp vestigial, I-articulate.
Gnathopod I coxa posteroventral corner notch absent; merus without posterodistal tooth; propodus palm convex, without posterodistal comer, posterodistal robust setae present. Gnathopod 2 coxa posteroventral corner notch absent; merus with subquadrate distoventral corner; propodus palm acute, convex, smooth, with distal hump bearing robust setae, without posteroventral corner; dactylus not reaching end of palm, apically falcate. Pereopod 5 carpus and propodus with few (or no) long slender setae along anterior margin. Pereopod 6 basis posterior margin subsigmoidal. posteroventral comer narrowly rounded or subquadrate; carpus and propodus with few (or no) long slender setae along anterior margin; propodus not expanded posterodistally. Pereopod 7 basis posterior margin subsigmoidal; propodus not expanded posterodistally.
227 Hoho cornishi n. sp.
Pleonites 1-3 not dorsally bicarinate; without dorsodistal spines. Epimeron 1 posteroventral corner with small acute spine. Epimeron 3 posteroventral corner with small acute spine. Urosomite 1 dorsally bicarinate. Uropod 3 rami longer than peduncle. Telson each lobe with 3 or more apical/subapical robust setae.
Etymology. Named for Keith Cornish who provided tremendous technical support and common sense in the early stages ofthis project.
Habitat. Living on the green algae Caulerpaflexili, Caulerpa caetoides, Halimeda sp.; the red algae Amphiroa anceps, Chondria succulenta, Galaxaura marginata, Halopteris platyeena, Jania natalensis, Melanthalia polydactyli, Peyssonelia nova hollandiae, Scinaia tsinglanensis; the brown algae Ecklonia radiata, Dictyota dichotoma, Halopteris platycena, Phaceloearpus apodus, Sargassum sp., Sporochnus radiciformis, Zonaria sp.; 0-15 m.
Distribution. New South Wales: Cook Island, Tweed Heads; Split Solitary Island, Muttonbird Island, Solitary Islands; Bare Island, Botany Bay; Jervis Bay; Kiola; Grasshopper Island, Wasp Island, Brulee Island, Tollegate Island, Jimmy's Island, Bateman's Bay.
Remarks. Hoho eornishi n. sp. is the first species ofthe genus to be described since the Hoho was established by Lowry & Fenwick, 1983. Hoho cornishi is distinguished by the shape ofthe distal hump on the palm ofgnathopod 2 propodus, usually with 5 and 7 robust setae on either side, and the subsigmoidal posterior margin ofpereopods 6 - 7. Of the three described species ofHoho (H carteta (J.L. Barnard, 1972a); H hirtipalma Lowry & Fenwick, 1983; H marilla (J. L. Barnard, 1972a», H. eornishi is most similar to H. hirtipalma, sharing a thick fringe ofsetae on the palm ofgnathopod 2. Hoho cornishi appears to have less robust setae on the margins ofthe pereopods than H. hirtipalma.
228 Hoho cornishi n. sp.
Figure 65: Hoho cornishi n. sp., Holotype, male, 12.5 mm, AM P71909, Muttonbird Island, Coffs Harbour. Scales for Hd and Ep represent 0.5 mm, all remainder represent 0.2 mm.
229 Hoho cornishi n. sp.
Figure 66: Hoho cO/onishi n. sp., Holotype, male, 12.5 mm, AM P71909, Muttonbird Island, cons Harbour. Scale represent 0.5 mm.
230 Gammaropsis legoliath n. sp.
6.3.5 Family Photidae Boeck 1871
Gammaropsis Liljeborg 1855
Gammaropsis is a problematic genus with several intergrading subgenera, some ofwhich are apparently polyphyletic (J.L. Barnard 1973; Barnard & Karaman 1991; Conlan 1993). Krapp-Schickel & Myers (1979) divided the Mediterranean regional fauna of Gammaropsis into two groups based on a collective oftoothed or smooth characters of the pleonites and urosomites, and the epistome length. This division has been further applied to Irish species by Myers & McGrath (1982). This classification, however, was not able to be extended to the southern hemisphere fauna (Myers 1985, 1995; Appadoo & Myers 2004), which have a combination oftoothed and smooth characters, including the Australian species described here. Myers (1985) recognised the need for more taxonomic work in this group.
The new species described below belongs in the subgenus G. (Gammaropsis) based on a reduced gnathopod 2 carpus, multiarticulate accessory flagellum and uniform anterior coxa. There are now eight species ofGammaropsis recorded from Australia (Lowry & Stoddart 2003); all are in the subgenus Gammaropsis, except for G. (Paranaenia) dentifera (Haswell, 1879).
Gammaropsis (Gammaropsis) legoliath n. sp.
Figure 67 - Figure 69
Type material. Holotype male, 4.1 mm AM P71912, Muttonbird Island, Solitary Islands, New South Wales, 300 lTS 153°10'E, Sargassum sp., 8 m, L.E. Hughes, 21 May 2002. Paratypes: female 5.4 mm, AM P71913; male, 3.4 mm, AM P71914; 22 specimens, AM P71915, type locality.
231 Gammaropsis legoliath n. sp.
Additional material examined. New South Wales: 20 specimens, AM P71916, North West Solitary Island, Solitary Islands, 30° 1'7"S 153° 18' 11 "E, 60 g artificial substrate, 10 m, L.E. Hughes, 30 January 2003.28 specimens, AM P71917, South Solitary Island, Solitary Islands, 300 12'0"S 153°15'48"E, Lobophora sp., Pterocladia sp., Zonaria sp. and mixed brown algae, 8 m, L.E. Hughes, 29 July 2003. 27 specimens, AM P71918, Korffs Islet, Solitary Islands, 30°19'8"S 153°9'12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003. Many specimens, AM P70434, NW side of Brush Island, 35°31 '39"S 1500 24'58"E, Zonaria diesingiana (alga), 16.2 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 9 February 2003, NSW stn 2029. Many specimens, AM P70442, NW side of Brush Island, North of Bateman's Bay, 35°31 '39"S 1500 24'58"E, Zonaria diesingiana (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson. 9 February 2003, NSW stn 2033. Many specimens, AM P70453, NW side of Brush Island, 35°31 '39"S 1500 24'58"E, Ecklonia radiata (alga), 12 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 9 February 2003, NSW stn 2026. 6 specimens, AM P70505, SW side of Grasshopper Island, 35°38'01 "S 150° 19'51 "E, Callophycus tridentifer (alga), II m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 10 February 2003, NSW stn 2036. Many specimens, AM P70444, SW side ofGrasshopper Island, NE of Point Upright, 35°38'0 I"S 150° 19'51 "E, Peyssonnelia novae holliandiae (alga), 13 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 10 February 2003, NSW stn 2038.5 specimens, AM P70510, NE of Point Upright, SW side ofGrasshopper Island, 35°38'01 "S 150° 19'51 "E, Peyssonnelia novae holliandiae (alga), 13 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 10 February 2003, NSW stn 2038. 20 specimens, AM P70509, west side of Wasp Island, north of Bateman's Bay, 35°40'02"S 150° 18'29"E, Melanthalia polydactylis (alga), 16 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 10 February 2003, NSW stn 2045. 10 specimens, AM P70506, west side of Wasp Island, north of Bateman's Bay, 35°40'02"S 1500 18'29"E, Curdiea crassa (alga), 16 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 10 February 2003, NSW stn 2043. Many specimens, AM P70446, west side of Wasp Island, north of Bateman's Bay, 35°40'02"S 1500 18'29"E, Curdiea crassa (alga), 16 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 10 February 2003, NSW stn 2043.
232 Gammaropsis legoliath n. sp.
Many specimens, AM P70438, west side of Wasp Island, north of Bateman's Bay, 35°40'02"S 1500 18'29"E, Peyssonnelia novae holliandiae (alga), 16 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 10 February 2003, NSW stn 2047.7 specimens, AM P70448, NW side of Tollgate Islands, Bateman's Bay, 35°44'46"S 150° 15'27"E, macroalga Caulerpa cactoides, 12 m, G.D. Wilson, N. Vee, 29 October 2002, NSW stn 2013.4 specimens, AM P70449, NW side ofTollgate Islands, Bateman's Bay, 35°44'50"S 1500 15'32"E, macroalga Caulerpaflexilis, 7.9 m, G.D. Wilson, AJ. Millar, 28 October 2002, NSW stn 2001. 14 specimens, AM P70450, NW side of Tollgate Islands, Bateman's Bay, 3544'46"S 150°15'28"E, macroalga Galaxaura marginata, 12 m, G.D. Wilson, AJ. Millar, N. Vee, 28 October 2002, NSW stn 2005. Many specimens, AM P70454, NW side ofTollgate Islands, Bateman's Bay, 35°44'50"S 1500 15'32"E, macroalga Chondria succulenta, 7.9 m, G.D. Wilson, AJ. Millar, 28 October 2002, NSW stn 2003. 10 specimens, AM P70502, NW side of Tollgate Islands, Bateman's Bay, 35°44'46"S 150°15'28"E, macroalga Sporochnus radiciformis, 12 m, G.D. Wilson, AJ. Millar, N. Vee, 28 October 2002, NSW stn 2006. Many specimens, AM P70503, NW side ofTollgate Islands, Bateman's Bay, 35°44'50"S 1500 15'32"E, macroalga Colpomenia sinuosa, 7.9 m, G.D. Wilson, AJ. Millar, 28 October 2002, NSW stn 2002. 24 specimens, AM P70436, NW side ofTollgate Islands, Bateman's Bay, 35°44'46"S 150°15'28"E, macroalga Scinaia tsinglanensis, 12 m, G.D. Wilson, AJ. Millar, 28 October 2002, NSW stn 2004. 5 specimens, AM P70435, west side ofNorth Tollgate Island, Bateman's Bay, 35°44'50"S 150°15'29"E, Halopteris platycena (alga), 11 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 8 February 2003, NSW stn 2022. Many specimens, AM P70501, west side ofNorth Tollgate Island, Bateman's Bay, 35°44'55"S 150° 15'27"E, Chondria succulenta (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 11 February 2003, NSW stn 2053. 5 specimens, AM P70452, west side ofNorth Tollgate Island, Bateman's Bay, 35°44'55"S 1500 15'28"E, Caulerpa cactoides (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 11 February 2003, NSW stn 2052. Many specimens, AM P70445, west side ofNorth Tollgate Island, 35°44'55"S 1500 15'28"E, Stypopodiumflabelliforme (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, G.D. Wilson, 11 February 2003, NSW stn 2055.7 specimens, AM P70507, Gutters, north
233 Gammaropsis legoliath n. sp.
of Burrewarra Point, 35°49'49"S 150°14'02"E, macroalga Stypopodium flabelliforme, 24 m, G.D. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1992.8 specimens, AM P70447, Gutters, north of Burrewarra Point, 35°49'52"S 1500 14'05"E, macroalga Codium lucasii, 23 m, G.D. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1997. Many specimens, AM P70440, gutters, north of Burrewarra Point, 35°49'49"S 150°14'02"E, macroalga Pachymenia prostrata, 24 m, G.D. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1996. 2 specimens, AM P70508, Gutters, north of Burrewarra Point, 35°49'49"S 150°14'02"E, branching hydroid in same bag as Pachymenia prostrata, 24 m, G.D. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1995.4 specimens, AM P70504, East Wall, north of Burrewarra Point, 35°50'01 "S 150° 14'1 O"E, macroalga Corallina berteri, 25 m, G.D. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1987. Many specimens, AM P70437, East Wall, north of Burrewarra Point, 35°50'01 "S 150° 14' 1O"E, macroalga Curdiea crassa with small amount ofMartensia australis, 25 m, G.D. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1986. Many specimens, AM P70351, East Wall, north of Burrewarra Point, 35°50'01 "S 150°14' 1O"E, macroalga Peyssonelia nova-hollandiae, 25 m, G.D. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1985.3 specimens, AM P70439, north of Broulee Island, 35°51 '20"S 1500 11'38"E, macroalgaPhacelocarpus apodus, 16 m, G.D. Wilson, A.J. Millar. N. Vee, 30 October 2002, NSW stn 2017. 5 specimens, AM P70451, north of Broulee Island, 35°51 '20"S 150°11 '38"E, macroalga Lobophora variegata, 16 m, G.D. Wilson, A.J. Millar, N. Vee, 30 October 2002, NSW stn 2016. 2 specimens, AM P70441, south of Broulee Island, 35°50'49"S 150° 11 '05"E, Amphiroa anceps & Jania natalensis, 8 m, G.D. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1989. 1 specimen, AM P70443, North Kianinny Gutter, Tathra, 36°44'04"S 149°59'12"E, Pterocladia lucida (alga), 5 m, P.B. Berents, J. Eu, A.J. Millar, G.D. Wilson, 12 February 2003, NSW stn 2060.
Description of holotype. Head. Eyes distally situated, wholly or partly in lateral cephalic lobes; lateral cephalic lobes triangular, apically subacute, anteroventral margin moderately recessed, moderately excavate. Antenna 1 shorter than antenna 2,
234 Gammaropsis legoliath n. sp.
with many long slender setae, peduncle article 3 shorter than article 1; accessory flagellum mutiarticulate, without aesthetascs. Antenna 2 peduncle similar to peduncle of antenna 1. Labrum epistome present. Mandibular palp article 3 subequal in length to article 2, spatulate with mostly apical setae. Labium mandibular processes acute or sub acute. Maxilla 1 inner plate without long apical setae, with a setae along medial margin. Gnathopod 1 coxa anteroventral comer rounded, without posteroventral spine. Gnathopod 2 symmetrical; coxa evenly rounded, subequal in length to coxa 1, without ventral spine; carpus not very reduced, triangular. Pereopod 3 coxa subequal in length and breadth. Pereopod 5 coxa anterior lobe much deeper than coxa 6; basis posterior margin smooth. Pereopod 6 slightly longer than pereopod 7; basis to propodus enlarged as rectolinear column; basis posterior margin smooth. Pereopod 7 basis posterior margin smooth; carpus more than 0.6 times length of propodus. Pleonite 3 without dorsal teeth. Epimeron 3 with posteroventral notch. Urosomite 1 with dorsal spine. Urosomite 2 without dorsal spines. Urosomite 3 with dorsal spine. Uropod 1 with well developed distoventral spur; inner ramus shorter than peduncle. Uropod 2 with vestigial distoventral spur; inner ramus subequal with peduncle. Uropod 3 peduncle longer than rami; inner ramus longer than outer ramus, inner ramus without apical setae; outer ramus I-articulate.
Female (sexually dimorphic characters). Based on paratype female 5.4 mm, AM P7I9I3. Pereopod 6 basis to propodus not enlarged.
Etymology. The name refers to the grossly enlarged pereopod 6, derived from the historical giant Goliath to reflect the size ofthe male pereopod 6 appendage.
Habitat. Living on the green algae Caulerpa cactoides, Caulerpaflexilis, Codium lucasii; the red algae Amphiroa anceps, Cal/ophycus tridentifer, Chondria succulata, Coral/ina berteri, Curdiea crassa, Galaxaura marginata, Jania natalensis, Martensia australis, MeIanthalia polydactylis, Pachymenia prostrate, Peyssonnelia novae hol/iandiae, Pterocladia lucida, Scinaia tsinglanensis and the brown algae Colpomenia
235 Gammaropsis legoliath n. sp.
sinuosa, holdfasts ofEcklonia radiata. Halopteris platycena, Lobophora variegate, Phacelocarpus apodus, Sargassum sp.; Sporochnus radiciformis, Zonaria diesingiana; 0 15 m.
Distribution. New South Wales: North Solitary, North West Solitary, South Solitary, Split Solitary, Muttonbird Island, Korffs Islet, Solitary Islands.
Remarks. As with many other Gammaropsis species, the palm of male gnathopod 2 in G. legoliath goes through several stages ofdevelopment, with the final palmar structure very different from that ofjuveniles. The other secondary sexual characteristic in males, the enlargement of pereopod 6, takes place before the development ofthe concave palm, and results in a male with gnathopod 2 with a convex crenulate palm and grossly enlarged pereopod 6.
The distinct adult male gnathopod 2 propodus formation of G. legoliath with a concave palm and raised blunt tooth is seen in two other Australian species, G. crassipes (Haswell, 1880) and G. thomsoni (Stebbing, 1888). In G. thomsoni gnathopod 2 is asymmetrical, the dominant gnathopod differs from G. legoliath in a second blunt proximal tooth on the propodus and a tooth on the inner margin ofthe dactylus, which corresponds with a sinus on the propodus. In G. crassipes and G. legoliath the gnathopods are symmetrical, and both have the peculiarly enlarged pereopod 6. In Haswell's (1880) original description ofG. crassipes the drawings ofthe enlarged pereopod 6 are a little exaggerated/overstated from the type material. Stebbing (1906) discussed G. crassipes and noted that pereopod 6 in his specimens was "not as stout as Haswells"'. Although most definitely enlarged, the form of pereopod 6 in G. crassipes is enlarged so that it appears to belong to a much larger specimen than the one it is attached to. The articles ofthe pereopod are enlarged but not expanded. In G. legoliath the large pereopod 6 is developed into a stout columnar limb with each article expanded and ofthe same width. In G. legoliath the posterior margin ofthe basis of pereopods 6-7 is straight
236 Gammaropsis legoliath n. sp.
while in G. crassipes they are sinusoidal. Gammaropsis legoliath is also much smaller than G. crassipes (~ 4 mm vs 8 mm+).
The characteristic large leg ofG. legoliath and G. crassipes are not new to the Photidae. Five species ofthe closely associated genus Photis also display a gross enlargement of the sixth pereopod: P. brevicaudata Stebbing, 1888; P. elephantis J.L. Barnard, 1962a; P. nigrocula Lowry, 1979; P. phaeocula Lowry, 1979 and P. trapherus Thomas & Barnard, 1991. In Photis the articles ofpereopod 6 remain proportional except for the merus, which may be expanded and/or elongate.
237 Gammaropsis legoliath n. sp.
Figure 67: Gammaropsis legoliath n. sp., Holotype, male, 4.1 mm, AM P71912, Muttonbird Island, Corts Harbour. Scales represents 0.2 mm.
238 Gammaropsis legoliath n. sp.
( P3 \ \,,- \. ~\
i ( t ~ ~
Figure 68: Gammaropsis legoliath n. sp., Holotype, male, 4.1 mm, AM P71912, Paratype, female "a", 5.4 mm, AM P71913, Muttonbird Island, cons Harbour. Scales represents 0.2 mm.
239 Gammaropsis legoliath n. sp.
Figure 69: Gammaropsis legoliath n. sp., Paratype, male "a", 3.4 mm, AM P71914, Paratype, female, 5.4 mm, AM P71913, Muttonbird Island, cons Harbour. Scales represents 0.2 mm.
240 Telsosynopia trifidilla n. sp.
6.3.6 Family Synopiidae Dana 1852
Telsosynopia Karaman 1986
J. L. Barnard (1972) discussed the genus Synopia and suggested that S. variabilis might warrant its own genus based on the entire telson. Karaman (1986) took the opportunity to erect the subgenus Synopia (Telso:-,ynopia) following the work by Andres (1984) who described two additional species ofSynopia, S. triangula and S. rotunda, with entire telsons. The subgenus Synopia (Synopia) was also established for the remaining Synopia species with cleft telsons. Karaman (1986) designated Synopia variabilis SpandL 1923 as the type-species for S. (Telsosynopia), and defined the subgenus as "with characters of the genus Synopia except, telson entire and mandibular (sic) more or less triturative". This subgenus, however, went unnoticed by later workers discussing Synopia with entire telsons. Barnard & Thomas (1989) reassessed the molar of T. variablis (as S. variabilis) as triturative. Ortiz & Lalana (1997) described S. paravariabilis, a fourth species of Synopia with an entire telson.
Telsosynopia Karaman, 1986 is elevated to generic rank here, based on the entire telson. The mandibular molar "more or less triturative", originally used to help diagnose the subgenus Telsosynopia, is ineffective in separating Synopia from Telsosynopia because all species are now known to have a triturative molar. Species composition
Species composition. Telsosynopia contains five species: T. paravariabilis (Ortiz & Lalana, 1997) new combination; T. rotunda (Andres, 1984); T. triangula (Andres, 1984); T. trifidilla n. sp. and T. variabilis (Spandl, 1923).
241 Telsosynopia trifidil/a n. sp.
Telsosynopia trifidilla n. sp.
Figure 70 - Figure 71
Type material. Holotype female, 5.5 mm, AM P71919. Paratype female, AM P71920, North West Solitary Island, Solitary Islands, New South Wales, 30°1'7"S 153° 18' II"E, 60 g artificial substrate, 10m, L.E. Hughes, 30 January 2003.
Additional material examined. New South Wales: 3 specimens, AM P71921, Southwest Solitary Island, Solitary Islands, 30°9'48"S 153° 13'48"E, airlift of mixed low turfing algae, 8 m, L.E. Hughes, 11 September 2003.3 specimens, AM P71922, Muttonbird Island, Solitary Islands, 30° ITS 153° IO'E, Sargassum sp., 6 m, L.E. Hughes, 21 May 2002. I specimen, AM P71923, Korffs Islet, Solitary Islands, 30° 19'8"S 153°9' 12"E, air! ift ofAmphiroa anceps and low turfing algae, 8 m, L.E. Hughes, 26 May 2003. 8 specimens, AM P71924, eastern side of Bare Island, Botany Bay, Sydney, 33 °59'58"S 151 °13'84"E, 60 g plastic artificial substrate, 6 m, L.E. Hughes, 12 March 2004. 1 specimen, AM P70348, Gutters, north of Burrewarra Point, Bateman's Bay, 35°49'49"S 1500 14'02"E, from macroalgaStypopodiumflabel/iforme, 24 m, G.D. Wilson. AJ. Millar, N. Vee, 27 October 2002, NSW stn 1992. 1 specimen, AM P70352, East Wall, north of Burrewarra Point, Bateman's Bay, 35°50'01 "s 1500 14'10"E, macroalga Coral/ina berteri, 25 m, G.D. Wilson. AJ. Millar, N. Vee, 25 October 2002, NSW stn 1987.
Type locality. North West Solitary Island, Solitary Islands, New South Wales, 30°1.116'S 153°18.184'E.
Description. Based on holotype female. 5.5 mm, AM P71919. Head protuberant. Eyes present; accessory eyes present, 2-5 ommatida. Antenna 1 length shorter than pereon, peduncle article 1 not elongate or bearing dorsodistal tooth; peduncle article 2
242 Telsosynopia trifidilla n. sp.
not elongate or bearing dorsodistal tooth; flagellum 9-20 articulate. Antenna 2 flagellum 28 articulate (or more). Mandibular palp present; molar triturative, columnar. Maxilliped foliaceaous. Gnathopod 1 simple; coxa not tapering distally; basis longer than carpus. Gnathopod 2 simple; dactylus vestigial terminal setae short, less than or subequal to length ofdactylus. Pereopod 4 coxa smaller than coxa 3, subtriangular. Pereopod 5 basis subovoid, expanded. Pereopod 7 basis subrectangular. Uropod 3 greatly exceeding length ofuropods 1-2. Uropod 3 peduncle short. Telson entire; subequal or longer than uropod 3 peduncle; setae lining lateral margin, apical margin trifid.
Etymology. Named from the trifid telson and articulate assistance provided by Helen Stoddart.
Habitat. Living on the red algae Amphiroa anceps, Corallina berteri; and the brown algae Sargassum sp.; Stypopodiumflabelliforme; 0-15 m.
Distribution. New South Wales: North West Solitary, Southwest Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Bare Island, Botany Bay; Bateman's Bay, Australia.
Remarks. In material from Bare Island, Botany Bay, male specimens had an elongate antenna 1 flagellum with more than 20 articles, while the female antenna 1 flagellum was consistently 9-articulate from all locations reported. This sexually dimorphic character has previously been recorded for one other species in the genus, T variabilis, by J.L. Barnard (1965: fig. 9b).
Within Telsosynopia, T paravariabilis, T trifidilla and T variabilis have a trifid apical margin ofthe telson. Telsosynopia trifidilla differs from other species in having gnathopod I basis longer than carpus; the apical setae on the gnathopod 2 dactylus short,
243 Telsosynopia trifidilla n. sp.
similar to the condition in T variabilis; pereopods 3-4 with the merus and carpus longer and more slender than those of T paravariabilis and T variabilis; coxa of pereopod 4 is a distinct subtriangular shape as in T triangula and the pereopod 5 basis is expanded as in T triangula and T variabilis.
Telsosynopia trifidilla is only the fourth synopiid described from Australian waters,and the first ofthe genus Telsosynopia. The closely related genus Synopia is also represented by a single species in Australia, S. ultramarina Dana, 1853, recorded from Great Barrier Reef, Queensland (K.H. Barnard 1931). Telsosynopia trifidiUa is a similar size to S. ultramarina, but easily separated by the cleft telson.
244 Telsosynopia trifidilla n. sp.
Figure 70: Telsosynopia trijidilla n. sp., Holotype, female, 5.5 mm, AM P71919, North West Solitary Islands. Scales for Hd represent 0.5 mm, all remainder represent 0.1 mm.
245 Telsosynopia trifidilla n. sp.
Figure 71: Telsosynopia trifidilla n. sp., Holotype, female, 5.5 mm, AM P71919, North West Solitary Islands. Scales represent 0.5 m.
246 Amphipod taxonomy
6.4 Discussion
In all, 97 species ofamphipod including 8 new species are reported here from the Solitary Islands region. 27 amphipod families were recorded, with the Melitidae the most dominant family represented by nine genera, and was also the most speciose family. The Ischyroceridae with ~six genera was the second most speciose group.
The amphipod biodiversity reported here represents extensive sampling ofthe epifaunal and algal-associated amphipods in the Solitary Islands. Many other habitats in the Solitary Islands region however, remain under-sampled for amphipod fauna. Soft bottom and estuarine environments, in particular, may hold many more unrecorded or new species. Further, the caprelliod fauna ofthe Solitary Islands will be documented by Takeuchi & Lowry (in prep.) as part ofa broader study into the Caprellids ofNew South Wales, Australia. The ampithoid fauna ofthe region will also be formally described in up and coming works by Peart (in prep.).
Many species occuring in the region appear intermittently in samples, contributing to transient faunal assemblage with low abundances (see section 5.4). An intuitive guess is that many more ofthese species occur in the Solitary Islands, and in the area more transient species will be recorded through continued research.
With the publication ofthe catalogue ofAustralian amphipods (Lowry & Stoddart, 2003), Australia has one ofthe most taxonomically up-to-date amphipod faunas in the world (J.K. Lowry, pers. comm.). The majority ofspecies reported in this study were described species. These existing descriptions are a result ofthe large body of work centred around the Sydney region ofNew South Wales. This could indicate that amphipod fauna in the Solitary Islands is largely temperately affiliated with a faunal assemblage similar to that ofthe south-east coast ofAustralia. The further 87 lots of Australian Museum material identifed to the new species described above, which are from more southerly NSW locations, would support a largely southern temperate association. However, this may be
247 Amphipod taxonomy
a result ofthe focus of study on algal dwellers, with algal habitats predominating in temperate regions; or represent a sampling bias, with more amphipod collection made from temperate regions than subtropical and tropical areas ofAustralia (Lowry & Stoddart, 2003).
Aside from aforementioned work co-ordinated by J.K. Lowry from the northern end of the Great Barrier Reef, the southern section ofthe Great Barrier Reef is as yet relatively unstudied (but see K.H. Barnard, 1931; Berents, 1983). As it stands, eight species from the northern Great Barrier Reefare also recorded from the Solitary Islands. Further, records from Cook Island, northern New South Wales, only one and halfdegrees north of the Solitary Islands, show 14 ofthe 24 species recorded also occur in the Solitary Islands (Australian Museum collection records). The Cook Islands and wider southern subtropical zone may represent an important location, as it is the extreme distribution limit for the large temperate macroalgae habitat ofEcklonia radiata. an important dominant algal habitat in the Solitary Islands.
This early indication ofshared fauna with more northern localities highlights the alternate possibility that the amphipod assemblage in the Solitary Islands region may be only a part of a larger range distribution, with temperate affiliated amphipod species eventually found to have a broader eastern coast distribution range in Australia. Further research into subtropical latitudes is clearly required before biogeographic distribution of amphipod faunal assemblage in the Solitary Islands and eastern Australia can be comprehensively understood.
248 General Discussion
CHAPTER SEVEN
General discussion
The aim ofthis project was to develop a method to facilitate the collection and documentation ofthe epifaunal amphipod fauna ofthe Solitary Islands and to describe new species ofamphipod collected during the study. An experimental approach was used with the objectives of: 1) exploring variation in amphipod assemblages with depth; 2) investigating the influence ofdeployment length on amphipod colonization of Artificial Substrate Units (ASUs); 3) exploring variation in amphipod assemblage recruiting to ASUs ofdiffering habitat architecture. From these results an optimal 'sampling package' ofASU types was assembled which maximised the efficiency ofamphipod collections. The ability ofthis sampling package to recruit assemblages that adequately represent local and regional fauna wa9 tested within the Solitary Island and at the additional location of Bare Island in Botany Bay, Sydney. In this synthesis, I will discuss: i) the use ofASUs for sampling biodiversity, including further research required and potential applications ofthe methods developed here using a sequential, experimental approach; and ii) how the data gathered using ASUs has helped to enhance the knowledge of amphipod ecology and biodiversity, especially within the Solitary Islands region.
7.1 Biodiversity assessment through ASU sampling packages
ASUs were used as a surrogate sampling method for collecting amphipods, as they provide a greater control over experimental design (Gee & Warwick, 1996; Smith & Rule, 2005; Underwood & Chapman, 2006). ASUs are a habitat ofstandardized structure and volume, reducing variability between samples (Costello & Thrush, 1991). Artificial Sampling Units have been successfully used in benthic and intertidal habitats to test a range of recruitment and community interaction hypotheses (Ghelardi, 1971; Myers &
249 General Discussion
Southgate, 1980; Costello & Thrush, 1991; Smith & Rule, 2002; Edgar & Klumpp, 2003; Rule, 2004).
In this study, ASUs were seen to be an effective method for collecting amphipod fauna. Not only did ASUs sample a large range ofamphipod species, as a sampling package this tool was a more efficient means of collecting algal-dwelling amphipods than the collection of natural habitats. This outcome is largely thought to be the result ofan ASU representing 'new space' in the environment, without the space pre-emptive influence of other taxa (Smith & Rule, 2002), coupled with the rapid colonization capabilities ofthe highly mobile amphipod epifauna. These factors provide an initial colonization advantage to this taxon, and facilitate their collection using ASUs. This rapid assessment ASU sampling package method, being four replicates offive ASU types deployed at 10 m for four weeks, can be applied to catalogue amphipod biodiversity across a range of locations and thus has broad application as a biodiversity assessment method.
As a one-offdeployment, the sampling package can provide a baseline assessment ofthe biodiversity ofamphipods in an area. Repeated deployment over time at the same site may provide a method for monitoring epifaunal amphipods that is not always achievable with natural habitats which, by and large, fluctuate in presence and in cover over seasonal and broader temporal scales (Edgar, 1983a).
Using a standardized method was important, not only for comparisons of heterogeneous benthic habitats within the Solitary Islands, but also for potential widespread future application ofthis method as a tool for collecting amphipod epifauna. ASU sampling packages provide a consistent habitat for comparisons across sites. This method allows a more rigorous investigation into assemblages and location trends, and is a means of value-adding to the taxonomic information gathered, which is an important feature to the concept ofthis work. Taxonomic inventorying is often seen as archaic, despite being a critical aspect of biodiversity studies (Valdecasas & Comacho, 2003; Martens & Seger, 2005). Such an experimental approach to data collection will hopefully provide a novel
250 General Discussion
opportunity to facilitate taxonomic/ecological collaboration for biodiversity research and provides a potential avenue for attracting funding for taxonomy.
Further research into the types ofhabitats ASU sampling packages mimic is now needed to understand more about the recruiting assemblages. Work by Smith & Rule (2002) has commenced to investigate this aspect, though the results to date suggest little congruence between kitchen scourer assemblages and surrounding natural habitats within the Solitary Islands. It is possible that ASUs represent a truly artificial environment. Following on from this, it would also be interesting to explore the development ofsampling packages to target specific amphipod taxa, trophic or functional groups. These 'designer' sampling packages would have application for monitoring specific facets ofthe amphipod assemblage at a site and such an approach would potentially be useful in the assessment ofspecific impacts and in environmental modelling.
7.2 Biodiversity of epifaunal amphipod of the Solitary Islands
Amphipods are a dominant component of marine invertebrate assemblages and, despite the considerable experimentation into subtidal habitats ofAustralia (Schreider, 1998), much ofAustralia's amphipod fauna remains unstudied (Lowry & Stoddart, 2003). Identification ofamphipod fauna is time consuming, somewhat difficult, and requires specialist knowledge, which often limits the treatment of amphipods in ecological studies (Underwood & Chapman, 2006; an exception being Edgar, 1983a-d & Hirst, 2003; 2007) .. However, identification of many groups ofsmaller marine invertebrates, including amphipods, is improving through the development of interactive keys, which provide a pictorially guided, condensed, multiple-choice identification tool (Lowry & Springthorpe, 2000-onwards; Glasby & Fauchald, 2003). Pursuing identification oftaxa to species level is particularly important for amphipods, as they are by five-fold, the most diverse group within the peracarid crustaceans, and maintain a high diversity at finer taxonomic levels (Kensley, 1998; Myers, 1999).
251 General Discussion
The amphipod fauna in the Solitary Islands has been investigated largely through community ecological studies. These works include experimentation into: the use ofkelp holdfast as a potential habitat for monitoring anthropogenic disturbance (Smith & Simpson, 1992; Smith, 1993; 1996; Smith et al., 1996); and recruitment ofthe invertebrate assemblages to ASUs (pot scourers) exploring trends across spatial, temporal and depth variables (Smith & Rule; 2002; Rule, 2004; Rule & Smith, 2005). The Solitary Islands are a region ofparticular interest for ecological studies, as it lies within the tropical - temperate convergence for the east coast ofAustralia. The Solitary Island Marine Park, which protects 71,000 hectares in the region, is the first and largest marine park in New South Wales. The work presented here is the first project to study specifically the amphipod epifauna ofthe Solitary Islands.
A total of97 amphipod species were recorded in this study (Table 38). Twenty-seven families were identified with the dominant family, the Melitidae, represented by nine genera; this was also the most speciose family. Most amphipod species occurred across a number ofdifferent algal habitats and on a range of ASU types. This agrees with numerous other studies which have found ubiquitous habitat relationships for the majority ofepifaunal amphipod fauna (Russo, 1988; Russo, 1997; Schreider, 1998; Edgar, 1983a). Some habitats, however, are thought to be more favourable than others in terms of providing space and protection from predators (Edgar, 1983c,d; Hacker & Steneck, 1990; Jacobi & Langevin, 1996). Further, the relationships ofamphipods with host-algae may be more complex than originally assumed, with neighbouring algal habitats also playing a role in habitat choice (Poore, 2004; 2005). Many ofthe amphipods recorded in this study are considered rare or transient species, and the implications ofthis on assemblages, and for collecting biodiversity, is considered below with respect to habitat. Variation in amphipod assemblage with location recorded in this work (section 4.5) supports previous finding by Smith (1993) and Rule (2004) which recorded high levels ofvariation at the smallest spatial scales examined, an hypothesis widely supported in marine sublittoral studies (Fraschetti et aI., 2005). As the ASU sampling package was more representative
252 General Discussion
ofbiodiversity at a regional (rather than local) scale within the Solitary Islands (section 5.3), this further highlighted the importance ofchoosing appropriate spatial scales for desired research outcomes (Fraschetti et aI., 2005; Anderson et aI., 2005).
7.3 Variation in amphipod assemblages on a habitat scale
Results ofthe work presented here strongly support the concept that amphipods are highly mobile and assemblages are not static (Fincham, 1974; Russo, 1988; Edgar, 1992; Martin-Smith, 1994; Jacobi & Langevin, 1996; Costello & Myers, 1996; Benoit el aI., 1998; Taylor, 1998). Investigating how this applies to ecological concepts is difficult to reconcile. At least some ofthe amphipods colonizing ASUs were recruiting from over 10 m away, as was inferred from their absence on habitats in the adjacent area (taken within a 10m radius) (section 5.3). Other studies have reported amphipod movements of up to 15 m in 24 hours (Virstein & Curran, 1986; Tanaka & Leite, 2004). Studies which have investigated mobility in algal-dwelling amphipods have shown remarkably high rates of dispersal, even over short time frames. In an enclosed mesocosm study, 1 - 20/0 ofthe standing stock ofamphipod and isopod fauna emigrated daily (Christie & Kraufvelin, 2004). In field-based studies, movement ofamphipods on macroalgae have been recorded at 1 - 2%, 20% and as much as 60% turnover within 24 hours (Edgar, 1992; Martin-Smith, 1994; Taylor, 1998). Fincham (1974), found 80% offauna captured in light traps suspended in the water column were algal-dwelling amphipod species. High daily turnover rates also imply that, over relatively short periods oftime, amphipods may cover large distances.
In this study, assemblages on different ASU types were variable, with a high degree of shared fauna (section 4.4). This supported the hypothesis that amphipods can exist on a range of habitats (Poore & Steinberg, 1999; Poore & Hill, 2005) and the relationship between assemblages was seen to be "neither simple nor direct" (Russo, 1990). Factors including differences in habitat architecture, and unmeasured factors such as predation
253 General Discussion
and patch size, would have influenced recruiting fauna, however, more research is needed to investigate these factors. Assemblages seen here, and elsewhere, demonstrate that a high proportion of fauna is either rare or transient, as defined by a low abundance or temporary presence in an assemblage (Costello & Myers, 1996; Chapman, 2006). In terms ofbiodiversity assessment, such factors dictate that repeated sampling within a region is required to adequately document fauna (Chapman, 2006).
7.4 Conclusions and future research
The outcomes ofthis project were the successful development ofa rapid assessment ASU sampling package, which consists of four replicates of five ASU types (Onion bags, Shower poofies, Kitchen Scourers, Rope fibre and Astro turt) deployed at 10m for four weeks, which is an effective and efficient tool for cataloguing and monitoring amphipod biodiversity. Continued research into the application ofthis ASU sampling package in other benthic environments, for example tropical and cold temperate regions, would further refine the use ofASUs as a monitoring tool. More detailed experimentation into the influence ofvarious aspects of complex ASU types (such as interstitial space, texture and surface-area) is required in order to further understand their influence on recruiting assemblages. This avenue of research may also facilitate the refinement of ASU designs to target specific taxa or groups (tropic, functional) for - a 'designer' ASU package tool. A further outcome ofthis work was the taxonomic description of nine new species and a new record ofamphipod from the Solitary Islands.
In conclusion, amphipods are an important part ofthe food chain, that are sensitive to changes in the environment and have potential as biodiversity indicators ofthe wider marine benthic community (Thomas, 1993; Edgar, 1992; Marsden & Rainbow, 2004). Amphipods are highly mobile and many fauna present within the epifaunal assemblages are transient or rare. Capturing the biodiversity ofthis fauna is therefore best achieved through a broadly repeated or ongoing approach. This rapid assessment ASU sampling
254 General Discussion
package for targeting amphipod fauna represents an efficient, standardized monitoring method for gathering biodiversity information on this poorly known group. In this way this ASU sampling package approach provides a documented successful experimental, ecological method for taxonomic investigations, and an alternate means for attracting funding and recognition for taxonomic research.
255 Literature cited
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288 Appendix 1
Results ofSIMPER analysis for ASU type within Location
290 Appendix 1.1: Simper results for North Solitary Island comparison between ASU types; a) Onion and Pootie, b) Onion and Scourer. c) Pootie and Scourer. d) Onion and Rope, e) Pootie and Rope, t) Scourer and Rope, g) Onion and Astro, h) Pooties and Astra, i) Scourer and Astra, j) Rope and Astra. k) Onion and Holdfast, I) Pootie and Holdfast. m) Scourer and Holdfast. n) Rope and Holdfasts. 0) Astra and Holdfast. Note some species names have been abbreviated to fit with tables. Paradexamine moorhousei = Paradexamin moor. a) d) Spt:cies Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Onion Pootie Onion Rope Av. dissimilarity = 19.10 Av. dissimilarity = 24.42 Ampithoe caddi 1.00 0.50 14.61 14.61 Aora lypica 1.00 0.25 18.88 18.88 Dulichiel/a australis 0.50 0.75 13.90 28.50 Ericthonills forbesii 1.00 0.25 18.88 37.76 Gammaropsis legoliath 0.25 0.25 9.96 38.47 Paradexamine moor. 0.75 0.50 12.18 49.94 Stenothoe sp. 0.25 0.25 9.93 48.40 Dulichiel/a australis 0.50 0.00 11.77 61.71 Podocerus sp. 0.75 0.75 9.76 58.16 Podocerus sp. 0.75 0.75 9.35 71.07 Plumithoe quadrimanus 1.00 0.75 7.71 65.87 Gammaropsis legoliath 0.25 0.00 6.06 77.12 b) -!l Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund AV.Abund Contrib% Cum.% Onion Scourer Pootie Rope Av. dissimilarity =43.13 Av. dissimilarity =32.72 Plumithoe quadrimanus 1.00 0.00 15.60 15.60 Aora lypica 1.00 0.25 14.91 14.9\ Protohyale solitaire 1.00 0.00 15.60 31.21 Ericthoniusforbesii 1.00 0.25 14.91 29.82 "'1al/acoota euroka 1.00 0.33 \\.30 42.5\ Dulichiella australis 0.75 0.00 14.60 44.42 Paradexamine moor. 0.75 0.33 9.56 52.07 Amplthoe caddi 0.50 1.00 10.33 54.74 Dulichiella australis 0.50 0.33 7.68 59.75 Paradexamine moor. 1.00 0.50 9.64 64.38 Aora tvpica 1.00 0.67 6.46 66.22 Podocerus sp. 0.75 0.75 7.31 71.69 c) f) Species Av.Abund Av.Abund Contrib% Cum.% Species AV.Abund Av.Abund Contrib% Cum.% Poofie Scourer Scourer Rope Av. dissimilarity = 41.96 Av. dissimilarity = 50.06 Paradexamine moor. 1.00 0.33 \2.46 12.46 Plumithoe quadrimanus 0.00 1.00 17.25 17.25 .'vIal/acoota euroka 1.00 0.33 12.46 24.91 Protohyale solitaIre 0.00 1.00 \7.25 34.5\ Plumithoe quadrimanus 0.75 000 12.04 36.95 Mal/acoota euroka 0.33 1.00 12.77 47.28 Protohyale solitaire 0.75 0.00 \2.04 48.99 Aora typica 0.67 0.25 9.09 56.37 Dlilichiella australis 0.75 0.33 10.32 59.31 Ertcthomusforbesl/ 0.67 0,25 9.09 65.45 Ampithoe caddi 0.50 0.67 8.53 67.84 Paradexanllne moor. 0.33 0.50 852 73.97
291 g) j) Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Onion Astra Rope Astro
Av. dissimilarity =39.57 Av. dissimilarity =39.97 Paradexamine moor. 0.75 0.00 10.41 10.41 Aora typrca 0.25 0.75 4.22 10.57 Ericthonius forbesii 1.00 0.25 10.36 20.77 Plumithoe quadrimanus 1.00 0.50 3.36 18.98 Plwnithoe quadrimanus 1.00 0.50 7.03 27.80 Paradexamine moor. 0.50 0.00 3.28 27.19 Protohyale solitaire 1.00 0.50 6.85 34.65 Parawaldeckia yamba 0.25 0.50 3.28 35.39 Dulichiel/a australis 0.50 0.25 6.85 41.49 Protohyale solitaire 1.00 0.50 3.26 43.53 Parawaldeckia yamba 0.00 0.50 6.85 48.34 ...!i..ricthonius forbesii 0.25 0.25 2.40 49.53 h) k) Species AV.Abund Av.Abund Contrib% Cum.% Species AV.Abund Av.Abund Contrib% Cum.% Poofie Astro Onion Holdfast Av. dissimilarity =44.32 Av. dissimilarity = 5.47 Paradexamine moor. 1.00 0.00 13.00 13.00 Ericthonius forbesii 1.00 0.00 13.61 13.61 Ericthonius forhesii 1.00 0.25 9.70 22.70 Ampithoe caddi 1.00 0.00 13.61 27.23 Dulichiel/a australis 0.75 0.25 8.18 30.88 Mal/acoota ellroka 1.00 0.25 10.60 37.83 Ampithoe caddi 0.50 0.75 6.73 37.61 Plwnithoe quadrimanus 1.00 0.25 10.60 48.43 Pilimithoe quadrimanlls 0.75 0.50 6.53 44.14 Paradexamine moor. 0.75 0.00 10.22 58.65 Protohyale solitaire 0.75 0.50 6.47 50.60 2rotohyale solitaire 1.00 0.50 7.59 66.25 i) ....!l Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund AV.Abund Contrib% Cum.% Scourer Astra Poofie Holdfast Av. dissimilarity =51.61 Av. dissimilarity = 5.43 Alal/acoota euroka 0.33 1.00 10.85 10.85 Paradexamine moor. 1.00 0.00 14.64 14.64 Ericthonius forbesii 0.67 0.25 7.97 18.81 Ericthof1/us forbesii 1.00 0.00 14.64 29.29 Protohyale solitaire 0.00 0.50 7.55 26.37 Mal/acoota euroka 1.00 0.25 11.44 40.73 Plumithoe qlladrimanus 0.00 0.50 7.27 33.64 Dlilichiel/a australis 0.75 0.00 11.25 51.98 Parawaldeckia yamba 0.00 0.50 7.27 40.91 Plumithoe quadrimanus 0.75 0.25 8.91 60.89 Ampithoe caddi 0.67 0.75 6.86 47.77 Protohyale solitaire 0.75 0.50 7.58 68.47
292 ~ Species Av.Abund Av.Abund Contrib% Cum.% Scourer Holdfast Av. dissimilarity =51.03 Podocerus sp. 1.00 0.50 14.66 14.66 Ericthoniusforbesii 0.67 0.00 12.93 27.59 Ampithoe caddi 0.67 0.00 12.93 40.52 Aora typica 0.67 0.75 12.08 52.60 Protohyale solitaire 0.00 0.50 10.15 62.74 ,\1allacoota euroka 0.33 0.25 8.37 71.12 n) Species Av.Abund Av.Abund Contrib% Cum.% Rope Holdfast Av. dissimilarity = 3.74 Ampithoe caddi 1.00 0.00 18.62 18.62 Plumlthoe quadrlmanus 1.00 0.25 14.69 33.31 Mal/acoota euroka 1.00 0.25 14.69 48.01 Aora typica 0.25 0.75 11.15 59.15 Protohyale solitaire 1.00 0.50 10.76 69.91 Podocerus sp. 0.75 0.50 9.46 79.37
0) Species AV.Abund Av.Abund Contrib% Cum.% Astro Holdfast Av. dissimilarity =59.41 ,\1allacoota euroka 1.00 0.25 11.48 11.48 Ampithoe caddi 0.75 0.00 10.92 22.40 Podocerus sp. 1.00 0.50 7.74 30.14 Protohyale solitaire 0.50 0.50 7.37 37.51 Plumithoe quadrimanus 0.50 0.25 7.23 44.74 Parawaldeckia yamba 0.50 0.00 7.18 51.92
293 Appendix 1.2: Simper results for North West Solitary Island comparison between ASU types; a) Onion and Pootie, b) Onion and Scourer, c) Pootie and Scourer, d) Onion and Rope, e) Pootie and Rope, t) Scourer and Rope, g) Onion and Astra, h) Poofies and Astro, i) Scourer and Astra, j) Rope and Astra, k) Onion and Holdfast I) Pootie and Holdfast m) Scourer and Holdfast n) Rope and Holdfasts, 0) Astra and Holdfast Note some species names have been abbreviated to tit with tables, Paradexamine moorhousei = Paradexamin moor.
a) Species Av.Abund Av.Abund Contrib% Cum.% ..!l Onion Poofie Species AV.Abund Av.Abund Contrib% Cum.% Av. dissimilarity = 57.98 Poofie Scourer Ampithoe sp.1 0.00 0.75 8.60 8.60 Av. dissimilarity = 53.84 Ericthonius forbesii 0.67 0.00 7.79 16.39 Ischyrocerid sp. A 0.00 1.00 I 1.55 11.55 Podocerus sp. 0.33 1.00 7.57 23.96 Shoemakerella sp. 0.00 1.00 11.55 23.10 Quadrimaera reishi 0.67 0.00 7.04 3].00 Gammaropsis legoliath 0.75 0.00 8.50 31.61 Ampithoe kava 0.33 1.00 7.04 38.04 Amp/thoe sp.1 0.75 0.25 7.56 39.17 Dulichiel/a australis 0.33 0.75 6.52 44.56 Plumithoe quadrimanus 0.50 000 6.32 45.48 Plumithoe quadrimanus 0.00 0.50 6.15 50.71 Dulichiella australis 0.75 0.50 6.04 51.52 .\.1al/acoota euroka 0.00 0.50 5.64 56.34 Mal/acoota euroka 0.50 0.00 5.81 57.33 Ericthonius forbesii 0.50 0.00 5.23 62.57 b) Species AV.Abund AV.Abund Contrib% Cum.% ...!!l Onion Scourer Species Av.Abund Av.Abund Contrib% Cum.% Onion Rope Av. dissimilarity = 54.72 Shoemakerella sp. 0.00 1.00 13.30 13.30 Av. dissimilarity = 68.31 Podocerus sp. 0.33 1.00 9.01 22.31 Aora typica 0.00 0.75 7.57 7.57 Gammaropsis legoliath 0.67 0.00 8.29 30.60 Ericthonius forbesii 0.00 0.75 7.57 15.13 Ischyrocerid sp. A 0.33 1.00 8.29 38.89 Aora hebes 1.00 0.25 7.57 22.70 Ericthonius forbesii 0.67 0.25 8.15 47.04 Ampithoe caddi 0.00 0.75 7.42 30.12 Quadrimaera reishi 0.67 0.25 7.53 54.57 Ampithoe sp.] 0.00 0.75 7.26 37.39 Ampithoe kava 0.33 0.75 7.37 61.94 Gammaropsis legoliath 0.67 000 6.14 43.52 Dulichiel/a australis 0.33 0.50 6.21 68.15 Quadrimaera reishi 0.67 0.00 6.14 49.66 Ampithoe kava 0.33 1.00 6.14 55.79
294 Gammaropsis legoliath 0.67 0.50 5.79 60.65 e) ~ Species Av.Abund AV.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Poofie Rope Poofie Astro Av. dissimilarity =51.43 Av. dissimilarity = 51.46 Aora typica 0.00 0.75 8.67 8.67 Shoemakerella sp. 0.00 1.00 10.81 10.81 Gammaropsis legoliath 0.75 0.00 8.29 16.97 Protohyale rubra 0.25 1.00 8.27 19.07 Dulichiella australis 0.75 0.00 8.10 25.06 Gammaropsis cf. 0.00 0.75 7.74 26.81 Ampit/we caddi 0.25 0.75 7.00 32.06 atlantica Aora hebes 0.75 0.25 6.95 39.02 Ampithoe sp.1 0.75 0.25 6.83 33.64 Plumithoe quadrimanus 0.50 0.00 6.11 45.13 Plwnithoe quadrimanus 0.50 0.25 5.65 39.29 Ericthonius forbesii 0.50 0.75 5.91 51.04 Protohyale solitaire 0.00 0.50 5.64 44.93 Mal/acoota euroka 0.50 0.50 5.63 56.67 Gammarops/s legoliath 0.75 0.50 5.58 50.51 Dulichiel/a australis 0.75 0.50 5.49 56.00 t) Species AV.Abund Av.Abund Contrib% Cum.% .J! Scourer Rope Species Av.Abund Av.Abund Contrib% Cum.% Av. dissimilarity = 60.78 Scourer Astra Aora typica 0.00 0.75 8.13 8.13 Av. dissimilarity = 44.86 Ericthonius forbesii 0.00 0.75 8.13 16.26 Protohyale rubra 0.00 1.00 13.63 13.63 Ischyracerid sp. A 1.00 0.25 8.13 24.39 Ischyracerid sp. A 1.00 0.25 10.20 23.84 Ampithoe caddi 0.00 0.75 7.98 32.37 Gal1lmarOpSls cf. 0.25 0.75 8.44 32.27 Shoemakerel/a sp. 1.00 0.25 7.81 40.18 atlantica Ampithoe sp.1 0.25 0.75 6.72 46.91 Protohyale solitaire 0.00 0.50 7.15 39.42 Aora hebes 0.75 0.25 6.56 53.47 Dul/elliella australis 0.50 0.50 6.73 46.16 Podocerus sp. 1.00 0.50 5.23 58.70 Podoeerus sp. 1.00 0.50 6.66 52.82 Gammaropsis legoliath 0.00 0.50 6.28 59.10 g) Parawaldeekia yamba 0.00 0.50 6.28 65.38 Species Av.Abund Av.Abund Contrib% Cum.% Onion Astra j) Av. dissimilarity = 55.97 Species Av.Abund Av.Abund Contrib% Cum.% Shoemakerel/a sp. 0.00 1.00 11.40 11.40 Rope Astra Gammaropsis 0.00 0.75 8. " 19.51 Av. dissimilarity = 58.89 cf.atlantiea Aora typica 0.75 0.00 7.46 7.46 Eriethonius forbesii 0.67 0.00 7.92 27.43 Erictholll/ls forbesii 0.75 0.00 7.46 14.91 Protohyale rubra 0.33 1.00 7.92 35.36 Ampithoe eaddi 0.75 0.00 7.33 22.25 Ampithoe kava 0.33 1.00 7.18 42.53 Slloemakerella sp. 0.25 1.00 7.20 29.44 Quadrimaera reishi 0.67 0.25 6.50 49.03 Gammaropsis cf. 0.00 0.75 6.92 36.37 Protohyale solitaire 0.33 0.50 5.83 54.86
295 atlantica Shoemakerella sp. 1.00 0.50 6.51 41.70 Aora hebes 0.25 0.75 6.28 42.65 Linguimaera hamigera 0.00 0.50 6.27 47.96 Ampithoe sp. I 0.75 0.25 6.01 48.66 Dulichiella australis 0.50 0.00 5.49 53.45 Protohyale solitaire 0.25 0.50 4.97 53.62 Ampithoe kava 0.75 0.75 5.15 58.60 k) ~ Species AV.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Onion Holdfast Rope Holdfast Av. dissimilarity = 59.78 Av. dissimilarity = 64.73 Paradexamine moor. 1.00 0.25 9.26 9.26 Aora hebes 0.25 1.00 7.89 7.89 Quadrimaera reishi 0.67 0.00 7.84 17.10 Ampithoe caddi 0.75 0.00 7.74 15.63 Ericthoniusforbesii 0.67 0.25 7.58 24.69 Gammaropsis legoliath 0.00 0.75 7.69 23.31 Podocerus sp. 0.33 0.75 7.44 32.13 Ampithoe sp. I 0.75 0.00 7.57 30.89 Ampithoe kava 0.33 0.75 7.04 39.17 Paradexamine moor. 1.00 0.25 7.51 38.40 Aora typica 0.00 0.50 6.42 45.59 Ericthonius forbesii 0.75 0.25 6.52 44.92 Linguimaera hamigera 0.00 0.50 6.17 51.77 Protohyale rubra 0.50 0.25 5.09 50.01 Shoemakerella sp. 0.00 0.50 6.17 57.94 ..!!!docerus sp. 0.50 0.75 5.09 55.09
I) ...2L Species Av.Abund Av.Abund Contrib% Cum.% Species AV.Abund AV.Abund Contrib% Cum.% Pootie Holdfast Astro Holdfast Av. dissimilarity = Av. dissimilarity = 55.70 56.78 Paradexamine moor. 1.00 0.25 8.36 8.36 Ampithoe sp.1 0.75 0.00 8.68 8.68 Protohyale rubra 1.00 0.25 8.36 16.72 Paradexamine moor. 1.00 0.25 8.34 17.01 Gammaropsls cf. atlantica 0.75 0.00 8.06 24.78 Dulichiella australis 0.75 0.00 8.10 25.11 Gammaropsis legoliath 0.50 0.75 5.90 30.68 Plwnithoe quadrimanus 0.50 0.00 6.19 31.31 Protohyale solitatre 0.50 0.25 5.81 36.49 Aora typica 0.00 0.50 5.75 37.05 .\1allacoota euroka 0.50 0.00 5.69 42.74 ,.lora rypica 0.00 0.50 5.76 42.25 Linguimaera hamigera 0.00 0.50 5.56 48.30 Shoemakerella sp. 1.00 0.50 5.76 48.01 Shoemakerella sp. 0.00 0.50 5.56 53.86 Podocerus sp. 0.50 0.75 5.60 53.61 m) Species AV.Abund Av.Abund Contrib% Cum.% Scourer Holdfast Av. dissimilarity = 56.17 Gammaropsis legoliath 0.00 0.75 Y.64 9.64 Ischyrocerid sp. A 100 0.25 9.64 19.28 Paradexamine moor. 1.00 0.25 9.40 28.68 Aora typica 0.00 0.50 6.51 35.19
296 Appendix 1.3: Simper results for Muttonbird Island comparison between ASLJ types: a) Onion and Pootie. b) Onion and Scourer. c) Poofie and Scourer. d) Onion and Rope, e) Pootie and Rope. t) Scourer and Rope, g) Onion and Astra. h) Pooties and Astra. i) Scourer and Astra. j) Rope and Astra. k) Onion and Holdfast, I) Pootie
and Holdfast. m) Scourer and Holdfast. n) Rope and Holdfasts. 0) Astro and Holdfast. a) c) Species Av.Abund AV.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Onion Pootie Pootie Scourer Av. dissimilarity = 51.93 Av. dissimilarity = 34.67 Paradexamine moor. 0.25 1.00 7.86 7.86 Amaryl/is kamata 0.75 0.50 6.68 6.68 Dulichiel/a australis 0.25 1.00 7.86 15.73 Quadrimaera reishi 0.50 0.50 6.64 13.32 Amaryllis kamala 0.00 0.75 7.56 23.28 Gammaropsis hasl1ielli 0.50 0.50 6.53 19.85 Podocerus sp. 0.25 1.00 7.49 30.77 Ischyrocerid sp. A 0.50 0.50 6.53 26.37 Quadrimaera reishi 0.25 0.50 5.29 36.07 Shoemakerella sp. 0.50 0.00 6.47 32.84 Ischyrocerid sp. A 0.75 0.50 5.29 41.36 Gammaropsis legoliath 0.50 0.25 6.29 39.13 Aora hebes 0.50 1.00 5.23 46.59 Prolohyale solitaire 0.25 0.50 6.16 45.30 Shoemakerella sp. 0.00 0.50 4.99 51.58 Tepidolpleustes coffsia 0.50 0.00 6.13 51.43 Gammaropsis legoliath 0.25 0.50 4.83 56.41 Paradexamine moor. 1.00 0.50 5.94 57.37 b) .J!l Species Av.Abund AV.Abund Contrib% Cum.% Species Av.Abund A\'.Abund Contrib% Cum.% Onion Scourer Onion Rope Av. dissimilarity = 49.87 Av. dissimilarity = 50.26 Dulichiella australis 0.25 1.00 9.64 9.64 Amp/thoe sp.1 0.25 1.00 11.45 11.45 Podocerus sp. 0.25 1.00 9.10 18.74 Mal/acoola kameruka 0.75 0.25 8.49 19.94 Paradexamine moor. 0.25 0.50 6.67 25.41 Paradexamine moor. 0.25 0.50 7.70 27.63 Aora hebes 0.50 1.00 6.41 31.82 Aora hebes 0.50 0.75 7.38 35.01 Ischyrocerid sp. A 0.75 0.50 6.39 38.21 Tethygenew mega/. 0.25 0.50 6.85 41.86 Garnrnaropsis haswelli 0.00 0.50 5.82 44.03 Amp/thae kava 0.25 0.50 6.85 48.71 Amaryllis kamata 000 0.50 5.82 49.85 Mal/acoata euroka 0.50 0.25 6.55 55.26 Quadrimaera reishi 0.25 0.50 5.66 55.51 Podocerus sp. 0.25 0.25 5.24 60.50 Bernlos ephipiurn 0.25 0.50 5.66 61.17 Ericthonius rodneyi 0.25 0.00 4.53 65.03
297 e) -E2 Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Poofie Rope Onion Astro Av. dissimilarity -57.46 Av. dissimilarity = 65.88 Dulichiel/a australis 1.00 0.00 8.99 8.99 Gammaropsis 0.00 1.00 9.18 9.18 Ampithoe sp.1 0.25 1.00 7.11 16.10 dallantica Podocerus sp. 1.00 0.25 7.04 23.14 Shoemakerella sp. 0.00 1.00 9.18 18.35 AmQ/:vllis kamala 0.75 0.00 6.71 29.85 Aora typica 1.00 0.25 7.15 25.50 .\.1al/acoola kameruka 0.75 0.25 5.37 35.21 Plumithoe quadrimanus 0.00 0.75 5.90 31.41 Ischyrocerid sp. A 0.50 1.00 4.83 40.04 Ischyrocerid sp. A 0.75 0.50 4.87 36.27 Quadrimaera reishi 0.50 0.00 4.83 44.87 Mal/acoola kameruka 0.75 0.50 4.78 41.05 Shoemakerella sp. 0.50 0.00 4.43 49.30 ,.lora flebes 0.50 1.00 4.78 45.83 Ampilhoe kava 0.25 0.50 4,35 53.65 Mal/acoola euroka 0.50 0.75 4.62 50.45 Dulie/nel/a australis 0.25 0.50 4.31 54.76 t) Species Av.Abund Av.Abund Contrib% Cum.% ...2l Scourer Rope Species Av.Abund AV.Abund Contrib% Cum.% Poofie Astro Av. dissimilarity = 59.63 Av. dissimilarity =55.69 Ampithoe sp.1 0.00 1.00 10.08 10.08 Gammaropsis cf 0.00 1.00 Dulichiel/a australis 1.00 0.00 10.08 20.16 8.02 8.02 atlanlica Podocerus sp. 1.00 0.25 7.95 28.11 Podocerus sp. 1.00 0.25 14.39 Mal/acoota kameruka 0.75 0.25 5.88 33.99 6.38 Aora typica 1.00 0.25 6.17 20.56 Ischyrocerid sp. A 0.50 1.00 5.31 39.29 Amaryllis kamala 0.75 0.00 5.99 26.55 Paradexarnine moor. 0.50 0.50 4.95 44.24 Plwnithoe quadrimanus 0.00 0.75 5.41 31.96 Ampilhoe kava 0.25 0.50 4.79 49.03 Mal/acoola euroka 0.25 0.75 4.70 36.67 Gammaropsis haswel/i 0.50 0.00 4.77 53.80 Paradexamine moor 1.00 0.50 4.53 41.20 AmQ/:vl/is kamata 0.50 0.00 4.77 58.58 Dulichiella australis 1.00 0,50 4.25 45.44 Mal/acoola kameruka 0.75 0.50 4.17 49,61
298 i) Species Av.Abund Av.Abund Contrib% Cum.% Scourer Astro k) Species Av.Abund Av.Abund Contrib% Cllm.% Av. dissimilarity = 60.81 Onion Holdfast Gammaropsis 0.00 1.00 8.38 8.38 cfatlantica Av. dissimilarity = 55.40 ShoemakereI/a sp. 0.00 1.00 8.38 16.76 Aora typica 1.00 0.25 11.04 11.04 Podocerus sp. 1.00 0.25 6.72 23.48 Ampitlwe kava 0.25 1.00 10.72 21.76 Plumithoe quadrimanus 0.00 0.75 5.55 29.03 Mal/acoota kameruka 0.75 0.25 8.16 29.92 Aora typica 0.75 0.25 5.43 34.46 Aora hebes 0.50 1.00 7.15 37.07 .'vfal/acoota euroka 0.25 0.75 4.91 39.37 Podocerus sp. 0.25 0.50 6.63 43.69 Dulichiel/a australis 1.00 0.50 4.50 43.86 Mal/acoota euroka 0.50 0.25 6.10 49.79 Paradexamine moor. 0.50 0.50 4.33 48.20 1schyracerid sp. A 0.75 0.75 5.40 55.19 Mal/acoota kameruka 0.75 0.50 4.32 52.52 Ericthonius rodneyi 0.25 0.25 5.32 60.51 -!2..uadrimaera reishi 0.25 0.25 4.48 65.00 j) Species AV.Abund Av.Abund Contrib% Cum.% Jl Rope Astra Species Av.Abund Av.Abund Contrib% Cum.% Av. dissimilarity = 71.58 Pootie Holdfast 1.00 8.25 8.25 Gammaropsis 0.00 Av. dissimilarity = 54.54 cfatlantica Aora typica 1.00 0.25 7.63 7.63 Ampithoe sp.1 1.00 0.00 8.25 16.50 Paradexamtne moor. 1.00 0.25 7.63 15.27 ShoemakereI/a sp. 0.00 1.00 8.25 24.75 Dulichiel/a australis 1.00 0.25 7.45 22.72 Plumithoe quadrimanus 0.00 0.75 5.34 30.09 Ampitlwe kava 0.25 1.00 7.17 29.88 Aora typica 0.75 0.25 5.08 35.17 Ammyl/is kamala 0.75 0.25 6.20 36.08 .\1al/acoola euroka 0.25 0.75 4.76 39.94 Mal/acoota kameruka 0.75 0.25 5.92 42.00 Ischyrocerid sp. A 1.00 0.50 4.49 44.43 Quadnmaera reishi 0.50 0.25 5.02 47.02 Paradexamine moor. 0.50 0.50 4.23 48.66 Podocerus sp. 1.00 0.50 4.98 52.00 .'vfal/acoota kameruka 0.25 0.50 3.92 52.57 Ischyrocerid sp. A 0.50 0.75 4.92 56.92
299 m) 0) Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Scourer Holdfast Astro Holdfast
Av. dissimilarity = 52.39 Av. dissimilarity = 60.71 Ampithoe kava 0.25 1.00 9.30 9.30 Gammaropsis cf.atlantica 1.00 0.00 9.93 9.93 Dulichiel/a australis 1.00 0.25 9.14 18.44 Ampithoe kava 0.25 1.00 8.05 17.99 Aora typica 0.75 0.25 7.80 26.23 Shoemakerel/a sp. 1.00 0.25 7.98 25.96 .\1al/acoota kameruka 0.75 0.25 6.96 33.19 Mallacoota euroka 0.75 0.25 5.68 31.65 Paradexamine moor. 0.50 0.25 6.39 39.58 Plumithoe quadrimanus 0.75 0.25 5.65 37.30 Podocerus sp. 1.00 0.50 6.10 45.68 Ischyrocerid sp. A 0.50 0.75 5.08 42.38 Ischyrocerid sp. A 0.50 0.75 5.93 51.61 Podocerus sp. 0.25 0.50 4.81 47.19 Amaryl/is kamata 0.50 0.25 5.61 57.22 Dulichiel/a australis 0.50 0.25 4.66 51.85 Gammaropsis haswel/i 0.50 0.00 5.52 62.75 Mal/acoota kameruka 0.50 0.25 4.48 56.32 n) Species Av.Abund AV.Abund Contrib% Cum.% Rope Holdfast
Av. dissimilarity = 56.31 Ampithoe sp. I 1.00 0.00 12.96 12.96 Aora typica 0.75 0.25 8.07 21.03 Ampithoe kava 0.50 1.00 7.07 28.10 Paradexamine moor. 0.50 0.25 6.91 35.01 Podocerus sp. 0.25 0.50 6.26 41.28 Tethygeneia mega/. 0.50 0.00 5.89 47.17 Mal/acoota kameruka 0.25 0.25 5.01 52.18 Mal/acoota euroka 0.25 0.25 4.24 56.42 Plumithoe quadrimanus 0.00 0.25 4.04 60.46
300 Appendix 1.4: Simper results for Korffs Islet comparison between ASU types; a) Onion and Pootie, b) Onion and Seourer, e) Poatie and Seaurer, d) Onion and Rape. e) Paatie and Rope, t) Seourer and Rope. g) Onion and Astra, h) Paoties and Astra. i) Seourer and Astra. j) Rope and Astra. k) Onion and Haldfast. I) Pootie and Haldfast, m) Seourer and Holdfast, n) Rope and Holdfasts, a) Astro and Holdfast.
a) c) Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Onion Poofie Pootie Scourer Av. dissimilarity = 47.66 Av. dissimilarity = 37.72 Shoemakerella sp. 0.00 1.00 9.21 9.21 Tepidolpleustes coffsia 1.00 0.00 11.38 11.38 Ampithoe caddi 0.75 0.00 7.06 16.27 Ampithoe sp. 1 1.00 0.25 8.66 20.04 Tepidolpleustes coffsia 0.25 1.00 7.06 23.33 Quadrimaera reishi 0.25 0.75 7.19 27.22 Dulichiel/a australis 0.00 0.75 6.87 30.20 Amaryllis kamata 0.75 0.25 6.97 34.19 Amaryllis kamata 0.00 0.75 6.75 36.95 Aora typica 0.75 0.50 5.85 4004 Paradexamine moor. 0.00 0.75 6.75 43.71 Mal/acoota malua 1.00 0.50 5.81 45.85 Podocerus sp. 0.25 0.75 5.81 49.52 Protohyale solitaire 0.50 1.00 5.79 51.64 J1allacoota malua 0.50 1.00 4.72 54.24 Aora hebes 1.00 0.50 5.70 57.34 .\1al/acoota kameruka 0.50 1.00 4.72 58.96 Hem/os ephipeiuJ11 0.50 0.50 5.70 63.04
b) d) Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Onion Scourer Onion Rope
Av. dissimilarity = 54.27 Av. dissimilarity = 59.27 Ampithoe caddi 0.75 0.00 7.38 7.38 AII/pithoe caddi 0.75 0.00 8.64 8.64 Ampithoe sp.1 1.00 0.25 7.32 14.71 Gammaropsis legoliath 0.75 0.00 8.64 17.28 Du/ichiella australis 0.00 0.75 7.32 22.03 Mal/acoota euroka 1.00 0.25 8.56 25.84 Quadrimaera reishi 0.00 0.75 7.08 29.11 Ericthonius forbesli 0.75 0.25 6.86 32.69 Paradexamine moor. 0.00 0.75 6.93 36.03 Aora typica 1.00 0.50 6.13 38.82 Gammaropsis legoliath 0.75 0.25 6.23 42.26 Mal/acoota ma/ua 0.50 1.00 5.80 44.62 Aora typica 1.00 0.50 5.17 47.43 Mal/acoota kameruka 0.50 0.75 5.65 50.26 .\1allacoota kameruka 0.50 1.00 4.94 52.37 Aora hebes 0.50 0.25 5.65 55.91 Shoemakerella sp. 0.00 0.50 4.92 57.29 Protohyale solitaire 0.75 0.50 5.58 61.49
301 e) g) Species Av.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Abund Contrib% Cum.% Poofie Rope Onion Astro Av. dissimilarity = 51.97 Av. dissimilarity = 53.70 Shoemakerel/a sp. 1.00 0.25 7.74 7.74 Ammyl/is kamata 0.00 1.00 8.15 8.15 Dulichiel/a australis 0.75 0.00 7.44 15.18 Gammaropsis cfatlantica 0.00 1.00 8.15 16.31 Ericthoniusforbesii 1.00 0.25 7.37 22.56 Ampithoe sp.1 1.00 0.25 6.39 22.69 Ammyllis kamata 0.75 0.00 7.30 29.85 Shoemakerel/a sp. 0.00 0.75 6.39 29.08 Tepidolpleustes coffsia 1.00 0.25 7.22 37.08 Ampithoe caddi 0.75 0.00 6.25 35.33 Aora hebes 1.00 0.25 7.22 44.30 Eusiroides monoculoides 0.00 0.75 5.75 41.08 Podocerus sp. 0.75 0.25 6.34 50.64 Quadrimaera reishi 0.00 0.75 5.75 46.83 Mal/acoota euroka 0.75 0.25 6.29 56.92 Bemlos ephippium 0.25 0.75 5.19 52.02 Paradexamine moor. 0.75 0.25 6.20 63.13 Mal/acoota malua 0.50 0.75 4.13 56.15
f) h) Species Av.Abund Av.Abund Contrib% Cum.% Species AV.Abund Av.Abund Contrib% Cum.% Scourer Rope Poofie Astro Av. dissimilarity =61.23 Av. dissimilarity = 42.37 Dulichiel/a australis 0.75 0.00 7.99 7.99 Gammaropsis efatlantica 0.00 1.00 8.76 8.76 ."vIal/acoota euroka 1.00 0.25 7.99 15.99 Tepidolpleustes co.IJsia 1.00 0.00 8.76 17.52 Quadrimaera reishi 0.75 0.00 7.66 23.65 Ampithoe sp.1 1.00 0.25 6.82 24.35 Ampithoe sp.1 0.25 0.75 6.70 30.35 Mal/acoota kameruka 1.00 0.25 6.82 31.17 Paradexamine moor. 0.75 0.25 6.40 36.75 Podocerus sp. 0.75 0.00 6.54 37.71 Ericthoniusforbesii 0.75 0.25 6.37 43.12 Eusiroides nlOnocliloides 0.00 0.75 6.24 43.95 ."vIal/acoota mailia 0.50 1.00 5.38 48.50 Dulichiel/a australis 0.75 0.25 5.58 49.53 Shoemakerel/a sp. 0.50 0.25 5.31 53.81 Paradexamine moor. 0.75 0.25 5.53 55.06 Protohyale solttaire 1.00 0.50 5.24 59.05 Quadrimaera reisht 0.25 0.75 5.38 60.44
302 !L- Species Av.Abund Av.Abund Contrib% Cum.% Scourer Astra k) Av. dissimilarity = 48.47 Species Av.Abund Av.Abund Contrib% Cum.% Gammaropsis 0.00 1.00 8.83 8.83 Onion Holdfast datlantiea Av. dissimilarity = 58. 12 Mal/acoota kameruka 1.00 0.25 6.91 15.74 Mal/acoota euroka 1.00 0.00 10.89 10.89 Gammaropsis legoliath 0.25 1.00 6.81 22.55 Ampithoe caddi 0.75 0.00 8.43 19.32 Amaryl/is kamata 0.25 1.00 6.53 29.08 Gammaropsis legoliath 0.75 0.00 8.43 27.75 Eusiroides monoeuloides 0.00 0.75 6.23 35.31 Podoeerus sp. 0.25 0.50 5.44 33.19 Dulichiel/a australis 0.75 0.25 5.73 41.04 Ericthoniusforbesii 0.75 0.50 5.44 38.64 Paradexamine moor. 0.75 0.25 5.54 46.58 Ampithoe sp.1 1.00 0.50 5.44 44.08 Aora typiea 0.50 0.75 4.51 51.08 Protohyale solitaire 0.75 0.50 5.44 49.52 J1al/aeoota malua 0.50 0.75 4.47 55.55 Shoemakerel/a sp. 0.00 0.50 5.44 54.97 Mal/acoota kameruka 0.50 0.50 5.44 60.41 j) Species Av.Abund Av.Abund Contrib% Cum.% ..!l Rope Astro Species Av.Abund Av.Abund Contrib% Cum.% Av. dissimilarity = 70.99 Poofie Holdfast Amaryllis kamata 0.00 1.00 7.31 7.3 I Av. dissimilarity = 52.48 Gammaropsis 0.00 1.00 7.31 14.61 Tepidolpleustes coffsia 1.00 0.00 9.58 9.58 cfatlantiea Mal/acoota malua 1.00 0.25 7.25 16.83 Gammaropsls legoliath 0.00 1.00 7.3 I 21.92 Mal/acoota euroka 0.75 0.00 7.14 23.97 .\.1al/acoota euroka 0.25 1.00 5.58 27.50 AmQ/)I/lis kamata 0.75 000 7.01 30.98 Bemlos ephippium 0.00 0.75 5.53 33.03 Paradexamine moor. 0.75 0.00 7.01 37.98 Eriethonius forbesii 0.25 1.00 5.39 38.42 Dulichiella australis 0.75 0.25 5.94 43.92 Aora hebes 0.25 1.00 5.28 43.70 Bemlos ephippiul1l 0.50 0.25 4.91 48.83 Eusiroides monoculoides 0.00 0.75 5.08 48.78 Podocerus sp. 0.75 0.50 4.79 53.62 Quadrimaera reishi 0.00 0.75 5.08 53.87 Ericthoniusforbesii 1.00 0.50 4.79 58.42
303 m) ....2l Species AV.Abund Av.Abund Contrib% Cum.% Species Av.Abund Av.Ablind Contrib% ClIm.% Scourer Holdfast Astro Holdtast
Av. dissimilarity = 57.78 Av. dissimilarity = 62.82 Mallacoota euroka 1.00 0.00 \0.60 10.60 Amaryllis kamata 1.00 0.00 7.99 7.99 Paradexamine moor. 0.75 0.00 7.61 \8.2\ Gammaropsis 1.00 0.00 7.99 15.99 Dulichiel/a australis 0.75 0.25 6.66 24.87 cf.atlantica Quadrimaera reishi 0.75 0.25 6.51 31.38 Gammaropsis legoliath 1.00 0.00 7.99 23.98 Aora typica 0.50 1.00 5.78 37.16 Mal/acaata ellroka 1.00 0.00 7.99 31.97 Aora hebes 0.50 0.75 5.3\ 42.48 Ellsiroides manoeu/oides 0.75 0.00 5.58 37.55 Ampithoe sp.l 0.25 0.50 5.30 47.78 Mal/acaota ma/ua 0.75 0.25 5.05 42.60 Protohyale solitaire 1.00 0.50 5.30 53.07 Bern/as ephippium 0.75 0.25 5.05 47.66 .\1al/acaota kameruka 1.00 0.50 5.30 58.37 QlIadrimaera reishi 0.75 0.25 4.77 52.43 Podocerlls sp. 0.00 0.50 4.00 56.43 n) Species AvAbund Av.Abund Contrib% Cum.% Rope Holdfast
Av. dissimilarity = 63.59 J1al/acoata ma/ua 1.00 0.25 9.74 9.74 Aora hebes 0.25 0.75 7.77 17.51 Aora typica 0.50 1.00 7.18 24.69 Ericthoniusforbesil 0.25 0.50 6.40 31.10 Podocerus sp. 0.25 0.50 6.40 37.50 Ampithoe sp.l 0.75 0.50 6.40 43.91 Protohya/e salitalre 0.50 0.50 6.40 50.31 Shoemakere//a sp. 0.25 0.50 6.40 56.71 .\1al/acoata kameruka 0.75 0.50 6.40 63.12
304 Appendix 2
List ofspecies for regional taxonomic distinctness analysis
305 Appendix 2.1: Restricted master list ofspecies used in regional analysis of taxonomic distinctness.
Species Species Amaryllis kamata Liljeborgia aequabilis Ampithoe caddi Liljeborgia polonius Ampithoe kava Linguimaera hamigera Ampithoe sp. 2 Linguimaera octodens Ampithoe sp. 3 Mallacoota euroka Ampithoe sp. 4 Mal/acoota kameruka Ampithoe sp.1 Mallacoota malua Aora hebes Paradexamine moorhousei Aora typica Paradexamine thadalee Bemlos australis Parawaldeckia sp. Bemlos ephipium Parawaldeckia yamba Bemlos fasciata Perampithoe sp. Bemlos trudis Photis cfdolichommata Ceradocus ramsayi Plumithoe quadrimanus Ceradocus serratus Podocerus sp. Dulichiella australis Polycheira cfantarctica Elasmopus arrawarra Protohyale rubra Elasmopus warra Protohyale solitaire Ericthonius forbesii Protohyale pusilia Ericthonius rodneyi Protohyale yake Eusiroides monoculoides Telsosynopia trifidilla Gammaropsis cfatlantica Tepidolpleustes coffsia Gammaropsis dentifer Quadrimaera reishi Gammaropsis haswelli Shoemakerella sp. Gammaropsis legoliath Stenothoe valida Hoho cornishi Tethygenia megalopthalma Imphimedia sp. Waldeckia sp. Ischyrocerid sp.t Ischyrocerid sp.2
306 Appendix 2.2: List ofspecies removed from regional analysis oftaxonomic distinctness due to Ii fe history not condusive to colonizing ASU habitats, low abundance records for the area or being only found in limited historic records. Life Low Historic Species history abundance record Ammyl/is sp. x Ambicholestes magel/ani x x Chevalia cf. aviculae x Cymadusa sp. x Cyproidea sp. x lschyrocerus longimanus x Jassa slatteryi x Lembos sp. x Leucothoe commensalis x Leucothoe gooweera x Leucothoe sp. x Paraleucothoe novaehol/andiae x x Ptilohyale crassicornis x x Sebs sp. x Ventojassa helenae x x 1Tentojassa zebra x x
307 Appendix 3
Lauren E. Hughes & James K. Lowry
New Species ofAmphipod (Crustacea: Peracarida) from the Solitary Islands, New South Wales, Australia
Published in Zootaxa June 2006
This paper was extracted from Chapter Six
308 ~ Zootaxa 1222: I-52 (2006) ISSN 1175-5326 (print edition) ~ www.mapress.comlzootaxa/ ZOOTAXA Copyright © 2006 Magnolia Press ISSN 1175-5334 (online edition)
New species of Amphipoda (Crustacea: Peracarida) from the Solitary Islands, New South Wales, Australia
LAUREN E. HUGHES' & JAMES K. LOWRy2
I School o/Environmental Science and Natural Resources Management, University ofNew England, National Marine Science Centre, PO Box J32/, Co.f/~· Harbour, New South Wales, 2450, Australia. E-mail: [email protected] : Cnlstacea section, Australian Museum, 6 College Street, Sydney, New South Wales, 20/0, Australia. E-mail: [email protected]
Table of contents
Abstract 2 Introduction 2 Material and methods 3 Hyalidae Bulycheva, 1957 3 Protohyale Bousfield & Hendrycks, 2002 5 Protohyle pusilla sp. nov 5 Protohyale solitaire sp. nov 8 Ischyroceridae Stebbing, 1899 11 Siphonoecetini Myers & Lowry, 2003 11 Ericthonius Milne Edwards, 1830 11 Ericthonius forbesii sp. nov 12 Ericthonius rodneyi sp. nov 16 Liljeborgiidae Stebbing, 1899 20 Liljeborgia Bate, 1862 20 Liljeborgia polonius sp. nov. . 20 Melitidae Bousfield, 1973 24 Elasmopus Costa, 1853 24 Elasmopus arrawarra sp. nov. . 24 Hoho Lowry & Fenwick, 1983 29 Hoho cornishi sp. nov 29 Photidae Boeck 1871 33 Gammaropsis Liljeborg 1855 33 Gammaropsis (Gammaropsis) legoliath sp. nov 34 Pleustidae Bucholz, 1874 40
Accepted by S. Ahyong: 30 Mar. 2006; published: / Jun. 2006 ZOOTAXA Austropleustinae Bousfield & Hendrycks, 1994 40 Tepidopleustes Karaman & Barnard, 1979 40 Tepidopleustes coffsiana sp. nov 41 Synopiidae Dana, 1852 44 Telsosynopia Karaman, 1986 44 Telsosynopia trifidilla sp. nov 44 Acknowledgements 48 References 48
Abstract
Nine new species ofepibenthic amphipods are described from the Solitary Islands, mid-north coast, New South Wales, Australia. Material was collected on natural habitats and from a range of small plastic artificial substrates. Protohyale pusilia (Chevreux, 1907) is reported from Australia for the first time. The subgenus Telsosynopia Karaman, 1986 is given generic status and Regalia juliana Lowry & Springthorpe, 2005 is transferred to Tepidopleustes in the pleustid subfamily Austropleustinae. New species include: Protohyale solitaire sp. nov. (Hyalidae); Ericthonius rodneyi sp. nov. and Ericthonius forbesii sp. nov. (lschyroceridae); Liljeborgia polonius sp. nov. (Liljeborgiidae); Elasmopus arrawarra sp. nov. and Hoho cornishi sp. nov. (Melitidae); Gammaropsis legoliath sp. nov. (Photidae); Tepidopleustes coffsiana sp. nov. (Pleustidae); and Telsosynopia trifidilla sp. nov. (Synopiidae).
Key words: Amphipoda, new species, Hyalidae, Ischyroceridae, Liljeborgiidae, Melitidae, Photidae, Pleustidae, Synopiidae, Elasmopus, Ericthonius, Gammaropsis, Liljeborgia, Protohyale, Telsosynopia, Tepidopleustes, taxonomy
Introduction
The Solitary Islands on the north coast of New South Wales are situated in a transition zone oftropical and temperate waters. The region supports a high algal diversity including stands ofthe large kelp Ecklonia radiata, together with hard and soft corals and other large sessile invertebrates on a rocky hard bottom substrate. Within these habitats epiphytic amphipods are the dominant invertebrate group, occupying a range of niches. Ninety seven species were collected during a study into the recruitment ofamphipods to artificial substrates from this region (Hughes, unpublished). Among these species, a number of commonly occurring taxa were found to be new to science and are described here: Protohyale solitaire sp. nov. (Hyalidae); Ericthonius rodneyi sp. nov. and Ericthonius forbesii sp. nov. (Ischyroceridae); Liljeborgia polonius sp. nov. (Liljeborgiidae); E lasmopus arrawarra sp. nov. and Hoho cornishi sp. nov. (Melitidae); Gammaropsis legoliath sp. nov. (Photidae); Tepidopleustes coffsiana sp. nov. (Pleustidae); and Telsosynopia trifidilla sp. nov. (Synopiidae). Protohyale pusilla (Chevreux, 1907) is reported for the first time since its original description from French Polynesia. The new
2 © 2006 Magnolia Press HUGHES & LOWRY record from the Solitary Islands is supported with illustrations ofthe new material. Regalia ZOOTAXA juliana Lowry & Springthorpe, 2005, recently described from Julian Rock on the north C!~p coast of New South Wales, is transferred to Tepidopleustes in the pleustid subfamily Austropleustinae. The subgenus Telsosynopia Karaman, 1986 (Synopiidae) is elevated to generic level.
Material and Methods
Samples from natural and artificial habitats were collected on scuba from the Solitary Islands, mid-north coast, New South Wales (Fig. 1). Artificial substrates consisted of a range ofsmall plastic units including: pot scourers; rope fibre; onion bags; shower poofies; astro turf and mimic kelp holdfasts. Material was fixed in 10% formalin and later preserved in 70% ethanol. The taxonomic descriptions were generated from several DELTA (Dallwitz 2005) databases. Characters in bold type are diagnostic. Descriptions of hyalid species were generated from a Delta database to world hyalid genera and species. Species ofProtohyale were diagnosed against all other species in the genus. Descriptions ofEricthonius species were generated from an ischyrocerid database and were diagnosed against all other species in the genus. The description of the new liljeborgiid is generated from a database to 15 other species of Liljeborgia, all of which have the same dorsal serration pattern, considered to be synapomorphic. Descriptions of Elasmopus and Hoho species were generated from the world Melitidae database. Elasmopus was diagnosed against other Australian Elasmopus and the new species of Hoho was diagnosed against all known species in the genus. The description of Gammaropsis legoliath was generated from a partial database to 32 species. Telsosynopia was diagnosed from a world Synopiidae generic database and the new species was diagnosed against all known species of Telsosynopia. Material described in this study is lodged at the Australian Museum, Sydney (AM) and Museum national d'Histoire naturelle, Paris (MNHN). The following abbreviations are used on the plates: A, antenna; Ep, epimeron; G, gnathopod; Hd, head; L, labium; Mn, mandible; Mxp, maxilliped; Mx, maxilla; P, pereopod; T, telson; U, uropod; Ur, urosomite; I, left; f, right.
Hyalidae Bulycheva, 1957
Serejo (2004) revised the higher classification of the talitridan amphipods based on cladistic analysis. The Talitroidea Rafinesque, 1815 is elevated to infraorder (Talitrida) and the Hyalidae is considered to be a sister taxon ofthe Dogielinotidae Gurjanova, 1953. The subfamilies of the Dogielinotidae include Dogielinotinae, Gurjanova, 1953, Hyalellinae Bulycheva, 1957 and Najniinae, J.L. Barnard, 1972 (new status). The
AMPHIPODA © 2006 Magnolia Press 3 ZOOTAXA Hyalidae is divided into two subfamilies, Hyacheliinae Bousfield & Hendrycks, 2002 and Hyalinae Bulycheva, 1957. This classification differs from Bousfield & Hendrycks (2002) who also included the Kuriinae as a subfamily of Hyalidae, which has now been elevated to family (Kuriidae) and superfamily (Kurioidea) status based on Serejo (2004). Bousfield & Hendrycks (2002) revised the genera of the Hyalinae (of Serejo 2004) and established five new genera, Apohyale, Protohyale, Ptilohyale, RufJohyale and Serejohyale. In Australia, the genera Apohyale, Parhyale and Ptilohyale are each represented by single species, Apohyale media (Dana, 1853), Parhyale longicornis (Haswell, 1879) and Ptilohyale crassicornis (Haswell, 1879). The genus Neobule Haswell, 1879, described from Australia, remains unidentifiable owing to the poor descriptive state ofthis taxon (Bousfield & Hendrycks 2002; Lowry & Stoddart 2003; Serejo 2004).
2; 4cr~; .. _ . Plover Island
~~ J -, tif;:~, •I \'firii 10km
North Solitary
North West Solitary
Wooigooiga Reef
South West Solitary
South Solitary
Cotfs Harbour Split Solitary
Muttonbird Island
Korffs Islet
FIGURE 1. Map ofstudy location, Solitary Islands, New South Wales, Australia.
4 © 2006 Magnolia Press HUGHES & LOWRY The remaining Australian hyalids fall into the genus Protohyale, which is divided into ZOOTAXA four subgenera Boreohyale Bousfield & Hendrycks, 2002, Diplohyale Bousfield & Hendrycks, 2002, Leptohyale Bousfield & Hendrycks, 2002, and Protohyale Bousfield & Hendrycks, 2002. This subgeneric classification was only partially applied to the Australian species, with only six of the then known eleven Australian "Hyales" species placed in subgeneric groups. The omitted species and the two species described here do not conform to the proposed Protohyale subgeneric classification. Therefore, the subgeneric classification is not applied to the Australian Protohyale species.
Protohyale Bousfield & Hendrycks, 2002
Protohyale pusilla (Chevreux, 1907) comb. nov. (Figs. 2-3) Hyale pusilla Chevreux, 1907: 415; 1908: 506-510, fig. 23-25.
Type material Holotype male 2.5 mm, MNHN-Am5773, Lagoon, Mangareva Island, Gambier Archipelago, French Polynesia, on coralline algae and Udotea sp., 5 m. Additional material examined. I male 4.6 mm, AM P71873; 3 specimens, AM P71874, North Solitary Island, Solitary Islands, 29°55'27"S 153°23' 18"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 17 February 2003, 1 specimen, AM P71875, Northwest Solitary Island, Solitary Islands, 30° 1'7"S 153° 18' 11 "E, 60 g plastic artificial substrate, 10m, L.E. Hughes, 30 January, 2003.
Description Based on male 4.6 mm, AM P71873. Head. Eyes round. Antenna 2 less than or equal to halfbody length, not densely setose. Maxilla 1 palp present, long, extending beyond end ofouter plate. Gnathopod 1 subchelate; coxa about as long as broad; propodus similar length to carpus (less than 1.5 as long as carpus), broad, length less than 2.5 x width, expanded distally, palm acute, with v-shaped distal sinus. Gnathopod 2 subchelate, carpus not lobate, not projecting between merus and propodus, without setae on the posterior margin; propodus enlarged relative to carpus, palm acute, sculptured, with two proximal humps, with robust setae, with fine simple setae, mid-palm with bowl-shaped hollow, with deep dactylar socket, comer of palm with large robust setae. Pereopods 3-4 with large robust striated setae on propodus. Pereopods 3-7 propodus distal robust seta not fusiform; dactylus with very thin seta on posterior margin. Pereopods 5-7 propodus with large robust striated setae, robust setae subequal in length. Pereopod 7 basis about as broad as long. Epimeron 3 posteroventral comer produced. Uropod 1 peduncle with enlarged distolateral robust seta. Uropod 3 uniramous, peduncle sides parallel, with distal robust
AMPHIPODA © 2006 Magnolia Press 5 ZOOTAXA setae, ramus subequal or shorter than peduncle. Telson longer than or subequal to uropod 3 peduncle, cleft, subtriangular, apically rounded.
Habitat Living on the green alga Chlorodesmis sp.; the brown algae Sargassum sp. and the holdfasts of Ecklonia radiata; and the red algae Amphiroa anceps, Delisea pulchra, Haliptilon roseum, Jania verrucosa; 0-15 m.
~ ~
FIGURE 2. Protohyale pusilla (Chevreux, 1907), male, 4.6 mm, AM P71873, North Solitary Island, Solitary Islands. Scales: Hd 0.5 mm; Ep 0.2 mm; all others 0.1 mm.
6 © 2006 Magnolia Press HUGHES & LOWRY ZOOTAXA C!~~
J
FIGURE 3. Protohyale pusilla (Chevreux, 1907), male, 4.6 mm, AM P71873, North Solitary Island, Solitary Islands. Scales: 0.2 m.
Distribution Gambier Archipelago, French Polynesia; North Solitary Island, South Solitary Island, Korffs Islet, Solitary Islands, Australia.
AMPHIPODA © 2006 Magnolia Press 7 ZOOTAXA Remarks This is the first record of Protohyale pusilla since its initial description by Chevreux (1907) and additional description in Chevreux (1908). The Solitary Islands material represents a large range extension from the type locality, Gambier Archipelago, French Polynesia, so the holotype was obtained for comparison. The largest Solitary Islands specimen is larger (male 4.6 mm) than those of Chevreux (1908) (male 2.6 mm). In the Solitary Islands specimens, the distal locking setae of pereopods 3-7 are subequal. The distal locking setae ofpereopod 5-7 in the type material is originally described with distal setae shorter than proximal, but given the small difference in relative size of the robust setae observed on the holotype, this character is now considered subequal for the type material. The character state •distal setae shorter than proximal' is reserved for species such as Apohyale media (Dana, 1853) where the difference in robust setal size is marked. The posterodistal lobe of pereopods 5-7 merus of the original material is more strongly developed than in the Solitary Islands specimens. Neither of these two pereopod characters, along with the larger size of the Solitary Islands specimens, are presently considered significant enough to separate the material here as a new species. Both Protohyale pusilla and P. wi/ari (1. L. Barnard, 1974) have a distinctive bowl shaped hollow on the mid-palm ofgnathopod 2. This structure is also weakly present in P. grenfelli (Chilton, 1916). In P. pusilla the gnathopod 2 palm proximal margin has two humps bearing setae, while in P. wi/ari the palm is smooth. In P. pusilia gnathopod 1 propodus palm uniquely has a V-shaped incision at the distal end.
Protohyale solitaire sp. nov. (Figs 4-5)
Type material Holotype male, 9.4 mm, AM P71881. Paratypes: 15 specimens, AM P71882, North Solitary Island, Solitary Islands, 29°55.450'S 153°23.306'E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 23 May 2003. Additional material examined. New South Wales: 1 specimen, AM P70430, N side of Arrawarra Headland, N of the Solitary Islands, 30 0 03'48"S 153°12'54"E, on Sargassum sp., 4 m, R.A. Peart, M. Rule, S. Richards, 21 September 1999, NSW stn 1670. 12 specimens, AM P71883, Woolgoolga reef, 30 0 TS 153°13 'E, on Jania verrucosa, 6 m, L.E. Hughes, 22 July 2003. 16 specimens, AM P71884, Southwest Solitary Island, Solitary Islands, 30 0 9'36"S 153°13'36"E, on Lobophora sp, 8 m, L.E. Hughes, 11 September 2003. 2 specimens, AM P71885, Split Solitary Island, Solitary Islands, 30 0 14'S 153°10'E, on Amphiroa anceps, 8 m, L.E. Hughes, 6 June 2002.
8 © 2006 Magnolia Press HUGHES & LOWRY Description ofholotype ZOOTAXA Head. Eyes round. Antenna 2 less than or equal to half body length, not densely @ setose. Maxilla 1 palp present, long, extending beyond end ofouter plate.
FIGURE 4. Protohyale solitaire sp. nov., holotype male, 9.4 mm, AM P71881, North Solitary Island, Solitary Islands. Scales: Hd, Ep 0.5 mm; all others 0.2 mm.
AMPHIPODA © 2006 Magnolia Press 9 ZOOTAXA
FIGURE 5. Protohyale solitaire sp. nov., holotype male, 9.4 mm, AM P71881, North Solitary Island, Solitary Islands. Scales: 0.2 mm.
Gnathopod 1 subchelate; coxa about as long as broad; propodus distinctly longer than carpus, broad, length less than 2.5 x width, propodus expanded distally, without v shaped distal sinus, palm acute. Gnathopod 2 subchelate; carpus not lobate, not projecting between merus and propodus, without setae on the posterior margin; propodus enlarged
10 © 2006 Magnolia Press HUGHES & LOWRY relative to carpus, palm acute, smooth, with robust setae, lined with fine simple setae, ZOOTAXA mid-palm without bowl-shaped hollow, with shallow dactylar socket, comer of palm with C!~~ large robust setae. Pereopods 3-4 with large robust striated setae on propodus. Pereopods 3-7 propodus distal robust seta not fusiform, with very thin seta on posterior margin. Pereopods 5-7 propodus with robust striated setae, distal robust seta shorter than proximal seta. Pereopod 7 basis about as broad as long. Epimeron 3 posteroventral comer produced. Uropod 1 peduncle with enlarged distolateral robust seta. Uropod 3 uniramous, peduncle sides parallel, with distal robust setae, ramus subequal or shorter than peduncle. Telson shorter than uropod 3 peduncle, cleft, subtriangular, apically rounded.
Etymology Named 'solitaire' after the type locality, the Solitary Islands.
Habitat Living on the green alga Chlorodesmis sp.; the brown algae Sargassum sp. and the holdfasts ofEcklonia radiata; and the red alga Amphipora anceps; 0-15 m.
Distribution New South Wales: Northwest Solitary, Woolgoolga Reef, Southwest Solitary, Split Solitary, Solitary Islands, Australia.
Remarks Protohyale solitaire is most similar to P. maroubrae (Stebbing, 1899) and P. rubra (Thomson, 1879), with the gnathopod 2 palm lined with setae. In P. maroubrae and P. rubra the palm is straight with robust setae and the distal palm comer is defined as a lip that forms from a dactylar socket. In P. solitaire the palm is convex with slender setae, and the dactylus closes into a very shallow socket that is only observable from the medial surface. In P. solitaire the propodus is distally expanded with an acute palm, while in P. rubra the gnathopod 1 propodus is elongate with an acute palm. Protohyale maroubrae is very distinct with gnathopod 1 'mitten-like' with a transverse palm.
Ischyroceridae Stebbing, 1899
Siphonoecetini Myers & Lowry, 2003
Ericthonius Milne Edwards, 1830
The genus Ericthonius has undergone much revision (see Myers & McGrath 1984). Lowry & Berents (1996) considered Ericthonius and Pseudoericthonius to be paraphyletic taxa in
AMPHIPODA © 2006 Magnolia Press 11 ZOOTAXA a cladistic analysis of the 'Ericthonius' group, which established the Cerapus and Siphonoecetes clades as sister taxa. Myers & Lowry (2003) analysed Ericthonius as part of a much larger corophiidean phylogeny and placed the 'Ericthonius' group in the Photoidea: Ischyroceriidae: Ischyroceriinae: Siphonoecetini. In a review of the North-East Atlantic Ericthonius species Myers & McGrath (1984) divided the genus into two groups. The two new Australian species are part of 'group one' in which the gnathopod 2 coxa has stridulating ridges, coxa 2 is widely separated from coxa 1 and 3, and gnathopod 2 carpus has two teeth or one in some hyperadults. The six Australian species of Ericthonius can be divided based on the gnathopod 2 basis, broad in E. brevicarpus Vader & Myers, 1996, E. coxacanthus Moore, 1988 and E. tacticus Moore, 1988 and thin and elongate in E. forbesii sp. nov., E. pugnax Dana, 1852 and E. rodneyi sp. nov.
Ericthoniusforbesii sp. nov. (Figs 6-8)
Type material Holotype male, 6.0 mm, AM P71886, Muttonbird Island, Solitary Islands, 30° ITS 153°10'E, on the brown alga Ecklonia radiata holdfasts, 8 m, L.E. Hughes, 23 May 2002. Paratypes: 1 female, 6.0 mm, AM P71887; 1 male, 4.8 mm, AM P71888; 1 male, 4.8 mm, AM P71889; 1 male, 5.5 mm, AM P71890; 23 specimens, AM71891, type locality. Additional material examined. New South Wales: 8 males, 8 females, AM P71892, North Solitary Island, Solitary Islands, 29°55'27"S 153°23' 18"E, 60 g plastic artificial substrate, 10m, L.E. Hughes, 17 February 2003. 2 males, 7 females, AM P71893, Korffs Islet, Solitary Islands, 300 19'8"S 153°9'12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003. Many specimens, AM P71894, E side of Bare Island, Botany Bay, Sydney, 33°59'58"S 151 °13 '50"E, 60 g plastic artificial substrate, 6 m, L.E. Hughes, 12 March 2004. 2 specimens, AM P70425, NE Botany Bay, 33°58' 11 "s 151 °12 '39"E, sand, 3 m, NSW State Pollution Control Commission, 9 December 1976, stn 32. 3 specimens, AM P70427, NW Botany Bay, 33°58'S 151°10'E, sand, 3.4 m, NSW State Pollution Control Commission, 22 November 1976, stn 12. 42 specimens, AM P25548, W of La Perouse, Botany Bay, 33°59'30"S 151 °13 'E, sand, 7 m, NSW State Pollution Control Commission, 20 December 1976, stn 43. 1 specimen, AM P49051, Weeney Bay, Botany Bay, 34°01'18"S 151°09'42"E, mud, 1 m, A.R. Jones, A. Roach, 30 March 1995. 1 specimen, AM P70423, E of Brighton-Ie-Sands, Botany Bay, 33°58'S 151 °1O'E, mud, 7 m, NSW State Pollution Control Commission, 18 November 1976, stn 11. 1 specimen, AM P70428, E ofBrighton-Ie-Sands, BotanyBay, 33°58'20"S 151°10'14"E, muddy sand, 7 m, NSW State Pollution Control Commission, 3 December 1976, stn 26. 1 specimen, AM P70424, SW Botany Bay, off Ramsgate, 33°59'S 151°10'E, mud, 4.9 m, NSW State Pollution Control Commission, 23 November 1976, stn 20. 1 specimen, AM P70426, S of
12 © 2006 Magnolia Press HUGHES & LOWRY Banksmeadow, Botany Bay, 33°58'47"S 151°12'16"E, mud, 8 m, NSW State Pollution ZOOTAXA Control Commission, 13 December 1976, stn 37. Many specimens, AM P23111, 1 km E of @ Burwood Beach, 32°57'31"S 151°44'43"E, bottom fine sand, 26 m, 18 December 1975.3 specimens, AM P23112, 1.5 km E ofBurwood Beach, 32°57'31"S 151°44'43"E, bottom mud and gravel, 28 m, 18 December 1975.
Description ofholotype Head. Eyes large (114 or more of head length). Maxilla 1 inner plate setae along medial margin. Mandibular palp article 3 distinctly shorter than article 2. Gnathopod 1 basis posterior margin without knob-like process; carpus distinctly longer than propodus; propodus very wide (less than 1.5 times as long as wide). Gnathopod 2 carpochelate; coxa 2 widely separated from coxa 1 and 3, rounded, distinctly broader than deep, ventral margin with stridulating nodules or ridges; carpus with distinct free posterior expansion, margin extended into a single carpal tooth or with two carpal teeth; propodus subequal to or shorter than carpus, rectangular; dactylus with cluster of distal setae. Pereopods 3-4 basis expanded, flask shaped. Pereopods 5-7 basis expanded, dactyli with accessory spines on anterior margins. Pereopod 5 merus without posterior distal lobe. Uropod 1 peduncle without distoventral corona of cuticular spines; outer ramus with more than 2 apical robust setae. Uropod 2 peduncle with several dorsal robust setae. Uropod 3 peduncle short (length 2 x or less breadth), expanded proximally. Female (sexually dimorphic characters). Based on paratype female 6.0 mm, AM P71887. Gnathopod 2 subchelate; carpus margin with a blunt extension; propodus very long (more than 2.5 times as long as carpus), palm acute, with distinct comer or spine on posterior margin, posterodistal margin with robust setae; dactylus without cluster ofdistal setae.
Etymology Named for Alistair Roderick Forbes, who supported the technical aspects of this project.
Habitat Living on the green alga Halimeda sp.; the brown algae Lobophora sp., Sargassum sp. and the holdfasts of Ecklonia radiata; the red algae Amphiroa anceps, Haliptilon roseum; and in association with the tunicate Herdmania momus; and on sand, mud and gravel bottoms; 0-15 m.
Distribution New South Wales: North Solitary, Northwest Solitary, South Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Bare Island.
AMPHIPODA © 2006 Magnolia Press 13 ZOOTAXA
FIGURE 6. Ericthoniusforbesii sp. nov., holotype male, 6.0 mm, AM P71886, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
14 © 2006 Magnolia Press HUGHES & LOWRY ZOOTAXA
FIGURE 7. Ericthonius forbesii sp. nov., holotype male, 6.0 mm, AM P71886; GI & G2, female, 6.0 mm, AM P71887, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
Remarks See remarks for Ericthonius rodneyi sp. nov.
AMPHIPODA © 2006 Magnolia Press 15 ZOOTAXA
FIGURE 8. Ericthonius forbesii sp. nov., paratypes G2 "a", hyperadult male, 5.5 mm, AM P71890; G2 "b", male, 4.8 mm, AM P71889; G2 "c", juvenile male, 4.8 mm, AM P71888, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
Ericthonius rodneyi sp. nov. (Figs 9-10)
Type material Holotype male, 4.5 mm, AM P7l895, Muttonbird Island, Solitary Islands, New South Wales, 300 17'S 153°10'E, on the brown alga Ecklonia radiata holdfasts, 8 m, L.E. Hughes, 23 May 2002. Paratypes: 1 female, 4.5 mm, AM P71896; 3 males, AMP71897, type locality. Additional material examined. New South Wales: 1 male, 6 females, AM P71898, Southwest Solitary Island, Solitary Islands, 30 0 9'48''S 153°13'48"E, on the brown alga E cklonia radiata, 10m, L.E. Hughes, 11 September 2003. 1 male, 9 females, AM P71899, Korffs Islet, Solitary Islands, 300 19'8''S 153°9'12"E, on the brown alga Ecklonia radiata holdfast,8 m, L.E. Hughes, 5 August 2003.
16 © 2006 Magnolia Press HUGHES & LOWRY Description ofholotype ZOOTAXA Head. Eyes small (less than 1/4 head length). Maxilla 1 inner plate with 2-3 setae. @ Mandibular palp article 3 medially broad, distinctly shorter than article 2.
FIGURE 9. Ericthonius rodneyi sp. nov., holotype male, 4.5 mm, AM P71895, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
AMPHlPODA © 2006 Magnolia Press 17 ZOOTAXA
FIGURE 10. Ericthonius rodneyi sp. nov., holotype male, 4.5 mm, AM P71895; 01 & 02, paratype, female, 4.5 mm, AM P71896, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
Gnathopod 1 basis posterior margin without knob-like process; carpus distinctly longer than propodus; propodus very wide (less than 1.5 times as long as wide). Gnathopod 2 carpochelate; coxa 2 widely separated from coxa 1 and 3, about as deep as
18 © 2006 Magnolia Press HUGHES & LOWRY broad, rounded, ventral margin with stridulating nodules or ridges; carpus with distinct ZOOTAXA free posterior expansion, margin with two carpal teeth; propodus subequal to or shorter C!~~ than carpus, short, rectangular; dactylus without cluster of distal setae. Pereopods 3-4 basis expanded, flask shaped. Pereopods 5-7 basis expanded; dactyli with accessory spines on anterior margins. Peraeopod 5 basis without posterodistal lobe. Uropod 1 peduncle without distoventral corona ofcuticular spines; outer ramus with 1-2 apical robust setae. Uropod 2 peduncle without dorsal robust setae. Uropod 3 peduncle short (length 2 x or less breadth), expanded proximally. Female (sexually dimorphic characters). Based on paratype female 4.5 mm, AM P71896. Gnathopod 2 subchelate; propodus long (less than 2.5 times as long as carpus), palm acute, with distinct comer or spine on posterior margin, posterodistal margin without robust setae.
Etymology Named for Professor Rod Simpson, the inaugural director of the National Marine Science Centre.
Habitat Living on the green algae Halimeda sp.; the brown algae Lobophora sp., Sargassum sp. and the holdfasts of Ecklonia radiata; the red algae Amphiroa anceps; and in association with the tunicate Herdmania momus, 0-15 ffi.
Distribution New South Wales: Northwest Solitary; Southwest Solitary; Split Solitary; Muttonbird Island; Korffs Islet; Solitary Islands.
Remarks Ericthonius forbesii and E rodneyi have no anteroproximal projection on the gnathopod 2 coxa. This separates them from E brevicarpus and E coxacanthus which have subquadrate and subtriangular proximal projections respectively on coxa 2. E ricthonius tacticus (and also E coxacanthus) have a row of 4 robust setae along the posterior margin of the gnathopod 2 carpus not present in other species of Australian E ricthonius. Both Eforbesii and E pugnax (males) have long setae at the distal end ofthe dactylus of gnathopod 2 that separates them from E. rodneyi. In E. forbesii and E. pugnax the gnathopod 2 carpal projection ofmale hyperadults is developed as a single carpal spur, but this is as yet unknown for E. rodneyi. Ericthonius forbesii may be distinguished from E pugnax by the posterior distal lobe on the basis ofpereopod 5 in E pugnax. Ericthonius forbesii and E. rodneyi appear to be most similar to each other in the male gnathopod 2 coxa shape, basis width and carpal lobe. However, both species, can be
AMPHIPODA © 2006 Magnolia Press 19 ZOOTAXA readily distinguished by eye size; E. forbesii has a large orange eye (more than 1/4 of the head length) and E. rodneyi has a small brownish eye. In Ericthonius forbesii, maxilla 1 has many setae on the inner plate, the apical margins of the uropod 1 rami and the peduncle ofuropod 2 have more robust setae than E. rodneyi. Erichtoniusforbesii also has the long distal setae on the gnathopod 2 dactylus in males, and adult specimens are slightly larger (6.0 mm) than E. rodneyi (4.5 mm). Gravid females were observed in E. rodneyi which helped distinguish it from juvenile E. forbesii. In both species, stridulations are present on the male and female gnathopod 2 coxa, unlike E. coxacanthus and E. tacticus, in which stridulations are only present in the male.
Liljeborgiidae Stebbing, 1899
Li/jeborgia Bate, 1862
Li/jeborgia polonius sp. nov. (Figs 11-12)
Type material Holotype male, 7.2 mm, AM P71900, Muttonbird Island, Solitary Islands, New South Wales, 300 1TS 153°10'E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 25 May 2003. Paratypes: 1 female, 6.5 mm, AM P7190 1; 1 specimen, AM P71902, type locality Additional material examined. New South Wales: 3 specimens, AM P71903, Northwest Solitary Island, Solitary Islands, 30°1'7"S 153° 18' 11 "E, 60 g plastic artificial substrate, 10m, L.E. Hughes, 23 May 2003. 3 specimens, AM P58595, Bulk Liquids Berth, Botany Bay, 33°58'30"S 151°12'36"E, piling scraping, 7 m, NSW Fisheries, 19 October 1998. 1 specimen, AM P70353, NW side of Tollgate Islands, Batemans Bay, 35°44'45"S 150 0 15'26"E, Ecklonia radiata holdfast & attached rock, 12 m, G.D. Wilson, A.J. Millar, 29 October 2002, NSW stn 2011. 3 specimens, AM P70431, 50 m E of Sullivan's Reef, Ulladulla, 35°21 '20"S 150 0 29'22"E, Ecklonia radiata holdfasts between boulders, 23 m, A. Murray, 5 May 1997, NSW stn 1328. 1 specimen, AM P70432, Golf Course Bommie, 500 m NE of Ulladulla Head, 35°20'29"S 150°29' 12 "E, Cluster of sponges under ledge, 15 m, K. Attwood, 2 May 1997, NSW stn 1295. 2 specimens, AM P70433, Golf Course Bommie, 500 m NE of Ulladulla Head, 35°20'29"S 150°29' 12 "E, grey & orange sponges on side oflarge boulder, 15 m, K. Attwood, 2 May 1997, NSW stn 1297. Description of holotype. Head. Eye very large (> 1/3 of head length). Lateral cepahlic lobe apically truncate. Mandibular palp article 3 much shorter than article 2. Gnathopod 1 basis posterior margin setose; dactylus inner margin with 9 teeth. Gnathopod 2 basis posterior margin with rows of thick setae; propodus palm evenly convex; dactylus inner margin 7-10 teeth. Pereopods 3-4 propodus with row of8-9 small
20 © 2006 Magnolia Press HUGHES & LOWRY robust setae, with distal seta similar to proximal setae. Pereopod 5 basis posterior margin ZOOTAXA serrate. Pereopods 5-6 basis subrectangular; dactylus apically subacute, dactylus short. Pereopod 7 propodus with long fine setae. Pleonite/urosomite dorsal serration formula (3-3-0-1-1).
UL I~ / \ ;' \ 1\/ I r 1V / Mn l
,
FIGURE 11. Liljeborgia polonius sp. nov., holotype male, 7.2 mm, AM P71900, Muttonbird Island, Solitary Islands. Scales: U, T 0.5 mm; Hd 0.2 mm; all others 0.1 mm.
AMPHIPODA co 2006 Magnolia Press 21 ZOOTAXA
FIGURE 12. Liljeborgia polonius sp. nov., holotype male, 7.2 mm, AM P71900, Muttonbird Island, Solitary Islands. Scales: 0.5 mm.Description ofholotype
Epimeron 3 with single postereoventral tooth, posteriodistal comer with notch. Uropod 1 inner ramus with 4-5 robust setae; outer ramus with 4-5 robust setae. Uropod 2 inner ramus with 4-5 robust setae; outer ramus 3--4 robust setae. Uropod 3 rami longer than peduncle; inner ramus with 4-6 robust setae; outer ramus 3-4 robust setae. Telson apical margin equal, apical setae well developed, single pair ofrobust apical setae.
22 © 2006 Magnolia Press HUGHES & LOWRY Female (sexually dimorphic characters). Based on paratype female 6.5 mm, AM ZOOTAXA P71901. Pereon. Pleonite/urosomite dorsal serration formula (0-0-0-0-0). C!~p
Etymology Named after the Shakespearian character Polonius (from Hamlet), as a play on his words "Neither a borrower (lumper) nor a lender (splitter) be", reflecting the difficulty in making decisions on species taxonomy within the Liljeborgia. Used as a noun in apposition.
Habitat Living on the brown algae Sargassum sp. and the holdfasts ofEcklonia radiata; and in association with sponges and the tunicate Herdmania momus; 0-15 m.
Distribution New South Wales: Northwest Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Batemans Bay.
Remarks Two species of Liljeborgia have been described from Australian waters, L. dubia (Haswell, 1879) and L. aequabilis (Stebbing, 1888). Liljeborgia dubia is very distinctive with numerous serrate body features, including uni-, tri- and quindentate dorsal serrations on the pleon and urosome and a heavily serrate posterior margin on the basis ofpereopods 5-7. This new species most closely resembles L. aequabiIis, which is aligned with a group of 16 other liljeborgids based on dorsal serration pattern. A 3-3-0-1-1 dorsal serration formula is present in 17 species of Liljeborgia: L. aequabilis; L. akaroica (subspecies L. a. akaroica and L. a. maria) Hurley, 1954; L. dellavallei Stebbing, 1906 (= L. mixta Schellenberg, 1925); L. enigmatica Ledoyer, 1986; L. eurycradus Thurston, 1974; L. geminata J. L. Barnard, 1969b; L. japonica Nagata, 1965; L. kinahani Bate, 1862; L. longicornis Schellenberg, 1931; L. macrodon Schellenberg, 1931; L. marcinobrio J.L. Barnard, 1969b; L. octodentata Schellenberg, 1931; L. petrae Lyons & Myers, 1991; L. polonius sp. nov., L. psaltrica Krapp-Schickel, 1975; L. serrata Nagata, 1965 and L. serratoides Tzvetkova, 1967. Liljeborgia polonius is similar to L. akaroica, L. geminata and L. petrae, which all have 4 or 5 serrate teeth on the dactylus of gnathopod 1, and telsonic lobes with even apices (telson unknown for L. petrae). Liljeborgia polonius has a single tooth and small sinus on the posterior distal corner of the epimeron 3 as does L. petrae, while in L. geminata and L. akaroica, the epimeron 3 corner is developed with one and two teeth respectively, and a large sinus. Liljeborgia polonius is most similar to L. petrae from the Gulf of Aqaba, Red Sea. Liljeborgia polonius differs by the 9 robust setae on the anterior margin ofthe propodus of
AMPHIPODA © 2006 Magnolia Press 23 ZOOTAXA pereopods 3 and 4, while L. petrae has only 4 robust setae. Liljeborgia polonius also has more robust setae on the peduncle ofthe uropods 1-3. Based on the geographic isolation of L. polonius from L. petrae and the above morphological differences, L. polonius is considered as a new species. Nagata (1965) remarked on a similar situation with L. serrata (from Seto Inland Sea, Japan) and L. macrodon (from southern South America), where species were morphologically very similar but widely separated geographically. Having grouped a set of Liljeborgia species on the basis of dorsal serrations, it is worth noting this character is somewhat unstable (Lyons & Myers 1991). The number of dorsal serrations in an individual is known to develop with age, and the dorsal serration pattern of a species may vary geographically. J.L. Barnard (1962b) and Ledoyer (1986) provide tables of dorsal serration formulae for several species but both emphasize that these should be used cautiously in initial identifications. In the species at hand the 3-3-0-1-1 pattern is stable for adult males examined. Strikingly, dorsal serrations are absent in females and juveniles examined. This sexually dimorphic character easily separates female L. polonius from female L. petrae and other closely affiliated species.
Melitidae Bousfield, 1973
Elasmopus Costa, 1853
Elasmopus arrawarra sp. nov. (Figs 13-14)
Type material Holotype male, 6.8 mm, AM P71904, Muttonbird Island, Solitary Islands, New South Wales, 30 0 1TS 153°10'E, Amphiroa anceps, 8 m, L.E. Hughes, 25 May 2003. Paratypes: female, 5.4 mm, AMP71905; 18 specimens, AM P71906, type locality. Additional material examined. New South Wales: 9 males, 7 females, AM P71907, Northwest Solitary Island, Solitary Islands, 30°1 '7"S 153°18'1l"E, on Chlorodesmis sp., 10m, L.E. Hughes, 23 May 2003. 18 specimens, AM P71908, Korffs Islet, Solitary Islands, 300 19'8"S 153°9'12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003. 1 specimen, AM P54991, South Ledge, Cook Island, 28°11 '39"S 153°34'38"E, yellow green sponge and crinoid, 12 m, A.R. Parker, 9 June 1993, NSW stn 853. 3 specimens, AM P54990, Cook Island, NE of Mary's Rock, 28°11 '25"S 153°34'47"E, orange frilly bryozoan, 19 m, R.T. Springthorpe, 8 June 1993, NSW stn 816. 24 specimens, AM P57117, Solitary Islands Jetty, Solitary Islands, 30 0 18'24"S 153°08'30"E , worm tubes encrusted with sponge on jetty pilings, 6 m, R. T. Springthorpe, 9 March 1992, NSW stn 725. 1 specimen, AM P57257, Solitary Islands Jetty, Solitary Islands, 30 0 18'24"S 153°08'30"E, Diopatra tubes at base of jetty pilings, 8.5 m, SJ. Keable, 9
24 © 2006 Magnolia Press HUGHES & LOWRY March 1992, NSW stn 727. Many specimens, AM P56657, Solitary Islands Jetty, Solitary ZOOTAXA Islands, 30 0 18'24"S 153°08'30"E, Pyurapraeputialis on jetty pilings, 8 m, P.B. Berents @ SJ. Keable, 9 March 1992, NSW stn 733. Many specimens, AM P56656, Solitary Islands Jetty, Solitary Islands, 300 18'24"S 153°08'30"E, arborescent sponge on jetty pilings, 7 m, SJ. Keable, 9 March 1992, NSW stn 735. Several specimens, AM P57258, Solitary Islands Jetty, Solitary Islands, 30 0 18'24"S 153°08'30"E, coral scrapings on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 726. Many specimens, AM P56658, Solitary Islands Jetty, Solitary Islands, 300 18'24"S 153°08'30"E, orange sponge on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 729. 3 specimens, AM P57259, Solitary Islands Jetty, Solitary Islands, 30 0 18'24"S 153°08'30"E, finger sponge on jetty pilings, 4 m, R.T. Springthorpe, 9 March 1992, NSW stn 734. 19 specimens, AM P57118, Solitary Islands Jetty, Solitary Islands, 30° 18'24"S 153°08'30"E, coral scrapings on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW stn 738. 1 specimen, AM P56659, Boambee Creek, Sawtell, 30 0 20'24"S 153°05'30"E, exposed flat of sandy mud, 0 m, Australian Museum party, 8 March 1992, NSW stn 723.
Description ofholotype Head. Lateral cephalic lobe broad, truncated, anteroventral corner rounded. Antenna 1 longer than antenna 2; peduncular article 1 longer than article 2, with 1 robust seta on posterior margin. Antenna 2 peduncular article 2 cone gland reaching at least to end ofpeduncular article 3; article 4 subequal to article 5. Mandible palp article 1 about twice as long as broad, not produced distally; article 2 subequal to article 3. Gnathopod 1 coxa anteroventral comer produced, rounded; propodus palm acute, convex. Gnathopod 2 left and right gnathopods subequal in size; merus with rounded distoventral comer; propodus setose, without medial depression, with well developed mid-palmar process, palm extremely acute, convex, with groups of anterodistal fine setae, without posteroventral comer, posterodistal robust setae absent, distal tooth apically subacute; dactylus closing along palm, not reaching end of palm. Pereopod 5 carpus and propodus with few (or no) long slender setae along anterior margin. Pereopod 6 basis posterior margin convex, castelloserrate. Pereopod 7 basis posteroventral corner narrowly rounded or subquadrate, with posterior margin castelloserrate. Epimera 1-2 margin without spines above posteroventral comer. Epimeron 1 posteroventral comer with small acute spine. Urosomite 1 without dorsal carina. Uropod 3 rami distally truncated; inner ramus shorter than outer ramus; outer ramus longer than peduncle, I-articulate. Telson apical conical extension absent, each lobe with 3 or more apical/subapical robust setae. Female (sexually dimorphic characters). Based on paratype female 5.4 mm, AM P71905. Gnathopod 2 propodus palm acute, with sparse robust setae, palm defined by posterodistal robust setae.
AMPHIPODA © 2006 Magnolia Press 25 ZOOTAXA
FIGURE 13. Elasmopus arrawarra sp. nov., holotype male, 6.8 mm, AM P71904, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
26 © 2006 Magnolia Press HUGHES & LOWRY ZOOTAXA C!~~ (-) \ P7
FIGURE 14. Elasmopus arrawarra sp. nov., holotype male, 6.8 mm, AM P71904; GI & G2, paratype female, 5.4 mm, AM P71905, Muttonbird Island, Solitary Islands. Scale: 0.2 mm.
Etymology Named from the field station where part ofthis study was based.
AMPHIPODA © 2006 Magnolia Press 27 ZOOTAXA Habitat Living on the green algae Chlorodesmis sp., Halimeda sp.; the red algae Delisea pulchra, Haliptilon roseum, Hypnea sp., Pterocladia sp.; the brown algae Dictyota sp., Dilophus marginatus, Lobophora sp., Sargassum sp. and holdfasts of Ecklonia radiata,; barnacle rubble; sponges; coral; sandy mud; bryozoans; and in association with the tunicates Herdmania momus and Pyura praeputialis; and in tubes of the polychaete Diopatra sp.; 0-15m.
Distribution New South Wales: Cook Island, Tweed Heads; Northwest Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Sawtell.
Remarks Krapp-Schickel & Ruffo (1990) recognised a group of Elasmopus defined by a dense fringe of setae along the palm ofgnathopod 2 and a castelloserrate margin on the basis of pereopods 6 and/or 7: Elasmopus canarius Krapp-Schickel & Ruffo, 1990; E. crenulatus Berents, 1983; E. laufolii Myers, 1986; E. pectinicrus J.L. Barnard, 1955; E. serricatus J.L. Barnard, 1969a; E. spinibasus Sivaprakasam, 1970 and E. yunde Barnard, 1974. Elasmopus arrawarra sp. nov., along with E. lapu Myers, 1985, E. nanshaensis Ren, 1998 and E. steelei Appadoo & Myers, 2003, also belong in this group. Elasmopus arrawarra, E. canarius E. nanshaensis, E. serricatus can be further separated by the mid-palmar process on the propodus ofgnathopod 2. In E. lapu, and E. yunde the mid-palmar process is present as a long low ridge. The distal palmar tooth on the propodus ofgnathopod 2 in E . canarius and E. nanshaensis is truncate while in E. arrawarra and E. serricatus the tooth is subacute. Elasmopus arrawarra is distinguished from all other species by the combination ofthe produced anterior comer ofthe gnathopod 1 coxa, the gentle parabolic shape ofthe mid-palmar process on gnathopod 2, the convex posterior margin ofthe basis ofpereopod 6, the castelloserrate posterior margin ofthe basis ofpereopods 6--7, the weak sharp tooth on the posteriodistal comer of epimera 1-3 and the inner ramus of uropod 3 without medial setae. In comparison to other Australian species ofElasmopus, E. arrawarra is superficially similar to E. warra Kelaher & Lowry, 2002 in the form of the gnathopod 2 with the acute distal palmar tooth, however the pereopods 6--7 basis in E. warra is not castelloserrate, and the setae along pereopods 6-7 are much longer than in E. arrawarra. Elasmopus crenulatus has a crenulate posterior margin on the basis of the pereopods 6-7 and a well developed distal tooth on the propodus of gnathopod 2, but does not have a mid-palmar process on the propodus of gnathopod 2. The arrangement of robust setae on the acute distal tooth on the palm ofgnathopod 2 is also different from that of E. arrawarra.
28 © 2006 Magnolia Press HUGHES & LOWRY Hoho Lowry & Fenwick, 1983 ZOOTAXA C!~p Hoho cornishi sp. nov. (Figs 15-16)
Type material Holotype male, 12.5 mm, AM P71909, Muttonbird Island, Solitary Islands, New South Wales, 30 0 1TS 153°10'E, Sargassum sp., 8 m, L.E. Hughes, 21 May 2002. Paratypes: 6 males, AM P7191 0, type locality. Additional material examined. New South Wales: Many specimens, AM P71911, E side ofBare Island, Botany Bay, Sydney, 33°59'58"S 151°13'84"E, 60 g plastic artificial substrate, 6 m, L.E. Hughes, 12 March 2004. 1 specimen, AM P57245, NE of Mary's Rock, Cook Island, 28°11 '25"S 153°34'47"E, Zonaria sp., 17 m, R.T. Springthorpe, 8 June 1993, NSW stn 814. 1 specimen, AM P57246, NE of Mary's Rock, Cook Island, 28° 11 '25"S 153°34'47"E, orange frilly bryozoan, 19 m, R.T. Springthorpe, 8 June 1993, NSW stn 816. 2 specimens, AM P57247, North Ledge, Cook Island, 28°11 '26"S 153°34'40"E, Zonaria sp., 10 m, L. Albertson, 8 June 1993, NSW stn 817. 1 specimen, AM P57248, South Ledge, Cook Island, 28°11 '39"S 153°34'38"E, rock, 15 m, K.B. Attwood, 9 June 1993, NSW stn 851. 7 specimens, AM P57249, 100 m NW of Split Solitary Island, 300 14'S 153°10'48"E, red algae, 17 m, S.J. Keable, 7 March 1992, NSW stn 682. 8 specimens, AM P57250, 100 m NW of Split Solitary Island, 30 0 14'S 153° 10'48"E, mixed red algae, 17 m, S.J. Keable, 7 March 1992, NSW stn 693. 2 specimens, AM P57242, 100 m NW of Split Solitary Island, 300 14'S 153°10'48"E, Halimeda sp., 17 m, S.J. Keable, 7 March 1992, NSW stn 689. 12 specimens, AM P57243, 100 m NW of Split Solitary Island, 300 14'S 153°10'48"E, brown algae, 17 ill, S.J. Keable, 7 March 1992, NSW stn 694. 1 specimen, AM P42820, Moe's Rock, S of Jervis Bay, 35°09'S 150 0 4YE, foliose bryozoan, 18 m, J.K. Lowry & R.T. Springthorpe, 29 June 1981, NSW stn 55. 1 specimen, AM P52794, Moe's Rock, S of Jervis nay, 35°09'S 1500 45'E, foliose bryozoan, 18 m, J.K. Lowry, R.T. Springthorpe, 29 June 1981, NSW stn 55. 1 specimen, AM P42816, off Snapper Point, Kioloa, 35°34'30"S 150°22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15.1 specimen, AM P42817, offSnapper Point, Kioloa, 35°34'30"S 150 0 22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15. 1 specimen, AM P42818, off Snapper Point, Kioloa, 35°34'30"S 150 0 22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15. 1 specimen, AM P42819, off Snapper Point, Kioloa, 35°34'30"S 150 0 22'E, algal washings from sandstone slab with coarse gravel, 20 m, J.K. Lowry, R.T. Springthorpe, 25 April 1981, NSW stn 15. 1 specimen, AM P42815, N side of O'Hara Head, Kioloa, 35°34'S 150 0 23'E, coralline algae, 2 m, P.B. Berents, J.K. Lowry, R.T. Springthorpe, P.M. Berents, 25 April 1981, NSW stn 12. 1 specimen, AM P70342, NE of Point Upright, SW side of Grasshopper
AMPHIPODA © 2006 Magnolia Press 29 ZOOTAXA Island, 35°38'01"S 150°19'51 "E, mixed algae, 11 m, P.B. Berents, J. Eu, A.J. Millar, GO. Wilson, 10 February 2003, NSW stn 2037. Many specimens, AM P70343, W side ofWasp Island, N ofBatemans Bay, 35°40'02"S 1500 18'29"E, Melanthaliapolydactylis (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, GO. Wilson, 10 February 2003, NSW stn 2045. Many specimens, AM P70334, N of Burrewarra Point, East Wall, 35°50'01"S 150°14' 10"E, from macroalga Peyssonelia nova-hollandiae, 25 m, GO. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1985. Many specimens, AM P70333, N of Broulee Island, 35°51 '20"S 150°11 '38"E, from macroalga Phacelocarpus apodus, 16 m, GO. Wilson, A.J. Millar, N. Vee, 30 October 2002, NSW stn 2017. 2 specimens, AM P70341, S of Broulee Island, 35°50'50"S 150°11 '05"E, from macroalga Halopteris platycena, 8 m, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1990. 5 specimens, AM P70337, S of Broulee Island, 35°50'49"S 150°11 '05"E, from corallines Amphiora anceps & Jania natalensis, 8 m, GO. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1989. I specimen, AM P70336, NW side of Tollgate Islands, Batemans Bay, 35°44'46"S 150 0 15'28"E, from macroalga Sporochnus radiciformis, 12 m, GO. Wilson, A.J. Millar, N. Vee, 28 October 2002, NSW stn 2006.2 specimens, AM P70335, NW side of Tollgate Islands, Batemans Bay, 35°44'46"S 150 0 15'28"E, from macroalga Galaxaura marginata, 12 m, GO. Wilson, A.J. Millar, N. Vee, 28 October 2002, NSW stn 2005. 5 specimens, AM P70338, NW side ofTollgate Islands, Batemans Bay, 35°44'50"S 1500 15'32"E, from macroalga Caulerpaflexilis, 7.9 m, GO. Wilson, A.J. Millar, 28, October 2002, NSW stn 2001. 2 specimens, AM P70339, NW side ofTollgate Islands, Batemans Bay, 35°44'46"S 150 0 15'28"E, from macroalga Scinaia tsinglanensis, 12 m, GO. Wilson, AJ. Millar, 28 October 2002, NSW stn 2004. I specimen, AM P70340, NW side of Tollgate Islands, Batemans Bay, 35°44'45"S 1500 15'26"E, from macroalga Ecklonia radiata holdfast and rock, 12 m, GO. Wilson, A.J. Millar, 29 October 2002, NSW stn 2011. Many specimens, AM P70345, W side of North Tollgate Island, Batemans Bay, 35°44'55"S 150 0 15'27"E, Chondria succulenta (alga), 9 m, P.B. Berents, J. Eu, A.J. Millar, GO. Wilson, II October 2003, NSW stn 2053. 2 specimens, AM P70344, W side of North Tollgate Island, 35°44'55"S 150 0 15'28"E, Caulerpa cactoides (alga), 9 m, P.B. Berents, J. Eu. AJ. Millar, GD. Wilson, II February 2003, NSW stn 2052. 2 specimens, AM P70332, N of Jimmy's Island, 35°48'56"S 150 0 14'06"E, from macroalga Dictyota dichotoma, 16 m, GO. Wilson, A.J. Millar, N. Vee, 29 October 2002, NSW stn 2010.
Description ofholotype Head lateral cephalic lobe broad, truncated, anteroventral comer rounded. Antenna 1 peduncular article I longer than article 2, with 4 or more robust setae along posterior margin. Antenna 2 peduncular article 4 longer than article 5. Mandibular palp vestigial, I-articulate. Gnathopod 1 coxa posteroventral comer notch absent; merus without posterodistal tooth; propodus palm convex, without posterodistal comer, posterodistal robust setae
30 © 2006 Magnolia Press HUGHES & LOWRY present. Gnathopod 2 coxa posteroventral comer notch absent; merus with subquadrate ZOOTAXA distoventral comer; propodus palm acute, convex, smooth, with distal hump bearing robust setae, without posteroventral comer; dactylus not reaching end of palm, apically falcate. Pereopod 5 carpus and propodus with few (or no) long slender setae along anterior margin. Pereopod 6 basis posterior margin subsigmoidal. posteroventral comer narrowly rounded or subquadrate; carpus and propodus with few (or no) long slender setae along anterior margin; propodus not expanded posterodistally. Pereopod 7 basis posterior margin subsigmoidal; propodus not expanded posterodistally.
FIGURE 15. Hoho cornishi sp. nov., holotype male, 12.5 mm, AM P71909, Muttonbird Island, Solitary Islands. Scales: Hd, Ep 0.5 mm; all others 0.2 mm.
AMPHIPODA © 2006 Magnolia Press 31 ZOOTAXA
FIGURE 16. Hoho cornishi sp. nov., holotype male, 12.5 mm, AM P7l909, Muttonbird Island, Solitary Islands. Scale: 0.5 mm.
Pleonites 1-3 not dorsally bicarinate; without dorsodistal spines. Epimeron 1 posteroventral comer with small acute spine. Epimeron 3 posteroventral comer with small acute spine. Urosomite 1 dorsally bicarinate. Uropod 3 rami longer than peduncle. Telson each lobe with 3 or more apical/subapical robust setae.
32 cD 2006 Magnolia Press HUGHES & LOWRY Etymology ZOOTAXA Named for Keith Cornish who provided tremendous technical support and common @ sense in the early stages ofthis project.
Habitat Living on the green algae Caulerpa jlexili, Caulerpa cactoides, Halimeda sp.; the red algae Amphiroa anceps, Chondria succulenta, Galaxaura marginata, Halopteris platycena, Jania natalensis, Melanthalia polydactyli, Peyssonelia nova-hollandiae, Scinaia tsinglanensis; the brown algae Ecklonia radiata, Dictyota dichotoma, Halopteris platycena, Phacelocarpus apodus, Sargassum sp., Sporochnus radiciformis, Zonaria sp.; 0-15 m.
Distribution New South Wales: Cook Island, Tweed Heads; Split Solitary Island, Muttonbird Island, Solitary Islands; Bare Island, Botany Bay; Jervis Bay; Kiola; Grasshopper Island, Wasp Island, Brulee Island, Tollegate Island, Jimmy's Island, Batemans Bay.
Remarks Hoho cornishi sp. nov. is the first species ofthe genus to be described since the Hoho was established by Lowry & Fenwick, 1983. Hoho cornishi is distinguished by the shape ofthe distal hump on the palm ofgnathopod 2 propodus, usually with 5 and 7 robust setae on either side, and the subsigmoidal posterior margin of pereopods 6-7. Of the three described species of Hoho (H. carteta (J.L. Barnard, 1972a); H. hirtipalma Lowry & Fenwick, 1983; H. marilla (J. L. Barnard, 1972a)), H. cornishi is most similar to H. hirtipalma, sharing a thick fringe of setae on the palm of gnathopod 2. Hoho cornishi appears to have less robust setae on the margins ofthe pereopods than H. hirtipalma.
Photidae Boeck 1871
Gammaropsis Liljeborg 1855
Gammaropsis is a problematic genus with several intergrading subgenera, some of which are apparently polyphyletic (J.L. Barnard 1973; Barnard & Karaman 1991; Conlan 1993). Krapp-Schickel & Myers (1979) divided the Mediterranean regional fauna of Gammaropsis into two groups based on a collective oftoothed or smooth characters ofthe pleonites and urosomites, and the epistome length. This division has been further applied to Irish species by Myers & McGrath (1982). This classification, however, was not able to be extended to the southern hemisphere fauna (Myers 1985, 1995; Appadoo & Myers 2004), which have a combination of toothed and smooth characters, including the Australian species described here. Myers (1985) recognised the need for more taxonomic work in this group.
AMPHIPODA © 2006 Magnolia Press 33 ZOOTAXA The new species described below belongs in the subgenus G. (Gammaropsis) based on a reduced gnathopod 2 carpus, multiarticulate accessory flagellum and uniform anterior coxa. There are now eight species of Gammaropsis recorded from Australia (Lowry & Stoddart 2003); all are in the subgenus Gammaropsis, except for G (Paranaenia) dentifera (Haswell, 1879).
Gammaropsis (Gammaropsis) legoliath sp. nov. (Figs 17-19)
Type material Holotype male, 4.1 mm AM P71912, Muttonbird Island, Solitary Islands, New South Wales, 30 0 1TS 153°10'E, Sargassum sp., 8 ill, L.E. Hughes, 21 May 2002. Paratypes: female 5.4 mm, AM P71913; male, 3.4 mm, AM P71914; 22 specimens, AM P71915, type locality. Additional material examined. New South Wales: 20 specimens, AM P71916, Northwest Solitary Island, Solitary Islands, 30° 01 '7"S 153° 18'11 "E, 60 g artificial substrate, 10 m, L.E. Hughes, 30 January 2003.28 specimens, AM P71917, South Solitary Island, Solitary Islands, 30 0 12'0"S 153° 15'48"E, Lobophora sp., Pterocladia sp., Zonaria sp. and mixed brown algae, 8 m, L.E. Hughes, 29 July 2003. 27 specimens, AM P71918, Korffs Islet, Solitary Islands, 300 19'8"S 153°9'12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003. Many specimens, AM P70434, NW side of Brush Island, 35°31 '39"S 1500 24'58"E, Zonaria diesingiana (alga), 16.2 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 9 February 2003, NSW stn 2029. Many specimens, AM P70442, NW side of Brush Island, N of Batemans Bay, 35°31 '39"S 150°24'58"E, Zonaria diesingiana (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 9 February 2003, NSW stn 2033. Many specimens, AM P70453, NW side of Brush Island, 35°31 '39"S 150 0 24'58"E, Ecklonia radiata (alga), 12 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 9 February 2003, NSW stn 2026. 6 specimens, AM P70505, SW side of Grasshopper Island, 35°38'01"S 150° 19'51 "E, Callophycus tridentifer (alga), 11 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 10 February 2003, NSW stn 2036. Many specimens, AM P70444, SW side of Grasshopper Island, NE of Point Upright, 35°38'01"S 150°19'51 "E, Peyssonnelia novae holliandiae (alga), 13 m, P.B. Berents, J. E u, A.J. Millar, GD. Wilson, 10 February 2003, NSW stn 2038.5 specimens, AM P70510, NE of Point Upright, SW side of Grasshopper Island, 35°38'01"S 150 0 19'51"E, Peyssonnelia novae holliandiae (alga), 13 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 10 February 2003, NSW stn 2038. 20 specimens, AM P70509, W side of Wasp Island, N of Batemans Bay, 35°40'02"S 150° 18'29"E, Melanthalia polydactylis (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 10 February 2003, NSW stn 2045. 10 specimens, AM P70506, W side of Wasp Island, N of Batemans Bay, 35°40'02"S 150 0 18'29"E, Curdiea crassa (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 10 February
34 © 2006 Magnolia Press HUGHES & LOWRY 2003, NSW stn 2043. Many specimens, AM P70446, W side of Wasp Island, N of ZOOTAXA Batemans Bay, 35°40'02"S 150 0 18'29"E, Curdiea crassa (alga), 16 m, P.B. Berents, J. E @ u, AJ. Millar, GD. Wilson, 10 February 2003, NSW stn 2043. Many specimens, AM P70438, W side of Wasp Island, N of Batemans Bay, 35°40'02"S 150 0 18'29"E, Peyssonnelia novae holliandiae (alga), 16 m, P.B. Berents, J. Eu, A.J. Millar, GD. Wilson, 10 February 2003, NSW stn 2047.7 specimens, AM P70448, NW side ofTollgate Islands, Batemans Bay, 35°44'46"S 150° 15'27"E, macroalga Caulerpa cactoides, 12 m, GD. Wilson, N. Vee, 29 October 2002, NSW stn 2013.4 specimens, AM P70449, NW side of Tollgate Islands, Batemans Bay, 35°44'50"S 150 0 15'32"E, macroalga Caulerpaflexilis, 7.9 m, GD. Wilson, AJ. Millar, 28 October 2002, NSW stn 2001. 14 specimens, AM P70450, NW side of Tollgate Islands, Batemans Bay, 35°44'46"S 150 0 15'28"E, macroalga Galaxaura marginata, 12 m, GD. Wilson, AJ. Millar, N. Vee, 28 October 2002, NSW stn 2005. Many specimens, AM P70454, NW side of Tollgate Islands, Batemans Bay, 35°44'50"S 1500 15'32"E, macroalga Chondria succulenta, 7.9 m, GD. Wilson, AJ. Millar, 28 October 2002, NSW stn 2003. 10 specimens, AM P70502, NW side ofTollgate Islands, Batemans Bay, 35°44'46"S 150 0 15'28"E, macroalga Sporochnus radiciformis, 12 m, GD. Wilson, AJ. Millar, N. Vee, 28 October 2002, NSW stn 2006. Many specimens, AM P70503, NW side of Tollgate Islands, Batemans Bay, 35°44'50"S 150 0 15'32"E, macroalga Colpomenia sinuosa, 7.9 m, GD. Wilson, A.J. Millar, 28 October 2002, NSW stn 2002. 24 specimens, AM P70436, NW side of Tollgate Islands, Batemans Bay, 35°44'46"S 150 0 15'28"E, macroalga Scinaia tsinglanensis, 12 m, GD. Wilson, AJ. Millar, 28 October 2002, NSW stn 2004. 5 specimens, AM P70435, W side of North Tollgate Island, Batemans Bay, 35°44'50"S 1500 15'29"E, Halopteris platycena (alga), II m, P.B. Berents, 1. Eu, AJ. Millar, GD. Wilson, 8 February 2003, NSW stn 2022. Many specimens, AM P70501, W side of North Tollgate Island, Batemans Bay, 35°44'55"S 1500 15'27"E, Chondria succulenta (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, GD. Wilson, 11 February 2003, NSW stn 2053. 5 specimens, AM P70452, W side of North Tollgate Island, Batemans Bay, 35°44'55"S 150 0 15'28"E, Caulerpa cactoides (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, GD. Wilson, 11 February 2003, NSW stn 2052. Many specimens, AM P70445, W side of North Tollgate Island, 35°44'55"S 150 0 15'28"E, Stypopodiumflabelliforme (alga), 9 m, P.B. Berents, J. Eu, AJ. Millar, GD. Wilson, 11 February 2003, NSW stn 2055. 7 specimens, AM P70507, Gutters, N of Burrewarra Point, 35°49'49"S 150 0 14'02"E, macroalga Stypopodiumflabelliforme, 24 m, GD. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1992. 8 specimens, AM P70447, Gutters, N of Burrewarra Point, 35°49'52"S 1500 14'05"E, macroalga Codium lucasii, 23 m, GD. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1997. Many specimens, AM P70440, gutters, N of Burrewarra Point, 35°49'49"S 150 0 14'02"E, macroalga Pachymenia prostrata, 24 ill, GD. Wilson, A.J. Millar, N. Vee, 27 October 2002, NSW stn 1996. 2 specimens, AM P70508, Gutters, N of Burrewarra Point, 35°49'49"S 1500 14'02"E, branching hydroid in same bag as Pachymenia prostrata, 24
AMPHIPODA © 2006 Magnolia Press 35 ZOOTAXA m, GD. Wilson, AJ. Millar, N. Vee, 27 October 2002, NSW stn 1995. 4 specimens, AM P70504, East Wall, N of Burrewarra Point, 35°50'01"S 150 0 14'10"E, macroalga Corallina berteri, 25 m, GD. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1987. Many specimens, AM P70437, East Wall, N of Burrewarra Point, 35°50'01"S 150° 14' 10"E, macroalga Curdiea crassa with small amount of Martensia australis, 25 m, GD. Wilson, AJ. Millar, N. Vee, 25 October 2002, NSW stn 1986. Many specimens, AM P70351, East Wall, N of Burrewarra Point, 35°50'01"S 150° 14' 10"E, macroalga Peyssonelia novahollandiae, 25 m, GD. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1985. 3 specimens, AM P70439, N of Broulee Island, 35°51 '20"S 150°11 '38"E, macroalga Phacelocarpus apodus, 16 m, GD. Wilson, AJ. Millar, N. Vee, 30 October 2002, NSW stn 2017. 5 specimens, AM P70451, N of Broulee Island, 35°51 '20"S 150°11 '38"E, macroalga Lobophora variegata, 16 m, GD. Wilson, A.J. Millar, N. Vee, 30 October 2002, NSW stn 2016.2 specimens, AM P70441, S of Broulee Island, 35°50'49"S 150°11 '05"E, Amphiroa anceps & Jania natalensis, 8 m, GD. Wilson, A.J. Millar, N. Vee, 25 October 2002, NSW stn 1989. 1 specimen, AM P70443, North Kianinny Gutter, Tathra, 36°44'04"S 149°59'12"E, Pterocladia lucida (alga), 5 m, P.B. Berents, 1. Eu, AJ. Millar, GD. Wilson, 12 February 2003, NSW stn 2060.
Description ofholotype Head. Eyes distally situated, wholly or partly in lateral cephalic lobes; lateral cephalic lobes triangular, apically subacute, anteroventral margin moderately recessed, moderately excavate. Antenna 1 shorter than antenna 2, with many long slender setae, peduncle article 3 shorter than article 1; accessory flagellum mutiarticulate, without aesthetascs. Antenna 2 peduncle similar to peduncle of antenna 1. Labrum epistome present. Mandibular palp article 3 subequal in length to article 2, spatulate with mostly apical setae. Labium mandibular processes acute or sub-acute. Maxilla 1 inner plate without long apical setae, with a setae along medial margin. Gnathopod 1 coxa anteroventral comer rounded, without posteroventral spine. Gnathopod 2 symmetrical; coxa evenly rounded, subequal in length to coxa 1, without ventral spine; carpus not very reduced, triangular. Pereopod 3 coxa subequal in length and breadth. Pereopod 5 coxa anterior lobe much deeper than coxa 6; basis posterior margin smooth. Pereopod 6 slightly longer than pereopod 7; basis to propodus enlarged as rectolinear column; basis posterior margin smooth. Pereopod 7 basis posterior margin smooth; carpus more than 0.6 times length of propodus. Pleonite 3 without dorsal teeth. Epimeron 3 with posteroventral notch. Urosomite 1 with dorsal spine. Urosomite 2 without dorsal spines. Urosomite 3 with dorsal spine. Uropod 1 with well developed distoventral spur; inner ramus shorter than peduncle. Uropod 2 with vestigial distoventral spur; inner ramus subequal with peduncle. Uropod 3 peduncle longer than rami; inner ramus longer than outer ramus, inner ramus without apical setae; outer ramus I-articulate.
36 © 2006 Magnolia Press HUGHES & LOWRY Female (sexually dimorphic characters). Based on paratype female 5.4 mm, AM ZOOTAXA P71913. Pereopod 6 basis to propodus not enlarged. @
~ ~".<''''' } ~Mn
FIGURE 17. Gammaropsis legoliath sp. nov., holotype male, 4.1 mm, AM P71912, Muttonbird Island, Solitary Islands. Scales: 0.2 mm.
AMPHIPODA © 2006 Magnolia Press 37 ZOOTAXA
FIGURE 18. Gammaropsis legoliath sp. nov., holotype male, 4.1 mm, AM P71912, paratype female "a", 5.4 mm, AM P71913, Muttonbird Island, Solitary Islands. Scales: 0.2 mm.
38 © 2006 Magnolia Press HUGHES & LOWRY Etymology ZOOTAXA The name refers to the grossly enlarged pereopod 6, derived from the historical giant C!~p Goliath to reflect the size ofthe male pereopod 6 appendage.
Habitat Living on the green algae Caulerpa cactoides, Caulerpa jlexilis, Codium lucasii; the red algae Amphiroa anceps, Callophycus tridentifer, Chondria succulata, Corallina berteri, Curdiea crassa, Galaxaura marginata, Jania natalensis, Martensia australis, Melanthalia polydactylis, Pachymenia prostrate, Peyssonnelia novae holliandiae, Pterocladia lucida, Scinaia tsinglanensis and the brown algae Colpomenia sinuosa, holdfasts ofEcklonia radiata. Halopteris platycena, Lobophora variegate, Phacelocarpus apodus, Sargassum sp.; Sporochnus radiciformis, Zonaria diesingiana; 0-15 m.
Distribution New South Wales: North Solitary, Northwest Solitary, South Solitary, Split Solitary, Muttonbird Island, Korffs Islet, Solitary Islands.
Remarks As with many other Gammaropsis species, the palm of male gnathopod 2 in G legoliath goes through several stages ofdevelopment, with the final palmar structure very different from that of juveniles. The other secondary sexual characteristic in males, the enlargement ofpereopod 6, takes place before the development ofthe concave palm, and results in a male with gnathopod 2 with a convex crenulate palm and grossly enlarged pereopod 6. The distinct adult male gnathopod 2 propodus formation ofG legoliath with a concave palm and raised blunt tooth is seen in two other Australian species, G crassipes (Haswell, 1880) and G thomsoni (Stebbing, 1888). In G thomsoni gnathopod 2 is asymmetrical, the dominant gnathopod differs from G legoliath in a second blunt proximal tooth on the propodus and a tooth on the inner margin ofthe dactylus, which corresponds with a sinus on the propodus. In G crassipes and G legoliath the gnathopods are symmetrical, and both have the peculiarly enlarged pereopod 6. In Haswell's (1880) original description of G crassipes the drawings of the enlarged pereopod 6 are a little exaggerated/overstated from the type material. Stebbing (1906) discussed G crassipes and noted that pereopod 6 in his specimens was "not as stout as Haswells". Although most definitely enlarged, the form of pereopod 6 in G crassipes is enlarged so that it appears to belong to a much larger specimen than the one it is attached to. The articles of the pereopod are enlarged but not expanded. In G legoliath the large pereopod 6 is developed into a stout columnar limb with each article expanded and ofthe same width. In G legoliath the posterior margin of the basis of pereopods 6-7 is straight while in G crassipes they are sinusoidal. Gammaropsis legoliath is also much smaller than G crassipes (~4 mm vs 8 mm+).
AMPHIPODA © 2006 Magnolia Press 39 ZOOTAXA
FIGURE 19. Gammaropsis legoliath sp. nov., Paratype, male "a", 3.4 mm, AM P71914, paratype female, 5.4 mm, AM P71913, Muttonbird Island, Solitary Islands. Scales: 0.2 mm.
The characteristic large leg of G legoliath and G crassipes are not new to the Photidae. Five species of the closely associated genus Photis also display a gross enlargement of the sixth pereopod: P brevicaudata Stebbing, 1888; P elephantis J.L. Barnard, 1962a; P nigrocula Lowry, 1979; P phaeocula Lowry, 1979 and P trapherus Thomas & Barnard, 1991. In Photis the articles of pereopod 6 remain proportional except for the merus, which may be expanded and/or elongate.
Pleustidae Bucholz, 1874
Austropleustinae Bousfield & Hendrycks, 1994
Tepidopleustes Karaman & Barnard, 1979
Type species Parapleustes barnardi Ledoyer, 1972, original designation.
Diagnosis Head rostrum moderately developed (0.25-0.5 x length of article 1). Labrum deeply notched, asymmetrical. Mandible molar conical, non-triturative. Maxilliped palp article 4
40 to 2006 Magnolia Press HUGHES & LOWRY reduced or absent. Gnathopods 1-2 similar in size and shape; carpi not contilevered; ZOOTAXA subequal to or longer than propodi. Pleonites 1-2 dorsally carinate. Pleonites 1-3 with @ strong to weak lateral ridges. Epimeron 3 posterior margin serrate to smooth; posteroventral comer rounded or subquadrate. Uropod 3 peduncle without large ventrodistal disk supporting rami; endopod longer than exopod.
Species composition Tep idopIeustes barnardi (Ledoyer, 1972); Tepidopleustes coifsiana sp. nov.; Tepidopleustes honomu (J.L. Barnard, 1970); Tepidopleustes juliana (Lowry & Springthorpe, 2005).
Remarks Tepidopleustes juliana was originally misplaced in the genus Regalia with which it bears a superficial resemblance. Notably, however, T. juliana lives in a different habitat from other species of Regalia - shallow water algal bottoms instead of deep water bottoms. Although the lateral ridging on the urosome of T. juliana resembles that of Regalia species, the labium of the former is widely separated, a distinctive feature of pleustid amphipods. Including T. cofftiana and T. juliana in Tepidopleustes expands the generic concept slightly to include species with a smooth posterior margin on epimeron 3 and simple gnathopods. These become gradational characters for species such as T. honomus, which have a weakly serrate posterior margin on epimeron 3 and weakly subchelate gnathopods.
Tepidopleustes coffsiana sp. nov. (Figs 20-21)
Type material Holotype male, 3.5 mm AM P72060, 3 slides, no carcass, S side of Woolgooiga reef, off Woolgoolga Beach, N of Coffs Harbour, New South Wales, Australia, 300 07'05"S 153°13'00"E, encrusted Sargassum sp., 5 m, R. Peart & S. Richards, 23 September 1999. Additional material examined. New South Wales: Many specimens, AM P72061, Muttonbird Island, Solitary Islands, 300 17'S 153°10'E, Sargassum sp., 6 m, L.E. Hughes, 21 May 2002. Many specimens, AM P72062, South Solitary Island, Solitary Islands, 30 0 12'00"S 153°15'48"E, Lobophora sp., Pterocladia sp., Zonaria sp. and mixed brown algae, 8 m, L.E. Hughes, 29 July 2003. Many specimens, AM P72063, Korffs Islet, Solitary Islands, 30° 19'8"S 153°9' 12"E, 60 g plastic artificial substrate, 8 m, L.E. Hughes, 26 May 2003.
AMPHIPODA © 2006 Magnolia Press 41 ZOOTAXA Description ofholotype Head rostrum moderate. Antenna 1 accessory flagellum absent. Antenna 2 peduncular articles 4-5 brush setae absent. Labrum with deep notch, asymmetrical. Labium inner lobes small, narrow; outer lobes large, narrowly separated. Mandible lacinia mobilis present on both sides, right blade-like, left serrate; molar non-triturative; accessory setal row well developed (4 or more setae); palp with strong apical seta. Maxilliped ischial endite (outer plate) not longer than palp article 1; palp article 3 apex rounded, palp dactylus absent. Gnathopod 1 simple; similar in size to gnathopod 2; carpus subequal in length to propodus. Gnathopod 2 weakly subchelate.
FIGURE 20. Tepidopleustes coffsiana sp. nov., holotype male, 3.5 mm AM P72060, Woolgoolga Reef.
Pleonites 1-3 each with carina; with weakly developed lateral ridging. Epimeron 3 posterior margin smooth, posteroventral comer rounded. Urosome 2 not dorsally occluded. Uropod 3 inner ramus slighlty longer than outer ramus. Telson weakly cleft «30%).
Etymology Named for Coffs Harbour, near the type locality.
Habitat Living on the brown alga Sargassum sp., 0-15 m.
Distribution New South Wales: South Solitary Island, Muttonbird Island, Korffs Islet, Solitary Islands; Woolgoolga reef.
42 © 2006 Magnolia Press HUGHES & LOWRY ZOOTAXA @)
FIGURE 21. Tepidopleustes coffsiana sp. nov., holotype male, 3.5 mm AM P72060, Wooigooiga Reef. Scales: G 1, G2, P3, P7 0.2 mm; all others 0.1 mm.
Remarks Tepidopleustes coffiiana sp. nov. is most similar to T. juliana (Lowry & Springthorpe, 2005). Both of these species differ from other species in the genus by having a smooth posterior margin on epimeron 3. Tepidopleustes co.fJsiana differs from T.juliana mainly in the weakly developed lateral ridges on pleonites I to 3 and the weakly cleft telson.
AMPHIPODA © 2006 Magnolia Press 43 ZOOTAXA Synopiidae Dana, 1852
Telsosynopia Karaman, 1986
J. L. Barnard (1972) discussed the genus Synopia and suggested that S. variabilis might warrant its own genus based on the entire telson. Karaman (1986) took the opportunity to erect the subgenus Synopia (Telsosynopia) following the work by Andres (1984) who described two additional species of Synopia, S. triangula and S. rotunda, with entire telsons. The subgenus Synopia (Synopia) was also established for the remaining Synopia species with cleft telsons. Karaman (1986) designated Synopia variabilis Spandl, ]923 as the type-species for S. (Telsosynopia), and defined the subgenus as "with characters ofthe genus Synopia except, telson entire and mandibular (sic) more or less triturative". This subgenus, however, went unnoticed by later workers discussing Synopia with entire telsons. Barnard & Thomas (1989) reassessed the molar of T. variablis (as S. variabilis) as triturative. Ortiz & Lalana (1997) described S. paravariabilis, a fourth species ofSynopia with an entire telson. Telsosynopia Karaman, 1986 is elevated to generic rank here, based on the entire telson. The mandibular molar "more or less triturative", originally used to help diagnose the subgenus Telsosynopia, is ineffective in separating Synopia from Telsosynopia because all species are now known to have a triturative molar.
Species composition Telsosynopia contains five species: T. paravariabilis (Ortiz & Lalana, 1997) new combination; T. rotunda (Andres, 1984); T. triangula (Andres, 1984); T. trifidilla sp. nov. and T. variabilis (Spandl, 1923).
Telsosynopia trifidilla sp. nov. (Figs 22-23)
Type material Holotype female, 5.5 mm, AM P71919, Northwest Solitary Island, Solitary Islands, New South Wales, 300 01.116'S 153°18.184'E. Paratype female, AM P71920, Northwest Solitary Island, Solitary Islands, New South Wales, 30°01 '07"S 153° 18' 11 "E, 60 g artificial substrate, 10 m, L.E. Hughes, 30 January 2003. Additional material examined. New South Wales: 3 specimens, AM P71921, Southwest Solitary Island, Solitary Islands, 300 09'48"S 153° 13 '48"E, airlift of mixed low turfing algae, 8 m, L.E. Hughes, 11 September 2003. 3 specimens, AM P71922, Muttonbird Island, Solitary Islands, 30 0 1TS 153°10'E, Sargassum sp., 6 m, L.E. Hughes, 21 May 2002. 1 specimen, AM P71923, Korffs Islet, Solitary Islands, 300 19'8"S 153°9' 12"E, airlift ofAmphiroa anceps and low turfing algae, 8 m, L.E. Hughes, 26 May
44 © 2006 Magnolia Press HUGHES & LOWRY 2003. 8 specimens, AM P71924, E side of Bare Island, Botany Bay, Sydney, 33°59.58'S ZOOTAXA 151 °13.84'E, 60 g plastic artificial substrate, 6 m, L.E. Hughes, 12 March 2004. I @ specimen, AM P70348, Gutters, N of Burrewarra Point, Batemans Bay, 35°49'49"S 150 0 14'02"E, from macroalga Stypopodiumflabelliforme, 24 m, G.D. Wilson. A.J. Millar, N. Vee, 27 October 2002, NSW stn 1992. I specimen, AM P70352, East Wall, N of Burrewarra Point, Batemans Bay, 35°50'01"S 150 0 14'IO"E, macroalga Corallina berteri, 25 m, G.D. Wilson. AJ. Millar, N. Vee, 25 October 2002, NSW stn 1987.
Description ofholotype Head protuberant. Eyes present; accessory eyes present, 2-5 ommatida. Antenna 1 length shorter than pereon, peduncle article 1 not elongate or bearing dorsodistal tooth; peduncle article 2 not elongate or bearing dorsodistal tooth; flagellum 9-20 articulate. Antenna 2 flagellum 28 articulate (or more). Mandibular palp present; molar triturative, columnar. Maxilliped foliaceaous. Gnathopod 1 simple; coxa not tapering distally; basis longer than carpus. Gnathopod 2 simple; dactylus vestigial tenninal setae short, less than or subequal to length of dactylus. Pereopod 4 coxa smaller than coxa 3, subtriangular. Pereopod 5 basis subovoid, expanded. Pereopod 7 basis subrectangular. Uropod 3 greatly exceeding length of uropods 1-2. Uropod 3 peduncle short. Telson entire; subequal or longer than uropod 3 peduncle; setae lining lateral margin, apical margin trifid.
Etymology Named from the trifid telson and articulate assistance provided by Helen Stoddart.
Habitat Living on the red algae Amphiroa anceps, Corallina berteri; and the brown algae Sargassum sp.; Stypopodium flabelliforme; 0-15 m.
Distribution New South Wales: Northwest Solitary, Southwest Solitary, Muttonbird Island, Korffs Islet, Solitary Islands; Bare Island, Botany Bay; Batemans Bay, Australia.
Remarks In material from Bare Island, Botany Bay, male specimens had an elongate antenna 1 flagellum with more than 20 articles, while the female antenna 1 flagellum was consistently 9-articulate from all locations reported. This sexually dimorphic character has previously been recorded for one other species in the genus, T. variabilis, by J.L. Barnard (1965: fig. 9b).
AMPHIPODA © 2006 Magnolia Press 45 ZOOTAXA
//1\ \AI )1/\ ~U2 ~ I \ -- ?
FIGURE 22. Telsosynopia trijidilla sp. nov., holotype female, 5.5 mm, AM P71919, Northwest Solitary Islands, Solitary Islands. Scales: Hd 0.5 mm; all others 0.1 mm.
Within Telsosynopia, T. paravariabilis, T. trifidilla and T. variabilis have a trifid apical margin ofthe telson. Telsosynopia trifidilla differs from other species in having gnathopod 1 basis longer than carpus; the apical setae on the gnathopod 2 dactylus short, similar to the condition in T. variabilis; pereopods 3--4 with the merus and carpus longer and more
46 © 2006 Magnolia Press HUGHES & LOWRY slender than those of T. paravariabilis and T. variabilis; coxa of pereopod 4 is a distinct ZOOTAXA subtriangular shape as in T. triangula and the pereopod 5 basis is expanded as in T. ~ triangula and T. variabilis.
Ur
FIGURE 23. Telsosynopia trifidilla sp. nov., holotype female, 5.5 mm, AM P71919, Northwest Solitary Islands, Solitary Islands. Scales: 0.5 mm.
AMPHIPODA © 2006 Magnolia Press 47 ZOOTAXA Telsosynopia trifidilla is only the fourth synopiid described from Australian waters, and the first of the genus Telsosynopia. The closely related genus Synopia is also represented by a single species in Australia, S. ultramarina Dana, 1853, recorded from Great Barrier Reef, Queensland (K.H. Barnard 1931). Telsosynopia trifidilla is a similar size to S. ultramarina, but easily separated by the cleft telson.
Acknowledgements
The first author was funded by an Australian Postgraduate Award Scholarship and internal postgraduate funding from the University of New England. Much of this research was conducted as part of a Ph.D candidature by LEH under the supervision of AssoclProf. S.D.A. Smith (University of New England /National Marine Science Centre) and Prof. R.D. Simpson (University of New England /National Marine Science Centre). We would like to thank two anonymous reviewers for constructive comments on the manuscripts, Dr Danielle DeFay (Museum national d'Histoire naturelle, Paris) for the loan ofP. pusilla, Dr Alan Millar (Royal Botanical Gardens, Sydney) for identifying the algal samples collected south of Sydney and Dr R. Peart for the pencil drawings ofTepidopleustes coffsiana.
References
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