Taxonomy
A total of 563 species is listed from deep water (>50 m) from the New Zealand Exclusive Economic Zone (EEZ). The 347 most common or distinctive species are fully illustrated and their diagnostic features outlined. Two new species are described: Ammoscalaria georgescotti n. sp. and Spiroplectammina carteri n. sp. When used in combination with our twin publication on New Zealand’s shallow-water benthic foraminifera (Hayward et al. 1999), these two bulletins provide descriptive data and illustrations of the 504 most common and distinctive benthic foraminifera living in New Zealand marine and brackish environments.
Ecologic distribution of deep-water foraminifera
We use census counts (59,000 specimens) of 424 species in 264 samples to map the distribution of deep-sea (50-5000 m depth) benthic foraminifera around New Zealand. Using Q mode cluster analysis (chord similarity coefficient) of the full census data we identify 7 high-level ecologic associations (A-G), 6 of which can be further subdivided into 32 subassociations. The deepest association (A), dominated by Nuttallides umbonifera and Globocassidulina subglobosa occurs at mid-lower abyssal depths (>3500 m) east of New Zealand. The next deepest, C (Epistominella exigua - Alabaminella weddellensis), is widespread at lower bathyal – abyssal depths (>1200 m) off both sides of central and northern New Zealand, but does not extend into the subantarctic zone. Three bathyal associations are recognised, with D and E occurring right around New Zealand and B restricted to the subantarctic. Association D (Cassidulina carinata – A. weddellensis) is usually in deeper, less current-swept waters than E (Globocassidulina canalisuturata – Bolivina robusta). Association B (Trifarina angulosa – Ehrenbergina glabra) also occurs in deeper parts of the bathyal zone (900-2000 m) along the strong current-swept, south-eastern margin of the Campbell and Bounty Plateaux, beneath the Subantarctic Front. Associations F (Cassidulina carinata – Trifarina angulosa) and G (Cassidulina carinata – Bulimina marginata f. marginata) mostly occur at mid shelf-upper bathyal depths (50-600 m) around southern and northern New Zealand respectively, with the dividing line approximating the position of the Subtropical Front.
Q mode cluster analysis (Jaccard coefficient) of presence/absence data of the 424 species in the 264 samples allows the recognition of nine high-level associations (a-i), four of which are further subdivided into 13 subassociations. The distribution of these associations is similar, but not identical, to that based on the quantitative data, indicating that total species composition is just as important as the relative abundance of dominant species in defining these regions.
R-mode cluster analysis (Pearson product-moment correlation coefficient) of full census data for the 56 most common species allows the recognition of eight groups, whose distribution is most strongly linked to bathymetry and secondly to geography. Two abyssal-lower bathyal associations (dominated by Globocassidulina subglobosa and Trifarina angulosa) are most abundant off southern New Zealand; one from lower bathyal-abyssal depths (Epistominella exigua) occurs dominantly east of northern and central New Zealand; an outer shelf to mid bathyal association (Cassidulina carinata) is more common to the east and south than the west; and another from mid-lower bathyal depths (Uvigerina peregrina) dominantly occurs off the west coast of New Zealand. The remaining three are common all around New Zealand.
Canonical correspondence analysis was used to relate sample associations to a set of environmental "drivers" using the full census data. Bathyal and abyssal associations appear to be more strongly influenced by depth-related variables (e.g., bottom temperature, salinity, oxygen, carbon flux) and shallower associations by latitude-related differences in surface-water characteristics (e.g., temperature, surface phosphate, chlorophyll-a). Environmental variables that influence faunal patterns at abyssal and lower bathyal depths (Assocs. A-C) appear to be, in decreasing order: bottom current strength (mud percentage proxy), carbonate corrosiveness (fragmentation index and planktic % proxies), quality, quantity and seasonality of organic carbon flux to the sea floor (surface phosphate, sea surface temperature, spring and summer chlorophyll-a proxies), and possibly properties of bottom water masses (salinity proxy). Faunal patterns within bathyal associations (D, E) are most strongly influenced by organic carbon flux (surface phosphate proxy), bottom oxygen concentrations (bottom water measurements) and bottom current strength (mud percentage proxy). Latitude-related variables driving mid shelf-upper bathyal faunal patterns (Assocs. F, G) include water temperature and primary productivity in the overlying waters (chlorophyll-a and organic carbon flux proxies).
Species diversity
Based on calculations for individual samples, the species diversity (α, H) of New Zealand benthic foraminiferal faunas overall and at all depths down to mid abyssal (<3000 m) decreases from lower to higher latitudes. At lower abyssal depths (>3000 m) faunas from all regions have a similar range of species diversity. There are no consistent diversity or evenness trends related to depth. Faunal evenness (E) decreases from north to south around New Zealand with more dominance of opportunistic species in the south where nutrients and food supply are more seasonally pulsed.
Based on SHE analysis for biofacies identification (SHEBI), 16 communities were identified in the deep sea around New Zealand. The communities exhibit a north to south latitudinal trend with lnS, H and lnE decreasing to the south. In the north and in the south the communities show an increase in lnS and H with depth. There is no trend with depth in the east and west areas. The community structure in each community was compared using the log series as a null model. Each area exhibits a unique pattern of community structure. In the north-east, 4 communities are recognised and only one at mid-bathyal depths does not resemble a log series. In the west, 3 communities are identified and only the outer shelf does not resemble a log series. The east is very different, where 4 out of 5 communities do not resemble a log series with only the community at abyssal depths doing so. In the south, all 4 of the recognised communities resemble a log series. All abyssal communities resemble a log series and 5 of 6 communities deeper than 1300 m resemble a log series. The log series is characterised by a constant H and is interpreted as representing community stability.
Frequency of species occurrence
We found that the pattern of species occurrence in deep water and throughout New Zealand localities approximates a typical log series plot. As a consequence the vast majority of species occur rarely (37% of species occur in <2% of localities) and only a small number occur widely (4% occur in >50% of localities). Buliminid and rotaliid species and those that dominantly live in shallow water have the highest frequency of occurrence and carterinid, astrorhizid, lituolid, trochamminid and robertinid species the lowest occurrence frequency.
Species duration
We used the recorded regional stratigraphic ranges of 642 modern New Zealand species (both deep, shallow and brackish) to investigate species duration patterns. The percentage of extant species in each of the following orders having a New Zealand fossil record are: Carterinida 0%, Trochamminida 0%, Lituolida 5%, Astrorhizida 10%, Robertinida 20%, Spirillinida 22%, Textulariida 30%, Miliolida 36%, Lagenida 45%, Rotaliida 53%, and Buliminida 59%. Foraminifera that live dominantly in normal marine salinity, shallow (< 100 m) and deep (> 100 m) water have a similar proportion of species recorded fossil from New Zealand (38-42%), with a much lower proportion from brackish environments (11%), reflecting the poor fossil record from brackish settings.
Of the 249 extant species with recorded fossil ranges in New Zealand: 3% first appeared in the Cretaceous, 1% in Paleocene, 10% in Eocene, 16% in Oligocene, 47% in Miocene, 14% in Pliocene and 9% in Pleistocene. These species have a mean partial species duration of 20 million years, comparable with a mean of 21 myrs for benthic foraminifera from the Atlantic margin of north America. There is no major difference in the timings of first appearances nor mean partial species durations between deep- and shallow-water-dwelling species.
Eighty-one percent of commonly occurring species (in >25% of localities) have a fossil record (mean species duration 21 myrs) compared with 14% of rarely occurring species (mean species duration 24 myrs). Sixty percent of endemic species (mean species duration 13 myrs) have a New Zealand fossil record compared with 43% of cosmopolitan species (mean species duration in NZ of 24 myrs) – the reverse of North American Atlantic coast data. This indicates that endemic species have been more common in New Zealand waters than in the North Atlantic, possibly a reflection of New Zealand’s isolation.
Biogeography
Sixty-four percent of the 642 extant New Zealand benthic foraminiferal species (both deep, shallow and brackish) have a cosmopolitan distribution, compared with 9% (52 spp) endemic to New Zealand and a further 8% to each of the South-west Pacific (including Australia), West Pacific and Pacific regions. A slightly greater proportion of deep-water (>100 m) species (69%) have a cosmopolitan distribution than do shallower-water (<100 m) species (55%), with brackish species (92%) having the highest proportion. Just 3% of deep-water species (Sigmoilopsis finlayi, Siphonaperta crassa, Spiroloculina novozelandica, Ruakituria pseudorobusta, Jullienella zealandica) are endemic to New Zealand, but 16% of shallow-water species are.
Our analyses of species presence/absence data suggest that the benthic foraminiferal biogeography around New Zealand differs at different depths and in different water masses. Five provinces can be recognised in our shallowest faunas (inner-mid shelf) and these correspond well with those identified from molluscs. At mid-outer shelf and upper bathyal depths, only 2 provinces can be identified. With increasing depth, greater subdivision is again possible, with 3 provinces recognisable at mid-lower bathyal depths and 4 at abyssal depths. Twenty-four percent (mostly common species) of all 424 species in our quantitative deep-water data set occur in all four regions - north, west, east and south of New Zealand - 17% are restricted to the north, 16% to the east, 8% to the south and just 4% to the west. The main faunal differences between regions are in the numerous rarely occurring species.
Paleoenvironmental assessment
There are many environmental drivers of the modern ecologic distribution of foraminifera and these vary from place to place. In this and previous shallow-water studies of modern foraminifera we have been able to correlate the strength of some of these environmental variables with the relative abundance of various taxa or associations. These correlations can be used to provide estimates of the paleoenvironments in which fossil foraminiferal faunas accumulated that are of value to geological, paleoclimatic and paleoceanographic studies. This uniformitarian approach is most applicable to Quaternary and Neogene faunas but far less reliable further back in time in the Paleogene and Cretaceous.
Planktic foraminiferal percentages and the relative abundance of different planktic species (census counts) can be used to estimate oceanicity, paleo-sea surface temperature and to give an indication of water depth. The composition of benthic foraminiferal faunas by order may provide a general indication of the past environment, but the relative abundance of benthic genera or species and the recognition of faunal associations allow more detailed environmental assessments. In the deep sea these are predominantly of water depth, seasonal or sustained carbon flux, strong bottom currents, bottom oxygen concentrations or exposure to carbonate corrosive bottom waters. In shallow or brackish environments these are predominantly water depth, tidal elevation, salinity, water temperature and exposure to water turbulence. Although depth is not a driver of foraminiferal distribution, a number of environmental variables show general trends with respect to depth, which allow depth estimates to be extracted from faunal composition data. Charts and tables summarising the depth distribution around New Zealand of a number of genera and species are provided to assist in paleodepth assessments. At bathyal depths we have identified c.60 benthic genera and species that appear to have distinct upper depth limits to their distribution and these provide an additional method to help refine paleodepth estimates of Neogene deep-sea faunas.
A method for rapid paleoenvironmental assessments of fossil New Zealand Neogene faunas is outlined, based on a quick estimate of planktic foraminiferal percentage, benthic foraminiferal composition and identification of dominant benthic taxa.