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The Relationship Between Depth, Exposure and Pectoral Fin Aspect Ratio

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The Relationship Between Depth, Exposure and Pectoral Fin Aspect Ratio Environmental Biology of Fishes (2005) 72: 33–43 Ó Springer 2005 Distribution and abundance of labrids in northeastern New Zealand: the relationship between depth, exposure and pectoral fin aspect ratio Christopher M. Denny Leigh Marine Laboratory, P.O. Box 349, Warkworth, New Zealand (e-mail: [email protected]) Received 28 August 2003 Accepted 3 March 2004 Key words: labridae, temperate, wrasse, fetch, spatial, reef fish Synopsis Physical factors influencing the distribution and abundance of seven common labrid fishes were examined over four rocky reef locations in northeastern New Zealand. Depth and exposure for each species (both within and among sexes) were related to pectoral fin aspect ratio. Each of the four locations (two mainland and two island) displayed distinct labrid assemblages, which were consistent over time, likely due to the influence of the East Auckland Current. There was a consistent depth-related trend for most species, regardless of location. Several species also showed a sex related depth difference. There was also a trend for some species to be associated with certain levels of wave exposure. For most species, the relationship between pectoral fin aspect ratio and the above physical variables was not as strongly evident in this temperate assemblage as has been previously found in tropical reef fish systems. Although some species did follow the predicted shifts in fin aspect ratio with depth and/or exposure, the observed trends were unre- lated to fin aspect ratio for many other species. These findings suggest that wave exposure may not be as important for labrids on northeastern New Zealand reefs as it may be in tropical coral reefs systems. The lower fin aspect ratios for New Zealand labrids, compared to tropical labrids, suggest that New Zealand labrids represent a subset of the total pectoral fin diversity in the Labridae. Consequently, the potential for distinct trends in fin aspect ratio and physical variables to be evident may be reduced. Introduction studies examining the spatial distribution and abundance of fishes on temperate reefs are rela- Understanding the processes which are important tively rare (Larson & DeMartini 1984, Choat in structuring reef fish assemblages is a central goal & Ayling 1987, Francis 1996, Brook 2002) with of reef fish ecologists. To determine these processes many studies investigating only a single species it is important to identify the distribution and (e.g. Lowry & Suthers 1998). Temperate fish abundance patterns of the relevant species over the assemblages have been shown to be influenced by spatial scales of interest. Previous studies have wave exposure (Thorman 1986), macroalgae cover shown that the abundance and distribution of (Ebeling & Laur 1985, Anderson 1994) or water coral reef fishes can be related to a variety of temperature (Stephens & Zerba 1981). In contrast habitat characteristics (see review in Jones & Syms to coral reef studies, workers have found no con- 1998). For instance, on coral reefs, reef fish sistent depth related trends (Bell et al. 1992, Cole abundance has been correlated with depth (Green et al. 1992). Furthermore, studies have failed to 1996, Richardson 1999), exposure (Russ 1984, find a strong correlation between reef fish density Williams 1991, Gust 2002) or live coral cover and habitat zones on coral (e.g. Green 1996, Gust (Bouchon-Navaro & Bouchon 1989). In contrast, 2002) and temperate reefs (Holbrook et al. 1993). 34 Recent studies have found correlations between waters in northern New Zealand (Francis &Evans pectoral fin morphology, swimming mode and 1993). The aim of this study was to examine if patterns of habitat use in labrid fishes (Bellwood depth and exposure influence the distribution and & Wainwright 2001, Fulton et al. 2001, Fulton & abundance of labrid fishes in northeastern New Bellwood 2002, 2004). Labrids rely on their pec- Zealand and the relationship between swimming toral fins during sustained swimming and generally ability (as examined through pectoral fin mor- use two thrust-producing mechanisms: drag-based phology) and these environmental variables. Po- rowing and lift-based flapping. Whilst the drag- tential differences between sexes in response to based mechanism provides good acceleration and these variables were also explored. This paper thrust at low speeds, lift-based thrust is mechani- makes the assumption that, in labrids, the fin as- cally much more efficient at higher speeds (Vogel pect ratio correlates with performance (Drucker 1994, Walker & Westneat 2000). A tendency to & Jenson 1996, Wainwright et al. 2002, Walker & predominantly use either mechanism, and the Westneat 2002). However, this assumption may associated implications for their swimming abili- not always hold true. ties, is reflected in the pectoral fin morphology of each species. Rounded fins (low aspect-ratio) are generally indicative of drag-based swimming at Methods slow speeds, whereas tapered fins (high aspect-ra- tio) indicate a tendency towards lift-based swim- Study locations ming and generally higher sustained swimming speeds (Drucker & Jenson 1996, Walker & West- Three locations were surveyed biannually in spring neat 1997, Wainwright et al. 2002, Walker & and autumn between 1999 and 2002; two island Westneat 2002). Recent studies have utilised this locations, the Poor Knights and Mokohinau Is- functional relationship to link swimming abilities lands, and a coastal location, Cape Brett. An in the labrid fishes to their patterns of distribution additional survey was conducted at Mimiwhang- across several spatial scales. For example, Fulton ata in autumn 2002 to give a further coastal et al. (2001) and Bellwood & Wainwright (2001) location for spatial comparisons (see Denny & revealed that labrids on the Great Barrier Reef Babcock (2004) for location details). General (GBR) with high pectoral fin aspect ratios domi- descriptions of coastal and island habitats in nated exposed high-energy locations, whereas northern New Zealand are given in Choat & Schiel species with low pectoral fin aspect ratios were (1982). All locations are influenced, to various relatively rare or absent from such locations. Such extents, by the East Auckland Current (EAC). The studies have suggested that the swimming abilities EAC, derived from the subtropical East Austra- of species may be limiting their access to high-en- lian Current, flows offshore parallel to the ergy locations, and may be a significant factor Northland coast and brings water that is clearer, influencing their habitat use and biogeography on and is often a few degrees warmer (2.5°C) than coral reefs, from regional (Bellwood & Wainwright nearby coastal areas (Stanton et al. 1997). 2001) to global scales (Bellwood et al. 2002). This work is an extension of a study that Study species investigated temporal changes in fish populations in response to the establishment of a no-take Labrids were selected as the study species in this marine reserve in northeastern New Zealand study as they are one of the most abundant and (Denny et al. 2003). During that study, fish widespread families of reef fish in New Zealand, assemblages were found to vary between locations, with 25 species recorded. Labrids also show little a result consistent with previous workers in this response to the presence of SCUBA divers which region (Choat & Ayling 1987, Brook 2002). These makes visual census techniques a reliable measure differences have been attributed mainly to broad- of their distribution, abundance and population scale hydrological patterns, particularly the East density (Samoilys & Carlos 2000). The seven spe- Auckland Current, that facilitates the dispersal of cies selected for study are the most abundant larvae from subtropical regions to the temperate labrid species in northern New Zealand; these were 35 Bodianus unimaculatus Gunther 1862 (pigfish), does not take into account differences in wave Coris sandageri Hector 1884 (sandagers wrasse), spectra among areas (Denny 1988) but does permit Notolabrus celidotus Bloch & Schneider 1801 tentative conclusions to be made (Thomas 1986). (spotty), Notolabrus fucicola Richardson 1840 The use of fetch as a proxy for exposure in (banded wrasse), Pseudolabrus luculentus Rich- northeastern New Zealand appears suitable as the ardson 1848 (orange wrasse), Pseudolabrus miles longest fetch and largest swells are from the Schneider & Forster 1801 (scarlet wrasse) and northeast/east. Suezichthys aylingi Russell 1985 (crimson cleaner- The following categories of wave exposure were fish). Labrids are usually sexually dichromatic, defined: Very Protected (VP, fetch < 300 km), each sex having a distinctive colour phase. The Protected (P, 301 and 1000 km), Semi-protected relationship between colour phase and sex was (SP, 1001–1500 km), Semi-exposed (SE, 1501– confirmed for the above species by histological 2500 km) and Exposed (E, fetch > 2501 km). examinations, except for N. fucicola where no The average wave height and direction at all relationship exists between colour phase and sex locations was estimated from data obtained from (Denny & Schiel 2002). the DATAWELL wave-rider buoy moored on the northern side of the Mokohinau Islands. The data Census technique for each record were obtained by taking the average of a 20-min record of the sea surface The distribution and abundance of labrids were height, which is transmitted to software running visually censused within 25 m long  5 m wide on a computer connected to the radio receiver at (125 m2) transects following Denny & Babcock the Leigh Marine Laboratory. (2004). Between 19 and 23 sites were surveyed at each location (except at Mimiwhangata where 16 Fin aspect ratio sites were surveyed), with nine transects being completed at each site. Each transect was con- The study species were captured at various sites ducted by first attaching a tape measure to the around northeastern New Zealand by handspear; substratum, then an observer swam for 25 m whilst B. unimaculatus (n ¼ 40), C. sandageri (n ¼ 23), N. counting all of the labrid individuals encountered celidotus (n ¼ 6), N. fucicola,(n ¼ 6), P. luculentus within 2.5 m either side of the transect line, and (n ¼ 9), P. miles (n ¼ 10) and S. aylingi (n ¼ 1).
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