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Prevalence of the commensal barnacle Xenobalanus globicipitis on cetacean species in the eastern tropical Pacific Ocean, and a review of global occurrence Item Type article Authors Kane, Emily A.; Olson, Paula A.; Gerrodette, Tim; Fiedler, Paul C. Download date 24/09/2021 07:02:11 Link to Item http://hdl.handle.net/1834/25470 395 Abstract—Distribution and preva- Prevalence of the commensal barnacle lence of the phoretic barnacle Xenobal- anus on cetacean species are reported Xenobalanus globicipitis on cetacean species for 22 cetaceans in the eastern tropical Pacific Ocean (21 million km2). Four in the eastern tropical Pacific Ocean, cetacean species are newly reported and a review of global occurrence hosts for Xenobalanus: Bryde’s whale (Balaenoptera edeni), long-beaked common dolphin (Delphinus capen- Emily A. Kane (contact author)1, 2 sis), humpback whale (Megaptera Paula A. Olson 2 novaeangliae), and spinner dolphin 2 (Stenella longirostris). Sightings of Tim Gerrodette Xenobalanus in pelagic waters are Paul C. Fiedler2 reported for the first time, and con- Email address for E. A. Kane: [email protected] centrations were located within three productive zones: near the Baja Cali- 1 Southampton College fornia peninsula, the Costa Rica Dome 239 Montauk Highway and waters extending west along the Southampton, New York 11968 10°N Thermocline Ridge, and near 2 National Marine Fisheries Service, NOAA Peru and the Galapagos Archipelago. Southwest Fisheries Science Center Greatest prevalence was observed on 8604 La Jolla Shores Dr. blue whales (Balaenoptera musculus) La Jolla, California 92037 indicating that slow swim speeds are Present address for contact author (E. A. Kane): Texas A&M University at Galveston not necessary for effective barnacle 5007 Avenue U settlement. Overall, prevalence and Galveston, Texas 77551 prevalence per sighting were gener- ally lower than previously reported. The number of barnacles present on an individual whale was great- est for killer whales, indicating that Xenobalanus larvae may be patchily distributed. The broad geographic distribution and large number of Barnacles of the superfamily Coronu- its hermaphroditic reproduction may cetacean hosts, indicate an extremely loidea live as obligate commensals on be synchronized with that of its host cosmopolitan distribution. A better sea turtles, cetaceans, sirenians, sea (Dollfus, 1968; Fertl, 2002). A five- understanding of the biology of Xeno- snakes, and crustaceans (Newman to six-month reoccurrence cycle has balanus is needed before this species and Ross, 1976). The monotypic Xeno- been reported for Xenobalanus (Van can be used as a biological tag. balanus globicipitis Steenstrup, 1851 Waerebeek et al., 1993; Orams and (herein referred to by genus) is spe- Schuetze, 1998), which may indicate cialized for living as a commensal on that its life span may be of similar whales and dolphins (Darwin, 1854). length or that occurrence is correlated The typical six-plate balanomorph with seasonal environmental condi- shell is small and is imbedded into tions. Xenobalanus has been reported the skin of the cetacean host. The on 30 cetacean species worldwide and membrane supporting the operculum has a prevalence ranging from 0.5% to is greatly elongated, so that externally 55% of individuals in each sighting. Xenobalanus resembles a pedunculate However, intensity is highly variable, barnacle. This species is most com- and there are some reports of greater monly observed on the trailing edges than 100 barnacles on a single host of the dorsal fin, pectoral flippers, and (Aznar et al., 2005). tail fluke of the host, although it has We examined the presence of Xeno- been reported in areas such as the ros- balanus on cetaceans in the eastern trum and the area between the teeth tropical Pacific Ocean (ETP). Based (Samaras, 1989). Xenobalanus does on photographs taken during research not receive nutrition from its cetacean cruises from 1977 through 2003, host and therefore is not considered mean prevalence, mean intensity, and Manuscript submitted 28 January 2008. Manuscript accepted 17 June 2008. a parasite. Instead, as a suspension- geographic distribution are described Fish. Bull 106:395–404 (2008). feeding cirriped, it uses the water flow for Xenobalanus on 22 host cetacean around swimming cetaceans and ben- species. In addition, peer-reviewed lit- The views and opinions expressed efits from being transported by its host erature on this subject is examined, or implied in this article are those of the author and do not necessarily (phoresis). This species is highly spe- updating a previous summary of the reflect the position of the National cialized to live on cetaceans (Seilacher, cetacean hosts of this barnacle (Raja- Marine Fisheries Service, NOAA. 2005) and it has been suggested that guru and Shantha, 1992). 396 Fishery Bulletin 106(4) 30°N 20°N 10°N 0°N 10°S 20°S 160°W 150°W 140°W 130°W 120°W 110°W 100°W 90°W 80°W 70°W Figure 1 Xenobalanus presence or absence for 445 cetacean sightings in the eastern tropical Pacific Ocean (ETP) in 2003 as determined from analysis of identification photographs. Dots (●) indicate cetacean sightings with no Xenobalanus observed; circles ( ) indicate sightings with one or more barnacles observed; the solid line indicates the border of the ETP study area. Presence or absence is overlaid on a background of graded shading representing the volume of chlorophyll-a (mg/m3) averaged from September to November 2003. Materials and methods In external appearance, Xenobalanus may be confused with the parasitic copepod Pennella balaenoptera (Ev- Data collection ans, 1994) or the stalked barnacle Conchoderma virga- tum (Ruppert et al., 2004). However, the much larger P. Cetaceans were photographed during a 2-ship, 4-month balaenoptera usually occurs along the flanks, whereas research cruise in 2003, covering 26,000 km of tran- Xenobalanus is generally found along the trailing edges sects surveyed for marine mammals (boundaries shown of the dorsal fin, pectoral flippers, and the fluke, as has in Figs. 1 and 2). Camera equipment included Canon been described for stranded and live cetaceans. Similar EOS 10D and D60 digital cameras (Canon USA, Lake to P. balaenoptera, C. virgatum requires a less specific Success, NY) with 75–300 mm image-stabilized zoom position for attachment, requiring any hard substrate and 400-mm fixed lenses. Date, latitude and longitude, (such as another barnacle, a tooth, or exposed bone), and cetacean species (as identified by trained cruise person- C. virgatum is considerably lighter in coloration than nel), and unique sighting number per cetacean group Xenobalanus. Digital photographic quality was sufficient were recorded with each photograph. In the laboratory, for accurate identification of the commensal Xenobala- additional data were recorded upon examination of pho- nus; no specimens were obtained for direct examination. tographs, including the number of usable photographs in During the cruise, typically many photographs were the sighting (as described below), number of individual taken for each sighting. For our study, usable photo- cetaceans identified in the sighting, number of individu- graphs had 1) to be in focus, 2) to be of sufficient resolu- als infested with Xenobalanus, and number of Xenobala- tion to identify a barnacle if present, and 3) to include nus present. If barnacles were clumped in such a way at least one cetacean dorsal fin. For large schools, only as to compromise the accuracy of the count, the maxi- one photograph per school was used in order to prevent mum number of discernible barnacles was recorded. The recounting individuals. For small schools, only photo- resolution of the digital photographs was such that in graphs of animals identifiable as individuals, either most cases, individual barnacles were easily identified. from field notes or from unique markings or pigmen- Kane et al.: Prevalence of Xenobalanus globicipitis on cetacean species in the eastern tropical Pacific Ocean 397 180°W 150°W 120°W 90°W 60°W 30°W 0° 30°E 60°E 90°E 120°E 150°E 180°E 3 4 60°N 5 60°N 2 1 6 8 11 10 9 30°N 19 30°N 15 7 18 0° 12 17 0° eastern tropical 13 Pacific Ocean (ETP) 20 30°S 14 30°S 16 60°S 60°S 23 22 21 N 24 180°W 150°W 120°W 90°W 60°W 30°W 0° 30°E 60°E 90°E 120°E 150°E 180°E Figure 2 Documented sightings of Xenobalanus worldwide on various cetacean hosts compiled from lit- erature review, and an outline of the current study area (eastern tropical Pacific Ocean). Ovals indicate the geographic region where barnacles have been reported and their size does not indicate intensity of infestation. Refer to Table 3 for the corresponding citations for each region. tation, were used. Photographs used represent a sub- that was based on data provided by the SeaWiFS Project, sample of individuals and did not include every animal NASA/Goddard Space Flight Center, and GeoEye. observed. Killer whales (Orcinus orca) were studied Several statistical tests were performed to determine in additional detail, by using photographs taken on significant differences among the data. A nonparamet- previous cruises dating back to 1977, and including ric Mann Whitney-U test was used to determine if the photographs from the ETP in the California and Mexico rates of prevalence differed between Mysticetes and killer whale catalog (Black et al., 1997). Odontocetes and a nonparametric Kruskal-Wallace test was used to determine if the prevalence rates differed Data analysis significantly among species. Chi-square goodness-of-fit test was used to determine if the number of barnacles Two measures of prevalence of Xenobalanus were calcu- per killer whale followed a Poisson distribution. After lated for each cetacean species with at least three sight- normalization, linear regression was used to determine ings with usable photographs.