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Use of Annual Density Banding to Estimate Longevity Eijiroh Nishi*1 and Moritaka Nishihira*2 Growth Bands of the Colonies Of

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Use of Annual Density Banding to Estimate Longevity Eijiroh Nishi*1 and Moritaka Nishihira*2 Growth Bands of the Colonies Of Fisheries Science 65(1) , 48-56 (1999) Use of Annual Density Banding to Estimate Longevity Eijiroh Nishi*1 and Moritaka Nishihira*2 *1Natural History Museum and Instit ute, Chiba, Aoba 955-2, Chuo, Chiba 260-8682, Japan *2Biological Institute , Graduate School of Science, Tohoku University, Sendai 980-8578, Japan (Received June 17, 1998) Growth bands of the colonies of the scleractinian coral Porites collected at Okinawan coral reefs were counted on Soft-X Ray micrographs, and age and growth rates of infaunal non-boring organisms were estimated indirectly from the annual coral-growth rings of the host coral. The organisms include a calcareous-tube polychaete Spirobranchus corniculatus (Grube), a sand-tube sabellariid polychaete, Idanthyrsus sp., a vermetid gastropod Dendropoma maxima Sowerby, a pecten bivalve Pedum spon- dyloideum Gmelin, and a coral barnacle Cantellius sp. The three tubicolous former species showed growth in various directions, and 15 to 20 years (S. corniculatus), 8 years (Idanthyrsus sp.), and 15 years (D. maxima) longevity were reliable estimates in this study. In contrast, the pecten bivalve and the coral barnacle showed lesser longevities, usually less than 7 years, and they grew vertically in almost the same direction as the host coral; thus, their growth rates can be estimated roughly by shell or slit-like hole length in relation to average host coral growth. We present here a new and practical method to esti mate age and longevity of non-boring coral infauna, and we discuss marine organism growth and lon gevity with emphasis on polychaetes. Key words: coral growth ring, infauna, age estimation , polychaete, Okinawa A major problem arising during the demographic analy coral skeleton overlying calcareous tubes. Soft X-radio- sis of marine organisms is the choice of parameters to be graphs of slabs cut along the growth axis of many massive measured and analyzed. These parameters must represent corals display alternating dark and light bands which out- a minimum variability for a given age. The best known line the former positions of the outer surface of the colony parameter for a demographic study is a set of size fre (Fig. 1). These dark and light bands represent variations in quency histograms. The growth parameters most often the density at which the skeleton was deposited . A pair of used are dry or fresh weight and body length or width. In bands-high density and low density (e.g., light plus dark tube building organisms, tube weight and size are the best bands)-represent one year's growth.7,14) parameter. The calcareous hard shells of molluscs and In the application of this method to coral infauna , there corals are known to show growth lines in the hard materi are 4 basic assumptions as follows: 1) organisms do not als depending on warm and cold seasons, tidal frequen bore into the coral skeleton , 2) they do not extend their cies, one-day activities, etc. (see Rhods and Lutz1)). These tubes or shells beyond the surface of the coral skeleton , 3) parameters must be manipulated with extreme care and a single individual occupies one tube, shell or hole , and it d with expert techniques. Examination of populations in the oes not move to other tubes , shells or holes, and 4) they field is a best way to study its growth, age and whole life are alive at the time of examination or only recently died. span, but in long-live organisms, this is hard to do. The in- For the last assumption, for example , tubes of Spirobran formation remaining on the hard structure of marine or chus and vermetids were used by crustaceans or small fish ganisms, particularly on soft-bodied worms such as poly after the worms' death.",") The life span of those tubes oc chaetes2-5) allows their life history data to be analyzed in a cupied by hole users could be estimated with age of an laboratory in some cases. original worm plus years (times) utilized by hole-users . Coral growth bands have been known to show annual Although the user of the empty tube or hole does not se. growth and the growth bands can be counted using Soft X- crete calcareous materials any more , the estimation of Rays,6-8) fluorescent-microscope,) and by laser densimet times used by the secondary space user is practically not al, ry.10) The information derived from coral growth bands ap- ways difficult. To know the age of the tube itself , not thf plies to many areas of oceanography, paleontology, worm, the tubes lacking original individual that had secret- etc. 8,11) ed the tube could be useful. Nishi and Nishihira12) demonstrated the method to esti mate the age of a tropical tubicolous polychaete Spirobran Materials and Methods chus corniculatus (described as S. giganteus, its taxonomy, see Nishi13) by counting the annual growth bands in the Over 40 massive coral colonies of Porites lutea with tube Corresponding address. Eijiroh Nishi, Natural History Museum and Institute, Chiba Aoba 955-2, Chuo , Chiba 260-8682, Japan. Tel: +81-43-222 8689 Fax: +81-43-266-2481 E-mail: [email protected] Age Estimation of Coral Infauna 49 Fig. 1. Sketch of images of coral growth bands and infaunas. Hp, Notaulax sp. (boring sabellid); Sc, Spirobranchus corniculatus; Is, Idanthyrsus sp.: Dm, Dendropoma maxima; Ps, Pedum spondiloidi um; Cs, Cantellius sp. worms, pecten shells, or coral-barnacles were collected at pears on the surface of the living tissues of coral (Smith"); Sesoko Island and Zampa Cape, Okinawa Island during Figs. 1 and 2A, B). 1994 to 1995. Only non-boring infauna species were stud Idanthyrsus sp. (Sabellariidae) dwells in packed sand- ied. After collection, corals were dried after rinsing with tubes and appear on dead coral colonies and on living mas synthetic detergent; thereafter, they were sliced into 2 to 5 sive colonies23) (Nishi and Kirtley, unpublished data: Figs. mm thick slices (maximum 10 mm thickness) at the por 1 and 2C, D). We studied only those individuals on living tion including the tubes or shells. The sliced plates were massive Porites colonies in the present study. then radiographed under Soft X-Rays (Softex, Hitachi 2) A vermetid gastropod Dendropoma maxima MB3). The exposure was 40 kVp, 3 mA, for 4 to 5 minutes Sowerby is a well known vermetid in coral reef area and (in a slice 3-4 mm thick). The source to subject distance frequently is found in dead and living corals in various was 50 cm. The areas of interest of the soft X-radiographs regions24,25)(Figs. 1 and 2E, F). Vermetids feed by a combi (generalized) is shown in Figs. I and 3 (partly drawn from nation of mucous nets and ctenidial cilia which collect photos of Nishi and Nishihira12)). detritus and other planktonic particles from the water.',) Usually we prepared one or two slices per worm, with a Dendropoma maxima is a solitary species and commonly maximum of 10 per worm, and the slice included the appears on massive coral species.") They are distributed whole or only a part of the tube (Figs. 1 and 3). If the slice on dead and living corals, but only those on living Porites included the whole tube or hole cut longitudinally as were studied. shown in Fig. 3A, the exact growth dates could be deter- 3) A scallop bivalve Bivalves living in scleractinian mined from the settlement to recent years. For pecten corals are also available as materials for age-estimation. bibalves and barnacles, we can cut the slice including its en- Pedum spondyloideum Gmelin attaches to living massive tire hole. If the slice included two openings of the same corals at a very early growth stage and subsequently tube as shown in Fig. 3B, the growth rate between the two becomes completely enclosed within the coral except for openings could be determined accurately. Some slices of its ventral region, which maintains an elliptical opening" tube of tubicolous species, however, contained only one (Figs. I and 2G, H). The growth rates of the coral and the opening clearly cut horizontally, so we could only com- scallop are precisely balanced, and the form of the scallop pare the size of recent tube openings on the coral surface is highly modified by its semi-enclosed existence to prevent with the opening that appeared in the slice. overgrowth by the coral, its upward growth results in an ex tremely elongate shell .21) Life Habits of Coral-Infauna Studied 4) A coral barnacle Some species of coral-associated 1) Tube worms (Polychaetous Annelida) Spirobran barnacle are buried in living colonies with a part of the chus corniculatus (Grube) is well known to be a coral in- shell apparent on the colony surface (Fig. 1 and Figs. 2K, habitant; it presently is in a taxonomically complicated L). Their growth is limited in vertical extent and their state. 13.17)Its life history and ecology have been studied by orifices usually appear on the surface. In Okinawan coral many researchers."-") The larvae settle on the exposed reefs, many species are known to live as coral coral skeleton near the living part of the coral and extend associates .28-30)Among the barnacles, Cantellius sp., which their tubes toward living tissue, then the tube is covered by was investigated in this study, is common in Porites colo the living coral tissues and the tube entrance (aperture) ap- nies (Nishi pers. obs.). The species has different forms in re- 50 Nishi and Nishihira Fig. 2, Photos of coral infauna. A-B, Spirobranchus corniculatus apertures (A) and tube (B); C-D, Idanthyrsus sp. aperture (C) and section of tube (D); E-F, Dendropoma maxi ma aperture (E) and internal plug (F); G-H, Pedum spondiloidium; J, K, L, Cantellius sp., aperture (J) and section of empty shell (K, L).
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