Trophic Relationships of Goatfishes (Family Mullidae) in the Northwestern Bawaiian Islands
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TROPHIC RELATIONSHIPS OF GOATFISHES (FAMILY MULLIDAE) IN THE NORTHWESTERN BAWAIIAN ISLANDS !THESIS SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFULLMENT OF THE REQU.IREl'!EN'fS FOR THE DEGREE O.P MASTER OF SCIENCE IN ZOOLOGY MAY 1982 by Carol T. Sorde.n Thesis committee: JulieB.. Brock, Chairman Ernst S. Reese John S. Stimson - i - We certify that we have read this thesis and that in our opinion it is satisfactory in scope and quality as a thesis for the degree of Master of Science in Zoology. Thesis committee Chairman - ii - lCKBOWLBDGEHEli"lS 'fhis thesis would not have been possible without ·the help of Stan Jazwinski and Alan Tomita wbo collected the samples a·t Midway, and 'fom Mirenda who identif.ied tbemolluscs. Many thanks to all my 'ft:iends in Hawaii and Alaska for all theit:: support, especially Stan Blum and Regie Kawamoto. "I am grateful to the members o.f my committee for encouragement and guidance, particularly my chairman, Dr. J. H. Brock, who gave ccntinued mot::al as well as academic suppot::t. Thanks also to Dr. J • .B. Randall fot: help with the taxonomy of l'Iulloide§, and Dr .E. A. Kay for help wi·th mollusc problems. This thesis is the result of research (Project No. NI/R-tl) supported in part by the university of Hawaii Sea Grant College Program under Institutional Grant Numbe.rs N1 79 11-D-00085 and N1 811A-D-00070, NOAA Office of Sea Grant, Department of Commerce. Further information on tbe original data may be obtai ned from the Hawaii Cooperative Fishery Research Unit, U.niversity of Hawaii. Last, but not least, I would like to thank my children, Renee, Geoff and Rachelle, who have been through a lot to give their mother an education. - iii - CONTENTS ACKNOWLEDGEMENTS . .. .. ,. ... ,. .. ....... iii LIST OF TABLES .. .. .. ,. .. .. '. .,. v LIST OF FIGURES • .. .. .. .. ". .. .. .. .. .. .. .. vi I. .INTBO.DlJCTION • • .. .. 1 II. MATEllIALS AND !!ETHODS .. .. '4 • • ,. .. .. .. .. 7 SUBSTRATUM SAMPLING PROCEDDIE • . .. .. 13 SAMPLE SITES .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. • • 14 III. RESULTS .. .. .. .. .. .. .. .. .. .. .. .. 18 Parupensus porpbyreus ....... .. .. • .. .. .. .. • 18 Mulloides vanicolensis .. .. .. .. .. .. .. .. .. .. • .. .. 19 Mulloides flavolineatus .. .. .. .. .. .. .. .. ...... • 21 Parupeneus multifasciatus ............ .. • .. 26 Parupeneus pleurostigma .. .. .. .. .. .. .. .. .. .. • .. 28 Overlap and Preference .. .. .. 31 Benthic samples .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 40 crustacea _ .. .. .. .. .. .. .. • .. 41 Other animal groups • .. .. .. .. ... .. .. .. .. .. 42 Polychaetes .. .. .. .. .. .. .. ... .. .. .. .. .. .. .. .. 42 Diversity and Similarity analysis .. • 45 IV. DISCUSSION • • .. • .. .. .. .. .. .. .. • .. • .. • .. .. .. .. 51 V. CONCLUSIONS .. .. .. • .. .. .. • .. • .. • .. • .. .. .. • .. .. 61 A~dix .£a.9~ A. SPECIES IN BENTHIC SAMPLES • .. .. .. • • .. • .. .. .. .. 65 BIBLIOGRAPHY .. • .. .. • .. .. .. .. .. .. .. • .. .. • .. .. .. .. .. .. 81 vii LIST OF TABLES 1. lumbers of each species of goatfisb collected .. .. .. 10 2. Ranks of food items of Parupeneus porpbyreus a .. .. .. 18 3. Banks 0.£ food items of nulloides vanicolensis " .. .. 20 4 .. Ranks of food items of Mulloides f la '10 Ii neat us · 22 5. Polychaete species in the diet of Mulloides flavolineatus .. .. .. .. " • .. .. .. • .. .. .. .. · . ,. .. 24 6.. Ranks of food items of Parupeneus multifasciatus .... 21 7. Banks of food items of Parupeneus pleurostigma • .. .. 30 8. Dietary overlap between goatfisb species at Midway .. 38 9. Characteristics of sand samples at Midway · '. • 41 10. Molluscs in benthic samples Station 1 · .. .. · .. .. .. 66 11. Molluscs in bentbic samples Station 10 .. .. .. .. .. • .. 68 12. Molluscs in bentbic samples station 14 .. • .. .. .. .. .. 69 13. Molluscs in bentbic samples Station 16 .. " .. • .. .. • 71 14. Molluscs in benthic samples station 21 .. .. 73 · .. .. " • 15. Animals in benthic samples Station 1 .. .. .. • .. .. • .. 75 16. Animals in bentbic samples Station 10. • .. .. " " .. .. 77 17. Animals in benthic samples Station 14 .. • • .. .. .. • 78 ··18.;- .- An-imalsinbentbic ·samples Station 16 •• · .. • • 19. Animals in benthic samples Sta tion 21 · . ., .. • 80 - viii - LIST OF FIGURES 1. location of sample sites at Midway Islands .. 2. Polychaete species in sand and guts Station 14 .. ;0 ;0 33 3. Polychaete species in sand and guts station 21 .. .. .. 36 4. Similarity analysis with qualit ati va data .. ,. .. .. • • 46 5 .. Similarity analysis with quanti tati va data " .. • • .. 49 - ix - I IITBODUCTIOII Sa:nd and rubble substrata associated with coral reefs may comprise a large percentage of the surface area of the reef (Thomassin 1918). The infauna of these areas are important food of reef dwellin<j ·fishesand invertebrates (Thomassin 1914) and in the trapp.ing and disturbance of sediments (Bailey-Brock 1(19). Analys.is of coral ree.f soft sediment communities has concent.rated on certaintaltonomic groups e.g- polychaetes (Bailey-Brock 1916), molluscs (Salvat 1961, 1910) and crustaceans (Thomassin 1914) •. Studies in the Indo-Pacific region describing community composition of coral reef sediments include Banner and Randall (1952), Salvat and Renaud-PIornant (1969), Gibbs (1978) and Edmondson (1946). The sand and coral infaunal communities of the atolls of the Northwestern Hawaiian Islands (NiHI) are not well known. Preliminary and incomplete data were obtained ·for the sessile benthos from a number of islands in the NiHI chain (Boucher pers. comm.), and the coral cryptofauna of French Frigate Shoals (Sorden 1980). Apart from some studies of micro-molluscs (Kay pers. comm.) the soft sediment - 1 - 2 communi·ty composition bas not. .been described from a.ny of the liRT. The inve.rtebrat.e infauna of soft sediment substrata provide a food resource for many species of coral reef fishes. Among these are a number of species of goatfishes (family Plullidae),whicb possess a pair of sensory barbels to aid in the location of prey amongst the sand. Goatfish barbels, located at the tip of the lower jaw, co.ntain abundant t.astebuds and are enervated by a la.rge branch of the facial nerve (Holland 1976, Suyebiro 1942) •... The barbels are used to detect potential prey in the sand, which is then dis·turbed by 'blowing' (Holland pers. comm. ), or digging with the pectoral fins (pe.rs. obs.). Holland (1976) found that the barbels were essential for the location of food by !~rlU!ene!1SlPorR.h.!.f~!!§. Taste buds a.re also abundant in sensory papillae in the buccal cavity. The mouth is protrusible, with small weak teeth; goatfish lack the crushing dentition found in some other fish families, e .. g_ .. Labridae (A1=Hussaini 1946,1947). Goatfish are often accompanied wbenfeeding by othe.r fish species that prey on the dist.urbed infauna (Robson 1968,1974). The family Plullidae is dist.ributed worldwide in t.ropical waters. All species are marine and carnivorous (Fowler 1933). Ele ven species belonging t.o t.hree genera are represented in the Hawaiian Islands (Gosline and Brock 1960, 3 Randall 1980). In the high Hawaiian Islands there is a commercial and subsistence fishery for goatfish, all species of which are highly esteemed as food. The early Hawaiians used species of Mulloides1 and EarJH!!neus both as food and as offerings to the gods (Titcomb 1912)., Goatfish are also a prey item for pisci vorous fish such as ulna (Ca.rangidae), and are an important diet item of seabirds, at least in the NIlHI (Harrison and Hida 1980). The diet of mnllid species has been studied int.he west Indies (Randall 1961), Puerto Rico (Bauer 1981), India (Thomas 1969) , Madagasca.r (Harmelin-Viv.ien 1919), and Japan (Suyehiro 1942). Pacific studies include Hiatt and strasburg (1960) at Enewetak,Marshall Islands, and Hobson (1914) off Kona, Hawaii. Mahi (1969) did a comprehensive study of the diet of RAIU!!1!~'§l!2!W!.I!!.§both juveniles a.nd adults. Five species of goatfish were included in the large trophic study of fishes of the Kana coast, Hawaii (Hobson 1974). Somewhat incomplete data are available on the diet of lyl!.oides!lSlvol!pefl\u§" from the NYHI (Okamoto and Kawamoto 1980) and ji. " flavolin!.~.1B§, ji.!anicol~n§i§, from Oahu (Kluegel 1921)., There are no data on the diet of fSlrUjeneJ1..e eleures!ig.!A from any location, er of t Dr. Philip Heemstra has found tha·tthere was no need for the establishment of the genus Mulloidicht~I~and that Mulloides·· is the valid genus name (through J •.' E. Randall pars.cemm.) 4 ..f. !U!l ti fasciatlls from NiHI. The mullids include both diu.rnal e.9-. Parup!neu.§ .. !ultil.a~c!atus,. and nocturnal or crepuscular feeders e.g. ,aull.9iges v ani~ol§nsi§, .fa~UBeJ1e!,}s a9.'l!h!!i~Um (Ho bson 1974,. Kahi 1969, Rarmelin-Vivien 1979) •.. The major food items consumed by most species are crabs, shrimps,. amphipods and other small crustaceans,. polychaetes, and bivalve and gastropods molluscs. Species which eat fish include fiu:uj!eneJ!§ luteys (Barmelin-Vivien 1(79) and I. £b'!;Is§,ydr9~ (Hobson 197q). However, diet composition appears to vary with locale and size o.f fish. Within one spec.ies the important p.rey i temsmay differ between islands, or between sites on the same island. In the Marshall Islands fish were the dominant prey of 1. liavolineat.!1:3(Hiatt and Strasburg 1960)where.as at Kona, Hawaii this species ate mostly bivalve molluscs, polychaetes and amphipods {Hobson 19714). l!tenoEYssp. vas one of the major prey of .f. porphl:~'§ at Kaneohe, Oahu but