BULLETIN OF MARINE SCIENCE, 45(2): 316-337, ]989
COMPARISON OF MYODOCOPID OSTRACODES IN TWO ZONES OF THE BELIZE BARRIER REEF NEAR CARRIE BOW CAY WITH CHANGES IN DISTRIBUTION 1978-1981
Anne Constant Cohen
ABSTRACT The number of myodocopid species (51) in the vicinity of Carrie Bow Cay, Belize, is the highest yet reported for any geographic area of similar size. Replicate sampling in two different sand-bottomed zones, a shallow (1.5 m) lagoon site and a deeper (18-30 m) fore-reef site, from 1978-1981, shows that myodocopid species and families differed between the zones, and that species and family composition varied from year to year, possibly due to disturbance by and recovery from a hurricane. Myodocopids were less abundant, but species diversity greater and yearly changes much smaller in the deeper, less disturbed fore-reef site than in the shallower lagoon site which is subject to greater storm disturbance. Some changes in species and family abundance following a storm may be related to differences in life-history strategies.
With one exception (Kornicker, 1958), previous collections ofmyodocopids on coral reefs have consisted of descriptions of a few species and/or have been from single samples made in each of a relatively few locations (Poulsen, 1962; 1965; Hartmann, 1964; Kornicker and King, 1965; Hall, 1985). Kornicker (1958) dis- cussed the systematics and distribution of myodocopids (21 species) on the Ba- hama Bank near Bimini, an island in the northern part of the range of Caribbean coral reefs and with reefs less developed than on the barrier reef off Belize. My study is part of a larger research project begun in 1972 by the Smithsonian Institution at Carrie Bow Cay in the central province of the 100-mile-Iong coral barrier reef lying 10-12 miles off the coast of Belize (Riitz1er and Macintyre, 1982). The central province has the highest coral species abundance and sea level reef development on the barrier reef (Burke, 1982). In 1978 Belize was hit by Hurricane Greta, providing an opportunity to study the effects of a major dis- turbance on the stability and persistence of the myodocopid fauna. Field studies on abundance of two or more species in any community covering a number of generations are rare, but important (Connell and Sousa, 1983). "The finding of nonstable conditions on a local scale and/or over the course of a few turnovers has important ecological and evolutionary implications. Disturbances causing local nonstable or persistent conditions are frequently essential for the stability or persistence of species on larger scales" (Connell and Sousa, 1983). This study is the first survey of the myodocopid fauna on a well developed coral reef. It is also the first analysis of distribution of the myodocopid fauna in relation to reef zonation, temporal fluctuations, and life-history.
METHODS
Collection Site. -Specimens were collected from a variety of habitats in the vicinity of Carrie Bow Cay (16°48'N, 88°05'W) mostly from 1976 to 1981 (Fig. I). Samples used for numerical comparisons were taken during 1978-1981 in the following two reef zones near a fixed transect line crossing the barrier reef just north of Carrie Bow Cay: the sand and rubble zone of the lagoon (Figs. 2, 3A) and the sand trough of the outer fore-reef (Figs. 2, 3B). These two sites were chosen for three reasons: (I) the fixed reef transect line had been previously photographed, mapped and divided into units defined and described on the basis of dominant biological and geological features (Riitzler and Macintyre, 1982); (2) although the two zones differed in reef zonation and depth, each had broad sandy bottoms
316 COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 317
o km. Tobacco Range ~:>:..
0:.:
o Q) ...... <~:t?~;··· II]Shoals ~ Sand Bores & Patch Reefs Figure I. Map of bamer reef complex in the vicinity of Came Bow Cay; area of larger map located on inset by rectangle [From Riitzler and Macintyre, 1982; used with permission of Riitzler]. which could be easily sampled by the same method; and (3) in preliminary samples many myodocopid ostracodes were found in these two sites. The sand and rubble sampling zone is about 1.5 m deep, with a bottom of poorly sorted sediment ranging in size from silt to gravel and 1.0-1.4 m thick over a hard bottom (Riitzler and Macintyre, 1982). The area sampled in the sand trough is about 18-30 m deep with substrate that "is poorly sorted, silt size to very coarse sand sediment plain. This sediment is mainly very fine to fine sand, but coarser material consisting of Halimeda plates, mollusks, benthic foraminiferans, and echinoids is scattered throughout. Probings indicated that sediment" is "more than 12 m in the axis of the trough" (Riitzler and Macintyre, 1982). To investigate local ranges of myodocopids, additional nonquantitative samples were taken from (I) other zones along the fixed reef transect line; (2) the following areas in the vicinity of Came Bow Cay: lagoon and back-reef around Came Bow Cay (sand cay) and South Water Cay (mangrove cay), south shores and a channel through Twin Cays (mangroves), South Water Cut (channel through the 318 BULLETIN OF MARINE SCIENCE, VOL. 45, NO.2, 1989 !! £ ::l J i ,j ~ ~~l ! J § 1 Ho; 'l; ;c .. " ::;; .. " ;: }>." .. }>.'" " ! § " 'l; ~ j . ~ ).- .." '";>-" '" , • • I ~ ! J j v;- ~~ I ~ 0: , ~ & § >.~~ ~c c I~i!.i §~ ! ~ i j ~ ! j ~ i £ ~ m~ ~Ui~~ O:u ~ "N • ~ ;~ Q; j CD .• i ~ i. ~ 0:~ ~ '" " S ."<; u ! 1; Q; ., L,-, , ~ a:" ,~ '. ";' ..... ! ". -..;: ·lp.rI~·.'.'". ::...· [lJ~:::::: " '.' ' CJ ."o ...-,:: .... :..... CD «>! cl~! 0 0 3 ••jg o lr !! N COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 319 Figure 3. A. Photo of bottom in sand and rubble zone ofIagoon, depth 1.5 m. B. Photo of bottom in sand trough of outer fore-reef, depth 30 m, showing part of movable transect line. barrier reef), Curlew Bank (submerged cay), lagoon patch reefs and adjacent sea grass (in deep lagoon in barrier reef complex), Ragged Cay and adjacent bank (western or inner edge of barrier reef complex); and (3) on the mainland: subtidal shores of northern (sand beach) and just north of (mangroves and sea grass) Dandriga. Nonquantitative collections were made by the author in May 1976; April 1977; January 1978; June and October 1979; and June 1981. Other ostracodes collected at the study site and deposited in the National Museum of Natural History (USNM) were also used to complete data on indi vidual species ranges (listed in Cohen, 1987). Additional data on the local range of Skogsbergia lerneri were obtained from baited traps. Traps were left overnight (about 1700-0900) in the lagoon 320 BULLETIN OF MARINE SCIENCE, VOL. 45, NO.2, 1989 and back-reef(one in June 1979, two in June 1981), during the day (0900-1700) in the back reef (one in June, 1981), and for about 24 h (two in the lagoon and back-reef, one in the spur and groove in January 1978; one in the lagoon and one in the spur and groove in June 1979; one each in the lagoon, back-reef, spur and groove, sand trough and outer ridge in October, 1979; one in the back-reef in 1981). Sampling Methods. -Samples were made by diving, usually with SCUBA, except in very shallow water where collections were made while wading. In depths accessible by diving this method is superior to dredging, because the area sampled can be chosen and the sampling monitored visually. Coring was tried, but found to yield very few ostracodes and therefore not used. Quantitative samples in the two study areas were made by dragging a rectangular hand net (16 cm high, 22 cm wide, mesh opening 0.2 mm) along the surface of the sediment sequentially along both sides of a movable 3-m transect line. The net lightly scraped the surface sediment so that the net sampled the upper 1-2 em of sediment and 14-15 em of water above it thus sampling the strata occupied by ostracods observed undisturbed and disturbed in dishes of sediment and seawater. The net was immediately twisted to trap material at the distal end and enclosed in a plastic bag. Quantitative comparisons are based only on samples made with the movable transect, first used in 1978. Single quantitative samples were taken in each of the two zones in 1978. At least three quantitative samples were taken in each zone during each subsequent field trip (June 1979, October 1979 and June 1981), and three per zone used in analysis of those dates (see below). An aquarium net was used in 1978, but replaced in 1979 with a net of the same dimensions as the aquarium net, but having a deeper bag. Net samples were decanted as described in Cohen (1983; 1987). Additional nonquantitative ostracode samples were made in surface sediment, rubble and by traps, but these collections were used only for life-history studies and in determining range limits of species. Sampling of rubble and by traps is described in detail in Cohen (1983; 1987). Sorting, Preparation and Identification of Ostracodes. - The preserved samples were stirred vigorously, drawing the lightweight ostracodes to the surface and allowing them to be decanted. This procedure was repeated until no more ostracodes were found. The thoroughness of the method was checked by examining the remaining residue of two samples; not more than two to four additional ostracodes were found in these residues. Other residues were spot-checked from time to time to insure that all ostracodes were being extracted. Because one 1981 lagoon sample contained almost 2,000 myodocopid ostracodes, a plankton splitter was used to divide another 1981 lagoon sample in half. The number of ostracodes extracted from that half was doubled for comparison with other samples. From the three field trips 1979-1981, only three ofthe quantitative samples taken each time in the sand and rubble zone were fully sorted because of time constraints and large sample sizes. All of the sand trough transect samples (18 samples) were sorted, but only three quantitative samples from each date (1979-1981) were chosen for quantitative comparisons on the basis of matching collecting periods and thoroughness of sorting and identifying. Samples not used were those which had originally been live-sorted in Belize and the residue then discarded. Subsequent checking of preserved residues of other samples 1 had believed were thoroughly live-sorted showed that ostracodes remained in the residue. The abundance of juveniles collected throughout this study complicated species identification, but also presented an opportunity to investigate developmental aspects such as clutch and embryo size, instar number, juvenile taxonomic characters and, to some extent, development time and life history patterns. Juveniles representing all myodocopid families and many species were found in the samples. Live specimens were maintained and reared as described in Cohen (1983; 1987). Skogsbergia lerneri were reared from egg to adult; Rutiderma new species. A were reared from egg to second instar; partial molt series were obtained from several other species. Deposition of Specimens. - Material is deposited in the Los Angeles County Museum of Natural History (LACM). When taxonomic descriptions are published, holotypes and paratypes will be deposited in the LACM, and additional paratypes in the Smithsonian Institution (USNM). RESULTS Comparison of Two Ecological Zones.-LAGOON: SAND AND RUBBLE ZONE (l.5m depth). Twenty-nine species and 3,281 individual myodocopid ostracodes were taken from the 10 quantitative samples (Table 1). Two measures of species di- versity for the lagoon myodocopids were calculated using one pooled sample (10 samples combined): species richness, H' = 0.90 (Shannon-Weaver diversity index) and species evenness (equitability), E = 0.61. The most abundant species was Rutiderma dinochelatum Komicker, 1958 (1,045 specimens) (Fig. 4, Tables 1,2). COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 321 ..,. o o N ..• o o o + + +0 +0 + III +0 +0 +0 N~ N'" o •••N QC ••• o o o + + + + + + + + + + + 00 o r-- •.... IC o o o o o ..• •.... o '<1" ..,. ..• '<1" o o ..• o + + +0 +0 + '<1"+0 +0 +0 I.D•....•\0- +QC 'Or-- o '" _<'I o o -~ -M ..,. + + + + + + + + + .., o o o o .., <'I 0'> IC N N Q o o o o o N + •...... ,. + +<'I+IIl+N +0 +0 +<'1 +0 + +N r-- N 00 _ N ••• ..,.'"..• o o o Q + + + + + + + + + + + + + Q o V'o ..• ..• o o o o '<1" •••••• \0\0 V'oV'o 00 00 NN NN 00 -- -- 00 00 00 00 00 NN N Q ..,. 00 <:1'1 00 o ..• '<1" N o o QC + + + + + + + + +0 ...... 00""" 0000 '<1"00 ..,.<'1 ..,<:1'1 o Q •••00 CIO \0- .., _<'I o o N"" .••••• f'I") -("I') ...N + + + + + + + + + Q + o o o QC QC '" \0 Q..• ..,. III..• ..• o N Q d o o o + + N o o +Q +0 +0 +0 +0 ..,...,. + +.., +0 +0 Q o o o o Ill ••• ~~ 'O~ o o o ..• ", -N + + + + + + + + + + + + + + N <:> o o o o o.... o o ..• Q o o .., o o o o o Q +.., +N +0 +.., +\0 +.., +0 +0 +0 +0 +Q Q o o ICIIl 0'> \0 N o o o o Q <:I'IN <'I + + + + + + + + + .., <'I o + + o o o o Q Q \0 III QQ 00 00 00 00 IIIIII ..,..,..,'" 00 00 00 00 00 00 •••••• z Ii= - N N \c Q Q N o o o o -1-- +' .... ", t- o Q o + + +", +", +Q + +0 ", •....•+ Q N~ N N o o 1I'l~ 0 + + + + + + + + N Q •••.... o o I""- 0 II'l Q'I Q t- .... Q Q o ", '5" o o '" 0 o o + + + +Q + +0 +0 +0 +", +0 + Q'IQ'I + •....•\c'" \c'" +0 o o ••• 1""- Q N ••.•N o o o o 0 1I'l....•••• o + + + + + + + + + + + + + + + o o .... Q <:> <:> o o o o o o 0 ,.... o •....•...... 0000 QQ ••..••.. QQ •••••• NN -- 00 00 00 00 00 Q'IQ'I 00 '1""1'" ••• ,..... N •••...... o QC) QC) \0 o ...... Q Q o o o o + + +", +Q +Q QC)\c +0 N'" +0 +0 +0 o Q Q •... ", o _1'1 o o II'lQ'l o + + + + + + + + Q Q + \c o o o o c '" o" Q'I '" N .... :0" J> e .... '0 t- Q Q C o Q Q o O + o '0'" + + C -1-0- +", +0 ::::: ::::: +Q +Q +0 +0 +0 ~ ri~ o ...... Q Q o o o E 0- + + + + + + + + + + + Q'I Q'I + + o ...... Q Q o N o ' QC) t- 1'1 o o Q Q ••• 1'1 o o o o + + + +0 +0 +Q +Q ••••1I'l +N ,....\0 +0 +",+ +0 + o o Q Q \C ....•••• o ' '3 Q Ill'" :g=OS III - OJ -Ill_.- .-5 ~g, rfJ COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323 t- o o o o o o o o N "" O + o o +-- +0 +0 +0 +0 +0 +.,., +0 +0 +0 +0 +0 "00 o o o o ON +N +0 ~-"", o o o o o o N"" o o o + + + + + + + + + + + + + + o o o o o o o o o o o o o t'- o o o o o o '" o o + + + '" ",,'" +0 +0 +0 .,.,00 +0 +"" +0 +", +", +"" +0 o o o o o o o '" o .,., o +0 +0 +0 +0 +0 +0 +0 +", +0 +", +0 o o o o 00 - o o o o o + + + + + + + + + + + + o o o o o ..•t'- o o o o o 00 CI\CI\00 00 00 """" 00000000 OOOO--OONN-- o o o o o +0 +0 +0 +0 +0 o o o o o + + + + + + + + + + + o o o o o N o " '" 2" :D" .0 2 ..• 'I::l t'- " o o o 00 o o N o 0 'I::l" + " +-", +0 CO M +0 +0 +N +0 +0 +.,., + +0 + + o OM o o 0'> N o o o o '-0- 0 ~ ~~ ••• 1"4 E 0 + + + + + + + + + + + + 0 o ...•. o o o o N 0 o .,., 1"4 o 1"4 o o N o o o o o N o o o + + + + +00 +"" +0 +0 +0 +0 +0 + .,., +0 +0 +0 ~~ o o N o o o o o o o o + + + + + + + + + + + + + + + o ...•. o o + 00 o o o o o o o o o ..• '" .0 '";:l Z .5 ::.G_ E ."''"_ •...til0 o u U •..• '"0. •..•'" '" til ~~ '" 0. t:: ...'" 324 BULLETIN OF MARINE SCIENCE, VOL. 45, NO.2, 1989 Table 2. Comparison of most abundant species in 10 combined lagoon and 10 combined sand trough samples (1978-1981) Lagoon Sand trough Species (No.) (%) (No.) (%) S. lerneri 2 0.06 190 16.4 V. new species ST 0 0 95 8.2 V. new species SA 0 0 26 2.2 P. sex 2 0.06 36 3.1 P. new species N 791 24.5 0 0 P. muelleri 124 3.8 12 1.0 S. setisparsa 28 0.87 31 2.7 A. americana 3 0.09 34 2.9 A. monambon 30 0.93 8 0.69 H. paucichelatus 172 5.3 187 16.1 R. new species A 158 4.9 48 4.1 R. dinochelatum 1,045 32.4 25 2.2 R. hartmanni 88 2.7 I 0.09 R. new species B 104 3.2 21 1.8 E. new species KE 479 14.9 0 0 E. new species KG 112 3.5 143 12.3 E. new species R 19 0.59 9 0.77 E. new species P 22 0.68 21 1.8 E. new species B 0 0 141 12.1 E. new species MJ 0 0 31 2.7 E. new species MR 0 0 32 2.8 E. donabbotti 32 0.99 57 4.9 E. new species J 12 0.37 14 1.2 Total 3,223 100 1,162 100 Also abundant were Parasterope new species N (791 specimens) and Eusarsiella new species KE (479 specimens) (Fig. 4). Two species, R. dinochelatum and Parasterope new species N accounted for 56% of the myodocopids collected in that zone. The three most abundant species accounted for 71% of the myodocopids collected. The most abundant families in the sand and rubble zone were Ruti- dermatidae (5 species, 1,396 specimens) and Cylindroleberididae (8 species, 983 specimens, all but 14 of these belonging to the Cylindroleberidinae). The Philo- medidae were represented by only 172 specimens, all belonging to Harbansus paucichelatus (Kornicker, 1958). Species belonging to the Cypridinidae were very rare in the samples (two species, four specimens). Almost half of the specimens collected in the lagoon zone belong to species either not present (eight species) or represented by fewer than 10 specimens in the fore-reef samples. Two additional species, Rutiderma new species Land Eu- sarsiella new species MO were collected in nonquantitative sand samples from this zone (Table 3), but they were not found in the 10 samples used for numerical comparisons (Tables I, 2). The largest sample (973 myodocopids, June 1981) represents a density of about 737 . m-2 of surface sediment (973 divided by 0.22 . 6 m). OUTERFORE-REEF:SANDTROUGH(18-30 m depth). Thirty species and 1,188 individual myodocopid specimens were found in the 10 samples analyzed. Species diversity values were: H' = 1.16, E = 0.78. The most abundant species was Skogsbergia lerneri (Kornicker, 1958) (190 specimens) (Fig. 4, Tables I, 2). Also abundant were Harbansus paucichelatus (187), Eusarsiella new species KO (143), and E. new species B (141), which together with S. lerneri, accounted for 55% of COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325 1100 1100 1000 ~ 1000 900 ~ 900 800 MAJOR LAGOON SPECIES 8 800 MAJOR SAND TROUGH SPECIES Q. 700 CIl '" 700 600 ~ 0> 600 500 eJ ~ 500 400 ~ at ••00 300 ~ 300 200 ::;t 200 100 2.. 100 a a R.d. P.N. E.KE. S.I. H.p. E.KD. LB. V.ST. E.d. R.A. Figure 4. Comparison often combined samples in lagoon and ten combined samples in sand trough; histogram shows abundance of most common species in each zone (those species contributing 71- 72% of the total ostracodes collected in each zone). Species shown in lagoon histogram arc R. d. = Rutiderma dinochelatum. P. N. = Parasterope new species N, E. KE. = Eusarsiella new species KE. Species shown in sand trough histogram arc S. I. = Skogsbergia lerneri. H. p. = Harbansus pauci- che/allls. E. KG. = Eusarsiella new species KO, E. B. = E. new species B, V. ST. = Vargula n. sp. ST, E. d. = Eusarsiella donabbotti, R. A. = Rutiderma new species A. Bar key: black = Rutidermatidae, right diagonal = Cylindroleberididae, left diagonal = Sarsiellidae, cross-hatch = Cypridinidae, open = Philomedidae. the myodocopids collected in the samples. These four species along with three others, Vargula new species ST (95 specimens), Eusarsiella donabbotti new species (57), and Rutiderma new species A (48), accounted for 72% of the myodocopids collected in the analyzed samples. The five most abundant species belong to the families Cypridinidae (4 species, 347 specimens), Sarsiellidae (13 species, 459 specimens), and Philomedidae (3 species, 191 specimens), the families most abun- dant in the sand trough. Specimens belonging to the other two myodocopid fam- ilies were relatively rare: Rutidermatidae (6 species, 98 specimens) and Cylin- droleberididae (5 species, 93 specimens, with only 43 of these belonging to the Cylindroleberidinae). About half of the specimens collected belong to species either not present (10) or not represented by more than 10 specimens in the lagoon samples. Three additional species Vargula new species M, Parasterope new species E, and Ruti- derma new species K were found in nonquantitative samples from sand trough rubble (Table 3). Temporal Changes in Myodocopid Fauna.-LAGOON: SAND AND RUBBLE ZONE. There was no change in species representation (except that rarer species did not occur in all samples), but there was a shift in relative species, family, and total abundance in collections made from January 1978 to June 1981 (Table 1, Fig. 5). The Rutidermatidae was the most abundant family at the beginning (1978,263 specimens) and end (1981, 946 specimens) of the study, although the most abun- dant species in the family changed from Rutiderma new species A in January 1978, to R. dinochelatum in all subsequent samples. Rutiderma hartmanni and R. new species B increased in abundance following each sampling period. The largest sample was the single 1978 sample of 387 ostracodes. Ostracode abundance in single replicate samples declined in 1979 and increased in 1981. The single 1978 sample was larger than any of the six replicate samples collected in 1979 (Table 1), but smaller than each of the replicate samples of 1981. Few cylindroleberids were collected in 1978 (four species, five specimens in the single sample). The Cylindroleberididae (particularly Parasterope new species N, 184 specimens) was the most abundant family in the June 1979 samples (8 species, 326 BULLETIN OF MARINE SCIENCE, VOL. 45, NO.2, 1989 Table 3. List of myodocopid species collected only in nonquantitative samples from Belize (also listed: other published localities) Taxa Locality (and year) Cypridinidae Vargula new species M Belize only: sand trough (trap and rubble, 1979), lagoon patch reefs (1976), South Water Cut (1978), spur and groove, (1976), outer reef slope (1979) Cylindroleberididae Cylindroleberidinae Parasterope new species E Belize only: sand trough (rubble, 1978), plankton (1976, 5 males), reef crest (1978) Postasterope abaco Belize: plankton (1975, I male). Bahamas Postasterope new species B Belize only: sand and rubble, W Carrie Bow Cay (1978) Postasterope new species X Belize only: South Water Cut (1977), W side Carrie Bow Cay ( 1978) Bruuniella new species A Belize: back-reef (1974), inner slope of outer ridge (1976); spur and groove (1978). Bermuda Diaslerope new species D Belize only: W side Carrie Bow Cay (1978, A-I male) Asteropteroninae Asteropella maclaughlinae Belize: Carrie Bow Cay (1978), Blue Ground (1981). W Flori- da. Texas Actinoseta hummelincki Belize: back-reef (1974, 1976), spur and groove (1976, 1978), Twin Cays (1977), South Water Cut (1978). Florida. Vene- zuela. ?W Panama Rutidermatidae Alternochelata polychelata Belize: Avicennia mangroves (I male). Bahamas Rutiderma new species L Belize only: sand and rubble zone (1979), lagoon plankton ( 1978) Rutiderma new species KE Belize only: sand trough (rubble, 1978), outer fore-reef slope (1979, 1982), spur and groove (1979), back-reef (I 974) Sarsiellidae Eusarsiella new species MO Belize only: sand and rubble zone (I 979), South Water Cut (I 977) Source of localities located outside Belize: PO.'itaslerope abaca Kornicker, 1986; Bruume/la new species A. Komicker, 1981; Acrinoseta maclallghtinae Komicker, 1981 (Kornicker, 1986); Aclmoseta hllmmelillcki Komicker, 1981 (Kornicker, 1986). 253 specimens), and continued to increase in abundance subsequently, though not as rapidly as rutidermatids and sarsiellids. Representatives of the Sarsiellidae were most abundant in June 1981 samples (l0 species, 376 specimens) and least abundant in June 1979 (8 species, 70 spec- imens). The most abundant sarsiellid species in January 1978 was Eusarsiella new species P (17). Eusarsiella new species KE and E. new species KO increased in number in subsequent samplings. Three species of Eusarsiella were absent from the January 1978 sample, but present in later samples. Their absence in the 1978 sample may be related to smaller sample size (single replicate) as most of these species were rare in later samples. Six Eusarsiella species were absent in either the June or October 1979 collections but present in the June 1981 collections. The Philomedidae are represented in this zone only by Harbansus paucichelatus, which declined in number in 1979 and increased in 1981. The Cypridinidae were rare in all samples. FORE-REEF:SANDTROUGH.There was also no change in species composition in this sample area (except that some less abundant species did not occur in all samples), but there were shifts in relative species and family abundances between COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327 LAGOON LAGOON .. LAGOON LAGOON lOG FORE REEF ... FORE REEF CI Cd P R C,CdPRS co F'omlly ramllv JAN 1978 JUNE 1979 OCTOBER 1979 JUNE 1981 Figure 5. Comparison of relative abundance of myodocopid families in lagoon and sand trough at four sampling dates (1978-] 981). Histograms for January 1978 show number of individuals collected for each family in a single sample in each ofthc two zones. Histograms for 1979-1981 show mean and standard error of three samples from each zone at each date. Samples are same shown in Tab]e I. CI = Cylindro]eberididae; Cd = Cypridinidae; P = Philomedidae; R = Rutidermatidae; S = Sar- siellidae. January 1978 and June 1981. Initially there was an increase, then a decline in total specimen numbers (Fig. 5, Table 1). The smallest sample was 46 ostracodes collected in the single 1978 sample, fewer ostracodes than any of the subsequent nine samples. The highest number collected was in the combined three October 1979 samples (total 473), while about equal numbers were collected in the com- bined three samples in June 1979 (total 356) and combined three samples in June 1981 (total 313). The largest single sample was also collected in October 1979. The most abundant species in the sand trough, Skogsbergia lerneri, as well as the other three sand trough species that belong to the family Cypridinidae, was most abundant in the June 1979 collection and declined in subsequent collections. Each of the three June 1979 samples was larger than the single 1978 sample. Harbansus paucichelatus, the only abundant species of Philomedidae, was al- most as abundant as S. lerneri, increasing slightly during the study period until June 1981, when it declined considerably. The Rutidermatidae and the most abundant species of that family in the sand trough, Rutiderma new species A, both showed numerical increase over most of the study period and a slight decline in October 1979. Like all other rutidermatids in the sand trough, Rutiderma dinochelatum was rare, but was represented by 19 specimens in the June 1981 collection. The Sarsiellidae (and five sarsiellid species) were most abundant in the October 1979 collection and two of those species were only present then. Six species (including the two only present in October 1979) declined in June 1981, and were less abundant than in June 1979 as well. The most abundant species of the Sarsiellidae in the sand trough, Eusarsiella new species KO, as well as two other species, showed an increase in numbers over the entire study period. Eight species of sarsiellids were not collected in all four sampling periods. COMPARISONOF ABUNDANCEIN THETwo ZoNES. About three times as many myodocopids were collected in the 10 lagoon samples (3,281) as in the 10 fore- reef samples (1,188). Total abundance of ostracodes in each of the 10 samples (1978-1981) from each zone was significantly different between the two sites [rank sum test (Ambrose and Ambrose, 1981); T = 69, P < 0.05], and also significantly different between zones in each of the nine replicate samples collected at each site 328 BULLETIN OF MARINE SCIENCE, VOL. 45, NO.2, 1989 Table 4. Chi-square comparison of myodocopid abundance in three lagoon and three sand trough samples collected at each of three dates Lagoon Sand trough Date Observed Expected Observed Expected June 1979 91 137 100 54 197 257 162 102 181 197 94 78 Oct. 1979 238 278 150 110 294 295 118 1]7 109 225 205 89 June 1981 392 386 147 152 973 759 85 300 418 358 81 141 (1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose, 1981); chi-square = 592, eight degrees of freedom, P < 0.05]. Total abundance of individual species (Table 2) was significantly different between the two zones [chi-square test of independence (Ambrose and Ambrose, 1981), chi-square = 2,776,22 degrees of freedom, P < 0.005]. The total number of individuals and individuals/species varied widely not only between sample areas and periods, but also between samples collected during the same period and the same sample area (Tables 1, 5). Sample distributions were both skewed and showed kurtosis even after square root and log transformations (tested by methods in Remington and Schork, 1970: 219), making parametric statistical analysis inadvisable. Inequality of sample variance is probably due in part to failure to make collecting and sorting efforts equal, but the large differences suggest that the ostracodes are not distributed evenly in the sandy bottom within each zone. DISCUSSION Diversity. - The small area sampled on the Belize Barrier Reef in the vicinity of Carrie Bow Cay was characterized by a very high number (51) of myodocopid species relative to previously published totals. Twenty-six of the Belize myodo- copid ostracodes were undescribed (Cohen, 1987). Only 32 Caribbean myodo- copid species, seven from Belize, had been reported previously (Poulsen, 1962; 1965; Wilson, 1913; Kornicker and King, 1965; Kornicker and Cohen, 1978; Kornicker, 1978; 1981; 1983a; 1984a; 1984b; 1986a; 1986b; Cohen, 1983; 1987; Cohen and Morin, 1986). One hundred five species of marine podocopid ostra- codes have also been reported from Belize (Teeter, 1975). Thirty-three (65%) of the 51 species were found in the sand trough of the fore- reef in an area only about 300 m long and 50 m wide, and 29 species (57%) in the sand and rubble zone of the lagoon in an area only about 50 m long and 50 m wide. By comparison, only two species were reported by Komicker (1974) from multiple quadrant grab samples in Cape Cod Bay, Massachusetts. Five species of myodocopids were reported by Elofson (1941; 1969) and Komicker (1987) from repeated dredging in the Skagerak and adjacent areas off Sweden. Lie (1968) reported only four myodocopid species (two very common) in repeated samples at 8 stations in Puget Sound, Washington, U.S.A. Baker (1974, 1975) listed 25 myodocopid species (two very common) from 491 stations on the continental COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329 Table 5. Mean number of specimens, standard deviation and variance of three samples collected in each of two reef zones at each date. Lagoon Sand trough Date x SD Variance SD Variance June 1979 156 57 3,249 118 38 1,444 Oct. 1979 214 94 8,836 158 44 1,936 June 1981 595 329 108,241 104 37 1,369 shelf of southern California. Only about 25 myodocopid species have been re- ported off the whole coast of Japan and adjacent seas (Hanai et aI., 1977), but there are more undescribed species there (based on collections by me and by Hiruta, personal communication). Kornicker (1986b) reported 45 myodocopid species in the northern Gulf of Mexico. On the Bahama Bank off Bimini, Kornicker (1958) collected 21 myodocopid species by dredging over a 10 by 12 km area. There are no data regarding myodocopid diversity on other well developed coral reefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported from Lizard Island on the Great Barrier Reef, Australia (Komicker, 1981; 1982; 1983b; Hall, 1987). Sixteen of the 51 myodocopid species at Carrie Bow Cay belong to the Sarsiellidae. Comparison of Ecological Zones. - While many species have overlapping distri- butions in the reef area studied, a comparison of repeated benthic samples in the two different sandy reef zones shows that the zones differ in myodocopid species composition, particularly in relative abundance (i.e., number of individuals) of many species. Dominant macro-organisms also differ between the two zones (Rutzler and Macintyre, 1982), and Spracklin (1982) found little overlap of hy- droid species between the back-reef (adjacent to my lagoon study area) and fore- reef sand trough at Carrie Bow Cay. The two areas I sampled differ not only in species composition, but in relative abundance of individuals belonging to each of the myodocopid families as well. Comb-feeder cylindroleberid and predatory rutidermatid myodocopids were abundant in the shallow lagoon site, but uncommon at the deeper fore-reef site, while the scavenging cypridinids and predatory sarsiellids were more common in the fore-reef site. Cylindroleberidinae have been observed to burrow just under the surface ofthe sediment and form mucous-like tubes around themselves (Mul- Ier, 1893, and personal observation). Perhaps there is less food available for stationary feeders in the sand trough of the fore-reef than in the shallow lagoon area. However, while the most abundant fore-reef cypridinid, Skogsbergia lerneri, was almost absent from my lagoon sediment samples, it was abundant in baited traps set there at night, and other reef zones (Cohen, 1987) (Table 1, 6). It was also abundant in a rubble sample collected in daytime from the adjacent channel (South Water Cut) cutting through the barrier reef about 400 m from the lagoon sampling area, more sparsely present in sediment from a variety of ecological sites in the area (e.g., lagoon patch reefs and the mangrove-covered Twin Cays), and has been found from Panama to the Bahamas and Gulf of Mexico in the northwest Atlantic Ocean at depths of 1-130 m. All of the sediment samples were made during daytime, while trap samples were made day and night, but mostly at night. Only two Skogsbergia lerneri were collected in the back-reef in the single day trap sample, but night trap samples each contained about 100 to several thousand. 330 BULLETIN OF MARINE SCIENCE, VOL. 45, NO, 2, 1989 Apparently S. lerneri, a good swimmer (personal observation), rarely spends day- light hours in sediment of the sand and rubble zone in the lagoon, but at night swims some distance, possibly as much as 400 m, to feed there. Species abundant in both zones have distributions extending beyond Belize, while many species collected in specific reef zones have been found only in Belize. Species abundant in both zones (though relatively more abundant in one of the zones), are Harbansus paucichelatus, Rutiderma new species A, Eusarsiella new species KO, and to a lesser extent E. donabbotti, Synasterope setisparsa, R. din- ochelatum and Parasterope muelleri. All of these species occur in a number of other habitats in the vicinity of Carrie Bow Cay; five of them have been collected in other NW Atlantic localities (Table 6) and therefore may be considered rela- tively widespread species. On the other hand, of the five species restricted to collections in the back-reef and lagoon, Parasterope new species N, Postasterope new species (1 specimen), E. new species KE, E. new species R, and Chelicopia arostrata, only the last-mentioned species is known outside Belize. Five species were restricted to collections in the fore-reef and channel through the reef: Vargula new species SA, V. new species ST, Harbansus new species A, Euphilomedes species, and Eusarsiella new species MO. None of these species have been collected elsewhere and the last three are rare in Belize. Temporal Changes.-I hypothesize that Hurricane Greta (September 1978) af- fected the shallow lagoon site more strongly than the deeper site. I also hypothesize that differences between myodocopids of the lagoon and fore-reef sites in partic- ular, and that the high number of myodocopid species occurring in the reef area at Carrie Bow Cay in general, are related to storm disturbance. Although species composition remained fairly stable in the two zones, relative abundance of in- dividuals of species and families changed during the 3 years, particularly in the lagoon samples (Table 1, Fig. 5). While total abundance increased considerably in the lagoon site in each collection period following the hurricane, total abundance in the fore-reef site first increased slightly and then declined slightly in 1981. The smaller fore-reef fluctuations may be a reflection of lesser disturbance, or they may be insignificant. The fore-reef site has a smaller number of ostracodes, a higher species diversity and smaller changes in relative abundance than the shallow lagoon site. A comparison of the effects of storm disturbance on ecological stability in the two study areas requires an estimation of relative force in the areas (Connell and Sousa, 1983). Evidence that the sand trough is less disturbed comes from the nature of its sediment. In the sand trough very fine sediment reaches a depth of 12 m at the axis of the trough, while sediment in the sand and rubble zone is poorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre, 1982). Storm disturbance in the sand trough is probably considerably less than in the lagoon because of the greater depth of the sand trough and its protected position between and below the inner and outer fore-reef ridges. I observed damage to corals in the shallow lagoon following Hurricane Greta. Kjerfve and Dinnel (1983) found that substantial waves from Hurricane Greta in 1978 approached Carrie Bow Cay from the SW, strongly affecting the lagoon. Riitzler and Macintyre (1982) found considerable damage and movement of the Acropora cervicornis coral usu- ally scattered about in the sand and rubble zone. Many lagoon ostracodes must have been washed away by the hurricane. In analysis of the sandy zone in the back-reef of Tobacco Reef, 3 km N of and similar to the lagoon site at Carrie Bow Cay, Macintyre et al. (1987) calculated that sand transport takes place during brief intervals of storm activity. Sand there was suspended and transported during COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331 Table 6. Myodocopid species collected in quantitative samples and also collected in nonquantitative samples from other Belize localities or reported from localities outside of Belize Taxon Nonquantitative sample locality (and published source of non-Belize localities) Cypridinidae Skogsbergia lerneri Belize: traps all major zones, rubble and algae in back-reef and Water Cut, Twin Cays, lagoon patch reefs. Bahamas, E Panama, Gulf of Mexico (Kornicker, 1984a) Vargula new species ST Belize only: patch reefs, South Water Cut rubble V. new species SA Belize only: edge of outer shelf of fore-reef Plerocypridina sex N. and S. Carolina, E and W Florida (Komicker, 1984a) Cylindroleberididae Parasterope new species N Belize only: lagoon Thalassia. back-reef, South Water Cut P. muelleri Belize: back-reef, spur and groove, reef slope, Twin Cays, patch reefs, plankton; English channel, West Indies, Mauritania, Medi- terranean, Florida, Bahamas, Bermuda (Kornicker, 1986b) Synaslerope setisparsa Bahamas (Kornicker, 1986b) S. new species A Belize only: patch reef, South Water Cut Amboleberis americana Belize: spur and groove, reef slope, South Water Cay; Texas, S. Carolina-Florida, Bahamas, Brazil, W Costa Rica, W Panama (Kornicker,198I) Asteropella monambon Belize: back-reef; Bahamas, Puerto Rico, Cuba (Kornicker, 1981) Aclinosela chelisparsa Belize: spur and groove, outer ridge, W Carrie Bow Cay, South Water Cut; Bahamas, W Florida, Bonaire, Venezuela, Panama, Cura~o (Kornicker, 1981) Philomedidae Harbansus paucichelalus Belize: patch reef, South Water Cut; N Carolina, Florida, Baha- mas, Gulf of Mexico (Kornicker, 1984b) Rutidermatidae Rutiderma new species A Belize: back-reef, Thalassia. plankton; Texas (Komicker, 1983a) R. dinochelatum Bahamas (Kornicker, 1983a) R. darbyi Belize: back-reef, spur and groove, lagoon patch reefs; N Carolina- W Florida, Bahamas (Kornicker, 1983a) R. harmanni Belize: South Water Cay; W Panama (Kornicker (\ 985) R. cohenae Belize: spur and groove, outer fore-reef slope; Bahamas, Florida Keys (Kornicker, 1983a) RUliderma new species B Belize only: lagoon patch reefs, back-reef, spur and groove, outer ridge, South Water Cut, lagoon plankton R. new species KO Belize only: spur and groove, lagoon plankton Sarsiellidae Dantya magnifica Belize only: outer reef slope Chelicopia arostrala Belize: back-reef; Florida, Bahamas (Kornicker 1986a) Eusarsiel/a uncus Florida, Bahamas (Kornicker, 1986a) E. new species KE Belize only: back-reef E. new species KO Belize only: lagoon patch reefs, Thalassia, spur and groove, reef slope, South Water Cut E. new species RU Belize only: back-reef E. new species C Belize only: inner outer ridge E. new species P Belize only: spur and groove E. new species MJ Belize only: sand and rubble zone E. new species MR Belize only: Ragged Cay, plankton E. donabbotti Belize only: lagoon patch reef, South Water Cut, Dangriga E. new species KN Belize only: back-reef plankton, Ragged Cay E. new species J Belize only: Twin Cays E. new species CL Belize only: Twin Cays 332 BULLETIN OF MARINE SCIENCE, VOL. 45, NO.2, 1989 Hurricane Greta, disturbed to the point of ripple formation during average storms, and undisturbed by tradewinds. Highsmith et a1. (1980) found that Hurricane Greta in 1978 caused fragmentation of coral, but increased numbers and redis- tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie Bow Cay, Belize, demonstrating that long-term reef calcification and growth rates may be highest on reefs periodically disturbed by storms of intermediate intensity. They found that Hurricane Greta caused breakage and scouring of corals in all zones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately 25 m. An analysis of ecological stability should use appropriate temporal and areal scales to examine the degree of constancy in (I) number of organisms, and! or (2) presence or absence of taxa (Connell and Sousa, 1983). Minimum temporal scale is one turnover or lifetime of the organism (Connell and Sousa, 1983). The study period from 1978-1981 greatly exceeds the lifetime of the most abundant species in the study, Skogsbergia lerneri. as this species was reared from egg to adult in a coral reef aquarium within 2 months and survived no longer than 2 months as an adult (adult females had two to three successive clutches) (Cohen, 1983). Some specimens of Rutiderma hartmanni A, partially reared under similar conditions, were brooded as embryos for 53 days, and developed from first to second instar in another 53 days (Table 7; Cohen, 1987). If duration of all four juvenile instars in Rutiderma is similar, as it is in Skogsbergia, complete devel- opment time from egg to adult would be about 9 months in Rutiderma. Areal scale in the two study sites probably exceeds the minimum space needed for generational replacement of the ostracodes. All instars of the most abundant species were usually found in single replicates and during each sampling period. Shifts in abundance of dominant taxa occurred not only at the species level, but at higher taxonomic levels, i.e., generic and familial. These alterations may be correlated with characters (perhaps synapomorphies) shared by all of the species belonging to a higher taxon. Possible characters include those affecting dispersal and life history tactics and feeding strategies. When one of these characters has relatively more importance in natural selection, selection could act at the higher taxonomic level as a byproduct of selection at the individual level (all included individuals possessing that same character state). Following the hurricane the most abundant ostracode species included only myodocopid taxa with swimming juveniles, Parasterope new species N in the lagoon and the cypridinid Skogsbergia lerneri in the fore-reef site. Both species are better swimmers even as adults (personal observation) than the rutidermatids, and to a lesser extent than sarsiellids, the two families which were more abundant before and 3 years after the hurricane. In the Rutidermatidae only adults have swimming hairs on the exopod of the second antenna (Kornicker 1985; Cohen, 1987), the limb used for swimming. In the other three families all instars have swimming hairs on that limb. After emerging from the brood chamber, instars of Rutiderma were not observed to swim for 70 days, and then swam for only a few millimeters (Cohen, 1987). Rutidermatidjuveniles displaced by a storm could only crawl and swim slowly, while swimming juveniles of other myodocopid taxa could regain a favorable habitat more rapidly by swifter swimming. However Rutiderma new species A was the dominant species in the lagoon site (and Eu- sarsiella new species P the dominant sarsiellid) only in the year before the hur- ricane. The dominant species 3 years after the hurricane was a different rutider- matid, R. dinochelatum (and sarsiellid E. new species KE). Predatory species were more abundant before and 3 years after Hurricane Greta, while just after the hurricane comb-feeders and scavengers were relatively more important. Specimens of Rutiderma and Eusarsiella often have whole copepods, COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333 c· O)~ .c",O' CONCLUSIONS Myodocopid species were diverse in this small area, predominant species and families differed between the sand trough of the fore-reef and the sand and rubble zone of the lagoon, and species and family composition varied from year to year. Further, myodocopids were less abundant overall, but displayed greater species richness and smaller yearly changes, in the samples from the deeper, less disturbed fore-reef site, than in the shallower lagoon site, I hypothesize that this temporal and zonal variation was correlated both with zonal differences in relative physical disturbance by Hurricane Greta, and with taxonomic differences (at the family, not only the specific level) in life history and functional morphology. Taxa col- lected in greatest abundance 9 months after Hurricane Greta include the fast- developing Skogsbergia lerneri (a cypridinid), characterized by large clutch sizes, COHEN: OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335 juveniles with swimming hairs, and scavenging diet. Less abundant in collections 9 months after the hurricane, but very abundant 3 years later, were slower-de- veloping rutidermatids with smaller clutch sizes, juveniles lacking swimming hairs, and predatory diet. Also more abundant and recovering more quickly than its congeners was the only sarsiellid species with double clutches. The data from this study are unfortunately limited, but seem to agree with the theories that diversity is related to interaction of physical disturbance with biological factors (Huston, 1979; Sousa, 1984) and highest diversity to intermediate levels of dis- turbance (Connell, 1978). ACKNOWLEDGMENTS This paper is dedicated to the memory of Donald P. Abbott who launched and encouraged me in the study of invertebrate zoology. I am grateful also to L. S. Kornicker for encouragement, training and advice in the study of myo- docopid ostracodes in general and in this research in particular. I thank R. Brusca and an anonymous reviewer for many helpful comments on the manuscript, and G. Hendler and A. Jahn for particular ideas, but responsibility for final content is mine alone. For providing laboratory space, advice and encouragement, I thank K. Riitzler and the Division of Crustacea, Smithsonian Institution, particularly T. E. Bowman, B. Kensley, and R. B. Manning, and at the Allan Hancock Foundation, University of Southern California, R. C. Brusca (now at the San Diego Museum of Natural History). This is contribution number 239, Caribbean Coral Reef Ecosystems (CCRE), Smithsonian Insti- tution, partly supported by a grant from the Exxon Corporation. Field work in 1981 was also supported by a grant from the Lerner Fund, American Museum of Natural History. I thank the Mead Foundation for a grant providing funds for a word processor and a trip to study collections at the Smithsonian Institution. Many provided collecting assistance: I particularly thank M. Carpenter, B. Kensley, R. Larson, K. Riitzler, J. D. Thomas, C. A. Child, G. Bretchko, S. Lewis, T. Rath, and J. Ferraris. T. Stebbins instructed me in computer graphics. For invaluable help in moving collections from Wash- ington, D. C. to California I thank C. S. 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ADDRESS: Division of Life Science, Los Angeles County Museum of Natural History, 900 Exposition Boulevard. Los Angeles. California 90007 (affiliation: Biology Dept .. University of California. Los Angeles).