Heteropod and Thecosome (Mollusca: Gastropoda) Macroplankton in the Florida Straits
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BULLETIN OF MARINE SCIENCE, 49(1-2): 562-574, 1991 HETEROPOD AND THECOSOME (MOLLUSCA: GASTROPODA) MACROPLANKTON IN THE FLORIDA STRAITS Harding B. Michel and John F. Michel ABSTRACT Previous areal records of hete:ropod and thecosome molluscs obtained with single- and graded-mesh plankton nets and the Isaacs-Kidd Midwater Trawl (IKMT) are compared with a collection of late juveniles and adults from the Florida Current caught by a MOCNESS- 10 trawl. Sequential depth intervals from the surface to a maximum of 750 m were quan- titatively sampled on transects between Palm Beach, Florida, and Settlement Point, Grand Bahama Island, Bahamas, in August 1978 and February 1979. Of the eight heteropod and 15 thecosome species or formae captured, the most numerous were, in order of abundance, the heteropods Atlanta peroni. Pterotrachea hippocampus. P. coronata and A. inc/inata, and the thecosomes Cavolinia uncinata forma uncinata. Diacria trispinosa and Clio pyramidata. Data on species abundance relative to earlier reports and gear types and on vertical distribution during day and night are presented. No notable evidence of deep-water occurrence was found. Reports of the occurrence and distribution of pelagic gastropods in the Rorida Current and the contributing currents from the Caribbean Sea and the Gulf of Mexico have been based primarily on collections made with relatively fine-meshed plankton nets. These sample populations of smaller species but often do not capture specimens oflarger ones such as the heteropds Pterotrachea coronata and P. hippocampus, and the thecosomes Clio recurva and Cavolinia uncinata. One purpose of this paper is to give quantitative evidence of incomplete sampling of these forms that, at least as adults, may avoid plankton nets, and also are poorly collected by slow-moving trawls, such as the IKMT, which pass small and inter- mediate sizes and tend to crush larger animals. Another is to describe the com- parative abundance of the larger heteropod and thecosome species obtained in the trawl designed to sample mesopelagic fishes. The literature cited excludes most historical works and consists mainly of recent efforts to approximate abundance and vertical distribution of species. Concerning heteropods, Taylor and Berner (1970) reported relative numbers collected in the Gulf of Mexico by three methods: a l-m net (mesh size 752 /otm),a "Bongo" sampler (mesh size 505 /otm)and a 10-foot IKMT ('14" stretch liner). Michel and Foyo (1976) sampled Caribbean zooplankton in multilevel horizontal tows of75 cm modified "Discovery" nets (graded mesh sizes 3.2 mm, 1.6 mm and 110 /otm), and they recorded numbers of heteropods caught in the comparable tows. Thecosomes have been more thoroughly studied in this area than heteropods. Wormelle (1962) examined samples collected in the Rorida Straits off Miami with the same type of Discovery net described in the previous paragraph. Catches obtained from the Caribbean Sea by Michel and Foyo (1976) formed the basis for Haagensen's thesis (1976). In their zoogeographic and taxonomic review, Be and Gilmer (1977) discussed sampling inadequacies associated with net and mesh size, towing speed and duration, and they referred to Wells' (1973) findings of effects of mesh size on estimates of euthecosome density in the Caribbean off Barbados. Van der Spoel produced extensive compendia on thecosomes (1967) and on thecosomes and heteropods (1976). In addition to a thorough systematic review, these works contain information on anatomy and world-wide distribution. Formae established by van der Spoel, of course, do not appear in the papers of Wormelle 562 MICHEL AND MICHEL: HETEROPOD AND THECOSOME MACROPLANKTON 563 (1962) and Taylor and Berner (1970) and in the review ofheteropods by Thiriot- Quievreaux (1973). In Haagensen (1976) and the present work, formae were assigned in all clear cases. Lalli and Gilmer's (1989) beautifully illustrated and detailed work on the biology of holophanktonic gastropods brings together modem knowledge of the kinds of heteropods and thecosomes and their life styles. METHODS The present data were obtained from collections made by Richard H. Backus and James E. Craddock, Woods Hole Oceanographic Institution, Woods Hole, MA. The MOCNESS-lO trawl was described by Backus and Craddock (1982), who used it to sample mesopelagic fishes. On two cruises of R/V OCEANUS,transects were made between Palm Beach, Florida, and Settlement Point, Grand Bahama Island, Bahamas (Table 1). After removing fishes from these tows, Backus and Craddock kindly sent the senior author the remainder of the catch. The circular openings of the MOCNESS-I 0 net are about 3 mm in diameter. The area of the mouth when fishing is approximately 10 m2• Speed during fishing varied from ca. 1.3-2.9 kt. Bottom depths extended from 770 to 360 m, east to west, and fishing depths covered multiple ranges from the surface to 750 m (Table 1). Because the objective was to collect mesopelagic fishes, greater depths and ranges were sampled in the daytime than at night, particularly during the hours around midnight. This sampling bias and the lack of replication permit evaluation of vertical distribution only in general terms. In 1979, heteropods and thecosomes were sorted from the original buffered formalin preservative, used after collection in 1978 and 1979, and stored in 70% alcohol. The condition of the material remains excellent. So few pseudothecosomes and gymnosomes were found in the samples that identification of most species was not attempted. Total counts of the other species and formae were made. RESULTS AND DISCUSSION A list of the species collected by MOCNESS-IO follows. In Tables 2 and 3, historical and present records are compared. The abundance of heteropod and euthecosome species obtained from the depths sampled at each station appears in Tables 4, 5 and 6. List Of Species Class Gastropoda Subclass Prosobranchia Order Heteropoda Family Atlantidae Oxygyrus keraudreni (Lesueur, 1817) Atlanta inclinata Souleyet, 1852 Atlanta peroni Lesueur, 1817 Family Carinariidae Cardiapoda placenta (Lesson, 1830) Carinaria lamarcki Peron and Lesueur, 1810 Family Pterotracheidae Pterotrachea coronata Niebuhr, 1775 Pterotrachea hippocampus Philippi, 1836 Pterotrachea scutata Gegenbaur, 1855 Subclass Opisthobranchia Order Thecosomata Suborder Euthecosomata Family Cavoliniidae Styliola subula (Quoy and Gaimard, 1827) Clio pyramidata Linnaeus, 1767 forma lanceolata (Lesueur, 1813) Clio recurva (Children, 1823) Cuvierina columnella (Rang, 1827) forma atlantica van der Spoel, 1970 Diacria trispinosa (de Blainville, 1821) 564 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991 Table I. Station data, OCEANUS cmises 49 and 55 between Palm Beach, Florida, and Settlement Point, Grand Bahama Island OcEANUS 49 Bottom Station Lat. Long. Time Sample depth depth Water filtered number Date N W R (m) (m) (m') 99-0 6 Aug 78 26°42' 79"08' 2103-2258 0-200 580 50,000 99-1 6-7 Aug 78 26°42' 79"08' 2258-0057 200-570-200 580 58,000 99-2 7 Aug 78 26°31' 79°12' 0057-0333 0-200 580 83,000 100-0 7 Aug 78 26°44' 79°23' 0502-0640 0-270 630 41,000 100-1 7 Aug 78 26°44' 79°23' 0640-1010 270-600-280 630 100,000 100-2 7 Aug 78 26°47' 79°25' 1010-1107 290-0 630 33,000 101-0 7 Aug 78 26°42' 79°31' 1231-1320 0-400 770 16,000 101-1 7 Aug 78 26°42' 79°31' 1320-1657 400-750-400 770 107,000 101-2 7 Aug 78 26°38' 79°31' 1657-1755 400-0 770 39,000 102-0 7 Aug 78 26°44' 79°40' 1929-2040 0-520 565 22,000 102-1 7 Aug 78 26°44' 79°48' 2040-2202 550-100 565 47,000 102-2 7-8 Aug 78 26°52' 79°40' 2202-0004 100-0 565 61,000 103-0 8 Aug 78 26°45' 79°44' 0106-0246 0-70 450 58,000 103-1 8 Aug 78 26°45' 79°44' 0246-0425 70-150-80 450 56,000 103-2 8 Aug 78 26°53' 79°46' 0425-0600 80-0 450 60,000 104-0 8 Aug 78 26°42' 79°31' 0836-0940 0-350 770 28,000 104-1 8 Aug 78 26°42' 79°31' 0940-1350 350-710-350 770 134,000 104-2 8 Aug 78 26°37' 79°32' 1350-1444 350-0 770 24,000 105-0 8 Aug 78 26°42' 79°31 ' 1525-1613 0-350 770 22,000 105-1 8 Aug 78 26°42' 79°31 ' 1613-1857 350-700-350 770 85,000 105-2 8 Aug 78 26°39' 79°32' 1857-1941 350-0 770 30,000 106-0 8 Aug 78 26°43' 79°31' 2045-2147 0-200 770 30,000 106-1 8 Aug 78 26°43' 79°31' 2147-2358 210-90 770 74,000 106-2 8-9 Aug 78 26°43' 79°31 ' 2358-0212 90-0 770 76,000 OcEANUs 55 143-0 17 Feb 79 26°41' 79°36' 1405-1521 0-400 720 32,000 143-1 17 Feb 79 26°41" 79°36' 1521-1635 400-650 720 30,000 143-2 17 Feb 79 26°41 ' 79°36' 1635-1720 650-400 720 27,000 143-3 [no specimens] 1720-1754 400-300 720 21,000 143-4 17 Feb 79 26°41' 79°36' 1754-1918 300-150 720 50,000 143-5 17 Feb 79 26°38' 79°36' 1918-2029 150-0 720 46,000 144-0 17 Feb 79 26°44' 79°36' 2122-2301 0-650 680 36,000 144-1 [no specimens] 2301-2356 650-400 680 31,000 144-2 17-18 Feb 79 26°44' 79°36' 2356-0054 400-160 680 33,000 144-3 18 Feb 79 26°44' 79°36' 0054-0148 160-100 680 31,000 144-4 18 Feb 79 26°44' 79°36' 0148-0301 100-50 680 42,000 144-5 18 Feb 79 26°40' 79°39' 0301-0413 50-0 680 35,000 145-0 18 Feb 79 26°43' 79°47' 1327-1430 0-350 360 25,000 [extrapolated] 145-1 18 Feb 79 26°43' 79°47' 1430-1506 340-320 360 19,000 145-2 18 Feb 79 26°43' 79°47' 1506-1557 320-360-310 360 24,000 145-5 18 Feb 79 26°37' 79°51' 1557-1734 310-0 360 50,000 146-0 18 Feb 79 26°42' 79°35' 1906-2207 0-680 750 70,000 146-1 [no specimens] 2207-2324 680-250 750 43,000 146-2 18 Feb 79 26°42' 79°35' 2324-2341 250-200 750 10,000 146-3 18-19 Feb 79 26°42' 79°35' 2341-0020 200-150 750 23,000 146-4 19 Feb 79 26°42' 79°35' 0020-0111 150-100 750 29,000 146-5 19 Feb 79 26°36' 79°30' 0111-0222 100-0 750 42,000 147-0 19 Feb 79 26°35' 79°29' 0312-0419 0-100 710 33,000 147-1 19 Feb 79 26°35' 79°29' 0419-0534 100-200 710 37,000 147-2 19 Feb 79 26"35' 79°29' 0534-0711 200-650 710 39,000 147-4 19 Feb 79 26"35' 79°29' 0711-0802 650-610-650 710 22,000 147-5 19 Feb 79 26"29' 79°27' 0802-1043 650-0 710 105,000 148-0 19 Feb 79 26"42' 79°21' 1230-1334 0-300 630 30,000 MICHEL AND MICHEL: HETEROPOD AND THECOSOME MACROPLANKTON 565 Table 1.