RECENT

MICROMORPH

BRACHIOPODS

FROM BARBADOS IMPLICATIONS

PALAEOECOLOGICAL

AND EVOLUTIONARY

by ULLA ASGAARD * & NIELS STENTOFT *

ABSTRACT

RI~SUMI~

Brachiopods were collected from 16 to 325m depth from reef and off-reef environments, off the west coast of Barbados (West Indies). The 15 species collected showed distinctive depth distributions, which are dependent in part on the distribution of preferred substrates and in part on the larval biology. Comparison with Upper Tertiary and early Quaternary micromorph brachiopod faunas from the Mediterranean area and similar forms from the uppermost Cretaceous and lowermost Tertiary white chalk and calcarenites from NW Europe indicates that the micromorph brachiopod faunas we know today already were established by the end of the Cretaceous.

Ces brachiopodes ont 6t6 r6colt6s entre 16 et 325m de profondeur, en domaine r6cifal et p6rir6cifal, au large des cbtes occidentales des Barbades (Antilles). Les 15 esp~ces montrent une r6partition bathym6trique distincte, d6pendant en partie de la r6partition des substrats favorables et de la biologie larvaire. Une comparaison avec les faunes de brachiopodes micromorphes du Tertiaire sup~rieur et du Quaternaire ancien de M6diterran6e et avec des formes similaires de la craie blanche et des calcarar6nites du Cr6tac6 terminal et du Tertiaire basal du NW de l'Europe, montre que les faunes de brachiopodes micromorphes modernes remontent/~ la fin du Cr6tac6.

* Geologisk Centralinstitut, Ostervoldgade 10, DK-1350 Kobenhavn K. Danemark.

Geobios, M6m. sp6cial n ~ 8

p. 29-33, 2 pl

Lyon, 1984

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30

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1 - D E P T H DISTRIBUTION

Substrates and sediments were collected with SCUBA on the fringe reef off Speightstown (-12 to e42 m) and from a transect out from Holetown and

Paynes Bay : barrier reef (SCUBA, -12 to -42 m), outer reef slope (Petersen grab, -55 to -92m) and offreef environment (dredge, -110 to -325m). The following brachiopods were found :

Depth alive

Depth dead 55 - 325m 60 - 3 2 5 m 110- 325m 2 4 0 - 325m 146 - 3 2 5 m 6 0 - 325m 25 - 70m 55 - 1 5 0 m 92 - 150m 6 0 - 172m 60,7219m ?-7285m 2 2 0 - 320m ? 2 4 0 - 325m

P I . : Fig. 1:8 1:9 1:4 2 : 6,7 1 : 6,7 1:1 2 : 4,5 1:5

1 :

Crania pourtalesi DALL Crania smooth sp. Crania costate sp. Cryptopora rectimarginata COOPER Gryphus bartletti (Dall) Terebratulina cailleti CROSSE Argyrotheca cf. bermudana DALE Argyrotheca shrammi (Crosse & Fischer) Argyrotheca rubrotincta (Dall) Argyrotheca barrettiana (Davidson) Argyrotheca crassa C O O P E R Argyrotheca sp. Megerlia cf. echinata (Fischer & Oehler0 Platidia anomioides (Scacchi & Philippi) Platidia clepsydra COOPER

55 - 325m 1 juv. 325m 2 4 0 - 325m 2 0 1 - 248m 90 - 201m 25 - 45m (16)55 - 140m 92 - 140m 92 - 140m 92 -?285m 190- 219m 219 - 250m 240 - 325m

2,3

2:8 2:1,2 1 : 10,11 2:3

The brachiopods fall into 6 groups according to depth. 1) Argyrotheca cf. bermudana occurred from 25 to -42m. 2) Argyrotheca schrammi occurred from 55 to -140m. 3) Argyrotheca rubrotincta and A. barrettiana occurred from -92 to -140m. 4) One (or may be two) Argyrotheca spp. and Platidia anomioides occurred in the interval between group 3 and the shallower limit of group 5.5) Cryptopora rectimarginata, Megerlia cf. echinata and Platidia clepsydra were found from about -220 to -325m (and deeper). Group 6 : Crania spp., Gryphus bartletti, Terebratulina cailleti and Argyrotheca crassa seem to have a wider range of depth.

A large, living Agariciafragilis colony was obtained from -55m, carrying a small population o f Argyrotheca schrammi and Crania pourtalesi. Other grabsamples from -55 to -73m contained bioclastic sand and gravel and larger corai-fragments, but no living brachiopods were found. From -73 to -172m, two demosponge substrates occurred : the orange, irregularly branched Agelas schmidti and a whitish, foliaceous Plakortis n. sp. that contained a symbiotic $iliquaria sp. The tubular branches of A. schmidti are about 1 cm in diameter and the skeleton incorporates grains of the sediment the sponge happens to rest on. Both to these grains and to the Siliquaria shells within the Piakortis sp. a rich epifauna is attached, including Argyrotheca

2 - SUBSTRATES From -25 to -42m Argyrotheca cf. bermudana was found on the undersides of living, foliaceous scleractinians, mainly Agaricia agaricites, but also on Mycetophyllia spp. and Montastrea annularis. Less commonly, live and dead, branched scleractinians also served as substrates, e.g. Porites astreoides, Mussa angulosa and Madracis mirabilis.

schrammi, A. rubrotincta, A. barrettiana, A. crassa and Terebratulina cailleti. In a few cases all 5 species
were found together on the same sponge. At -73, and down to -325m (and deeper) synsedimentary cementation has taken place. This has resulted in nodular lumps o f bioclastic material up to 10cm thick in the shallower part of the transects, while in the deeper part regular crustlike hardgrounds seem to

--31 m

be present (Stentoft, 1982). The lumps and crusts both bore a rich epifauna including brachiopods from groups 3, 4, 5 and 6.

Gryphus bartletti also was very small for the genus. The only two living specimens found were attached to pebbles smaller than their shells with short distally split pedicles. Dead shells of all sizes were common in the sediment samples.
Few live Crania spp. were found, and no dead ventral valves were observed on apparently suitable substrates. Dorsal valves were common in the sediment samples. They were all irregular in shape ; many of them show xenomorphic sculpture that indicates that the ventral valves were cemented with the entire surface to the substrates. Only two populations of Platidia anomioides were found. The close adpression to the substrate in this species is clearly to be seen from the irregularities of the figured specimens. The Argyrotheca spp. from group 4 are less brightly coloured than the shallow water species and tend to be much wider than long and have large endopunctae compared to the shallow water Argyrotheca spp.

3 - E C O L O G Y AND POPULATION STRUCTURE

The micromorph Argyrotheca spp. are fastened to the substrate by an extremely short and stout pedicle. The commissure is thus perpendicular to the substrate and the valves gape widely when the animal feeds. When observed in the laboratory A. cf. bermudana and A, schrammi were seen to swivel violently when disturbed by swimming crustaceans and ridding themselves of sediment. Argyrotheca spp. are known to brood their larvae, which results in a patchy distribution. In the case of A. cf. bermudana many m2 of apparently suitable substrates were found to be barren, and we were unable to confirm the distribution given by Logan (1975) ; no specimens were found on the dives at the fringe reefs off Holetown in -6 to -10m depth. The populations collected in late April and early May 1980 showed a bimodal distribution of mature specimens and recently settled individuals. Argycotheca cf. bermudana, A. schrammi and A. rubrotincta quickly attain full size and have a short lifespan. In the accompanying sediments very few shells of immature specimens were found, which would indfcate a low juvenile mortality. The deeper water Argyrotheca spp. have occasionally clear and crowded growth-lines near the commissure, indicating a longer lifespan. The brachiopods down to -73m were found in cryptic habitats on the undersides of foliaceous scleractinians or between branches of branched forms. Below -73m the brachiopods came out into the open fastened to sponges and cemented lumps and crusts. No caves were situated on the chosen reef transects and this might explain why no thecideans were found. Thecideans have been reported from reef-caves elsewhere in the Caribbean (Jackson & alii, 1971, Logan, 1977). The almost ubiquitous Terebratulina cailleti may also be considered a micromorph species since none of the specimens found exceeded 1 cm in length. It is possible that this species broods its larvae as does T. septentrionalis, but the material did not show a particularly low juvenile mortality as was the case for the smaller Argyrotheca spp. T. cailleti is furnished with j long, distally split pedicle. Specimens in the laboratory gave lively demonstrations of their ability to flap their valves and whip round on their pedicles when disturbed.

Cryptopora rectimarginata and Platidia clepsydra were the dominant living brachiopods found in the deeper dredges. They do not exceed 3ram in length and owing to the transparency of their shells they are easily overlooked. These species do not bore into the substrates as do the other brachiopods and are therefore easily dislodged. P. clepsydra does not live with its dorsal valve pressed against the substrate.
The quantitative analysis of living populations and death assemblages is not yet completed, but the results so far do not seem to deviate from similar studies in other regions (e.g. Noble & alii, 1976, Noble and Logan, 1981 and Steward, 1981). Constituent grain analyses made on selected samples show that where the brachiopods are common they do not exceed 2% of the grains and in many cases they constitute only 0.5% (Stentoft, 1982). Similar results were reached elsewhere by Caulet (1967) and Logan (1977).

4 - O T H E R RECENT M I C R O M O R P H B R A C H I O P O D FAUNAS

The Caribbean Sea exhibits the greatest diversity of micromorph brachiopods (Cooper, 1977) closely followed by the Mediterranean and tropical to subtropica~,Eastern Atlantic (Logan, 1979). Scattered records exist from the Red Sea, Persian Gulf, Indian Ocean and the Pacific (Thomson, 1927, Cooper, 1973a, 1973b, 1981, 1982), Gulf of California (Hertlein and Grant, 1944) and Bermuda (Logan, 1975).

32

All these faunas show a surprisingly similar composition to the Barbados fauna, having a dominance of Argyrotheca spp. in the sublittoral areas and Kraussinidae, Platidiidae and Cryptopora spp. taking over at the shallow end of the bathyal range. The similarity also extends to the morphology of the species. The Argyrotheca spp. of group 1 in most cases are smooth or have few and nearly obsolete costae, the greatest width being anterior to the hinge line. Argyrotheca spp. of groups 2 and 3 are variably costate and some forms have their greatest width at the hinge line. Most of the Argyrotheca spp. of group 4 are alate and furnished with large endopunctae. This parallelism in the circumtropical belt would seem to indicate a long history of shallow micromorph brachiopod faunas.

Chalk of the Campanian/Maastrichtian boundary of N.W. Germany and found that the Lower Maastrichtian species were present in the Upper Campanian. In contrast, however, Campanian nearshore and coastal sediments of Sweden have an extremely diverse brachiopod fauna which seems totally devoid of micromorph species (except thecideans). Further back in the Cretaceous the information on micromorph brachiopods so far is scanty, apart from on thecideans from shallow water environments (see Pajaud, 1974 for references). A rich fauna of minute cancellothyridids is found in the Turonian white chalk of northern France (M. Bagge Johansen pers. comm. 1983).

6 - T H E O R I G I N OF MODERN M I C R O M O R P H BRACHIOPOD FAUNAS

- FOSSIL M I C R O M O R P H B R A C H I O P O D FAUNAS

The Recent micromorph brachiopod faunas can be traced back to the Upper Tertiary, and at least the Mediterranean species have existed since then." Lower Tertiary shallow water brachiopods have been described from the Caribbean area, Western and Eastern U.S.A. the Mediterranean area, N.W. Europe and from the Indopacific. In the offshore sublittoral sediments of the Danian of Denmark and the nearshore sediments of Holland and Belgium a variety of Argyrotheca spp. and other micromorph brachiopods are found (Asgaard, 1968 and unpublished observations ; M. Bagge Johansen pers. comm., 1983). By the Danian the different morphological patterns we can recognize today in groups 1-4 already seem to be present. Steinich (1965) and Surlyk (1972) described the extremely diverse brachiopod fauna from the Maastrichtian white chalk of northern Europe with its 32 micromorph species. Surlyk (1972) divided the species in groups according to their morphology. 1) 20 minute species adapted to a life on very small substrates e.g. slender, branched bryozoans. In this group were cancellothyridids, megathyridids, platidiids and dallinids. 2) Two cancellothyridids, one terebratellid and one thecidean were hemispherical and secondarily free-living. Finally 3) two craniids and 3 thecideans were cemented by a small area to the substrates. However, the Maastrichtian nearshore sediments have not yet received a similar investigation. Surlyk (1982) described the brachiopods from the white

As mentioned above, shallow water micromorph brachiopod faunas seem to have existed in the warm seas since the Eocene. By late Cretaceous, a diverse micromorph brachiopod fauna had evolved which had adapted to a life in the open, fastened to small and rare substrates or free-lying on mud in a shallow offshore environment. Cenomanian to Campanian nearshore environments that we know of are dominated by large brachiopods (and the small thecideans). The thecideans had a cryptic mode of life at least since the Upper Jurassic (Pajaud, 1974). We consider that the cryptic mode of life of the non-thecidean micromorph brachiopods is a new adaptation by a fauna already well established (( in the open > below the > photic zone. Logan (1977, 1979) considered the factors encouraging a cryptic mode of life in brachiopods to be the availability of suitable sttbstrates, competition for space, larval photophobia, sedimentation and energy level, while predation hardly played any role. We would consider another factor to be highly relevant in this matter : that of the almost ubiquitous armies of regular echinoids grazing on all exposed surfaces in the photic zone of warm seas (Samarco, 1980, Hawkins, 1981). The active grazing on algae, particularly red calcareous encrusting algae, causes the removal of epilithic forms from the grazed surfaces. The disappearance of large brachiopods from the photic zone of the tropical seas by the end of the Cretaceous, therefore, might be linked to the evolution of the camarodont Echinoida and the appearance of Diadema at that time.

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33

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Acknowledgements

R.G. Bromley (Copenhagen) assisted in the fieldwork and offered valuable criticism of the manuscript. The Danish

Natural Science Research Council supported the fieldwork financially.

REFERENCES

ASGAARD U. (1968) - Brachiopod palaeoecology in Middle Danian limestones at Fakse, Denmark. Lethaia, Oslo, 1, 103-121. CAULET J.P. (1967) - Les sc~diments meubles tt brachiopodes de la marge continentale alg~rienne. Bull. Mus. natL Hist. nat., Pads, 39, 779-792. COOPER G.A. (1973a) - New Brachiopoda from the Indian Ocean. Smithsonian Contrib. Paleobiol., Washington D.C., 16, 1-43. COOPER G.A. (1973b) - Vema's Brachiopoda (Recent). Smithsonian Contrib. Paleobiol., Washington D.C., 17, 1-51. COOPER G.A. (1977) - Brachiopods from the Caribbean Sea and adjacent waters. Stud. trop. Oceanogr., 14, 1211, Coral Gables Florida. COOPER G.A. (1981) - Brachiopoda from the Southern Indian Ocean (Recent) Smithsonian Contrib. Paleobiol., Washington D.C., 43, 1-93. COOPER G.A. (1982) - New brachiopods from the southern hemisphere. Smithsonian Contrib. Paleobiol., Washington D.C., 41, 1-43. HAWKINS C.M. (1981) - Efficiency of organic matter absorbtion by the tropical Diadema antillarum PHILIPPI fed non-macrophytic algae. Journ. exp. mar. Biol. EcoL, 49, 245-253. HERTLEIN L.G. & GRANT U.S. (1944) - The Cenozoic Brachiopoda of Western North America. PubL Univ. California, Math. phys. Sci., Los Angeles, 3, 1-236. JACKSON J.B.C., GOREAU T.F. & HARTMAN W.D. (1971 ). Recent brachiopod-coralline sponge communities and their paleoecological significance. Science, 173, 623625. LOGAN A. (1975) - Ecological observations on the Recent articulate brachiopod Argyrotheca bermudana DALL from the Bermuda Platform. Bull. mar. Sci., 25, 186-204. LOGAN A. (1977) - Reef-dwelling articulate brachiopods from Grand Cayman, B.W.I. Proc. third internatl Coral Reef Syrup. Miami Florida, 1 (Biology), 88-93.

LOGAN A. (1979) - The Recent Brachiopoda of the Mediterranean Sea. Bull. Inst. ocean. Monaco, 72 (1434), l - l l 2 . NOBLE J.P.A., LOGAN A. & WEBB G.R. (1976) - The Recent Terebratulina community in the rocky subtidal zone of the Bay of Fundy, Canada. Lethaia, Oslo, 9, 117. NOBLE J.P.A. & LOGAN A. (1981) - Size-frequency distributions and taphonomy of brachiopods: a Recent model. Palaeogeogr. Palaeoclimatol. Palaeoecol., 36, 87-105. PAJAUD D. (1974) - i~cologie des Th~cid~s. Lethaia, Oslo, 7, 203-218. SAMARCO P.W. (1980) - Diadema and its relationship to coral spat mortality : grazing, competition and biological disturbance. Journ. exp. mar. Biol. Ecol., 45, 245272. STEINICH G. (1965) - Die artikulaten Brachiopoden der Rtlgener Schreibkreide (Unter-Maastricht). Paltlontol. Abhandl, Berlin, A2 (1), 1-220. STENTOFT N. (1982) - En sedimentologisk og faunistisk undersogelse af Holocaene for-revs karbonat-sedimenter ved Barbados. Unpubl. masters thesis, University of Copenhagen. STEWART I.R. (1981) - Population structure of articulate brachiopod species from soft and hard substrates. New Zealand Journ. Zool, 8, 197-207. SURLYK F. (1972) - Morphological adaptations and population structures of the Danish chalk brachiopods (Maastrichtian, Upper Cretaceous). Biol. Skr. kgl. Dan. Vidensk. Selsk, Copenhagen, 19 (2), 1-57. SURLYK F. (1982) - Brachiopods from the CampanianMaastrichtian boundary sequence, Kronsmoor (NW Germany). Geol. Jahrb, Hannover, A 61, 259-277. THOMSON J.A. (1927) - Brachiopod morphology and genera (Recent and Tertiary). New Zealand Board Sci. Art. Manual, Wellington N.Z., 7, 1-338.

PLATE 1

Fig. 1 - -

Argyrotheca cf. bermudana dorsal view, x 32.

Fig. 2,3 - -

Argyrotheca crassa dorsal view, x 8, cardinalia a n d b r a c h i d i u m , X 14.

Fig. 4 - -

Cryptopora rectimarginata dorsal view, x 18.

Fig. 5 - -

Argyrotheca barrdttiana dorsal view, x 18.

Fig. 6,7 - -

Terebratulina cailleti dorsal views o f n a r r o w a n d wide forms, x 6.

Fig. 8 - -

Brachial valve o f Crania pourtalesi, x 7.

Fig. 9 - -

Brachial valve o f Crania costate sp., x 7.

Fig. 10,11 - -

Platidia anomioides cardinalia a n d b r a c h i d i u m , x 42 : a n d b r a c h i a l valve, x 20.

N.B. Note varying scale.

Geoblos M(~m. special 8

pl. 1
U. Asgaard & N. Stentoft

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;:.~

~ ii i
~

,.:,~;
....... ... ....

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4 2

PLATE 2

Fig. 1,2 - -

Brachial and pedicle valve o f

Platidia anomioides, x 21 a n d x 16.

Fig. 3 - -

Platidiaclepsydra dorsal view, x 13. Argyrotheca schrammi o n its substrate, side view a n d dorsal view, x 35.

Fig. 4,5 - -

Fig. 6,7 - -

Brachial a n d pedicle valve o f

Gryphus bartletti, x 3 a n d x 2.5.

Fig. 8 - -

Details of w o r n brachial valve o f

Megerlia echinata, x 10.

N.B. Note varying scale.

Geobios M~m. special 8

pl. 2
U. Asgaard & N. Stentoft

x\
"".~'~::.,. ~. 9 -L

....... ~ " ~,
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