DIVERSITY AND ECOLOGICAL SIGNIFICANCE OF DEEP-BURROWING MACROCRUSTACEANS IN COASTAL TROPICAL WATERS OF THE AMERICAS (DECAPODA: THALASSINIDEA) DARRYL L. FELDER cology of thalassin- row construction by the laomediid idean shrimp in in- INFRAORDER: THALASSINIDEA Latreille, 1831 Axianassa australis Rodrigues and tertidal and subtidal Shimizu in Brazil (Dworschak and marine and estuarine habitats has Superfamily - Thalassinoidea Dana, 1852 Rodrigues, 1997) and recruitment and gained growing attention in literature Family - Thalassinidae Dana, 1852 growth of the callianassid Lepidoph- of the last decade. Most recent eco- thalmus sinuensis Lemaitre and Rodri- logical focus has been accorded to a Superfamily - Callianassoidea Dana, 1852 gues on the Caribbean coast of Co- limited set of genera assignable to * Family - Callianassidae Dana, 1852 lombia (Nates and Felder, 1999). Eco- several of the eleven families com- Callianassa sensu stricto, Biffarius, logical studies at the community and prising this infraorder (see * in box). Neotrypaea, Trypaea, Lepidophthalmus, ecosystems level form a larger body In accord with Tudge et al. (2000), Callichirus, Sergio, Nihonotrypaea, Glypturus, of recent work, which calls attention the preferred nomenclature does not Corallianassa, Eucalliax, Neocallichirus, to varied roles of thalassinidean popu- conform to that proposed by Sakai “C.” tyrrhena, “C.” candida, “C.” filholi, lations in determining assemblage (1999). Also, as used here, the abbre- “C.” truncata,“C.” kraussi, “C.” laurae structure and their involvement in viation “C.” indicates species that re- Family - Ctenochelidae Manning and Felder, 1991 biogeochemically modulated processes main treated under the genus Calli- Family - Callianideidae Kossmann, 1880 (Table II). Major contributions in this anassa sensu lato, despite recent * Family - Laomediidae Borradaile, 1903 arena over the last decade have cen- taxonomic restriction of that genus Axianassa, Jaxea tered on population dynamics, com- which excludes these species (see Family - Thomassiniidae de Saint Laurent, 1979 munity composition, bioturbation, re- Manning and Felder, 1991). * Family - Upogebiidae Borradaile, 1903 lated geochemical effects, nutrient cy- As here evident, Upogebia cling, and food webs. Among these, genera treated in modern ecological works on populations of (at least in works are for the most part calli- Superfamily - Axioidea Huxley, 1879 part) the callianassids Eucalliax jo- anassids, a single family that in- * Family - Axiidae Huxley, 1879 nesi (Heard) and Neocallichirus ra- cludes a large number of intertidal Axius, Axiopsis thbunae (Schmitt) in the British Virgin and shallow subtidal representatives. * Family - Calocarididae Ortmann, 1891 Islands (Murphy and Kremer, 1992), Recent treatments of thalassinidean Calocaris the callianassids Glypturus acan- reproductive ecology have focused on Family - Micheleidae Sakai, 1992 thochirus Stimpson, Neocallichirus recruitment, settlement stimuli, repro- Family -Strahlaxiidae Poore, 1994 grandimana (Gibbes), Corallianassa ductive output and environmental longiventris (A. Milne Edwards) and constraints on reproduction of calli- axiid Axiopsis serratifrons (A. Milne anassids, while those in physiological and nants in burrow waters for callianassids Edwards) in Belize (Dworschak and Ott, behavioral ecology have centered on ven- and members of several other families 1993) and the callianassid Lepidophthal- tilatory mechanisms, burrow construction (Table I). Scant attention among these mus sinuensis on the Caribbean coast of and metabolic constraints imposed by hy- has been afforded to tropical populations Colombia (Nates and Felder, 1998) do poxia and reduced nutrients or contami- of the Americas except in studies of bur- demonstrate the potential significance of KEYWORDS / Thalassinideans / Mudshrimp / Bioturbation / Burrowing / Recibido: 09/04/2001. Aceptado: 10/08/2001 Darryl L. Felder. Ph.D., Louisiana State University. Professor of Biology, University of Louisiana –Lafayette (ULL). Director of ULL–Laboratory for Crustacean Research. Associate Research Scientist, NMNH, Smithsonian Institution, Washington, DC. Address: Department of Biology, University of Louisiana, Lafayette, LA 70504- 2451, USA. e-mail: [email protected] 440 0378-1844/01/10/440-10 $ 3.00/0 OCT 2001, VOL. 26 Nº 10 TABLE I such populations in community and eco- SELECTED STUDIES ON ECOLOGY OF THALASSINIDEAN systems level dynamics of the American CRUSTACEANS SINCE 1990 tropics. One implication of these studies, be they focused on American Location Habitat Focus Taxa Authors tropical or other environments, is that deep burrowing thalassinidean shrimp are re- Japan intertidal sand settling, recruitment Nihonotrypaea 1 peatedly observed to dominate processes Japan intertidal sand brooding, development Nihonotrypaea 2 in a number of intertidal and subtidal Japan intertidal-sh subtidal larval dispersal Nihonotrypaea 3 spp. 3 habitats, where they often form dense ag- Japan intertidal-sh subtidal burrow construction Nihonotrypaea 2 spp. 4 gregations. Aggregation may be favored by Japan intertidal patterning, density Nihonotrypaea 5 larvae adapted for estuarine retention or Japan intertidal competition/snails Nihonotrypaea 6 abbreviated development; in at least some Japan intertidal competition/isopods Nihonotrypaea 7 cases, specific sedimentary cues favor Japan intertidal competition/eels Nihonotrypaea 8 dense and gregarious settlement of lar- Japan intertidal abundance, life history Nihonotrypaea 9 vae. Physiology and behavior may be Australia intertidal ventilation, respiration Biffarius, Trypaea 10 adapted to facilitate burrowing within Australia intertidal community interactions Trypaea 11 and exploiting of hypoxic or even anoxic Australia intertidal associated bivalves/redox Trypaea 12 sediments for nutrients, with controlling Australia intertidal sand burrow form, function Biffarius 13 consequences on populations of associated Australia intertidal sand food sources, burrows Biffarius, Trypaea 14 microbes, macrobenthos, rooted autotro- Australia intertidal sand bioturbation, carbon, Biffarius 15 phs, or planktonic autotrophs and het- microbes erotrophs of the overlying water column. New Zealand intertidal sand population dynamics “C.” filholi 16 Through their modulation of physiochemi- New Zealand intertidal sand bioturbation effects “C.” filholi 17 cal parameters within porewaters and the New Zealand intertidal sand effects on communities “C.” filholi 18 overlying water column, the activities of New Zealand intertidal sand reproduction “C.” filholi 19 thalassinidean populations may touch upon South Africa intertidal mudflat hydrodynamics Upogebia 20 virtually all aspects of productivity in shal- South Africa intertidal growth, production Upogebia 21 low marine systems. Regardless, the scope South Africa intertidal hydrocarbon accumulation “C.” kraussi 22 of this impact remains largely unmentioned South Africa intertidal harvest impacts “C.” kraussi, 23 in traditional treatments of coastal marine Upogebia ecology, and the diversity of thalassinide- South Africa intertidal-sh subtidal detrital food webs “C. kraussi 24 ans and their commensal associates re- South Africa- intertidal feeding, gut flora Upogebia 2 spp., 25 sponsible for these impacts remains mark- Australia-N Trypaea Neotrypaea, edly underestimated. Am/P “C.” kraussi Aside from compiling Red Sea sh subtidal organic cycling “C.” laurae 26 major ecological works on thalassinideans Mediterranean subtidal sediment oxygen flux “C.” truncata 27 from the last decade, the present paper Mediterranean subtidal biogeochemistry “C.” truncata 28 briefly treats in further detail three gen- Mediterranean intertidal-sh subtidal environ effects on larvae “C.” tyrrhena 29 era among these animals, all now known Mediterranean intertidal-sh subtidal facultative lecithotrophy “C.” tyrrhena 30 to occur widely and often in dense accu- Mediterranean intertidal-sh subtidal pollution, metabolic “C.” tyrrhena 31 mulations throughout the American trop- Mediterranean intertidal-sh subtidal reproductive output “C.” tyrrhena 32 ics, though species-level systematics in Mediterranean intertidal reproduction, growth Upogebia 33 this region remain in part unresolved. Re- Mediterranean intertidal-sh subtidal burrow construction “C.” tyrrhena, 34 cent and ongoing studies of selected spe- “C.” candida cies among these genera, some not previ- Mediterranean intertidal population dynamics “C.” tyrrhena, 35 ously published, are herein used to further “C.” candida exemplify ecological roles of thalassinide- UK-N Sea subtidal gill morph/function Upogebia 3 spp., 36 ans, especially in the context of environ- Calocaris Callianassa, mental issues bearing on populations of Jaxea, Axius the American tropics. UK-N Sea subtidal ventilation, respiration Calocaris 37 UK-N Sea subtidal sulfide adapt, metabolic Calocaris, Jaxea, 38 Populations of Lepidophthalmus Callianassa along Warm-Temperate to Tropical UK-N Sea subtidal burrowing Callianassa 39 Coastlines UK-N Sea subtidal burrowing, feeding adapt Callianassa, Jaxea, 40 Upogebia Emphasis on this genus UK-N Sea subtidal activity, bioturbation Callianassa 41 is warranted because of 1) its potentially UK-N Sea subtidal feeding, mechanics Callianassa 42 dominant role in ecosystem processes of UK-N Sea subtidal feeding, sorting, gut flora Upogebia 3 spp., 43 tropical estuaries, 2) its previously under- Calocaris Callianassa, estimated diversity and regional endemism Jaxea, Axius in these habitats,