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Reproductive Biology of a Wood-Boring Isopod, Sphaeroma Terebrans, with Extended Parental Care

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Reproductive Biology of a Wood-Boring Isopod, Sphaeroma Terebrans, with Extended Parental Care Marine Biology (1999) 135: 321±333 Ó Springer-Verlag 1999 M. Thiel Reproductive biology of a wood-boring isopod, Sphaeroma terebrans, with extended parental care Received: 9 December 1998 / Accepted: 24 June 1999 Abstract The isopod Sphaeroma terebrans Bate, 1866 in size while in the maternal burrow, and juveniles of burrows in aerial roots of the red mangrove Rhizophora similar sizes could also be found in their own burrows. mangle L. The burrows serve as shelter and as a repro- Males did not participate in extended parental care, ductive habitat, and females are known to host their since most of them left the females after copulation. ospring in their burrows. I examined the reproductive Many females that were born in the summer produced biology of S. terebrans in the Indian River Lagoon, a one brood in the fall and a second during winter/early shallow lagoon stretching for 200 km along the At- spring. Females that were born in the fall produced one lantic coast of Florida, USA. Reproductive isopods were brood during spring/early summer, but then probably found throughout the year, but reproductive activity died. Extended parental care in S. terebrans is short was highest in the fall and during late spring/early compared to other peracarid crustaceans. It is concluded summer. During the latter periods, large numbers of that this reproductive strategy in S. terebrans serves subadults established their own burrows in aerial roots. primarily to shelter small juveniles immediately after The average numbers of S. terebrans per root were high they emerge from the female body, when their exoskel- during the fall, but decreased during the winter and eton is still hardening and their physiological capabilities reached lowest levels at the end of the summer. Females are still developing. Thus, in S. terebrans, extended pa- reached maturity at a larger size than males, but also rental care probably aids to protect small juveniles from grew to larger sizes than the males. The average size of adverse physical conditions in their subtropical intertidal females varied between 8 and 10 mm, the average size of habitat. males between 6.5 and 8.5 mm. The number of embryos female)1 was strongly correlated with female body length. No indication for embryo mortality during de- Introduction velopment was found. Parental females (i.e. with juve- niles in their burrows) hosted on average between 5 and Extended parental care is known from peracarid crus- 20 juveniles in their burrows (range 1 to 59). Most ju- taceans inhabiting a variety of habitats, such as seagrass veniles found in female burrows were in the manca stage and algal beds (Brearley and Walker 1995, 1996; Aoki and 2 to 3 mm in body length. Juveniles did not increase 1997), shallow soft-bottoms (Mattson and Cedhagen 1989; Thiel et al. 1997), and hard-bottoms (Shiino 1978). The most long-lasting parental care has been found in peracarid species that inhabit burrows, tubes or nests. In Communicated by N.H. Marcus, Tallahassee these species, juveniles may experience parental care for several months (Thiel 1999a). In free-ranging species 1 M. Thiel that do not host their ospring in a parental dwelling Smithsonian Marine Station, 5612 Old Dixie Highway, but on their body, the parental care period is short, Fort Pierce, Florida 34946, USA usually not exceeding a few weeks (Aoki 1997; Thiel 1999a). For most peracarid species with extended pa- Present address: rental care, the reasons for the break-up of the parent± 1 Facultad Ciencias del Mar, Universidad Catolica del Norte, Larrondo 1281, ospring association are not known. Campus Guayacan, Parental dwellings may not be stable enough to per- Coquimbo, Chile mit long-lasting coexistence of parents and their o- Fax: 0056 51 209812 spring. Burrows or tubes in soft-bottoms, and nests e-mail: [email protected] in exposed seagrass or algal beds may be frequently 322 destroyed, resulting in the separation of parents and Careful observations by Messana et al. (1994) only their ospring. The stability of a parental dwelling could recently revealed that the wood-boring isopod Sphae- thus have a substantial in¯uence on the duration of roma terebrans engages in extended parental care, during parental care. Among the most stable dwellings in the which juveniles live with their parents in so-called family marine environment are those that are established in burrows. Females host their ospring in their burrows biotic microhabitats such as sponges, ascidians, bra- before these have fully developed their boring capabili- chiopods, bivalves, roots, rhizomes and wood. Several ties (Thiel 1999d). It is not well known how many ju- peracarid (and other) crustacean species are known to veniles live in one family burrow, or how large they grow raise their ospring within biotic microhabitats (Wol during this period of extended parental care. In several 1976; Vader and Beehler 1983; Gonzalez and Jaramillo other peracarid crustaceans boring in wood (Limnoria 1991; Conlan and Chess 1992; Messana et al. 1994). spp.: Menzies 1954, 1957), wood-like substances such as These biotic microhabitats may be suciently stable to kelp stipes (Peramphithoe stypotrupetes: Conlan and allow a long period of extended parental care, possibly Chess 1992; Chess 1993), or seagrass blades (Brearley resulting in the occurrence of very large ospring (Duy and Walker 1995, 1996), males have frequently been 1996; Thiel 1999b). However, many biotic microhabitats found to cohabit with females. These observations sug- are small (salps, bivalves, ascidians, roots and rhizomes), gest that male boring peracarids may be involved in and thus do not provide much space for large families. parental care to a higher degree than males of other Usually, only one or two parents and a single clutch of peracarid species. It is not unusual to ®nd pairs of fe- juveniles are found in these small biotic microhabitats male and male S. terebrans within one burrow (Estevez (Vader and Beehler 1983; Gonzalez and Jaramillo 1991). and Simon 1975; Estevez 1978; Messana et al. 1994), but Besides sponges, large pieces of wood constitute another it is not known whether males of this species actively biotic microhabitat in the marine environment that participate in parental care or not. In the present study, could accommodate large numbers of closely related I paid ample attention to collect S. terebrans from in- individuals. Wood is utilized both as habitat and as food dividual burrows, to obtain a better understanding of by a variety of dierent peracarid species (Menzies 1954; the reproductive biology of these wood-boring isopods. Eltringham and Hockley 1961; KuÈ hne and Becker 1964; John 1970; Rotramel 1975; Ribi 1982; Perry and Brusca 1989). Large pieces of wood often host numerous indi- viduals of similar sizes. Due to their destructive activities Materials and methods in arti®cial wood structures, a vast amount of research has been conducted on wood-boring peracarids (see Study area Clapp and Kenk 1963), but surprisingly few details are The study was conducted in the Indian River Lagoon, which ex- available about their reproductive biology. tends 200 km along Florida's Atlantic coast (27°32¢N; 80°20¢W). Sphaeroma terebrans is a wood-boring isopod, whose Sphaeromatid isopods occur in various intertidal and shallow main distribution is in mangrove forests around the subtidal habitats of the lagoon (Kensley et al. 1995). The lagoon was arti®cially modi®ed during the 1950s and 1960s as part of a world, where it preferentially dwells in the aerial roots of program designed to control mosquito populations. The extensive red mangrove trees (see e.g. Harrison and Holdich 1984; modi®cations included the construction of large impoundments Villalobos et al. 1985). This isopod also builds burrows that are temporarily ¯ooded and then drained (De Freese 1995; in other substrates such as dead wood and marshgrass Larson 1995). The formation of dams and reduced saltwater-ex- change within the impoundments substantially altered the intertidal rhizomes (Estevez 1994). It is generally assumed that the habitats in the lagoon (De Freese 1995). In many parts of the burrow serves as a shelter for the isopod rather than as a lagoon subjected to extensive remodeling, red mangrove trees food resource (John 1970). The size of a burrow gener- (Rhizophora mangle L.) now only grow in a narrow fringe along the ally does not exceed that of its inhabitant, and boring outer side of these impoundments. All collections referred to herein were made along these mosquito impoundments. The collection site activity decreases drastically after the isopod has estab- is 5 km north of the Fort Pierce Inlet. Water temperatures range lished a burrow in whatever substrate is available (John between 15 and 34 °C, and salinities between 17.5 psu (practical 1970). In experimental studies, S. terebrans apparently salinity units) during the winter months and 35 psu during the survived best in waters that contained planktonic algae summer months (SA Reed unpublished data). The oxygen levels in )1 (John 1970). The setation and morphology of the the lagoon water range between 2mll (late spring/summer) to 12 ml l)1 (fall), but usually vary between 5 and 8 ml l)1. The maxilliped and the ®rst three pereopods of S. terebrans water level in the lagoon is highest during the fall, and reaches low strongly suggest that this isopod ®lters its food out of the values during the summer months (Provost 1973). water in a similar manner as its boring congener S. quoyanum (Rotramel 1975). The aerial roots of red mangroves thus provide an ideal habitat for S. terebrans Sampling for several reasons, e.g. (1) the anterior parts of aerial The aerial roots of the red mangrove Rhizophora mangle L., con- roots are very soft (Gill and Tomlinson 1975), and it is taining burrows of Sphaeroma terebrans Bate, 1866, were collected therefore easy to establish burrows in them; (2) aerial monthly along the south-eastern shore of a shallow bay in the roots are frequently ¯ushed by water (during high tide), lagoon.
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