Lobophora variegata - IngentaConnect

BULLETIN OF MARINE SCIENCE, 68(2): 207–219, 2001

THE IMPORTANCE OF LOBOPHORA VARIEGATA (PHAEOPHYTA: DICTYOTALES) AS A HABITAT FOR SMALL JUVENILES OF PANULIRUS ARGUS (DECAPODA: PALINURIDAE) IN A TROPICAL REEF LAGOON Patricia Briones-Fourzán and Enrique Lozano-Álvarez ABSTRACT The fishery for the spiny lobster Panulirus argus in the area around Puerto Morelos is one of the least productive in the state of Quintana Roo (Mexican Caribbean coast). Density of postalgal juveniles of P. argus (i.e., juveniles ≥ 15 mm carapace length, CL) is low in the shallow areas of the reef lagoon at Puerto Morelos despite high annual indices of puerulus influx into the lagoon as measured with artificial collectors. We hypothesized that a population bottleneck might exist at the algal juvenile phase (juveniles 6–15 mm CL, residing in vegetated shallow areas), owing to a lack of suitable habitats for the settlement of incoming postlarvae and the subsequent survival of algal juveniles. At five sites within the reef lagoon, algal juveniles were collected with an epibenthic net during June 1995 (early summer) and March 1997 (late winter). In March, we estimated, in the same five sites, the density of two species of seagrass and five functional-form groups of macroalgae, as well as the biomass of Lobophora variegata (ruffled form), a drift phaeophyte, which grows in extremely dense mats over a number of substrates. Significantly more algal juveniles were obtained in March than in June but this temporal pattern was apparently not related to peaks in influx of postlarvae into the reef lagoon. The abundance and distribution of algal juveniles in March was not related to the density of seagrass or algae, but was significantly related to the biomass of L. variegata. The ruffled form of L. variegata is very abundant in seagrass meadows throughout the Caribbean; hence, it may represent an important habitat for pueruli and algal juveniles of P. argus in other areas as well. Density of algal juveniles in the reef lagoon was 146 ha−1 in June and 263 ha−1 in March. Considering a conservative average of 150 algal juveniles ha−1, and a residence of 2 mo in the algal habitats, the reef lagoon can harbor around 900 algal juveniles ha−1 in a year, a figure comparable to productive areas for lobster such as the Florida Keys, USA. Therefore, the lobster population bottleneck in Puerto Morelos does not occur at the algal-juvenile phase.

The spiny lobster Panulirus argus (Latreille) is a major fishing resource throughout the Caribbean Sea. In Mexico, it is heavily fished on the eastern margin of the Yucatán Peninsula (State of Quintana Roo). Along the coast of Quintana Roo, some lobster fishing areas are very productive, whereas in others P. argus is rather scarce. The area around Puerto Morelos (20°51'N, 86°55'W) is one of the latter. Because of poor catches in this area, fishermen must travel several kilometers to the north and south of Puerto Morelos to catch lobsters (Padilla-Ramos and Briones-Fourzán, 1997). After a protracted oceanic larval life, the final larval stage of P. argus metamorphoses into a postlarva called puerulus, which returns to the coast and settles in shallow, vegetated areas where it molts and takes residence for a few months. Reported settlement habitats for pueruli of P. argus include mangrove roots (Acosta and Butler, 1997), seagrass meadows (Buesa, 1965), and especially beds of rhodophyte macroalgae of the genus Laurencia (Marx and Herrnkind, 1985a). Laurencia is considered a prime habitat for settled pueruli and algal juveniles [i.e., juveniles ≤15 mm carapace length, CL, (Butler 207

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and Herrnkind, 1997)] along the Florida Keys, where it is very abundant. However, although Laurencia occurs throughout the Caribbean, the hard bottoms where this alga thrives are not equally common in the Caribbean, where most of the coasts typically have mangrove-fringed shorelines and seagrass beds (Acosta and Butler, 1997). The reef lagoon at Puerto Morelos extends from the coastline out to a coral reef that lies slightly diagonally at a distance of 700–1300 m. Mangroves have no direct contact with the sea because they are separated by a sandbar 100–200 m in width, which forms the shoreline of the reef lagoon. Depth in the lagoon ranges from 0.50). Monthly mean water temperatures in the reef lagoon during January 1995–March 1997 ranged from 24.5 to 30.1°C (Fig. 2). Daily water temperature ranged from 27.8 to 31.0 (mean: 29.0°C) in May–June 1995, and from 24.0 to 27.5 (mean: 25.2°C) during FebruTable 2. Size distribution and mean age per size class of algal juveniles of Panulirus argus in the reef lagoon at Puerto Morelos in summer and winter. Age was estimated on the basis of a mean size of pueruli at settlement of 6.0 mm carapace length (CL) (Negrete-Soto, 1994) and a growth rate of 2.0 mm CL per month in the summer and 1.32 mm CL per month in the winter (Lellis and Russell, 1990). Size class (mm CL) 6.1−7.0 7.1−8.0 8.1−9.0 9.1−10.0 10.1−11.0 11.1−12.0 12.1−13.0 13.1−14.0 14.1−15.0 15.1−16.0 16.1−17.0 17.1−18.0 18.1−19.0 Total

Early summer Frequency Mean age after (n) settlement (mo) 5 0.81 7 0.94 2 1.06 0 1.19 0 1.31 2 1.44 0 1.56 0 1.69 0 1.81 0 1.94 0 2.06 0 2.19 1 2.31 17

Late winter Frequency Mean age after (n) settlement (mo) 10 0.89 16 1.02 2 1.16 8 1.30 0 1.43 2 1.57 1 1.71 1 1.84 1 1.98 1 2.12 0 2.25 0 2.39 1 2.53 43

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Figure 2. Monthly mean water temperature (top line) and mean (± SE) number of postlarvae (pueruli) of Panulirus argus per collector (bottom set of lines) in the reef lagoon at Puerto Morelos, January 1995–March 1997.

ary–March 1997. Pueruli of P. argus at Puerto Morelos settle at a mean size of 6.0 mm CL (Negrete-Soto, 1994). Algal juveniles 6–11 mm CL kept at 30 and 24°C grow an average of 2.0 mm CL and 1.32 mm CL mo−1, respectively (Lellis and Russell, 1990). Therefore, algal juveniles ≤10.0 mm CL had an estimated age of 0.81–1.19 mo from settlement in our June 1995 samples, and 0.89–1.30 in March 1997 (Table 2). Peak settlements on artificial collectors in 1995 occurred in April, August, and November-December, whereas in 1996, two peaks were apparent in May and July (Fig. 2). These peaks did not relate to the disparate abundance of algal juveniles ≤10 mm CL in the summer and winter in the algal benthic habitats in the lagoon (i.e., the latter could not be directly related to the strength in the influx of pueruli 1–2 mo in advance as measured with artificial collectors). DISTRIBUTION OF ALGAL JUVENILES VS VEGETATION.—Density of the groups of vegetation differed per site (Table 3). Both species of seagrass were relatively dense in sites 1–3, but were absent in site 4. At site 5, S. filiforme was absent, and T. testudinum had the highest density but its blades were much shorter than in sites 1, 2 and 3. The macroalgal components also varied with site. The density of Penicillus-type and Halimeda algae was the highest in all sites, but differed widely among sites. For example, at site 5, only Halimeda spp. were present. The rest of the algal groups contributed little to plant density at all sites. The brown alga L. variegata had a high biomass in sites 1 and 2, and lower in site 3, whereas it was absent in sites 4 and 5. These results agree with the known distribution and density of vegetation in this reef lagoon (van Tussenbroek, 1994, 1995; Ruiz-Rentería et al., 1998). Because of the nocturnal behavior of P. argus, and the mode of operation of the epibenthic net, only the night samples were considered representative of the population of algal

1 2 3 4 5

Site

Density: mean number of foliar groups (seagrasses) or thalli (macroalgae) m−2 Seagrass Total Macroalgae Total T. testudinum S. filiforme seagrass Spongy algae Halimeda-type Penicillus-type Udotea-type macroalgae 134.4 504.8 639.2 12.0 20.8 35.2 4.0 72.0 119.2 440.8 560.0 12.8 159.2 38.2 1 .6 211.8 156.8 350.4 507.2 4.0 96.0 6.8 0.8 107.6 0 0 0 3.2 44.0 56.0 0 103.2 198.4 0 198.4 0 100.0 7.8 1.6 109.4

Biomass of drift algae (g m−2 dry weight) Lobophora variegata 25.02 24.66 5.03 0 0

Table 3. Density of two species of seagrasses and four functional-form groups of macroalgae, and biomass of drift algae (Lobophora variegata) in five sampling sites, reef lagoon at Puerto Morelos, Mexico, during March 1997 (late winter).

BRIONES-FOURZÁN AND LOZANO-ÁLVAREZ: DRIFT ALGAE AS HABITAT FOR JUVENILE SPINY LOBSTER

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Table 4. Regression coefficients and test of significance at α = 0.05 of individual regressions of density of seagrasses (Thalassia testudinum and Syringodium filiforme), density of macroalgae (several species), and biomass of the brown alga Lobophora variegata versus density of algal juveniles of the spiny lobster Panulirus argus in the reef lagoon at Puerto Morelos, Mexico (df = 3 in all regressions; n.s. = non significant; s. = significant). Regression Density of seagrasses vs density of juveniles Density of macroalgae vs density of juveniles Biomass of Lobophora variegata vs density of juveniles

r2 0.224 0.202 0.843

F 1.204 0.758 16.473

P > 0.25 (n.s.) > 0.50 (n.s.) < 0.025 (s.)

juveniles in the lagoon. In the mid-lagoon zone, trawls ranged in length from 48 to 64 m (mean ± SD = 54.4 ± 4.6 m), and in the backreef area from 50 to 66 m (56.7 ± 6.0 m). However, we considered 60 m as the average trawl length to be as conservative as possible in our estimates of the density of algal juveniles. Thus, the average area covered by the 10 trawls in each site was 342 m2 [net width (0.57 m) × length of trawl (60 m) × number of trawls (10)]. The density of algal juveniles at each site (Table 1) was separately related to the total density of seagrass, the total density of algal groups, and the biomass of L. variegata (Table 3), by fitting independent regressions after transforming all sets of data to log (x + 1). Only the relationship with the biomass of L. variegata was significant (Table 4). DISCUSSION Algal juveniles of P. argus inhabit vegetated habitats in shallow areas and are typically solitary (Marx and Herrnkind, 1985a). During the day, these juveniles remain either in the deepest recesses of the algae, beneath the algae (Butler and Herrnkind, 1994), or buried in the substrate, leaving only the antennae protruding (Calinsky and Lyons, 1983). During the night, these small juveniles roam their algal refuges and nearby areas to feed on small invertebrates. This behavior could explain the smaller numbers of algal juveniles obtained with the epibenthic net in our day samples, and warrants the use of only the night samples to estimate their density. Algal juveniles occurred throughout the reef lagoon at Puerto Morelos but they were more abundant in the mid-lagoon zone (sites 1–3). This suggests that the structural complexity of the seagrass-algal community, which was higher in the mid-lagoon zone, was important in the spatial distribution of the small lobster juveniles. Plant architecture (the tridimensional form of growth of a plant) is an important factor in the complexity of benthic habitats for invertebrate communities in shallow areas (den Hartog, 1982, and references therein; Stoner and Lewis, 1985; Edgar et al., 1994). Seagrass meadows in the Caribbean are dominated by one or more species of seagrass, but these plants are structurally simple, with either flat, elongated blades (T. testudinum), or elongated thin cylindrical shoots (S. filiforme). It is the associated macroalgae which add to the structural complexity of the benthic microhabitats in seagrass beds. Complexity in sites 1–3 was due mainly to the extensive, dense mats of L. variegata, which grows attached to a vast array of substrates, including other algae (Coen and Tanner, 1986; Rodríguez-Almazán, 1997). In the Florida Keys, Marx and Herrnkind (1985a) found that small juveniles of P. argus, up to 20 mm CL, were more abundant in or beneath red macroalgae (Laurencia spp.),


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which form intricately branched clumps. These algae also provided small juveniles with abundant prey in the form of small invertebrates. Thus, Marx and Herrnkind (1985b) concluded that the protection of otherwise susceptible young stages of P. argus from predators and physical stress while providing access to food enhanced the value of algae as refugium. This lead to the use of the term ‘algal juveniles’ for these small stages of P. argus (Butler and Herrnkind, 1997). In Cuba, Serpa-Madrigal and Areces (1995) found that algal juveniles of P. argus 7–12 mm CL showed a significant preference for the laminar chlorophyte alga Ulva fasciata over algae with cylindrical or flattened branches forming a polycryptic habitat (the rhodophytes Bryothamnion triquetrum, Acanthophora spicifera and Gracilaria mammillaris), and algae with fleshy stipes and thick, nude cylindrical branches (Kappaphycus alvarezii and K. striatum). These authors suggested that the convoluted laminar architecture of U. fasciata provided these juveniles with a better shelter against predators than algae with an interstitial architecture. Similarly, Holmlund et al. (1990) found that algae such as Padina gymnospora and Ulva spp. offered amphipods better protection against piscine predators than interstitial algae of the genera Hypnea, Gracilaria and Sargassum. The higher abundance of algal juveniles in the mid-lagoon zone at Puerto Morelos was related to the biomass of L. variegata. This brown alga is widely distributed and occurs from tropical to warm-temperate zones. It grows in three different morphological forms: (1) adherent crusts, (2) flat decumbent blades, and (3) erect, ruffled, often unattached blades (Norris and Bucher, 1982; de Ruyter van Steveninck et al., 1988). The ruffled form is characteristic of calm, shallow waters (Littler et al., 1989) and dominates Thalassia beds distant from reef structures, where grazing pressure is relatively low (Coen and Tanner, 1989). Only the ruffled form is found in the mid-lagoon zone at Puerto Morelos (Rodríguez-Almazán, 1997; Reyes-Zavala, 1998), where water is calmer compared to the backreef zone (Ruiz-Rentería et al., 1998). The ruffled form of L. variegata resembles ruffled loose-leaf lettuce (hence its name) with lobed margins on the broad blades (Littler et al., 1989), and often forms ball-like clumps as a consequence of continued lateral blade growth (Coen and Tanner, 1989). Therefore, the ruffled form of L. variegata has more surface area per unit weight than other algae with less foliar area. L. variegata occurs either attached to various substrates (including other algae) forming extensive and dense carpets (Rodríguez-Almazán, 1997), or as unattached individuals which roll along the sand bottom in large aggregations. In stormy weather, clumps of L. variegata may detach and drift to other parts of the lagoon where they remain as loose mats (Rodríguez-Almazán, 1997). Of the several functional groups of macroalgae in the Puerto Morelos reef lagoon, the most structurally complex in the mid-lagoon area are the drift algae. This group also includes the genera Dictyota and Laurencia but is overwhelmingly dominated by the ruffled form of L. variegata (Reyes-Zavala, 1998; van Tussenbroek and Reyes-Zavala, 1998). Although we studied only the vegetation in our sampling sites during the winter, L. variegata consistently exhibits the highest percentages in biomass throughout the midlagoon [up to 52% of total plant biomass, (Reyes-Zavala, 1998)]. In addition, sites in the lagoon dominated by L. variegata exhibit the highest abundances of small invertebrates that are prey to algal juveniles (Briones-Fourzán and Estrada-Olivo, 1998; MonroyVelázquez, 2000). Hence, L. variegata represents an important habitat for P. argus pueruli and algal juveniles throughout the reef lagoon at Puerto Morelos. The ruffled form of L.

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variegata is common in other parts of Quintana Roo (Aguilar-Rosas, 1990) and is one of the most conspicuous algae in Thalassia beds in Belize (Lewis and Wainwright, 1985); consequently, its importance as a habitat for small juveniles of P. argus may also be considerable in other areas of the Caribbean. In the Florida Keys, the extensive Laurencia habitat is used by successive classes of new P. argus recruits, once the algal juveniles outgrow the algal cover and move to other types of habitats with crevice-type shelters (e.g., sponges, solution holes, gorgonians, hard corals, rocky outcrops, etc.). Marx and Herrnkind (1985a) proposed that the size at which juveniles move from the algal habitat to the crevice-type shelters could be related to the abundance and distribution of the latter. For example, at Burnt Point, Florida, where crevice-type shelters abound, this change in habitat occurs at sizes of 15–20 mm CL (Marx and Herrnkind 1985a) whereas at Elliot Key, Florida, where crevice-type shelters are scarcer, juveniles do not change habitat until they reach sizes of 25–30 mm CL (Andrée, 1981). In the reef lagoon at Puerto Morelos, Lozano-Álvarez et al. (1998) found juveniles 12–15 mm CL already residing in crevice-type shelters, and in the present study, the number of algal juveniles >10 mm CL in the vegetated habitats was very small, indicating that few of them remain in the algal habitat after reaching 10 mm CL. Yet, crevice-type shelters are very scarce in the reef lagoon (Lozano-Álvarez et al., 1998, and unpubl. data). Therefore, instead of relating the change of habitat at such small sizes to a large abundance of crevice-type shelters, we believe it rather reflects the ephemerous nature and dynamics of the habitat for algal juveniles (patches of L. variegata) in this reef lagoon. The dynamics of the mats of L. variegata, growing in dense carpets in calm weather then clumps detaching in severe weather and drifting to other parts of the reef lagoon, could also underlie the temporal variations in the abundance of algal juveniles. Mats of detached algae remain stationary for days to weeks, and induce structural changes to the habitat. They can be either beneficial, increasing habitat complexity, or deleterious, by producing oxygen defficiency caused by algal respiration and bacterial activity (Norkko and Bonsdorff, 1996). It would be interesting to study the effect of unattached mats of L. variegata on the temporal abundance and spatial distribution of algal juveniles of P. argus. Our total night samples per season covered an estimated 1710 m2 [342 m2 per site × number of sites (5)]. By considering the total number of algal juveniles obtained at night in June (25) and in March (45), the overall density of algal juveniles throughout the reef lagoon was estimated as 146 ha−1 in early summer and 263 ha−1 in late winter. Marx and Herrnkind (1985a) estimated a density of 278 algal juveniles ha−1 during May–September in Burnt Point, Florida. Owing to year-round influx of pueruli and rapid growth of the algal stages of P. argus, turnover rates of algal juveniles in the reef lagoon may be high. Considering a conservative average density of 150 algal juveniles ha−1 and a residence of 2 mo in the algal habitats, the reef lagoon could harbor around 900 algal juveniles ha−1 in a year, again, similar to what Marx and Herrnkind (1985a) estimated in the Florida Keys (954 ha−1). Yet, the contrastingly low densities of postalgal juveniles in the reef lagoon at Puerto Morelos (0– 31 ha−1, Lozano-Álvarez et al., 1998) suggest that the population bottleneck does not occur at the algal juvenile phase, but rather at the postalgal phase, and that this could be related to the scarcity of crevice-type shelters for lobsters >12 mm CL in the reef lagoon. Currently, a research program involving the deployment of artificial shelters targeting postalgal juveniles is under way, in which preliminary results show a significant increase


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in density of postalgal juveniles in experimental sites with artificial shelters (LozanoÁlvarez et al., 1998, and unpubl. data). ACKNOWLEDGMENTS We greatly acknowledge the invaluable help in field and laboratory activities of F. Negrete-Soto. C. Barradas-Ortiz and V. Monroy-Velázquez provided much help at different stages of this study; the latter also shared with us data from her Master’s thesis. J. Estrada, E. Cadena, V. Castañeda, and P. Rangel aided in field work. F. Ruiz-Rentería kindly provided the water temperature data, and A. Banaszak reviewed the manuscript. This study was part of project No. 1171-N, funded by CONACYT (National Council for Science and Technology, Mexico) and UNAM (National University of Mexico). The Secretary of the Environment, Natural Resources and Fisheries (SEMARNAP, Mexico), issued permits No. 270295-310-03 and 030298-213-03 to conduct the samplings.

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DATE ACCEPTED: July 24, 2000.

ADDRESS: Universidad Nacional Autónoma de México, Instituto de Ciencias del Mar y Limnología, Unidad Académica Puerto Morelos. Ap. Postal 1152, Cancún, Q. R. 77500 Mexico. E-mail: .