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Floods, Habitat Hydraulics and Upstream Migration of Neritina Virginea (Gastropoda: Neritidae) in Northeastern Puerto Rico

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Floods, Habitat Hydraulics and Upstream Migration of Neritina Virginea (Gastropoda: Neritidae) in Northeastern Puerto Rico Caribbean Journal of Science, Vol. 41, No. 1, 55-74, 2005 Copyright 2005 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez Floods, Habitat Hydraulics and Upstream Migration of Neritina virginea (Gastropoda: Neritidae) in Northeastern Puerto Rico JUAN F. BLANCO1 AND FREDERICK N. SCATENA2 1Department of Biology, University of Puerto Rico, Rio Piedras Campus, P.O. Box 23360, San Juan, Puerto Rico 00931–3360. Corresponding author: [email protected], [email protected] 2Department of Earth and Environmental Sciences, University of Pennsylvania, Philadelphia, PA 19104–6313 ABSTRACT.—Massive upstream migrations of neritid snails (Neritidae: Gastropoda) occur in tropical and subtropical streams worldwide, but their seasonality and proximate causes are unknown. We monitored massive upstream migrations of Neritina virginea for 99 weeks, and conducted a detailed study of snail density, size, and hydraulic descriptors in lower Río Mameyes, northeastern Puerto Rico. The study assessed the 1) timing and seasonality of upstream migration, 2) size composition of migratory aggregations, 3) patterns of habitat use, and 4) role of floods on upstream migration. Massive upstream migrations (500–3000 ind/m2) were observed in 44 of 99 weeks of observation. While N. virginea aggregations occurred at random time intervals, they were clumped during rainy periods. Migratory aggregations consisted mostly of small individuals (5–7 mm). Greater mean density was consistently observed in a stable riffle than in an unstable run (115.7 and 17.8 ind/m2, respectively), but mean density increased and mean size reduced in both reaches during the first 7 upstream migratory events. N. virginea density and size dynamics differed between reaches as a function of habitat hydraulics. While juveniles used the stable riffle as a permanent habitat and preferred passageway, they also used an adjacent, unstable reach after storm events. Density variation was correlated with days postflood (>3.5 m3/s) in both reaches. Our observations indicated that massive upstream migrations of N. virginea juveniles occur at least once a month, presumably as habitat-dependent responses to floods. KEYWORDS.—Neritid snails, diadromy, physical habitat, disturbances, Neotropical streams INTRODUCTION Lyons 1993), Japan (Nishiwaki et al. 1991a; Hirata et al. 1992), French Polynesia (Resh In coastal and insular streams and rivers, et al. 1990, 1992; Liu and Resh 1997), and migrations between marine and fresh wa- Puerto Rico (Covich and McDowell 1996; ters (i. e., diadromy) are common among Pyron and Covich 2003). aquatic fauna (Ford and Kinzie 1982; Mc- Recently, mark-and-recapture studies in Dowall 1998). Many species of fish, shrimp, northeastern Puerto Rico suggested that crayfish, and crabs exhibit this type of mi- neritid gastropods are more active and gration (Baker 1978). Nevertheless, migra- travel longer distances during given peri- tory events have been less frequently re- ods of the year and that upstream migra- ported in gastropod mollusks; although it tion may be seasonal (Pyron and Covich is known that at least 13 families include 2003). However, other one-year mark-and- migratory species (Huryn and Denny recapture study on a neritid gastropod in 1997). Among tropical gastropods, the fam- southern Japan showed no seasonal occur- ily Neritidae comprises several freshwater rence of upstream migrations, or seasonal genera (subfamily Neritinae) whose indi- changes in mean distance movement viduals migrate upstream in massive ag- (Nishiwaki et al. 1991a). These findings gregations. Such migrations of freshwater contrast with another study in the same neritids were reported in Hawaii (Ford area, showing that maximum travel dis- 1979; Ford and Kinzie 1982), Costa Rica tance varies over the year, being greater (Schneider and Frost 1986; Schneider and during the period of high water tempera- ture between April and August (Hirata et al. 1992). Records of gastropod density and ms. received April 12, 2004; accepted November 15, egg laying in French Polynesia (Resh et al. 2004 1991, 1992) and Japan (Nishiwaki et al. 55 56 JUAN F. BLANCO AND FREDERICK N. SCATENA 1990b; Hirata et al. 1992) also suggest a sea- Measuring habitat stability in flashy sonal occurrence of such migrations, but tropical streams is logistically difficult. For- the controlling factors remain unknown. tunately, channel hydraulics may be used There are no additional long-term, high- to estimate the forces experienced by frequency studies dealing with upstream streambed elements and organisms (Now- migrations of neritid gastropods, although ell and Jumars 1984; Statzner et al. 1988; other aspects such as life history (Ford Davies and Barmuta 1989; Way et al. 1993). 1979), growth rates, and fecundity (Shi- If measured close to the streambed, stan- gemiya and Kato 2001), habitat selection dard Reynolds number (Re) and roughness (Liu and Resh 1997; Ohara and Tomiyama Reynolds number (Re*) indicate if micro- 2000), and predators (Teixeira 1994; Resh et flows are turbulent (Re>2000), laminar al. 1999) were studied elsewhere. (Re<500), rough (Re*>70), or smooth Schneider and Lyons (1993) proposed (Re*<70). Similarly, Froude number (Fr) in- that upstream migrations of neritids in a dicates if near-bed flows are supercritical Costa Rican stream were related with (i.e., erosive, Fr>1) or subcritical (i.e., depo- increased fish predation in the estuary. sitional, Fr< 1). Typically, flood stable habi- Small-sized individuals were more abun- tats have larger streambed elements, and dant within migratory groups, and they more turbulent and rough flows. Unstable were also more responsive to the presence habitats generally have fine-grained sub- of predators, as similarly observed in other strates and experience nearly laminar or freshwater gastropods (e.g., Alexander and smooth flows at baseline discharge (Nowell Covich 1991). The distribution of predatory and Jumars 1984; Davies and Barmuta 1989; fish (Allan 1995), and the quantity and Naiman 1998; Montgomery and Buffington quality of periphytonic food (Johnson and 1998; Matthaei et al. 1999a, b). Brown 1997; Biggs and Smith 2002) can also In this study we tested the following hy- be correlated with the spatio-temporal potheses: 1) upstream migration events of variations in discharge and water velocity. neritid gastropods are seasonal, 2) migra- Thus the occurrence of upstream migra- tory aggregations consist of small-sized in- tions might ultimately be a function of dividuals, 3) individuals use turbulent, stream discharge and channel hydraulics. rough flows as passages during upstream For example, laboratory experiments using migrations and as permanent habitats, and clams demonstrated that emigration is dis- 4) influence of flood regime on the distri- played only after increased water move- bution of neritid gastropods depends on ment, even when density-dependent com- habitat hydraulics and stability. petition is strong under slow water movement (Powers and Peterson 2000). In ATERIALS AND METHODS natural conditions, the flash flood distur- M bance can be an important control of stream Study organism community dynamics (Hart and Finelli 1999; Lake 2000). Several studies document The presence of the freshwater neritid that invertebrate abundance is a function of Neritina virginea (Linné 1758) in several is- the elapsed time after storm flows in both lands of the Caribbean has been noted in tropical and temperate streams (Grimm many studies, some from the middle of last and Fisher 1989; Flecker and Feifarek 1994; century (Russel 1941; Aguayo 1966; Hum- Ramírez and Pringle 1998). Recent studies frey 1971). Other species have also been re- also suggest that the effects of storm flow ported in the region (Russel 1941; Aguayo on benthic fauna are mediated by habitat 1966; Humfrey 1971), but may be color stability (reviewed by Lake 2000). Habitats variants of N. virginea (Cosel 1986; Diaz and experiencing greater scouring such as runs Puyana 1994; J. F. Blanco, unpublished and plane beds (Matthaei et al. 1999a, b) data). While the presence of N. virginea in show lower abundance and persistence of the Caribbean is well documented, massive benthos than more resistant riffle and pool upstream migrations have been recently habitat (e.g., Gjerløv et al. 2003). documented in two streams (e.g., Mameyes NERITINA VIRGINEA UPSTREAM MIGRATIONS 57 and Espíritu Santo) in northeastern Puerto 1771 of road PR Route 3 (18°22’27”N, Rico (Covich and McDowell 1996; Pyron 65°45’50” W, elevation: 5 m above sea level) and Covich 2003). over the Río Mameyes, where two reaches, separated by an elevated and stabilized is- land formed after the construction of the Study area bridge in 1982. Most of the river’s flow runs through a ∼11 m wide the main reach (MR: This study was conducted in a lower seg- a riffle at right and looking downstream). ment of Río Mameyes, draining the Lu- Channel depth is nearly constant across the quillo Experimental Forest (LEF), located in section (<40 cm), and the streambed con- northeastern Puerto Rico (Fig. 1a). The up- sists of mid-sized boulders (<50 cm) and per part of the watershed, managed by the cobbles. The right bank of the reach is a United States Forest Service, is covered by concrete-lined bridge abutment. Less water tropical wet forests (Scatena 1989). The flows through a side reach (SR) that occurs lower part of the watershed is suburban- on the opposite side of the bridge. This ized, but has extensive abandoned pastures reach is 3 m wide, less than 30 cm deep, (Ramos 2001). Río Mameyes is considered and is influenced by deflected flow from a the most conserved stream in Puerto Rico, channel bend located 5-m upstream. The and is gauged by the US Geological Survey streambed consists of cobbles in the deep- (USGS). The highest discharge is typically est part and of gravel in the shallowest part. observed during the two rainy seasons The MR and SR join about 40 m down- of the year: May and August-December stream, and this point becomes a decision- (Fig. 1b). making area for the migratory organisms The study site is located beneath Bridge moving upstream.
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