Covich A.P.

Migration patterns, densities, and growth of Neritina snails in Rio Espiritu Santo and Rio Mameyes, northeastern Puerto Rico

PYRON, MARK; COVICH, ALAN P. 2003. Migration Patterns, Densities, and Growth of Neritina punctulata Snails in Rio Espiritu Santo and Rio Mameyes, Northeastern Puerto Rico.. Caribbean Journal of Science, Vol. 39, No. 3, 338-347, .

Abstract: 
Snail size-frequency distributions in Rios Espiritu Santo and Mameyes, which drain the Luquillo Experimental Forest, Puerto Rico, showed that Neritina punctulata with shell lengths greater than 30 mm were the most abundant size class at upstream sites. The highest densities for all size classes were at the downstream sites. Growth rates were 0.015 mm/day for a large cohort (¡«25 mm shell length) and 0.035 mm/day for a small (¡«15 mm shell length). Minimum longevity estimates range from 3 to 7 years. Size distribution data suggest that snails migrate upstream. An August mark-recapture study resulted in most snails remaining at the release site, and some moved downstream. The greatest upstream distance reached was 200 m in 12 weeks. In a May mark-recapture study, when snails were observed moving in aggregations, the greatest distance moved was 200 m in 27 days, suggesting seasonality in movements and reproduction. Movement rates of snails in aggregates in May were between 0.08 and 0.17 cm/s. All of the snails marked in May moved upstream, and none were observed downstream from, or at the release site. We suggest that upstream movements can be explained by snails avoiding increased predation in the downstream reaches of these rivers from fishes, crustaceans, and birds

On the relative importance of pool morphology and woody debris to distributions of shrimp in a Puerto Rican headwater stream

Pyron, M., Covich, A.P. and Black, R.W. 1999. On the relative importance of pool morphology and
woody debris to distributions of shrimp in a Puerto Rican headwater stream. Hydrobiologia, 402:
207–215.

Abstract: 
In this paper, we report the sizes and distributional orientation of woody debris in a headwater rainforest stream in the Luquillo Experimental Forest (LEF), Puerto Rico. We also provide results of a 4-month study of a wood addition experiment designed to increase cover for benthic macroinvertebrates (freshwater shrimp). We added branch-sized woody debris to 20 pools in three streams.We trapped four species of freshwater shrimp (two species of benthic detritivores and two predatory shrimp species) during each of the 4 months following wood additions. An analysis of pool morphology (maximum depth, surface area and volume) provided a useful predictor of shrimp abundances. In general, numbers of shrimps increased with sizes of stream pools. A repeated measures ANOVA demonstrated no effect of woody debris additions on total numbers of shrimp per pool area. Two detritivore species (Atya lanipes, a filter feeder and Xiphocaris elongata, a shredder) decreased in abundance with increased woody debris and there was no statistical relationship between woody debris additions and predators (Macrobrachium carcinus and M. crenulatum). Small woody debris additions may have altered flow velocities that were important to filter-feeding Atya at the microhabitat scale, although the overall velocities within pools were not altered by wood additions. Lower numbers of Atya and Xiphocaris in two of the three streams may result from the occurrence of two predaceous fishes (American eel and mountainmullet) and more predatoryMacrobrachium in these streams. One likely interpretation of the results of this study is that the stream pools in these study reaches had sufficient habitat structure provided by numerous rock crevices (among large rocks and boulders) to provide refuge from predators. Addition of woody debris did not add significantly to the existing structure. These results may not apply to stream channels with sand and gravel substrata where crevices and undercut banks are lacking and where woody debris often plays a major role by providing structure and refuge.

Effects of drought and hurricane disturbances on headwater distributions of palaemonid river shrimp (Macrobrachium spp.) in the Luquillo Mountains, Puerto Rico

Covich, Alan P.; Crowl, Todd A.; Heartsill-Scalley, Tamara 2006. Effects of drought and hurricane disturbances on headwater distributions of palaemonid river shrimp (Macrobrachium spp.) in the Luquillo Mountains, Puerto Rico.. J. N. Am. Benthol. Soc., 25(1):99-107.

Abstract: 
Extreme events (hurricanes, floods, and droughts) can influence upstream migration of macroinvertebrates and wash out benthic communities, thereby locally altering food webs and species interactions. We sampled palaemonid river shrimp (Macrobrachium spp.), dominant consumers in headwaters of the Luquillo Mountains of northeastern Puerto Rico, to determine their distributions along an elevational gradient (274–456 m asl) during a series of disturbances (Hurricane Hugo in 1989, a drought in 1994, and Hurricane Georges in 1998) that occurred over a 15-y period (19882002).We measured shrimp abundance 3 to 6 times/y in Quebrada Prieta in the Espiritu Santo drainage as part of the Luquillo Long-Term Ecological Research Program. In general, Macrobrachium abundance declined with elevation during most years. The lowest mean abundance of Macrobrachium occurred during the 1994 drought, the driest year in 28 y of record in the Espiritu Santo drainage. Macrobrachium increased in abundance for 6 y following the 1994 drought. In contrast, hurricanes and storm flows had relatively little effect on Macrobrachium abundance.

Predator–prey interactions in river networks: comparing shrimp spatial refugia in two drainage basins

COVICH, A. P., T. A. CROWL, C. L. HEIN, M. J. TOWNSEND, AND
W. H. MCDOWELL. 2009. Predator-prey interactions in
river networks: comparing shrimp spatial refugia in two
drainage basins. Freshwater Biology 54:450–465.

Abstract: 
1. Analysis of drainage networks provides a framework to evaluate the densities and distributions of prey species relative to locations of their predators. Upstream migration by diadromous shrimp (Atya lanipes and Xiphocaris elongata) during their life cycle provides access to headwater refugia from fish predation, which is intense in estuaries and coastal rivers. 2. We postulate that geomorphic barriers (such as large, steep waterfalls >3.5 m in height), can directly limit the distribution of predatory fishes and, indirectly, affect the densities of their prey (freshwater shrimps) in headwater streams. 3. We compared densities of shrimp in pools above and below waterfalls, in four headwater tributaries in two river basins of the Luquillo Mountains of northeastern Puerto Rico. We measured shrimp densities twice a year over 8 years (1998–2005) in Prieta, Toronja, Bisley 3 and Bisley 5 streams, which differ in drainage network positions relative to steep waterfalls in Rı´o Espı´ritu Santo and Rı´o Mameyes. 4. Predatory fishes are absent in the Prieta and Toronja pools and present in Bisely 3 and in lower Bisley 5 pools. Atya lanipes and X. elongata rarely occur in the Bisley streams where predatory fishes are present but these shrimps are abundant in Prieta and Toronja, streams lacking predatory fishes. 5. The mean carapace length of X. elongata is longer in pools where fish are present (Bisley 3 and lower Bisley 5) than in pools lacking fish (Prieta, Toronja, Upper Bisley 5). The increased body size is primarily due to significantly longer rostrums of individuals in stream reaches with fish (below waterfall barriers) than in those reaches lacking fish (above waterfall barriers). Rostrum length may be an adaptation to avoid predation by visually feeding fishes. 6. Atya lanipes and X. elongata distributions and densities were predicted primarily by drainage network position relative to the presence or absence of predatory fishes. High, steep waterfalls effectively impeded fish from moving upstream and created a spatial refuge. Xiphocaris elongata may rely on size refugia (longer rostrum) to minimize predation where spatial refugia are lacking.

Effects of coupled natural and anthropogenic factors on the community structure of diadromous fish and shrimp species in tropical island streams

CATHERINE L. HEIN*, ANDREW S. PIKE†1 , JUAN F. BLANCO‡, ALAN P. COVICH§ , FREDERICK N. SCATENA†, CHARLES P. HAWKINS* AND TODD A. CROWL. Effects of coupled natural and anthropogenic factors on the community structure of diadromous fish and shrimp species in tropical island streams. Freshwater Biology. Vol 56, Is 5 pp 1002-1015.

Abstract: 
1. Overlapping river and road networks provide a framework for studying the complex interactions between natural and human systems, with river-road intersections as focal areas of study. Roads can alter the morphology of stream channels, pose barriers to freshwater fauna, provide easy access to streams for humans and non-native species and accelerate the expansion of urban development. 2. We determined what variables control the structure of diadromous fish and shrimp communities and assessed whether particular road crossings altered community structure in north-eastern Puerto Rico. We identified 24 sites that represented a range of river and road sizes across two catchments that drain El Yunque National Forest in Puerto Rico. 3. The location of natural barriers and the size of stream pools were the most important variables for predicting six of fifteen fish and shrimp distributions. Predatory fishes were predicted to be limited to areas in the river network below large, steep waterfalls, whereas adult shrimp Atya lanipes (Atyidae) were predicted to be present above these waterfalls. The fish Awaous banana was predicted to be present in pools >11.6 m wide, whereas the shrimp Xiphocaris elongata was predicted to be present in pools <10.4 m wide. The distributions of nine species were predicted poorly, but three of these species were common and three were rare. 4. Although urban and agricultural land covers were among the top three predictors of five species distributions, they were probably good predictors because they were correlated with the natural gradient. Further study is necessary to disentangle natural and anthropogenic gradients. 5. Road crossings, 10 of which were culverts, were not dispersal barriers for fishes or shrimps. On average, species were present both upstream and downstream from road crossings at 68% of sites where they occurred. Absences upstream or downstream from road crossings occurred at 16% of sites each and likely resulted from a failure to detect species.

Linking habitat stability to floods and droughts: effects on shrimp in montane streams, Puerto Rico

COVICH, A. P., T. A. CROWL, AND F. N. SCATENA. 2000. Linking
habitat stability to floods and droughts: effects on
shrimp in montane streams, Puerto Rico. Verhandlungen
der Internationalen Vereinigung fu¨ r theorestische und
angewandte Limnologie 27:2430–2434.

Abstract: 
Most previous studies on Caribbean flood and drought frequency have examined hydrological (e.g. MORRIS & VAZQUEZ 1990, GARCIA et al. 1996) rather than ecological effects (COVICH et al. 1998). We analyzed temporal and spatial distributions of rainfall, stream flow, and mean maximum pool depth over an 8-year period (1990–1997) to evaluate effects of variable flows. We examined the response of a biotic variable (the coefficient of variation in shrimp densities) to changes in water depth (coefficient of variation of maximum pool depth) along an elevational gradient.

Linking habitat stability to floods and droughts: effects on shrimp in montane streams, Puerto Rico

Covich A.P., Crowl T.A. & Scatena F.N. (2000) Linking
habitat stability to floods and droughts: effects on
shrimp in montane streams, Puerto Rico. Verhandlungen
der Internationalen Vereinigung fu¨r Theoretische und
Angewandte Limnologie, 27, 2430–2434.

Abstract: 
Most previous studies on Caribbean flood and drought frequency have examined hydrological (e.g. MORRIS & VAZQUEZ 1990, GARCIA et al. 1996) rather than ecological effects (COVICH et al. 1998). We analyzed temporal and spatial distributions of rainfall, stream flow, and mean maximum pool depth over an 8-year period (1990–1997) to evaluate effects of variable flows. We examined the response of a biotic variable (the coefficient of variation in shrimp densities) to changes in water depth (coefficient of variation of maximum pool depth) along an elevational gradient.

Non-Indigenous Bamboo along Headwater Streams of the Luquillo Mountains, Puerto Rico: Leaf Fall, Aquatic Leaf Decay and Patterns of Invasion

O'CONNOR, PAUL J.; COVICH, ALAN P.; SCATENA, F. N.; LOOPE, LLOYD L. 2000. Non-indigenous bamboo along headwater streams of the Luquillo Mountains, Puerto Rico: leaf fall, aquatic leaf decay and patterns of invasion. Journal of Tropical Ecology 16 :499-516.

Abstract: 
The introduction of bamboo to montane rain forests of the Luquillo Mountains, Puerto Rico in the 1930s and 1940s has led to present-day bamboo monocultures in numerous riparian areas. When a non-native species invades a riparian ecosystem, in-stream detritivores can be affected. Bamboo dynamics expected to in¯uence stream communities in the Luquillo Experimental Forest (LEF) were examined. Based on current distributions, bamboo has spread downstream at a rate of 8 m y-1. Mean growth rate of bamboo culms was 15.3 cm d-1. Leaf fall from bamboo stands exceeded that of native mixed-species forest by c. 30% over a 10-mo study. Bamboo leaves (k = -0.021), and leaves from another abundant riparian exotic, Syzygium jambos (Myrtaceae) (k = -0.018), decayed at relatively slow rates when submerged in streams in ®ne-mesh bags which excluded macro-invertebrate leaf processors. In a second study, with leaf processors present, bamboo decay rates remained unchanged (k = -0.021), while decay rates of S. jambos increased (k = -0.037). Elemental losses from bamboo leaves in streams were rapid, further suggesting a change in riparian zone / stream dynamics following bamboo invasion. As non-indigenous bamboos spread along Puerto Rico streams, they are likely to alter aquatic communities dependent on leaf input.

Effects of extreme low flows on freshwater shrimps in a perennial tropical stream

COVICH, A.P.; CROWL, T.A.; SCATENA, F.N. 2003. Effects of extreme low flows on freshwater shrimps in a perennial tropical stream.. Freshwater Biology 48, 1199-1206.

Abstract: 
1. Long-term data on rainfall suggests that perennial rainforest streams rarely are subject to drying of riffles or pools in the wet, non-seasonal Caribbean climate of Puerto Rico. Unusually low rainfall in 1994 caused some headwater riffles to dry out completely, resulting in isolated pools, reduced pool volumes and loss of access to microhabitats by benthic invertebrates. 2. From 1992 to 1998, shrimp populations were sampled bimonthly using baited traps in six pools along 1200 m (from 305 to 480 m in altitude) of Quebrada Prieta, a second-order headwater stream in the Luquillo Experimental Forest (Caribbean National Forest). 3. Following contraction of the smaller and shallower pools in the most upstream sectionof the stream, mean densities of the dominant shrimp (Atya lanipes) increased from 22 to 75shrimp m)2 of pool area during the 1994 drought year. A second common species(Xiphocaris elongata) increased from 5 to 14 shrimp m)2. A smaller percentage of adults of both species was gravid during the drought. 4. Following the 1994 drought (1995–1998), densities of both shrimp species and reproductive activity of Atya returned to predrought (1990–1993) levels. However, the reproductive activity of Xiphocaris remained lower than in the predrought period. 5. It is suggested that prolonged droughts, even in tropical rainforest biomes, may significantly alter aquatic communities through localised crowding effects resulting from habitat contraction, and lead to prolonged decreases in reproductive output. Consequently, major alterations in aquatic populations and communities would be predicted by current climate change scenarios of decreased total rainfall and increased variability.
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