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DATA SET IDENTIFIER: Indirect upstream effects of dams: consequences of migratory consumer extirpation in Puerto Rico

PROJECT TITLE: Ecological studies of dams and migratory fauna

PROJECT DESCRIPTION: The ecological roles of diadromous fauna (freshwater shrimps, fishes, and snails) in Puerto Rico were studied in the context of examining consequences of their loss from streams above large (height >15 m) dams. Four sub-projects were conducted:
(a) Indirect upstream effects of dams: consequences of migratory consumer extirpation in Puerto Rico. This sub-project examined the effects of decimation of migratory fauna populations on stream ecosystem structure above large dams. We compared streams above large dams to streams without large dams, in terms of relative abundances of migratory fauna and ecosystem components affected by migratory fauna in previous in situ experiments. Previous research indicated that: (1) all native fishes and shrimps in Puerto Rico are extirpated from habitats upstream of large dams without regular spillway discharge because they are diadromous, whereas streams above large dams with regular spillway discharge have greatly reduced abundances of diadromous fauna, and streams without large dams have relatively natural fish and shrimp assemblages (Holmquist et al. 1998); and (2) small-scale experimental exclusion of native shrimps and fishes decreases leaf decay rates and increases epilithic organic and inorganic matter, chlorophyll a, carbon, nitrogen, carbon-to-nitrogen ratio (C:N), and chironomid biomass (Pringle et al. 1999, Crowl et al. 2001, March et al. 2001, March et al. 2002). We examined whether sites above large dams had higher levels of epilithic coarse organic matter, algae, fine organic and inorganic matter, carbon, nitrogen, C:N, and non-decapod invertebrates, consistent with previous findings of small-scale experimental shrimp and fish exclusions.
(b) Do small-scale exclosure/enclosure experiments have relevance for large-scale extirpation of migratory fauna? This sub-project took advantage of large dams to examine effects of experimental scale. Alteration of migratory stream populations due to large dams represents a large-scale (i.e. whole-catchment), long-term (i.e. decades) "extirpation experiment." We compared this large-scale perturbation to four small-scale experiments (two exclusion experiments in sites with no large dams and two shrimp addition experiments in dammed sites) that we conducted in a subset of the sites sampled in sub-project (a). We examined whether small-scale experiments predicted effects at the large-scale in terms of direction and magnitude.
(c) Conservation and management of migratory fauna and dams in tropical streams of Puerto Rico. This sub-project was a review, in which we: (1) examined Puerto Rico's potential to serve as a window into the future of freshwater migratory fauna in tropical regions, given the island's extent and magnitude of dam development and the available scientific information on ecology and management of the island's migratory fauna, and (2) reviewed ecology, management and conservation of migratory fauna in relation to dams in Puerto Rico. Our review included a synthesis of recent and unpublished observations on upstream effects of large dams on migratory fauna and an analysis of patterns in free crest spillway discharge across Puerto Rican reservoirs.
(d) Ecological effects of non-migratory native and non-native fauna above large dams in tropical streams, Puerto Rico. This sub-project consisted of a set of electric exclusion experiments conducted above two of the dammed sites in sub-project a. Previous research demonstrated higher abundances of exotic fishes above large dams (Holmquist et al. 1998). Thus, these experiments examined whether exotic fishes affect stream ecosystem patterns and confound any interpretations in sub-project a regarding roles of migratory macroconsumers.

LTER CORE AREAS: Annotate all that apply (See online list)

Disturbance Patterns

LEF LTER 1 RESEARCH TOPIC: (Annotate all that apply) (See online list)

Disturbance

We define a data file as a component of a data set. A data set can have only one data file or more. Basically, different data files have different data structures or format .
DATA SET FILES (SUBSETS):

Data File No.	Data File Identifier	On-Line Filename	Starting Date	Periodicity of sample	End Period
1	Locations of high-gradient survey sites	highgradsites.txt	N/A	N/A	N/A
2	Characteristics of dams and dammed high-gradient survey sites	dam&sitecharacteristics.txt	N/A	N/A	N/A
3	Physical parameters at high-gradient survey sites	highgradphyshab.txt	June 2001	irregular	June 2003
4	Chemical parameters at high-gradient survey sites	highgradchem.txt	June 2001	irregular	June 2003
5	Benthic resources at high-gradient survey sites	highgradbenthicresources.txt	June 2001	irregular	June 2003
6	Macroconsumers (fishes, shrimps, crabs, tadpoles) in snorkeling and electroshocking samples from high-gradient survey sites	highgradmacroconsumers.txt	June 2001	irregular	June 2003
7	Non-decapod invertebrate biomass at high-gradient survey sites	highgradinverts.txt	June 2001	irregular	June 2003
8	Low-gradient site locations and physical and chemical parameters	lowgradphyshab&chem.txt	July 2000	irregular	August 2000
9	Benthic resources at low-gradient survey sites	lowgradbenthicresources.txt	July 2000	irregular	August 2000

RESEARCH LOCATION: Stream reaches above large dams in Puerto Rico.

INVESTIGATORS:

PRINCIPAL INVESTIGATORS	E-MAIL ADDRESS:
Effie A. Greathouse	effieg@gmail.com

OTHER RESEARCHERS	E-MAIL ADDRESS:
Catherine M. Pringle	cpringle@uga.edu
William H. McDowell	wmcdowell@lternet.edu
Jeff G. Holmquist

CONTACT PERSONS	Phone Number (Include area code)	E-MAIL ADDRESS:
Effie A. Greathouse	916-332-5771 (Whitney Masonry)	effieg@gmail.com

SOURCE OF FUNDING (SPONSOR): National Science Foundation (NSF) Luquillo LTER (BSR 8811902, DEB9411973, DEB0080538, DEB0218039), the NSF Doctoral Dissertation Improvement Grant program (DEB 0308543), the U.S. Department of Agriculture (grants 10-21-RR551-141, 10-21-RR250-109); and the NSF Graduate Fellowship program (fellowship to EAG)

DATA SET ABSTRACT: Large dams degrade the integrity of a wide variety of ecosystems, yet direct downstream effects of dams have received the most attention from ecosystem managers and researchers. We investigated indirect upstream effects of dams resulting from decimation of migratory freshwater shrimp and fish populations in Puerto Rico, USA, in both high- and low-gradient streams. In high-gradient streams above large dams, native shrimps and fishes were extremely rare, whereas similar sites without large dams had high abundances of native consumers. Losses of native fauna above dams dramatically altered their basal food resources and assemblages of invertebrate competitors and prey. Compared to pools in high-gradient streams with no large dams, pool epilithon above dams had 9 times more algal biomass, 20 times more fine benthic organic matter (FBOM), 65 times more fine benthic inorganic matter (FBIM), 28 times more carbon (C), 19 times more nitrogen (N), and 4 times more non-decapod invertebrate biomass. High-gradient riffles upstream from large dams had 5 times more FBIM than did undammed riffles but showed no difference in algal abundance, FBOM, or non-decapod invertebrate biomass. For epilithon of low-gradient streams, differences in basal resources between pools above large dams vs. without large dams were considerably smaller in magnitude than those observed for pools in high-gradient sites. These results match previous stream experiments in which the strength of native shrimp and fish effects increased with stream gradient. Our results demonstrate that dams can indirectly affect upstream free-flowing reaches by eliminating strong top-down effects of consumers. Migratory omnivorous shrimps and fishes occur throughout the tropics, and the consequences of their declines upstream from many tropical dams are likely to be similar to those in Puerto Rico. Thus, ecological effects of migratory fauna loss upstream from dams encompass a wider variety of species interactions and biomes than the bottom-up effects (i.e., elimination of salmonid nutrient subsidies) recognized for northern temperate systems.

DATA SET METHODS: See Greathouse et al. (2006) and Greathouse (2005). (full reference in the "REFERENCES" section below). Our main data set compares dammed and undammed streams that are high gradient (i.e., above waterfalls that block migration of predatory fishes). We used a natural experiment approach (sensu, Diamond 1986), sampling macroconsumers, benthic resources, and non-decapod invertebrates in seven stream reaches upstream from large dams and 10 undammed reaches (Fig. 1a). We refer to sites above large dams as ''dammed'' and sites with no large dams as ''undammed''; however, low-head dams (height ,15 m) may occur upstream or downstream from any of our sites. The 10 undammed sites were dominated by shrimp and Sicydium with no predatory fishes. Physical conditions of the seven dammed sites are characteristic of streams that are dominated by shrimps and Sicydium (Table 1); it is highly likely that they were dominated by shrimp and Sicydium prior to dam construction. High-gradient dammed sites were above large dams without regular ''free crest'' spillway discharge (i.e., water flowing over the sloping face of a dam; hereafter referred to as spillway discharge). A lack of regular spillway discharge was previously found to cause upstream extirpation of shrimps and native fishes (Holmquist et al. 1998). Thus, our highgradient stream survey was designed to compare sites where native shrimps and Sicydium were extirpated (dammed) and sites with relatively natural shrimp and Sicydium assemblages (undammed). We chose sites so that physical and chemical parameters would not confound our comparison of dammed vs. undammed rivers.

Sites were sampled once between June 2001 and June 2003, in haphazard order, at base flow over a reach at least 10 times the channel width. To sample epilithic chlorophyll a, fine (,1 mm) benthic inorganic matter (FBIM), fine benthic organic matter (FBOM), coarse (.1 mm) benthic organic matter (CBOM), and non-decapod invertebrates, we obtained randomly located samples from boulders/bedrock in each of the two major habitat types (pools and riffles that, on average, represented 44% and 54% of stream habitat, respectively). We also analyzed FBOM samples from pools for total C, N, and C:N (molar ratio). Chlorophyll a, FBIM, and FBOM were sampled using a suction device modified from Loeb (1981). Loeb samples were composited by habitat type (12 over three poo1s and 12 over three riffles) and brought back to the laboratory on ice for subsampling. A chlorophyll a subsample was filtered onto a precombusted glass fiber filter (Whatman GF/F, 0.7 mm), frozen, and analyzed fluorometrically according to standard methods (APHA 1985). A FBIM/FBOM subsample was filtered onto a precombusted, preweighed GF/F filter and dried at 508C for at least 24 h. FBIM/FBOM filters from riffles were weighed to the nearest 0.001 g, ashed at 5008C for 3 h, and reweighed to determine inorganic and ash-free dry mass (AFDM) per unit area. For FBIM/FBOM filters from pools, we cut the dried filter into two roughly equal sections and weighed each section to the nearest 0.001 g. One filter section was analyzed for C and N using a PerkinElmer CHN analyzer (PerkinElmer, Wellesley, Massachusetts, USA). The other section was analyzed for inorganic mass and AFDM, as described for FBIM/FBOM filters from riffles.

Three riffles were sampled for CBOM and non-decapod benthic invertebrates by scrub brushing invertebrates and benthic matter from a known area into a 210-mm hand net. In three pools, we used the ‘‘benthic block net’’ described by Greathouse and Pringle (2005) to sample CBOM quantitatively (i.e., mass per unit area) and non- decapod invertebrates semiquantitatively (i.e., biomass per unit effort). Invertebrate/CBOM samples were preserved with ethanol and processed following methods modified from Lugthart and Wallace (1992). We sieved samples in nested 1-mm and 250- mm sieves prior to sorting for macroinvertebrates and performing AFDM analyses on CBOM using methods described for FBIM/FBOM filters. Invertebrates were identified to the lowest practicable level (generally family for insects and class or order for other nondecapod invertebrates), and length–mass regressions were used to determine biomass.

All sites except U9 were electroshocked for macroconsumers (fishes, adult shrimps, crabs, tadpoles) by using a catch-per-unit- effort method (point abundance sampling) modified from Fie`vet et al. (1996). Two dip netters captured macroconsumers at 5–6 electroshocking points in pools. For riffles, point samples were collected by electroshocking over lengths of ;0.5–1 m into nets placed downstream (at four points per site, dip nets completely blocked flows of ;0.2–0.4 m wide; at two points per site, a 1-m seine net was placed in flows that were ;1–2 m wide). At site U9, electroshocking data from removal sampling in one riffle and one pool was normalized by shocking time and number of dip netters in order to obtain an estimate with effort similar to point sampling. In pools of a few dammed sites, we also observed rare occurrences of native shrimps and fishes that were not captured during electroshocking.

REFERENCES:
Greathouse, E. A., C. M. Pringle, W. H. McDowell, and J. G. Holmquist. 2006. Indirect effects of dams: upstream consequences of migratory fauna extirpation in Puerto Rico. Ecological Applications 16: 339-352.

Greathouse, E.A. 2005. Large dams and migratory biota affect tropical stream ecosystems at different scales in Puerto Rico. Ph.D. Dissertation, University of Georgia, Athens.

CROSS-REFERENCES (other data sets related to this one): LTERDBAS #138: Conservation and management of migratory fauna and dams in tropical streams of Puerto Rico; LTERDBAS #139: Indirect upstream effects of dams: consequences of migratory consumer extirpation in Puerto Rico;

STORAGE SITES(of data files): Data is maintained by Effie Greathouse.

INVESTIGATOR'S ASSIGNED KEYWORDS: amphidromy, benthic invertebrates, Decapoda, fishes, freshwater shrimps, migratory fauna, natural experiment, omnivorous macrobiota, primary consumer, tropical stream

LEF LTER OFFICIAL KEYWORDS (See table): OTHER PLOTS, HEADWATER STREAM, RIVER, WATER ABSTRACTION, DISTURBANCE, ALGAE, AQUATIC, DECAPODS, FISH, INSECTS, DISSERTATION, PEER REVIEWED JOURNAL

PUBLICATIONS:
Greathouse, E. A., C. M. Pringle, W. H. McDowell, and J. G. Holmquist. In press. Indirect effects of dams: upstream consequences of migratory fauna extirpation in Puerto Rico. Ecological Applications.

RESTRICTED ___ UNRESTRICTED _X_

SITES DESCRIPTIONS: We studied streams in mountainous regions of Puerto Rico, USA, that are characterized by only slight seasonal variation in temperature and rainfall (Larsen 2000). Flash floods (discharge increases up to 10-fold in ,1 h) occur throughout the year and wash out most accumulations of large woody debris. Land cover is mixed, with substantial recovery of tropical forest associated with declines in agriculture (e.g., pasture, coffee) and increases in low-density residential development over the past 30-60 years (Grau et al. 2003). Puerto Rican streams are dominated by migratory shrimps and fishes. Except for the catadromous American eel (Anguilla rostrata), all native freshwater shrimps and fishes are thought to be amphidromous, with adult females releasing larvae that passively drift downstream to the estuary before migrating back upstream as juveniles (Chace and Hobbs 1969, March et al. 1998, Nieves 1998). ALSO SEE DATA FILE 1.

VARIABLES (ATTRIBUTES):
D1,unnamed tributary of Carite reservoir,18,3,50.94,66,6,16.77

VARIABLES (ATTRIBUTES):

2File Name or # above (all in which the variable appears)	1	1	1	2
AbbreviationAbbreviation(as it appears on the data file)	londeg	longmin(')	longsec(')	Dam
NAME OF VARIABLE	Longitude degrees	Longitude minutes	Longitude seconds	Dam name
DEFINITION OF VARIABLE	The degrees of the site's longitude coordinates	The minutes of the site's longitude coordinates	The seconds of the site's longitude coordinates	The name of the large dam occurring below high-gradient study sites
UNIT	degrees	minutes	seconds
PRECISION			0.01
RANGE OR LIST OF VALUES	{66}	{0,.,56}	{3.34,.,58.44}	absolute, Carite, Garzas, Caonillas, Dos Bocas, Matrullas, Luchetti, Toa Vaca
DATA TYPE	integer	integer	decimal
MISSING DATA CODES	none	none	none

VARIABLES (ATTRIBUTES):

3File Name or # above (all in which the variable appears)	2	2	2	3, 8	3, 8
AbbreviationAbbreviation(as it appears on the data file)	spillwayelev(ft)	spillwayelev(m)	distanceupstream	order	link
NAME OF VARIABLE	Spillway crest elevation	Spillway crest elevation	Distance upstream from reservoir	Stream order	Stream link
DEFINITION OF VARIABLE	The elevation of the crest of the dam's free-crest spillway	The elevation of the crest of the dam's free-crest spillway	The distance between the site and the upstream end of the dam's reservoir	The Strahler stream order of the site based on USGS blue lines	The Shreve stream link of the site based on USGS blue lines. number of USGS blue line headwater streams draining into the site.
UNIT	feet above sea level	meters above sea level	km
PRECISION	0.1 ft a.s.l.	0.1 m a.s.l.	0.1 km
RANGE OR LIST OF VALUES	{295,.,1783.6}	{89.9,.,543.6}	{0.7,.,33.6}	{1,.,5}	{1,.,215}
DATA TYPE	decimal	decimal	decimal	integer	integer
MISSING DATA CODES	MGT indicates that the dam does not have a free-crest spillway and instead has a morning-glory tunnel spillway	MGT indicates that the dam does not have a free-crest spillway and instead has a morning-glory tunnel spillway	none	none	none

VARIABLES (ATTRIBUTES):

4File Name or # above (all in which the variable appears)	3, 8	3	3	3, 8	3
AbbreviationAbbreviation(as it appears on the data file)	gradient	slope	aspect	elevation	catchmentarea
NAME OF VARIABLE	Stream gradient	Field slope	Aspect	Elevation	Catchment area
DEFINITION OF VARIABLE	The ratio of tghe slope to gradient of the stream calculated from USGS maps. Calculated by dividing the slope meters by the gradient meters.	The ratio of tghe slope to gradient of the stream reach measured in the field. Calculated by dividing the slope meters by the gradient meters.	Aspect of the site	Elevation of the site, m above sea level	The catchment area of the site
UNIT			degree	meter	hectare
PRECISION	0.001	0.001	1	1	0.1
RANGE OR LIST OF VALUES	{0.004,.,0.179}	{0.045,.,0.152}	{0,.,270}	{90,.,810}	{64.1,.,559.4}
DATA TYPE	decimal	decimal	integer	integer	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

5File Name or # above (all in which the variable appears)	3	3	3	3, 8	3
AbbreviationAbbreviation(as it appears on the data file)	78ag	78development	bankfullwidth	wettedwidth	92forest
NAME OF VARIABLE	1977-1978 catchment land cover in agriculture	1977-1978 catchment land cover in development	Bankfull width	Wetted width	1991-1992 catchment land cover in forest
DEFINITION OF VARIABLE	The percentage of the catchment area in agricultural land cover	The percentage of the catchment area in development land cover	The width of the channel from bank to bank	The width of the wetted part of the stream on the day of measuring physical habitat	The percentage of the catchment area in forest land cover
UNIT			meter	meter
PRECISION	0.1	0.1	0.1	0.01	0.1
RANGE OR LIST OF VALUES	{0,.,99.9}	{0,.,8.8}	{3.0,.,12.0}	{1.25,.,31.03}	{55.3,.,97.9}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

6File Name or # above (all in which the variable appears)	3	3	3, 8	3	3
AbbreviationAbbreviation(as it appears on the data file)	rifflevel	poolvel	depth	92ag	92urban
NAME OF VARIABLE	Riffle water velocity	Pool water velocity	Water depth	1991-1992 catchment land cover in agriculture	1991-1992 catchment land cover in urban/barren
DEFINITION OF VARIABLE	Mean water velocity in riffles	Mean water velocity in pools	Mean water depth	The percentage of the catchment area in agricultural land cover	The percentage of the catchment area in urban/barren land cover
UNIT	metersPerSecond	metersPerSecond	meter
PRECISION	0.01	0.01	0.01	0.1	0.1
RANGE OR LIST OF VALUES	{0.29,.,1.02}	{-0.02,.,0.07}	{0.11,.,0.43}	{0.5,.,44.4}	{0,.,1.62}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	--- indicates no measurement of water velocity	--- indicates no measurement of water velocity	none	none	none

VARIABLES (ATTRIBUTES):

7File Name or # above (all in which the variable appears)	3	3	3, 8	3, 8	3
AbbreviationAbbreviation(as it appears on the data file)	78forest	78shrub	cancov	substrate	bedrock
NAME OF VARIABLE	1977-1978 catchment land cover in forest	1977-1978 catchment land cover in shrub	Canopy cover	Substrate coarseness	Percentage bedrock
DEFINITION OF VARIABLE	The percentage of the catchment area in forest land cover	The percentage of the catchment area in shrub land cover	Canopy cover over the stream reach	The mean of numbered codes for substrate sizes; see Greathouse et al. (in press) and Greathouse (2005) for details	The percent of substrate measurements that were bedrock
UNIT
PRECISION	0.1	0.1	0.1	N/A	0.1
RANGE OR LIST OF VALUES	{0,.,99.4}	{0,.,20.8}	{0,.,90.5}	{0.58,.,5.8}	{0,.,57.6}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

8File Name or # above (all in which the variable appears)	3	3	3	3	3
AbbreviationAbbreviation(as it appears on the data file)	boulder	cobble	coarsegravel	finegravel	sand
NAME OF VARIABLE	Percentage boulder	Percentage cobble	Percentage coarse gravel	Percentage fine gravel	Percentage sand
DEFINITION OF VARIABLE	The percent of substrate measurements that were boulder	The percent of substrate measurements that were cobble	The percent of substrate measurements that were coarse gravel	The percent of substrate measurements that were fine gravel	The percent of substrate measurements that were sand
UNIT
PRECISION	0.1	0.1	0.1	0.1	0.1
RANGE OR LIST OF VALUES	{5.3,.,69.0}	{1.4,.,28.0}	{6.3,.,24.4}	{0.9,.,20.8}	{2.1,.,30.2}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

9File Name or # above (all in which the variable appears)	3	3	3, 5, 7	4	4, 8
AbbreviationAbbreviation(as it appears on the data file)	silt	temp	Comments	conductivity	TDN
NAME OF VARIABLE	Percentage silt	Water temperature	Comments added for information management purposes	Water conductivity	Total dissolved nitrogen
DEFINITION OF VARIABLE	The percent of substrate measurements that were silt	Temperature of the stream water	Carries the information of the record whenever the value entered by the field technician was not numeric. For example, if the silt percent was less than .1, the value entered by the technician in the corresponding data field record was "<0.1". The latter was removed from the data field cell and a comment "silt<0.1;" was added to the Comments data field corresponding to that record.	Conductivity of the stream water in micro-Seimens per centimeter (uS/cm)	Total dissolved nitrogen in the stream water in micrograms nitrogen per liter (ug N/L)
UNIT		celsius		micro-Seimens per centimeter	microgramsPerLiter
PRECISION	0.1	0.1		1	1
RANGE OR LIST OF VALUES	{0,.,2.7}	{18.0,.,26.0}		{74,.,390}	{127,.,2118}
DATA TYPE	decimal	decimal	alphanumeric	integer	integer
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

10File Name or # above (all in which the variable appears)	4	4, 8	4	4, 8	4
AbbreviationAbbreviation(as it appears on the data file)	NO3	NH4	DON	SRP	DOC
NAME OF VARIABLE	Nitrate	Ammonium	Dissolved organic nitrogen	Soluble reactive phosphorus	Dissolved organic carbon
DEFINITION OF VARIABLE	Amount of dissolved nitrate in the stream water in micrograms nitrogen per liter (ug N/L)	Amount of dissolved ammonium in the stream water in micrograms nitrogen per liter (ug N/L)	Amount of dissolved organic nitrogen in the stream water in micrograms nitrogen per liter (ug N/L)	Amount of soluble reactive phosphorus in the stream water in micrograms phosphorus per liter (ug P/L)	Amount of dissolved organic carbon in the stream water ion micrograms carbon per liter (ug C/L)
UNIT	microgramsPerLiter	microgramsPerLiter	microgramsPerLiter	microgramsPerLiter	microgramsPerLiter
PRECISION	1	0.1	1	1	1
RANGE OR LIST OF VALUES	{92,.,2114}	{0.5,.,48.0}	{0,.,327}	{6,.,110}	{323,.,3360}
DATA TYPE	integer	decimal	integer	integer	integer
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

11File Name or # above (all in which the variable appears)	5, 9	5	5, 9	5, 9	5, 9
AbbreviationAbbreviation(as it appears on the data file)	poolchl	poolCBOM	poolFBOM	poolFBIM	poolcarbon
NAME OF VARIABLE	Chlorophyll a in pools	Coarse benthic organic matter in pools	Fine benthic organic matter in pools	Fine benthic inorganic matter in pools	Benthic carbon in pools
DEFINITION OF VARIABLE	Benthic chlorophyll a in pools in mg per square m	Coarse (>1mm) benthic organic matter in pools in g per square m	Fine (<1mm) benthic organic matter in pools in g per square m	Fine (<1mm) benthic inorganic matter in pools in g per square m	Benthic carbon in pools in g per square m
UNIT	milligramsPerSquareMeter	gramsPerSquareMeter	gramsPerSquareMeter	gramsPerSquareMeter	gramsPerSquareMeter
PRECISION	0.1	0.1	0.1	0.1	0.1
RANGE OR LIST OF VALUES	{2.0,.,87.5}	{0.4,.,55.5}	{1.5,.,181.9}	{2.5,.,1296.6}	{0.2,.,80.0}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):,

12File Name or # above (all in which the variable appears)	5, 9	5	5	5	5
AbbreviationAbbreviation(as it appears on the data file)	poolnitrogen	rifflechl	riffleCBOM	riffleFBOM	riffleFBIM
NAME OF VARIABLE	Benthic nitrogen in pools	Chlorophyll a in riffles	Coarse benthic organic matter in riffles	Fine benthic organic matter in riffles	Fine benthic inorganic matter in riffles
DEFINITION OF VARIABLE	Benthic nitrogen in pools in g per square m	Benthic chlorophyll a in riffles in mg per square m	Coarse (>1mm) benthic organic matter in riffles in g per square m	Fine (<1mm) benthic organic matter in riffles in g per square m	Fine (<1mm) benthic inorganic matter in riffles in g per square m
UNIT	gramsPerSquareMeter	milligramsPerSquareMeter	gramsPerSquareMeter	gramsPerSquareMeter	gramsPerSquareMeter
PRECISION	0.01	0.1	0.1	0.1	0.1
RANGE OR LIST OF VALUES	{0.04,.,9.00}	{2.0,.,132.8}	{<0.1,.,13.8}	{2.2,.,28.0}	{0.7,.,144.5}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

13File Name or # above (all in which the variable appears)	6	6	6	6	6
AbbreviationAbbreviation(as it appears on the data file)	poolsnorkelgoby	poolsnorkelcrab	poolxipho	poolatyid	poolmac
NAME OF VARIABLE	Sicydium plumieri in pool snorkeling	Epilobocera sinuatifrons in pool snorkeling	Xiphocaris elongata in pool electroshocking	Atyidae in pool electroshocking	Macrobrachium in pool electroshocking
DEFINITION OF VARIABLE	Density of Sicydium plumieri in pools, as measured by snorkeling a known area in no. per square m (individuals/square m)	Density of Epilobocera sinuatifrons in pools, as measured by snorkeling a known area in no. per square m (individuals/square m)	Number of Xiphocaris elongata in pools, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Atyidae in pools, as measured by point abundance electroshocking in mean no. per point sample ( individuals per point sample)	Number of Macrobrachium in pools, as measured by point abundance electroshocking in mean no. per point sample ( individuals per point sample)
UNIT	numberPerMeterSquared	numberPerMeterSquared	number	number	number
PRECISION	0.01	0.01	0.1	0.01	0.1
RANGE OR LIST OF VALUES	{0,.,2.93}	{0,.,0.14}	{0,.,25.8}	{0,.,8.17}	{0,.,2.4}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

14File Name or # above (all in which the variable appears)	6	6	6	6	6
AbbreviationAbbreviation(as it appears on the data file)	poolgoby	poolcrab	poolguppy	pooltad	rifflexipho
NAME OF VARIABLE	Sicydium plumieri in pool electroshocking	Epilobocera sinuatifrons in pool electroshocking	Poeciliidae in pool electroshocking	Tadpoles in pool electroshocking	Xiphocaris elongata in riffle electroshocking
DEFINITION OF VARIABLE	Number of Sicydium plumieri in pools, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Epilobocera sinuatifrons in pools, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Poeciliidae in pools, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of tadpoles in pools, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Xiphocaris elongata in riffles, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)
UNIT	number	number	number	number	number
PRECISION	0.1	0.01	0.01	0.01	0.01
RANGE OR LIST OF VALUES	{0,.,0}	{0,.,0.33}	{0,.,16.33}	{0,.,14.67}	{0,.,5.67}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

15File Name or # above (all in which the variable appears)	6	6	6	6	6
AbbreviationAbbreviation(as it appears on the data file)	riffleatyid	rifflemac	rifflegoby	rifflecrab	riffleguppy
NAME OF VARIABLE	Atyidae in riffle electroshocking	Macrobrachium in riffle electroshocking	Sicydium plumieri in riffle electroshocking	Epilobocera sinuatifrons in riffle electroshocking	Poeciliidae in riffle electroshocking
DEFINITION OF VARIABLE	Number of Atyidae in riffles, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Macrobrachium in riffles, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Sicydium plumieri in riffles, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Number of Epilobocera sinuatifrons in riffles, as measured by point abundance electroshoc in mean no. per point sample (individuals per point sample)king	Number of Poeciliidae in riffles, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)
UNIT	number	number	number	number	number
PRECISION	0.1	0.01	0.01	0.01	0.01
RANGE OR LIST OF VALUES	{0,.,15.0}	{0,.,2.83}	{0,.,1.33}	{0,.,1.33}	{0,.,16.33}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

16File Name or # above (all in which the variable appears)	6	7	7	7	7
AbbreviationAbbreviation(as it appears on the data file)	riffletad	poolgyrinid	poolchironomid	poolodonata	poolbaetid
NAME OF VARIABLE	Tadpoles in riffle electroshocking	Gyrinidae in pools	Chironomidae in pools	Odonata in pools	Baetidae in pools
DEFINITION OF VARIABLE	Number of tadpoles in riffles, as measured by point abundance electroshocking in mean no. per point sample (individuals per point sample)	Biomass of Gyrinidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of non-tanypodid and non-leaf-mining Chironomidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Odonata in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Baetidae in benthic block net samples from pools in mg ash-free dry mass per block net sample
UNIT	number	milligram	milligram	milligram	milligram
PRECISION	0.01	0.01	0.01	0.01	0.01
RANGE OR LIST OF VALUES	{0,.,0}	{0,.,26.74}	{0.03,.,17.74}	{0,.,10.70}	{0.03,.,10.07}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

17File Name or # above (all in which the variable appears)	7	7	7	7	7
AbbreviationAbbreviation(as it appears on the data file)	poolhydroptilid	poolcalamoceratid	poolcaenid	poolceratopogonid	poolleptophlebiid
NAME OF VARIABLE	Hydroptilidae in pools	Calamoceratidae in pools	Caenidae in pools	Ceratopogonidae in pools	Leptophlebiidae in pools
DEFINITION OF VARIABLE	Biomass of Hydroptilidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Calamoceratidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Caenidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Ceratopogonidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Leptophlebiidae in benthic block net samples from pools in mg ash-free dry mass per block net sample
UNIT	milligram	milligram	milligram	milligram	milligram
PRECISION	0.01	0.01	0.01	0.01	0.01
RANGE OR LIST OF VALUES	{0,.,11.94}	{0,.,10.21}	{0,.,9.93}	{0,.,9.38}	{0,.,2.03}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

18File Name or # above (all in which the variable appears)	7	7	7	7	7
AbbreviationAbbreviation(as it appears on the data file)	poolancylid	poololigochaetaA	poolother	rifflehydroptilid	rifflesimuliid
NAME OF VARIABLE	Ancylidae in pools	Oligochaeta in pools	Other taxa in pools	Hydroptilidae in riffles	Simuliidae in riffles
DEFINITION OF VARIABLE	Biomass of Ancylidae in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of Oligochaeta in benthic block net samples from pools in mg ash-free dry mass per block net sample	Biomass of other taxa in benthic block net samples from pools	Biomass of Hydroptilidae in riffles in mg ash-free dry mass per square meter	Biomass of Simuliidae in riffles in mg ash-free dry mass per square meter
UNIT	milligram	milligram	milligram	milligramPerSquareMeter	milligramPerSquareMeter
PRECISION	0.01	0.01	0.01	0.1	0.1
RANGE OR LIST OF VALUES	{0,.,4.44}	{<0.01,.,2.63}	{0.12,.,5.01}	{6.2,.,2082.4}	{0,.,1231.6}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

19File Name or # above (all in which the variable appears)	7	7	7	7	7
AbbreviationAbbreviation(as it appears on the data file)	rifflechironomid	riffleblephericerid	rifflecrambid	rifflehydropsychid	rifflebaetid
NAME OF VARIABLE	Chironomidae in riffles	Blephericeridae in riffles	Crambidae in riffles	Hydropsychidae in riffles	Baetidae in riffles
DEFINITION OF VARIABLE	Biomass of Chironomidae in riffles in mg ash-free dry mass per squaremeter	Biomass of Blephericeridae in riffles in mg ash-free dry mass per square meter	Biomass of Crambidae in riffles in mg ash-free dry mass per square meter	Biomass of Hydropsychidae in riffles in mg ash-free dry mass per square meter	Biomass of Baetidae in riffles in mg ash-free dry mass per square meter
UNIT	milligramPerSquareMeter	milligramPerSquareMeter	milligramPerSquareMeter	milligramPerSquareMeter	milligramPerSquareMeter
PRECISION	0.1	0.1	0.1	0.1	0.1
RANGE OR LIST OF VALUES	{1.2,.,215.8}	{0,.,290.8}	{0,.,402.2}	{0,.,236.6}	{0,.,104.6}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

20File Name or # above (all in which the variable appears)	7	7	7	7	7
AbbreviationAbbreviation(as it appears on the data file)	riffleodonata	rifflepsychodid	riffleempidid	riffleceratopogonid	riffleother
NAME OF VARIABLE	Odonata in riffles	Psychodidae in riffles	Empididae in riffles	Ceratopogonidae in riffles	Other taxa in riffles
DEFINITION OF VARIABLE	Biomass of Odonata in riffles in mg ash-free dry mass per squaremeter	Biomass of Psychodidae in riffles in mg ash-free dry mass per squaremeter	Biomass of Empididae in riffles in mg ash-free dry mass per squaremeter	Biomass of Ceratopogonidae in riffles in mg ash-free dry mass per square meter	Biomass of other taxa in riffles in mg ash-free dry mass per square meter
UNIT	milligramPerSquareMeter	milligramPerSquareMeter	milligramPerSquareMeter	milligramPerSquareMeter	milligramPerSquareMeter
PRECISION	0.01	0.01	0.01	0.01	0.01
RANGE OR LIST OF VALUES	{0,.,22.06}	{0,.,11.63}	{0,.,26.46}	{0,.,17.15}	{0.20,.,1.31}
DATA TYPE	decimal	decimal	decimal	decimal	decimal
MISSING DATA CODES	none	none	none	none	none

VARIABLES (ATTRIBUTES):

21File Name or # above (all in which the variable appears)	8, 9	8	4, 8	4, 8	4, 8
AbbreviationAbbreviation(as it appears on the data file)	streamname	drainage	lowDON	lowNO3	lowDOC
NAME OF VARIABLE	Stream name	Drainage	Dissolved organic nitrogen	Dissolved nitrate	Dissolved organic carbon
DEFINITION OF VARIABLE	The name of the stream site	The name of the drainage the stream site occurs in	Amount of dissolved organic nitrogen in the stream water	Amount of dissolved nitrate in the stream water	Amount of dissolved organic carbon in the stream water
UNIT			milligrams nitrogen per liter	milligrams nitrogen per liter	milligrams carbon per liter
PRECISION			0.01	0.01	0.01
RANGE OR LIST OF VALUES	Duey, Guajataca, Guanajibo, Guayanilla, Inabon, Jacaguas, La Plata, Limon, Matrullas, Tallaboa, Tanama, Toa Vaca	Arecibo, Duey, Guajataca, Guanajibo, Guayanilla, Inabon, La Plata, Manati, Tallaboa, Tanama, Toa Vaca	{0.08,.,0.24}	{0,.,1.93}	{0.42,.,2.35}
DATA TYPE	alphabetic	alphabetic	integer	integer	integer
MISSING DATA CODES	none	none	none	none	none

COMPUTATIONAL METHODS:

DATE OF LAST REVIEW: May 2, 2009

DATE OF LAST ENTRY: August, 2000

STAGE OF DATA SET MANAGEMENT (dates):
RECEIVED: December 12, 2005
ENTERED: February 1, 2006
FILED ON-LINE: February 3, 2006
REVIEWED BY RESEARCHER:

FILING MEDIA:
NAME OF DOCUMENTATION FILE: EcologicalApplicationsMetadataForm.doc, lterdb139.htm
NAME OF ON - LINE CATALOG: LTERDBAS
RECORD #: 139
DOCUMENT TYPE: binary data only
PRIORITY TO BE ENTERED: N/A