3. PARSING THE NETWORK
As with most methodologies,
some of the most important assumptions in Network Analysis are made at the outset.
They concern decisions as to what to choose as the primary elements of the network
and how these taxa are to be connected. In this project, these decisions are
simplified somewhat by the requirement that the network be comparable to the
ATLSS simulation. That is, each state variable of the ATLSS model must have
its counterpart in the accompanying network.
Network analysis, however, can
treat far more complicated webs of interaction than are possible using simulation
modeling. Keeping the simulation dynamics of many coupled processes from becoming
pathological is a difficult balancing act. Because NA does not deal explicitly
with dynamics, far greater taxonomic resolution becomes possible with this form
of analysis. As a result, some taxa in the ATLSS model will be represented by
several compartments in the graminoid network. In addition, compartments can
be added to the network with relative ease. For example, the creation of an
individual-based model for one of the lesser-known bird species (e.g. the swamp
sparrow) would entail significant effort. As will be described below, it requires
nowhere near as much work to include this compartment in the quantified trophic
web.
Although the primary reason
for creating these networks is to serve as a calibration benchmark for ATLSS,
it should be mentioned that each single network and its ensuing analysis could
also serve independent purposes. For example, some biologists will be curious
to know how the particular species that they are interested in, which might
not appear in ATLSS, will fare under the proposed hydrological scenarios. With
regard to trophic interactions at least, the results of NA should allow those
investigators and managers to make some educated guesses about how the excluded
population might change. For example, NA quantifies the direct and indirect
trophic interactions of each compartment with all other compartments in the
web. In particular, the stocks and activities of each compartment that do not
appear in ATLSS can be compared with all those that do participate through NA.
Any coefficients and ratios as may result can be applied to the predicted outputs
from ATLSS to estimate the accompanying trophic status of the non-included species.
In other words, after calibration NA can serve to expand the scope of predictions
from ATLSS.
From an ecological viewpoint,
virtually all the important trophic components (i.e., those comprising at least
5% of the standing biomass or activity) have been written into ATLSS. These
include detritus, microbes, aquatic macrophytes, mesoinvertebrates, macroinvertebrates,
piscivorous fish, planktivorous fish, etc.
As for the previous
three networks, the cypress wetland ecosystem, the Florida Bay ecosystem and
the mangrove ecosystem, it happened that data did exist to make at least crude
estimates of the stocks and activities of a number of some other ecosystems
members. Similarly, it was decided to include these "parallel" species in the
NA of the graminoid network.
In addition, in order to wind
up with more homogeneous and comparable networks, the choices that were made
in the three previous network models have been taken into consideration in drawing
up the list of components for the graminoid ecosystem. Because many of the species
that are resident in the mangroves and cypress systems feed in the graminoid
marshes, there will be interactions between these systems, and some redundancies
among their species. For example, most of the fishes and invertebrates listed
for cypress system are present as well in the graminoids. The same goes for
many of the mammals and birds in the cypress and the birds in the mangroves.
Conversely, the estuarine and marine species of the mangrove and Florida Bay
systems are not seen in the freshwater graminoid marshes. This results in a
lower number of compartments in the graminoid ecosystem than comprised any of
the preceding three systems. In deciding which elements to include in the network
and how these taxa are to be connected, the same resolution was used as was
employed with the previous three biotopes, in order that the elements and structure
of the four different network models will remain comparable.
For some categories, such as
invertebrates and fish, individual compartments for each of the more abundant
and better-documented species have been reserved, while the remaining species
were combined into single "other macroinvertebrates", "other
small fish" and "other large fish" compartments.
In many networks the
major lexical question involved how to treat the manifold species that comprise
some functional groups for which data at the level of individual species were
not available. To characterize these poorly resolved parts of the ecosystem,
it was decided to group the species into generalized compartments. Such was
the case in the graminoid network for mesoinvertebrates and passerine birds.
The same aggregation has been done for bacteria and protozoans in the water
column and sediments, which were assigned to a compartment called "living
POC" and "living sediments", respectively.
The completed networks both
consist of 66 separate components. Only those species closely associated with
the dominant patterns have been retained. Hence, only those primary producers,
invertebrates, fishes, reptiles, birds and mammals commonly found in the graminoid
system made the final list of species. Once completed, the list was aggregated
somewhat according to certain criteria, such as species sharing the same diet,
available data and the goals of ATLSS. For example, most species to be included
in ATLSS as individual-base models have been maintained as separate compartments
in the network.
The following is the final list
of components composing the graminoid ecosystem network, along with a very brief
description of what each compartment entails:
3.A Living Microbial
compartments
The
initial elements of the network are the living compartments of the microbial
loop. They have been divided as follows:
- LIVING SEDIMENTS. Includes all
bacteria and fungi living at the sediment level.
- LIVING POC. Bacteria and protozoans
that occur in the water column are all represented in this compartment.
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3.B Primary producers
A description of the dominant primary
producers is reported in the "study area" section of this report. They have
been divided as follows:
- PERIPHYTON. Mostly diatomaceous
forms of algae.
- MACROPHYTES. The macrophyte assemblage
includes sawgrass (Cladium jamaicense), two species that coexist with
sawgrass Peltandra virginica and Pontederia cordata.
Eleocharis spp. was most abundant within the wet prairie community,
while Hymenocallis palmeri, Panicum hemitomon, and Sagittaria lancifolia,
co-occur within the wet prairie community (Daoust
& Childers, 1999).
- UTRICULARIA. The vascular plants
included in the floating mat consist of two species: Utricularia chara
and Utricularia bacopa (not common). Utricularia is a carnivorous
plant that generally occurs in nutrient-poor habitats that are sunny and moist
(at least during the growing season), where they are often dominant (Givnish,
1989).
- FLOATING VEGETATION. The floating
vegetation includes all non-rooted plants, and a few that may or may not be
rooted. Examples include as pond lilies and duckweed.
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3.C Invertebrates
As might be expected, the invertebrates
of the Everglades graminoids are similar in species composition to those of
the Cypress system. However, better data were available for the construction
of the graminoid network, so that the invertebrate assemblage could be expanded
in the newer graminoid model.
- APPLE SNAIL. This compartment
contains only Pomacea paludosa, the apple snail. Other snails are found
in Aquatic Macroinvertebrates.
- FRESHWATER PRAWN. Contains only
the freshwater shrimp Palaemonetes paludosus.
- CRAYFISH. The Everglades are
populated almost exclusively by Procambarus alleni. There is some evidence,
however, that another species, Procambarus clarkii, which has been
introduced throughout Florida, may be present in some areas of the Everglades
(Franz and Franz 1990).
- MESOINVERTEBRATES. This is the
first of four general invertebrate compartments. It contains small aquatic
invertebrates, including animals from the following taxonomic groups: Oligochaeta,
Bosminidae, Chydoridae, Daphniidae, Macrothricidae, Sididae, Calanoid and
Harpacticoid copepods, Chironmidae. Fish larvae of comparable size are also
included in this compartment.
- OTHER MACROINVERTEBRATES. The
second of the general invertebrate compartment contains all aquatic invertebrates
larger than the mesoinvertebrates (just described) and smaller than the predatory
aquatic insects. Representative animals from the following taxonomic groups:
Trichoptera, Certaopogonidae, Tanypodinae, Gastropoda (excluding apple snails).
Fish larvae of comparable size are also placed in this compartment because
of their similar trophic stature.
- LARGE AQUATIC INSECTS. All large
predatory aquatic insects and larvae (e.g., dragonfly and damselfly nymphs)
found in the graminoid marshes.
- TERRESTRIAL INVERTEBRATES. This
compartment contains all the invertebrates that are active in non-aquatic
environments. There are very little quantitative data about these organisms,
and the data here have been extrapolated from tropical grasslands outside
the Everglades (Janzen and Schoener 1968). Organisms in this compartment will
include adult forms of members of the aquatic invertebrates (Odonates, Ephemeroptera)
and also any of the exclusively terrestrial insects (Lepidopterans, ants,
wasps, etc.).
- FISHING SPIDER. This compartment
contains only the fishing spider, Dolomedes triton, because of its
unique feeding habits (Jordan et al. 1994). The
other spiders are found in the Terrestrial Invertebrates compartment.
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3.D Fishes
The fishes in the graminoids
are generally the same species that were found in Cypress, however the fish
assemblage in the Cypress was divided into only three compartments, small, herbivorous-omnivorous
fish, small predatory fish and large, predatory fish. These assemblages could
be separated in the graminoid marshes due to significantly more quantitative
information available for this system.
- GAR. A member of the family Lepisosteidae,
Lepisoseus platyrhinchus, one of the top predatory fish species.
- SHINERS AND MINNOWS. This compartment
contains only the diminutive minnows of the family Cyprinidae, including such
species as: golden shiner (Notemigonus chrysoleucas), tailight shiner
(Notropis maculatus), coastal shiner (Notropis petersoni) and
pugnose minnow (Opsopoedus emiliae)
- CHUBSUCKERS. Lake chubsucker
(Erimyzon succetta) is a member of the family Catastomidae.
- CATFISH. Contains species from
the family Ictaluridae and Clariidae. Typical species are: white catfish (Ameiurus
catus), yellow bullhead (A. natalis), brown bullhead (A. nebulosus),
channel catfish (Ictalurus punctatus) and tadpole madtom (Noturus
gyrinus). The exotic walking catfish (Clarias batrachus) is the
only representative from the family Clariidae.
- FLAGFISH. One of the most abundant
fishes in the Everglades, the flagfish (Jordanella floridae) is a member
of the family Cyprinodontidae.
- TOPMINNOWS. The topminnow species
of the family Fundulidae. Included representatives are: Golden topminnow (Fundulus
chrysotus), banded topminnow (F. cingulatus) and lined topminnow
(F. lineolatus).
- BLUEFIN KILLIFISH. One member
of the family Fundulidae, the bluefin killifish (Lucania goodei), has
been given its own compartment, because it is so abundant.
- KILLIFISHES. This compartment
contains the remaining species of family Fundulidae, such as the Seminole
killifish (Fundulus seminolis) and the marsh killifish (F. confluentus).
- MOSQUITOFISHES. Consists of only
the Poecilid species, Gambusia affinis. The mosquitofish is another
highly abundant small fish species.
- POECILIDS. Contains the remaining
Poecilds, including the very abundant species least killifish (Heterandria
formosa) and sailfin molly (Poecilla latipinna). It also contains
an introduced Poecilid species, the pike killifish (Belonesox belizanus).
- PYGMY SUNFISH. The Centrarchids
(sunfishes) have been split into as many single species compartments as possible,
due both to their diverse feeding habits and to the large proportion of total
fish biomass that they comprise. This compartment is devoted to the Everglades
Pygmy Sunfish (Elasoma evergladei).
- BLUESPOTTED SUNFISH. Enneacanthus
gloriosus is one of the smaller Centrarchid species.
- WARMOUTH. The warmouth (Lepomis
gulosus) is one of the larger predatory species of sunfish in the Everglades.
- DOLLAR SUNFISH. Lepomis marginatus.
- REDEAR SUNFISH. The redear sunfish
(Lepomis microlophus) consumes primarily gastropods.
- SPOTTED SUNFISH. Lepomis punctatus.
- OTHER CENTRARCHIDS. Contains
any previously unmentioned Centrarchid, most notably the bluegill (Lepomis
macrochirus) and the black crappie (Pomoxis nigromaculatus).
- LARGEMOUTH BASS. Micropterus
salmoides is one of the top predator fishes.
- CICHLIDS. This compartment contains
the assemblage of introduced Cichlids: Oscar (Astronotus oscellatus),
peacock cichlid (Cichla ocellaris), black acaria (Cichlasoma bimaculatum)
mayan cichlid (Cichlasoma uropthalmus), blue tilapia (Tilapia aurea),
and spotted tilapia (Tilapia mariae).
- OTHER LARGE FISHES. This is the
first of two generic fish compartments and contains the rarer large freshwater
fish of the Everglades as well as those euryhaline species that sometimes
invade the freshwaters. Included are: Bowfin (Amia calva), American
eel (Anguilla rostrata), pickerels (Esox spp.) shads (Dorosoma
spp.), mullets (Mugil spp.) and snooks (Centropomis spp.)
- OTHER SMALL FISHES. This is the
second of two generic fish compartments and contains the rarer small freshwater
fish of the Everglades as well as those euryhaline species that are known
to invade freshwaters at various times of the year. Included are Silversides
(Menidia spp.), gobies (Gobiosoma spp. and Microgobius
spp.) and swamp darter (Etheostoma fusiforme).
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3.E Reptiles
and Amphibians
Although not composed of exactly
the same species, the list of reptile compartments resembles that of the cypress
ecosystem. Reptiles have been divided into four separate compartments: snakes,
lizard, turtles and alligators, most consisting of several species, with the
exception of one single- species taxon, devoted to the American alligator (Alligator
mississippiensis). Amphibians were divided into six separate compartments:
salamanders, salamander larvae, large frogs, medium frogs, small frogs and tadpoles
(Diffendorfer, et al. 1999).
- SALAMANDERS. The salamander compartment
includes the dwarf and greater sirens (Pseudobranchus striatus and
Siren lacertina), the two-toed amphiuma (Amphiuma means) and
the pensinsula newt (Notophthalmus viridescens) (Diffendorfer,
et al. 1999).
- SALAMANDER LARVAE. The larvae
of all salamander species present in the graminoids are lumped here.
- LARGE FROGS. Only one species
of large frog is found in the graminoids: the pig frog (Rana grylio)
(Diffendorfer, et al. 1999).
- MEDIUM FROGS. Medium frogs include
the southern toad (Bufo terrestris), green treefrog (Hyla cinerea),
Cuban treefrog (Osteopilus septentrionalis), Florida chorus
frog (Pseudacris nigrita), southern cricket frog (Acris gryllus),
squirrel treefrog (Hyla squirella) and southern leopard frog (Rana
sphenocephala) (Diffendorfer, et al. 1999).
- SMALL FROGS. Small frogs include
the oak toad (Bufo quercicus), eastern narrowmouth toad (Gastrophryne
carolinensis), greenhouse frog (Eleutherodactylus planirostris)
and the little grass frog (Limnaoedus ocularis) (Diffendorfer,
et al. 1999).
- TADPOLES. The tadpoles of all
the frogs in the graminoids are gathered into this compartment.
- TURTLES. Turtles include the
common musk turtle (Sternotherus odoratus), striped mud turtle (Kinosternon
baurii), eastern mud turtle (K. subrubrum subrubrum), Florida box
turtle (Terrapene carolina bauri), chicken turtle (Deirochelys reticularia),
peninsula cooter (Pseudemys floridana), Florida redbelly turtle (P.
nelsoni), red ear slider (P.scripta), Florida snapping turtle (Chelydra
serpentina) and the Florida softshell turtle (Apalone ferox) (Diffendorfer,
et al. 1999).
- SNAKES. Snakes found in the graminoids
include: the Florida water snake (Nerodia fasciata), Florida green
water snake (N. floridana), brown water snake (N. taxispilota),
striped crayfish snake (Regina alleni), Florida brown snake (Storeria
dekayi victa), black swamp snake (Seminatrix pygaea), eastern garter
snake (Thamnophis sirtalis), peninsula ribbon snake (Thamnophis
sauritus), southern ringneck snake (Diadophis punctatus), rough
green snake (Opheodrys aetivus), mud snake (Farancia abacura),
black racer (Coluber constrictor), eastern indigo snake(Drymarchon
corais couperi), red rat snake (Elaphe guttata), yellow rat snake
(E. obsoleta), Florida kingsnake (Lampropeltis getulus floridana),
scarlet kingsnake (L. triangulum elapsoides), scarlet snake (Cemophora
coccinea), cottonmouth (Agkistrodon piscivorus), dusky pygmy rattlesnake
(Sistrurus miliarius barbouri) and eastern diamondback rattlesnake
(Crotalus adamenteus) (Diffendorfer, et al, 1999).
- LIZARDS. Lizards in the graminoid
Everglades include the brown anole (Anolis sagrei), green anole (A.
carolinensis), ground skink (Scincella laterale), island glass
lizard (Ophisaurus compressus) and the five-lined skink (Eumeces
inexpectatus) (Diffendorfer, et al. 1999).
- ALLIGATORS. The American alligator
(Alligator mississippiensis).
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3.F Mammals
Mammals in the graminoids show
considerable overlap with those listed in the cypress wetland network.
- MUSKRAT. Colonies of muskrat
(Neofiber alleni) are abundant throughout the Shark River Slough and
the open glades of the ENP (Tilmant, 1975).
- MICE AND RATS. Species found
in the graminoids include the cotton mice (Peromyscus gossypinus),
cotton rats (Sigmodon hispidus), rice rats (Oryzomys palustris)
and the exotic house mouse (Mus musculus).
- RABBITS. The marsh rabbit (Silvilagus
paulstris).
- RACCOONS. Raccoons (Procyon
lotor) are very abundant in the graminoid wetlands.
- OPOSSUMS. The Virginia opossum
(Didelphis virginiana).
- OTTER. The river otter (Lutra
canadensis).
- MINK. The mink (Mustela vison).
- DEER. The white-tailed deer (Odocoileus
virginianus).
- BOBCAT. Lynx rufus.
- PANTHER. The Florida panther
(Felis concolor).
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3.G Birds
Taxonomy, feeding habits and
the availability of information were primary concerns in establishing the final
list of birds. Species of birds that feed in the graminoids, but do not roost
there, were not included, as they have already been accounted for in the cypress
and mangrove models. Therefore, the compartments given in this model are mostly
those not found in the other models. For lists of all species to be found in
the graminoid Everglades, see Stevenson and Anderson
(1994).
- GREBES. Only one species of grebe
is found in the graminoids, the pied-billed grebe (Podilymbus podiceps).
- BITTERNS. One species of bittern
breeds in the graminoids, the least bittern (Ixobrychus exilis).
- DUCKS. Eight species of Anseriformes
breed or roost in the graminoids: fulvous whistling duck (Dendrocygna bicolor),
green-winged teal (Anas crecca), mottled duck (Anas fulvigula),
mallard (Anas platyrhynchos), blue winged teal (Anas
discors), northern shoveler (Anas clypeata), ring necked ducks
(Aythya collaris) and the ruddy duck (Oxyura jamaicensis) (Stevenson
and Anderson, 1994).
- SNAILKITES. The snail kite (Rostrhamus
sociabilis) is the most important species of Accipitridae in this system.
- NIGHTHAWK. Only one species of
Caprimulgiformes roosts or breeds in the graminoids, the common nighthawk
(Chordeiles minor).
- GRUIFORMES. Including the American
coot (Fulica americana), limpkin (Aramus guarauna), king rail
(Rallus elegans), common moorhen (Gallinula chloropus) and Sandhill
crane (Grus canadensis pratensis) (Stevenson
and Anderson 1994).
- CAPE SABLE SEASIDE SPARROW. The
endangered Cape Sable seaside sparrow (Ammodramus maritimus mirabilis)
breeds in the graminoids.
- PASSERINES. The passerines that
roost or breed in the graminoids include the red winged blackbirds (Agelaius
phoeniceus), common yellowthroat (Geothlypis trichas), swamp sparrow
(Melospiza georgiana), eastern meadowlark (Sturnella
magna), boat-tailed grackle (Quiscalus major) and the common grackle
(Quiscalus quiscula) (Stevenson and Anderson
1994).
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3.H Detritus Compartments
Detritus has been divided into
three separate compartments:
- SEDIMENT CARBON. Includes all
the non-living organic carbon available in the first 10 centimeters of sediment.
- LABILE DETRITUS. Liable detritus
is organic matter in the water column that decomposes at a relatively fast
rate. As with the cypress network, we use the information reported in Dierberg
and Ewel (1986) for the decomposition rate of labile detritus (5.7/y.)
- REFRACTORY DETRITUS. Refractory
detritus represents organic matter that decomposes at a relatively slower
rate than labile detritus (0.17.y-1) (Dierberg
and Ewel, 1986).
Seasonality at this latitude is marked more by variations in freshwater input
than by changing temperature. Thus, two separate networks mimic seasonality:
one for the wet season (from June to November) when freshwater inputs are high
and another for the dry season (December to May), when inputs are relatively
low.
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