Phytoneuron

. Fertig. 2011. Epiphytes and the National Wetland Plant List. Phytoneuron 2011-16: 1-31.

EPIPHYTES AND THE NATIONAL WETLAND PLANT LIST

Robert W. Lichvar

U.S. Army Engineer Research and Development Center

Cold Regions Research and Engineering Laboratory

72 Lyme Road

Hanover, NH 03755-1290

Walter Fertig
Moenave Botanical Consulting
1117 West Grand Canyon Drive

Kanab, UT 84741

ABSTRACT

The National Wetland Plant List (NWPL) is a list of species that occur in wetlands in the
United States. It is a product of a collaborative effort of four Federal agencies: the U.S. Army Corps
of Engineers, the U.S. Environmental Protection Agency, the U.S. Fish and Wildlife Service, and the
Natural Resources Conservation Service. The NWPL has many uses, but it is specifically designed
for use in wetland delineation for establishing the extent of Federal jurisdictional of wetland
boundaries. To be listed in the NWPL, a plant must be rooted in soil, so there is a direct relationship
between a plant's occurrence and its preference for hydric soils. This relationship, coupled with the
plant's frequency of occurrence in wetlands, is used to place it in one of five categories representing
the probability that the plant occurs in a wetland. Many species are considered to be epiphytes, but
they represent various life forms, ranging from purely epiphytic to frequently occurring on the
ground. Based on a literature review of 192 species across the United States and its territories, we
determined which species fell into four categories of epiphytic life forms or are terrestrial and should
not be considered epiphytes. Of the 192 species reported as epiphytes, 33 were determined to be
terrestrial and 107 can grow on the ground for at least part of their life forms. Only these 140 species
will be retained in the NWPL. This review documents the process of evaluating which epiphytes
qualify for being retained on the NWPL. Documentation includes the literature and its review to
support retaining the species on the NWPL. The reasoning behind removing Cuscuta from the list is
also documented.

KEY WORDS: wetlands, wetland plants, epiphytes, National Wetland Plant List, wetland
delineation

The National Wetland Plant List (NWPL) is used in wetland delineation and restoration of
wetlands, as well as providing a resource of botanical information about wetland plants. Each species
determined to be a wetland plant has been placed in one of five rating categories representing the
estimated probability, or frequency, with which it is thought to occur in wetlands, as opposed to
nonwetlands, across its entire range (Table 1). These category assignments were developed through a
thorough review of the botanical literature and the best professional judgment of national and regional

Currently the NWPL is being revised under the administrative direction of the LIS. Army
Corps of Engineers with cooperation from the U.S. Fish and Wildlife Service (USFWS), the U.S.
Environmental Protection Agency, and the Natural Resources Conservation Service. The designated
list for wetland delineation under Section 404 of the Clean Water Act is the 1988 list (referred to here
as the "88 list") (Reed 1988). The NWPL was updated in 1996 (referred to here as the "96 list," as
posted in a USFWS draft web publication) (Reed 1998), but the update was never officially finalized.

Lichvar and Fertig: Epiphytes and the National Wetland Plant List 2

The current revision of the NWPL will be based on more precise scientific criteria than for previous
lists, it will reflect changes in botanical nomenclature, and it will be divided into new geographic

regions based on ecological rather than political boundaries. Proposed changes in wetland ratings
will be vetted by botanists and wetland ecologists on regional and national panels, states, academics,
and the public using a national database with a web interface. The revision of the NWPL includes an
ongoing effort to assess the entire flora of the United States and its territories to ensure that the list is
comprehensive and complete.

Epiphytes — plants that grow on or are attached to other living plants (Schimper 1888) — are
a complication for the NWPL. In preparing previous wetlands lists, the USFWS applied an
unpublished rule that no epiphytes were to be included because they are not rooted in the soil (Reed,
USFWS, pers. comm.). The U.S. Army Corps of Engineers (Environmental Laboratory 1987)
defined the hydrophytic vegetation community for wetland delineation purposes as 'the sum total of
macrophytic plant life that occurs in areas where the frequency and duration of inundation or soil
saturation produce permanently or periodically saturated soils of sufficient duration to exert a
controlling influence on the plant species present." Tinder this definition, plants not rooted in the soil,
which the USFWS assumed to include epiphytes, cannot be considered hydrophytes.

Epiphytes are a diverse taxonomic group, with species in at least 33 families and over 230
genera worldwide. When hemi- epiphytic plants are included — species that can be both epiphytic
and occur on the ground — the number of families increases to 83 and the number of genera to over
875 (Gentry & Dodson 1987). About 80% of all vascular epiphytes are monocotyledons (Kress
1989).

A review of the literature quickly reveals that epiphyte species have a variety of life forms
and occur in a variety of habitats (Gentry & Dodson 1987, Wagner et al. 1999, Acevedo -Rodriguez
2005), bringing the simplistic no-epiphyte wetland rule into question. One life form that may violate
the logic behind the no-epiphyte rule is that of hemi-epiphytes. This group contains two forms with
different life forms, sometimes described as primary and secondary epiphytes. Primary hemi-
epiphytes begin their life form as epiphytes and later become rooted in the ground. Secondary hemi-
epiphytes begin life rooted in soil and later assume an epiphytic life form and are no longer rooted in
the soil (Putz & Holbrook 1989). Some species in both these hemi-epiphytes groups may warrant
consideration as wetland species, and it is possible that epiphytes with other life forms should be
considered as well.

The current efforts to update the NWPL includes 192 epiphytes or hemi-epiphytes, primarily
because earlier wetland plant lists by the USFWS had assigned wetland ratings to 122 epiphytic
species, even though this was in opposition to their own basic rule not to include any epiphytes. The
discovery of these epiphytes on the list prompted further investigation to see if any epiphytes met the
rule of needing to be rooted in the ground.

To support the scientific quality of the NWPL, we compiled a draft list of various categories
of epiphytes, evaluated the literature to develop ecological profiles of their life forms, and compiled
literature references to support the groupings of epiphytes for further consideration as wetland
species. The information presented here will support the updating of the NWPL for epiphytes and
will provide background for those species that could considered to be wetland plants and that should
be evaluated for a wetland rating.

i Fertig: Epiphytes and the National V

Methods.

A list of potential epiphytes that may warrant further consideration as wetland plants was
obtained as a collaborative effort with BONAP (Biota of North America Program). Kartesz (in press)
tracks the flora of North America and maintains an extensive database of distribution and biological
attributes based on a national inventory of herbaria, scientific literature, and information from
recognized specialists in many groups of plants. The BONAP database already contains a list of
epiphytes known within the United States and its territories, but the list is limited to a general
category identified only as epiphytes.

To identify whether some part of an epiphyte's life form includes being rooted on the ground,
we sorted the epiphyte list into five categories. We reviewed 59 literature sources to determine the
life form of each of the potential epiphyte species and to place each species into one of the categories.

3. Facultative epiphytes of trees and terrestrial sites. These plants can occur either on tree trunks c

or on the ground in soil (but not on rocky cliffs or boulders). They are never restricted tc
epiphytic life form. This group includes both categories of hemi-epiphytes.

4. Facultative epiphytes of rock and terrestrial sites. These pla

in soil on the ground, but they are never true epiphytes on trt

■restrial or at least rooted in soil on the ground. This group
lianas, and species that lean on other plants for support at
maturity but are not rooted to the host plant.

Additionally, the genus Cuscuta (dodder) was evaluated as a possible epiphtye. This genus
was not on the 88 and 96 lists. However, during the current update process, many people have
submitted the suggestion that this genus and some of its species deserve wetland ratings. This genus
is scattered throughout most of North America and is frequently found in wetlands.

Results.

Within the continental USA, Puerto Rico, Hawaii, and the Marianas in the south Pacific
(Figure 1), the area covered by the NWPL, there are 192 species reported to be epiphytes by BONAP
(Table 1). Of these, 52 are obligate epiphytes of trees or rocky cliffs and do not qualify as potential
wetland plants. Thirty-three species are primarily terrestrial and rarely (if ever) have a true epiphytic
or epipetric life form. It is possible that some of these species may be wetland plants, but they need
to be assessed during the updating process of the NWPL. A total of 107 species were found to be
facultative epiphytes (or epipetric species) that also occur in various terrestrial environments and are
rooted in soil during part of their life form. These species need to be evaluated as part of the updating
process of the NWPL to determine if their frequency of occurrence in wetlands meets wetland
indicator standards.

Of the 122 species of epiphytes that had been assigned wetland indicator ratings on the 88
and 96 lists, 91 were categorized as facultative epiphytes in our review and 31 were categorized as
obligate epiphytes that did not occur on the ground (Table 2). Of the remaining 70 species that are
reported here as epiphytes, the USFWS had assigned many of these species into two other categories;

these categories were "No Occurrence (NO) in any USFWS region," which had 15 species, and "Not

i Fertig: Epiphytes and the National V

enough Information to make 2 determination (NI)," which had 55 species. Voucher specimens :
exist to verify the occurrence of the species in the NO group in various locations of the U.S.

Cuscuta (dodder).

Cuscuta, in the Convolvulaceae family, is a genus of annual parasitic plants with a worldwide
distribution. In the USA they occur in every state except Alaska (BONAP 2010), Cuscuta spp. are
considered holoparasites — they depend entirely on their hosts for water and nutrients (Albert et al.
2008). Most species lack chlorophyll, and for those that do have chlorophyll, photosynthesis
provides for only a small amount of the plant's needs. Cuscuta spp. are considered pests on a wide
variety of plants, many of agricultural significance.

Cuscuta seeds germinate on or near the soil surface in a variety of habitats. As the rootless,
leafless stem grows, it rotates and coils around any vertical object. If the object is a suitable host, the
Cuscuta stem secretes an adhesive substance and induces the host to do the same. Then Cuscuta
grows haustoria, which are root tips that penetrate the host tissue and provide a pathway for water,
nutrients, and other compounds. At this point, the Cuscuta plant becomes detached from the soil and
has no more contact with the ground throughout the rest of its life. Once established on a host,
Cuscuta grows rapidly and can spread easily to nearby hosts. Cuscuta plants flower from late spring
to fall, and each plant can produce thousands of small seeds, which can remain viable in the soil for
10 years or more.

Cuscuta's life form as a holoparasite and its ability to break connection from the soil shortly
after germination when it begins its parasitic phase supports its elimination from the NWPL as
previously interpreted in the unpublished rule of the USFWS that wetland plants need to be rooted in

Discussion.

During the process of updating the NWPL over the past three years, all 192 species of
epiphytes evaluated in this review were included on the update list, since over half of them had a
previous wetland ratings in 88 and 96 lists. Some species had already been rated as wetland plants
but had not been vetted for their life form to determine if they frequently occur on the ground, so we
retained all 192 reported epiphytes on the NWPL until the public input phase is over. After the
updating is complete but before the list is finalized, those species determined to be obligate epiphytes
(including epipetric species) will be dropped from the NWPL. The remaining facultative epiphytes,
including those that are epiphytic on trees and shrubs and on rocks, will be retained. All species
retained, based on this review of their life form, that received a wetland indicator status during the
review process will be included on the final NWPL.

ACKNOWLEDGMENTS

Funding for this study was provided by the Wetland Regulatory Assistance Program (WRAP)
of the U.S. Army Corps of Engineers. We thank John Kartesz of Biota of North America (BONAP)
for sharing the list of species in the United States and its territories that have been referred to as
epiphytes and for the use of Figure 1.

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Table 1. Wetland Indicator Status Ratings and their cardinal rating categories, as described i
the National List of Plant Species that Occur in Wetlands (Reed 1988).

ImlkMlor SI. iltiv ultltn-v i.iIkmii

% Occurrence in
Wetlands

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99

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67-99

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1-33

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Lichvar and Fertig — Epiphytes and the National Wetland Plant List

Figure 1. Density gradient, map of occurrences of epiphytes in the United States and its territories,
according to BONAP (2010) The figure shows the richness coefficient of epiphytes. Darker colors
indicate greater richness. Puerto Rico, Hawaii, Florida, and Alaska are not drawn to scale so that
differences in richness will he more apparent.

l-liirrslitSymlK.'sisufNA < BONA I' 21) I II

Legend

Grey = outside study area
Dark tan = no species reported
Light tan = 1-10 species
Chartreuse = 1 1-20 species

Lime green = 21-30 species
Kelly green = 31-60 species
Dark green = 71-80 species
Black = 81-100 species