. 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. 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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 Obligate (OBL) 99 I.kiiIi.iIiw WuI.nkI.I V\\ , 67-99 I.KIlll.lllWil V'l 34-66 I.kiiIi.iIiw 1 pl.iiul.l VI . 1-33 Upland (UPL) 1 H -8 t s if p ■ a 1 u il ii if 8-3 •3 1 ■c ,§ 1-ss ^ 7 1 1 t.E 8" 00 g g s. - is M ■a dcj II ■Si = 1 9 6 |< « ° 4 1 S.-8 •2 j § HI ill 1 1 1 i i ^ 1 e 6 « c tl1| *%>% I'll! ll •i E 1| 1 S g sl ^ .3 § 3) 4 ! 1 i 1| n § -C fa 1 s i'i = 4 -- f . |1| 1 1 s til ll 1 ||f a 1 .3 1 11 il "a ^ :&§> fr-g S(§ •3 4 = 1 P g £| 4 || If lj| ill X X X X X X X X 1 i 1 3 : « g III 1 Ig.sf J 1 ."J Hi a. > « a III s 3 a f -s-i ££3 1 III Its •8 Is 3 % I e2 1 « si 11 3 H 1 || iS "S. .3 jS I | |j 1 1 | | K i 1 1 5 8 1 a 11 SI 1 1 II £ 'So 1 s i E 5 "§ S3 II |] ~1 '? a; § 3 m |g 11 II 1 If 1! 1! id ■5 J "§| 1| |1§ it X X £ °> Is d s IS o '§■ d £ 5 l.l°J ^ > & 1 m 5 1 1 S3 1 1 c 111 1 11 ~s "^ ?■ 1 s '111 8.2 8- II IS . .1 d | .s I gM 1 a c^ d, > ^ h M _£± r>J | Jl|il«lf B-?8 &^| | 1 1 I'l "3 |1 '§ J | r ; P. 0=3 ■IflF 3: :&isijii l ii II 1.1 i E 5 1 .a S s |£ •« .3 .S E o 1 H ill i, 1 1 "s i'lP 1 •§■ i .3 '£ 8 1 g ii §£ s ,S £ £ 8 1 t || P ii s 8 1 1 1 1 J3 ,1 II 11- 111 I 1-3 -• v - S g S3 ■S S 1 Is s. j - ^ in '13 1 If II - _ S g 1 £ ii ■■; = tl § = i te || j| ll 1 1st f |a ^1"3 s g« 1 s 1'' --. = = it £ °> Is d s IS X X X X o '§■ d X X X £ 5 i is 3 a i a i .1 3 5.3-f .§ 1 M M fe d< > .3 a 111 5 g Jj g 111 3^ & 1 "S J J § g 11 8 1 ll g II mi- 1 c Hi t! 1 -8 1|i 1.1 1 ll : -'= . e-g's ||| I si 1.11 •£■ a g S3 S I = '| lis •lis III ■1 -g .8 ! s-f tj « <=- ^> 8 liJ s <§ 1 .3 1 i? s 1 n X X I i £ a i £ ll Is 3* I § I I Is if 1 N 3J f i 'S f. sts H STr*l Hi 1 i i " 1 1 '& 1. 6 "111. :§=!•§ i ?M 'S •§ ■& 'f,| | _ ■|| | § Lithophytic or terrestrial herb in moist and wet forests from sea level to above 1000 m (var. angustifolia) or on ridges, slopes, streams, banks, and edges of forest in rocky wet montane forests (var. simplicior) (Acevedo-Rodriguez & Strong 2005) X X X X 1! i a ££3 1 .s 11 < s J fa tq 1 11 HI 1 U op III f J J§ lllg J .11 1 I'M §•« - 111 l| & 1 j « J- 8:a 5 -, ~ = - - >, = lilt 5 Is- 1 § ^ 1 if "A 1 §•! g| 1 P X X 1 1 X ffi Hawaii, Florida, Puerto Rico, Virgin Islands ffi£^3 a .s mi o'S3 8 .3 ill S3 3 1 1 £?8 ■: : i £?1 1 1 51 51 Is %8 's-1 '§■§ 'g-S s8 § i E 5 1 1 1 S _ ll || g g || 1 IS is '8-.S || '£ -§ I t> 1 Is 11 &"£ g § if s 1 1 II '&! Id Pi H<2 ! s 1 3 S 8 f g 8 e 1 1 8 § % t§ g2 'S a || 1 ■s 1 f la g2 1 s s 1 8 1 1 f g H 2 8 a 1 | i 1 fa g 2 1 § - I ■IL '-Ron f 1 js g 'S ^ 8 1 1 1 3 g ^ •S « o ■lis 1-1 ill ill O Is d s IS X X X X X X X X X X o '§■ d £ 5 ffi is 3 ffi * * 3 s 3 ffi ffi ¥ 3 i 5 an S o| 60 S ml ll 1 '" 1 '1 1 ll a .3 ll S3 Jl 1 11 IS 1 c 1 H§ HI 6 ll 11 Si ol, 1 €> o | II 6 1 i E 5 f f! ■-I II J| -IS HI 8" || ||| "8-jg » i 11 it X £ °> Is d s IS X o '§■ d £ 5 i 1 M I Jfi ffi fe d, > ,3 5 n fi 1 1 1 c 1 1 1 1 ^n ill! ■ill S'Bb 111 1-8 & ■M sis ||| 111 1 g 1 o •S S3 g> 11! !-* - a li S S3 s lil ;! s^ 2 ? - ? "o S3 I -S '> = §-! ill! 111! Questionably epiphytic: "epipetric or terrestrial on rock ledges, talus slopes, and on soil in shade and exposed places" (Lellinger 1985). Montgomery and Wagner (1993) note all Dryopteris taxa in North America as being terrestrial, rarely growing on rock •L. a,l IS. 'I| 11 1 s S3 « 8. a 1 | X X X 1 III 1-a 8 %< 66B s 3 5-1 ■if M ll Jill l|j e|fe 111 1 1 f !l 8 -S 1 t a 1 1 1 » & 3 ^ •B 3 1° 1 s| j — s 8 <§ 1 s |l 1 s !■! !>•§ 14 ll '?< f 1 it X £ °> Is d s X X IS X X X X X X X X X o '§■ d £ X 5 ffi ffi is 3 i a s 3 S 3 is 1 i is I i 5 1 li 1 ll i 1 ll i | Li 1 ll -5 J2 ll 1 11 li 1 ll ll 1 c i I 53 "a 1 || 3 b 1 It §, 1 1 Ii S3 •§ 1 1 S3 ; ll :• "Si 1 o" & if S3 g. 1 |l "§..3 S3 I 1 ti 55 1. 1 "1- * if i E 5 Unlike all other species in the genus, Lomariopsis kunzeana rarely climbs trees and usually occurs in hammocks or limestone sinkholes (Moran 1993); Lellinger (1985) considers the species epipetric in limestone sinkholes ST 13 s § 3 i |1 a 8 | £ te fit =3 ! ^ 'C .2 S "S £,„ 1 if III 1 8;8|| .1 S !'§■ 1° °'% |& 'o 1 ft, ~ £ "5 = - 1 n Ms S 1 H Is si := § H 1 s M 1 1 1 'S S3 e 1 o S .a op ■;.'; --=. 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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