Environmental change in the terrestrial vegetation of Bermuda: Revisiting Harshberger (1905)

Abstract

In June of 1905 the pioneer ecologist John W. Harshberger visited Bermuda to investigate sand dune plants, survey the main plant formations and collect herbarium specimens. His specimens and observations are of significant value as a baseline to understand the extraordinary ecological change due to plant invasion that the island has experienced over the last 100 years. His observations on the vegetation types of Bermuda are contained in a six-page summary of a talk that he gave to the Academy of Natural Sciences of Philadelphia. He divided the terrestrial vegetation of the island into eight types. While five of these, mangrove swamp, salt marsh, brackish marsh, cliff rock, and sand dune, have vegetation that is lightly invaded and still recognizable from his descriptions, three others, cedar forest, limestone sinks forest, and scrub, have been altered by invasive plants beyond recognition except in a few small nature reserves that are actively managed against invasives. The seed rain from the introduced starling feeding on the fruits of invasive plants is heavy and ubiquitous. Any conservation of terrestrial vegetation now requires weeding against invasives in perpetuity, a “conservation treadmill” that needs to be budgeted for and carefully applied if conservation is to succeed.

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Acknowledgements

We thank Wendy Frith for logistical assistance and Dr. D. M. Percy for assistance in the field. Dr. Craig W. Schneider kindly provided comments on Harshberger’s algae. We are also indebted to Dr. Thomas Zanoni and one anonymous referee for their helpful reviews.

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Correspondence to Quentin Cronk.

Appendix 1

Appendix 1

Named localities and species mentioned by Harshberger (1905). Names of species are as given in the original text. For currently accepted plant names see Appendix 2.

Locality Terrestrial Plant Formation in 1905 Species mentioned (1905)
Walsingham
saltwater ponds
Marine: tidal because fed by underground channels Valonia utricularis, Caulerpa plumaris, C. racemosa, Colpomenia sinuosa
Castle Harbour Mangrove: dense growth of two mangroves in association (rare) Avicennia nitida, Rhizophora mangle, Penicillus capitatus
North Shore inland ponds Mangrove: swamps (inland) Avicennia nitida, Pluchea purpurascens, Sesuvium portulacastrum, Salicornia fruticosa, Heliotropium curassavicum, Cyperaceae spp.
Shelly Bay saltwater pools Mangrove: swamps (inland) Ulva lactuca, Avicennia nitida
Salt marsh along Harrington Sound (Wistowe) Salt marsh Salicornia fruticosa, Heliotropium curassavicum, Sesuvium portulacastrum, Cyperaceae spp., Stenotaphrum americanum, Avicennia nitida (rare)
Smith’s Parish Marsh Brackish marsh: devoid of trees, soil rendered brackish by underground channels Typha angustifolia, Baccharis heterophylla, Dichromena leucocephala, Scirpus lacustris, Osmunda cinnamomea,
Acrostichum aureum
Devonshire Marsh Brackish marsh: a large area with a few pools of water in the depressions, probably once a pond (by 1905 shaded by encroaching trees – Sabal blackburniana and Juniperus bermudiana) Juniperus bermudiana, Sabal blackburniana, Baccharis heterophylla, Pteris (Pteridium) aquilina caudata, Cladonia sp., Leucobryum sp., Typha angustifolia, Acrosticum aureum. Pools: Sphagnum spp., Proserpinaca palustris, Lemna minor [Coulter: Hydrocotyle asiatica, Herpestis monnieria, Mentha viridis, Dichromena leucocephala, Osmunda regalis and O. cinnamomea].
Pembroke Marsh Brackish marsh: similar to Devonshire Marsh Juniperus bermudiana, Sabal blackburniana around edge, Lemna minor in pool; [Lefroy: Nasturtium officinale (in water channels), Ascyrum crux-andreae, Kosteletzkya virginica, Waltheria americana (= indica), Eclipta erecta, Pluchea odorata, Ceratophyllum demersum, Kyllinga monocephala, Scirpus validus, Cladium occidentale (= germanicum) and Spiranthes brevilabrus].
Shelly Bay dunes Low dunes on North Shore with zonation briefly described Cakile aequalis, Tournefortia gnaphalodes, Scaevola Plumieri, Croton maritimus, Ipomoea pes-caprae, Stenotaphrum americanum, Juniperus bermudiana, Sisyrinchium bermudianum, Lepidium virginicum, Euphorbia buxifolia, Canavalia obtusifolia, Opuntia vulgaris, Yucca aloifolia (sandy hills)
Tucker’s Town dunes Dunes: here Harshberger describes Conocarpus erectus forming thickets, which in places protect the dune crest Scaevola plumieri, Solidago sempervirens, Borrichia arborescens, Dodonaea viscosa, Conocarpus erectus, Stenotaphrum americanum, Foeniculum vulgare
Paget dunes [prob. Elbow Beach] High dunes but highly invaded by exotics Exotic: Nerium oleander, Lantana camara, Lantana crocea, Native: Croton maritimus, Canavalia obtusifolia, Dodonaea viscosa, Borrichia arborescens, Passiflora suberosa
Cliff rock (general) General coastline of Bermuda: jagged rocks Borrichia arborescens, Conocarpus erectus, Solidago sempervirens, Lantana involucrata, Lantana camara, Euphorbia buxifolia, Stenotaphrum americanum, Juniperus bermudiana, Yucca aloifolia, Sabal blackburniana, Coccoloba uvifera, Sesuvium portulacastrum, Lippia nodiflora, Linaria elatine
Cedar forest (general) All hills not under cultivation are covered with Juniperus bermudiana and Sabal blackburniana. The undergrowth is heavily invaded but introduced trees only occasional Juniperus bermudiana, Sabal blackburniana, Lantana involucrata, Lantana camara, Nerium oleander, Lippia nodiflora, Solidago sempervirens, Sisyrinchium bermudianum, Adiantum bellum, Bryophyllum calycinum, Stenotaphrum americanum, Citharexylum quadrangulare, Buddleia americana, Crescentia cujete, Hura crepitans, Duranta plumieri, Ficus carica, Nicotiana glauca
Limestone sinks forest (general) The Walsingham Tract (and several places near Harrington Sound) have numerous depressions (sinks), and caves Trees and shrubs: Juniperus bermudiana, Sabal blackburniana, Celtis mississippiensis, Citharexylum quadrangulare, Calophyllum calaba, Citrus limonium, Ficus carica, Musa Cavendishii (planted), Melia azederach, Lantana camara, Lantana involucrata; herbs: Commelina agraria (=nudiflora), Peperomia obtusifolia, Bryophyllum calycinum, Stenotaphrum americanum, Asplenium trichomanes; lianes: Jasminum gracile, Rhus toxicodendron, Ipomoea purpurea, Cardiospermum halicacabum; [Lefroy, plants of limestone sinks: Ampelopsis (Parthenocissus) quinquefolia, Asplenium crenulatum, Callicarpa ferruginea, Asplenium myriophyllum, Dodonaea viscosa, Chiococca racemosa, Eugenia axillaris (=monticola), Elaeodendron xylocarpum, Caesalpinia (Guillandina) bonducella, Forestiera porulosa, Jasminum gracile, Ipomoea purpurea, Passiflora ciliata (=foetida), Jatropha curcas, Psilotum triquetrum, Peperomia obtusifolia, Pteris heterophylla, Psychotria undata, Trema (Sponia) Lamarckiana, Sicyos angulatus, Xanthoxylum clava-herculis, Triumfetta semitriloba]
Scrub (general) Typical of cleared and abandoned land, consisting almost entirely of exotic species Lantana camara, Lantana involucrata. Bryophyllum calycinum, Nerium oleander, etc.

Appendix 2

Plants mentioned by Harshberger (formations refer to the main habitat types referred to by Harshberger; “reported by Coulter” and “reported by Lefroy” indicate plants listed by Harshberger from other records). Currently accepted names are given in the first column. The second column notes the Harshberger name, if different (by synonymy or misidentification). Likely authorities are given for Harshberger names although none were given in the original paper. The final column lists collections by Harshberger, with herbarium locations of specimens given (in parentheses). This is not a complete list of his collections: specimens remain to be discovered, and species not in Harshberger (1905) are not listed here.

Currently accepted name Harshberger name, if different Affiliation Formation Harshberger specimens, all collected in 1905
Juniperus bermudiana L.   conifer sand dune (wind-swept form), cliff rock (wind-swept trees), cedar forest, scrub (relict), limestone sinks, brackish marsh (encroaching margins) June (MO)
Acrostichum danaeifolium Langsd. & Fisch. Acrostichum aureum L. fern brackish marsh (edge)  
Adiantum bellum T.Moore   fern cedar forest (rock crevices) 12 June, rock cliffs (MO, US); 16 June limestone sinks, Walsingham (MO)
Anopteris hexagona (L.) C.Chr. [Anopteris heterophylla auct., non (L.) Diels] Pteris heterophylla L. fern limestone sinks (reported by Lefroy)  
Asplenium heterochroum Kunze Asplenium trichomanes L. fern limestone sinks (sides of sinks) 13 June, Devil’s Hole (MO, US); 16 June, Walsingham (US)
Asplenium myriophyllum (Sw.) C.Presl   fern limestone sinks (reported by Lefroy)  
Diplazium laffanianum (Baker) C.Chr. Asplenium crenulatum [sensu Lefroy] fern limestone sinks (reported by Lefroy)  
Osmunda regalis L.   fern brackish marsh  
Osmundastrum cinnamomeum (L.) C.Presl Osmunda cinnamomea L. fern brackish marsh (frequent) 17 June, Smith’s and Devonshire marshes (MO, US)
Psilotum nudum (L.) P.Beauv. Psilotum triquetrum Sw. fern limestone sinks (reported by Lefroy)  
Pteridium caudatum (L.) Maxon Pteris (Pteridium) aquilina var. caudata (L.) Link fern brackish marsh (drier peaty soil) 17 June, Devonshire Marsh (US)
Avicennia germinans (L.) L. Avicennia nitida Jacq. flowering plant mangrove swamp (heads of bays, saltwater ponds), salt marsh (rare)  
Baccharis glomeruliflora Pers. Baccharis heterophylla Kunth flowering plant brackish marsh (in thickets, encroaching margins) June, Smith’s Parish Marsh (US)
Bacopa monnieri (L.) Wettst. Herpestis monnieri (L.) Kunth flowering plant brackish marsh (reported by Coulter) 14 June, North Shore roads (US)
Borrichia arborescens (L.) DC   flowering plant sand dune, cliff rock June, rocks along shore (US)
Bryophyllum pinnatum (Lam.) Oken Bryophyllum calycinum Salisb. flowering plant cedar forest (abundant in undergrowth, abandoned cellars), scrub, limestone sinks (bottoms of sinks)  
Buddleja madagascariensis Lam. Buddleja americana L. flowering plant cedar forest (occasional) 13 June, thickets near houses (MO)
Caesalpinia bonduc (L.) Roxb. Caesalpinia (Guillandina) bonducella (L.) Fleming flowering plant limestone sinks (reported by Lefroy)  
Cakile lanceolata (Willd.) O.E.Schulz Cakile aequalis L’Hér. ex DC. flowering plant sand dune (upper beach) 14 June, North Shore along roads (US); 13 June (US)
Callicarpa americana L. Callicarpa ferruginea Sw. flowering plant limestone sinks (reported by Lefroy)  
Calophyllum calaba L.   flowering plant limestone sinks  
Canavalia rosea (Sw.) DC. Canavalia obtusifolia (Lam.) DC. flowering plant sand dune 13 June, sand dunes, South Shore (MO, US)
Cardiospermum microcarpum Kunth Cardiospermum halicacabum L. flowering plant limestone sinks  
Cassine xylocarpa Vent. [Cassine laneana (A.H.Moore) J.Ingram if considered endemic sp.] Elaeodendron xylocarpum (Vent.) DC. flowering plant limestone sinks (reported by Lefroy) 16 June, limestone sinks, Walsingham (MO, US)
Celtis laevigata Willd. Celtis mississippiensis Bosc. flowering plant limestone sinks  
Centella asiatica (L.) Urb. Hydrocotyle asiatica L. flowering plant brackish marsh (reported by Coulter)  
Ceratophyllum demersum L.   flowering plant brackish marsh (reported by Lefroy)  
Chiococca alba (L.) Hitchc. [Chiococca bermudiana S.Br.] Chiococca racemosa L. flowering plant limestone sinks (reported by Lefroy) 15 June, limestone sinks, Walsingham (MO, US)
Citharexylum spinosum L. Citharexylum quadrangulare Jacq. flowering plant cedar forest (occasional), limestone sinks 16 June, limestone sinks, Walsingham (MO, US)
Citrus limon (L.) Osbeck Citrus limonium auct. flowering plant limestone sinks  
Cladium mariscus (L.) Pohl subsp. jamaicense (Crantz) Kük. Cladium occidentale (C. germanicum) Schrad. flowering plant brackish marsh (reported by Lefroy) June, Smith’s Parish Marsh (US)
Coccoloba uvifera (L.) L.   flowering plant cliff rock 14 June, Hotel Frascati [Flatts] (MO, US)
Commelina diffusa Burm.f. Commelina agraria Kunth flowering plant limestone sinks 4 June, Hotel Frascati [Flatts] (US)
Conocarpus erectus L.   flowering plant sand dune (incl. some thickets on dune crest), cliff rock (prostrate trees) 13 June, sand dunes South Shore (US)
Crescentia cujete L.   flowering plant cedar forest  
Croton punctatus Jacq. Croton maritimus Walter flowering plant sand dune (occasional clumps) 17 June, Tucker’s Town (US)
Cyperaceae spp.   flowering plant salt marsh  
Dodonaea viscosa Jacq.   flowering plant sand dune, limestone sinks (reported by Lefroy) 16 June, Walsingham (US)
Duranta erecta L. Duranta plumieri Jacq. flowering plant cedar forest 12 June, Hotel Frascati [Flatts] (MO, US)
Eclipta prostrata (L.) L. Eclipta erecta L. flowering plant brackish marsh (reported by Lefroy) 21 June, Pembroke Marsh (US)
Eugenia axillaris (Sw.) Willd. Eugenia monticola (Sw.) DC. flowering plant limestone sinks (reported by Lefroy)  
Euphorbia mesembryanthemifolia Jacq. Euphorbia buxifolia Lam. flowering plant sand dune, cliff rock  
Ficus carica L.   flowering plant cedar forest (abandoned cellars), limestone sinks 12 June, waste limestone (US)
Foeniculum vulgare Mill.   flowering plant sand dune  
Forestiera segregata (Jacq.) Krug & Urb. Forestiera porulosa
(Michx.) Poir.
flowering plant limestone sinks (reported by Lefroy)  
Heliotropium curassavicum L.   flowering plant mangrove swamp (with Avicennia in brackish swamps), salt marsh 12 June, salt marsh near Wistowe, Harrington Sound (MO, US)
Heliotropium gnaphaloides L. Tournefortia gnaphalodes (L.) R.Br. ex Roem. & Schult. flowering plant sand dune  
Hura crepitans L.   flowering plant cedar forest  
Hypericum hypericoides (L.) Crantz Ascyrum crux-andreae L. flowering plant brackish marsh (reported by Lefroy)  
Ipomoea indica (Burm.) Merr. Ipomoea purpurea (L.) Roth flowering plant limestone sinks (reported by Lefroy) 16 June, Walsingham (MO, US)
Ipomoea pes-caprae (L.) R.Br.   flowering plant sand dune 13 June, sand dunes (MO, US)
Jasminum simplicifolium G.Forst. subsp. australiense P.S.Green Jasminum gracile Andrews flowering plant limestone sinks (reported by Lefroy) 12 June, Hotel Frascati [Flatts] (US);13 June, limestone sinks, Walsingham (US)
Jatropha curcas L.   flowering plant limestone sinks (reported by Lefroy)  
Kickxia elatine (L.) Dumort. Linaria elatine (L.) Mill. flowering plant cliff rock  
Kosteletzkya virginica (L.) C.Presl ex A.Gray   flowering plant brackish marsh (reported by Lefroy)  
Kyllinga brevifolia Rottb. Kyllinga monocephala Thunb. flowering plant brackish marsh (reported by Lefroy)  
Lantana camara L.   flowering plant sand dune, cliff rock, cedar forest, scrub, limestone sinks 12 June, Flatt’s Village (US)
Lantana camara L. Lantana crocea Jacq. flowering plant sand dune  
Lantana involucrata L.   flowering plant cliff rock, scrub, limestone sinks, cedar forest 12 June, hills under junipers (US)
Lemna valdiviana Phil. Lemna minor L. flowering plant brackish marsh 21 June, Pembroke Marsh (MO)
Lepidium virginicum L.   flowering plant sand dune  
Limonium carolinianum (Walter) Britton Statice lefroyi Hemsl. flowering plant salt marsh (mentioned but apparently not seen by Harshberger)  
Melia azederach L.   flowering plant limestone sinks  
Mentha spicata L. Mentha viridis (L.) L. flowering plant brackish marsh (reported by Coulter)  
Musa acuminata Colla Musa cavendishii Lamb. flowering plant limestone sinks (planted)  
Nasturtium officinale R.Br.   flowering plant brackish marsh (in water channels - Lefroy)  
Nerium oleander L.   flowering plant sand dune, cedar forest, scrub 12 June, Flatts (MO, US)
Nicotiana glauca Graham   flowering plant cedar forest (abandoned cellars)  
Opuntia sp.   flowering plant sand dune  
Opuntia stricta (Haw.) Haw. [O. dillenii (Ker Gawl.) Haw., sensu Britton] Opuntia vulgaris Mill. flowering plant sand dune  
Parthenocissus quinquefolia (L.) Planch. Ampelopsis (Parthenocissus) quinquefolia (L.) Michx. flowering plant limestone sinks (reported by Lefroy)  
Passiflora foetida L. Passiflora ciliata Aiton flowering plant limestone sinks (reported by Lefroy)  
Passiflora suberosa L.   flowering plant sand dune 19 June, Paget dunes (MO, US)
Peperomia magnoliifolia (Jacq.) A. Dietr. [P. septentrionalis S.Br. if considered endemic sp.] Peperomia obtusifolia (L.) A.Dietr. flowering plant limestone sinks (rough rocks in bottoms and reported by Lefroy) 15 June, limestone sinks, Walsingham (US)
Phyla nodiflora (L.) Greene Lippia nodiflora (L.) Michx. flowering plant cliff rock, cedar forest 12 June (US)
Pluchea odorata (L.) Cass.   flowering plant brackish marsh (reported by Lefroy) 15 June, Shelly Bay (US)
Pluchea odorata (L.) Cass. Pluchea purpurascens (Sw.) DC. flowering plant mangrove swamp (with Avicennia in brackish swamps)  
Proserpinaca palustris L.   flowering plant brackish marsh  
Psychotria ligustrifolia (Northr.) Millsp. Psychotria undata Jacq. flowering plant limestone sinks (reported by Lefroy) 16 June 1906[5], Walsingham (US)
Rhizophora mangle L.   flowering plant mangrove swamp (heads of bays, saltwater ponds) 17 June, Walsingham (MO, US)
Rhynchospora colorata (L.) H.Pfeiff. Dichromena leucocephala Michx. flowering plant brackish marsh (reported by Coulter) 17 June, Smith’s Parish Marsh (MO, US)
Sabal bermudana L.H.Bailey Sabal blackburniana Glazebrook flowering plant brackish marsh (encroaching margins), cliff rock, cedar forest, limestone sinks 1905, North Shore (NY, US)
Sarcocornia perennis (Miller) A.J.Scott Salicornia fruticosa (L.) L. flowering plant mangrove swamp (with Avicennia in brackish swamps), salt marsh 12 June, small salt marsh at Wistowe, Harrington Sound (MO, US)
Scaevola plumieri (L.) Vahl   flowering plant sand dune  
Schoenoplectus tabernaemontani (C.C. Gmel.) Palla Scirpus validus Vahl. flowering plant brackish marsh (reported by Lefroy) 11 June, Smith’s Parish Marsh (US)
Schoenoplectus tabernaemontani (C.C.Gmel.) Palla Scirpus lacustris L. flowering plant brackish marsh  
Sesuvium portulacastrum (L.) L.   flowering plant mangrove swamp (with Avicennia in brackish swamps), salt marsh, cliff rock 12 June, small salt marsh at Wistowe, Harrington Sound (MO, US)
Sicyos angulatus L.   flowering plant limestone sinks (reported by Lefroy)  
Sisyrinchium bermudiana L. Sisyrinchium bermudianum auct. flowering plant sand dune, cedar forest 12 June (US)
Solidago sempervirens Michx.   flowering plant sand dune, cliff rock, cedar forest 13 June, Harrington Sound (NY, US)
Spiranthes brevilabrus Lindl.   flowering plant brackish marsh (reported by Lefroy)  
Stenotaphrum secundatum (Walter) Kuntze Stenotaphrum americanum Schrank flowering plant sand dune (incl. mats on the lee slopes), salt marsh, cliff rock (covering available soil), cedar forest (carpeting beneath cedars), limestone sinks (abundant) 12 June (MO, US)
Toxicodendron radicans (L.) Kuntze Rhus toxicodendron L. flowering plant limestone sinks (sides of sinks)  
Trema lamarckiana (Roem. & Schult.) Blume   flowering plant limestone sinks (reported by Lefroy)  
Triumfetta semitriloba Jacq.   flowering plant limestone sinks 16 June, Walsingham (MO, US)
Typha domingensis Pers. Typha angustifolia L. flowering plant brackish marsh 15 June, Devil’s Hole (MO)
Waltheria indica L. Waltheria americana L. (W. indica L.) flowering plant brackish marsh (reported by Lefroy)  
Yucca aloifolia L.   flowering plant sand dune, cliff rock 19 June, Paget Botanic Gardens (US)
Zanthoxylum flavum Vahl (Z. clavis-herculis sensu Lefroy) Xanthoxylum clava-herculis L. flowering plant limestone sinks (reported by Lefroy)  
Cladonia sp.   lichen brackish marsh (drier peaty soil)  
Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier   marine alga (brown) marine algal (saltwater ponds)  
Dictyopteris polypodioides (DC.) J.V. Lamour. Haliseris polypodioides (DC.) C.Agardh marine alga (brown) marine algal (surge algae)  
Dictyota bartayresiana J.V.Lamour.   marine alga (brown) marine algal (surge algae)  
Padina pavonica (L.) Thivy Padina pavonia (L.) J. V.Lamour. marine alga (brown) marine algal (surge algae, tidal pools, sandy bottoms)  
Sargassum natans (L.) Gaillon Sargassum bacciferum (Turner) C.Agardh marine alga (brown) marine algal (surge algae)  
Sargassum sp.   marine alga (brown) sand dune (washed ashore at high tide)  
Stypopodium zonale (J.V.Lamour.) Papenfuss Zonaria lobata C. Agardh marine alga (brown) marine algal (surge algae)  
Acetabularia crenulata J.V.Lamour.   marine alga (green) marine algal (tidal pools)  
Anadyomene stellata (Wulfen) C.Agardh Anadyomene flabellata J.V.Lamour. marine alga (green) marine algal (surge algae)  
Caulerpa chemnitzia (Esper) J.V.Lamour. Caulerpa racemosa (Forsskål) J.Agardh marine alga (green) marine algal (saltwater ponds)  
Caulerpa chemnitzia (Esper) J.V.Lamour. Caulerpa racemosa var. uvifera (C.Agardh) J. Agardh. marine alga (green) marine algal (tidal pools)  
Caulerpa cupressoides var. lycopodium Weber-van Bosse   marine alga (green) marine algal (sandy bottoms)  
Caulerpa mexicana Sond. ex Kütz. Caulerpa taxifolia (M. Vahl) C.Agardh marine alga (green) marine algal (sandy bottoms)  
Caulerpa mexicana Sond. ex Kütz.   marine alga (green) marine algal (tidal pools)  
Caulerpa sertularioides (S.G.Gmel.) M.Howe Caulerpa plumaris (Forsskål) C.Agardh marine alga (green) marine algal (saltwater ponds)  
Codium decorticatum (Woodw.) M.Howe Codium tomentosum Stackhouse marine alga (green) marine algal (surge algae)  
Halimeda incrassata (J.Ellis) J.V.Lamour. Halimeda tridens (J. Ellis & Solander) J.V. Lamour. marine alga (green) marine algal (surge algae, sandy bottoms)  
Halimeda tuna (J.Ellis & Solander) J.V.Lamour. Halimeda tuna marine alga (green) marine algal (surge algae)  
Neomeris annulata Dickie Neomeris dumetosa J. V.Lamour. marine alga (green) marine algal (surge algae)  
Penicillus capitatus Lamarck   marine alga (green) marine algal (sandy bottoms), mangrove swamp  
Ulva lactuca L.   marine alga (green) mangrove swamp (surge algae, tidal pools, with Avicennia in brackish swamps)  
Ulva rigida C.Agardh Ulva latissima auct. marine alga (green) marine algal (surge algae)  
Valonia macrophysa Kütz. Valonia utricularis (Roth) C.Agardh marine alga (green) marine algal (saltwater ponds)  
Digenea simplex (Wulfen) C.Agardh   marine alga (red) marine algal (tidal pools)  
Galaxaura rugosa (J. Ellis & Solander) J.V.Lamour. Galaxeura lapidescens (J. Ellis & Solander) J. V.Lamour. marine alga (red) marine algal (surge algae)  
Halymenia bermudensis Collins & M.Howe Rhodymenia palmata (L.) Greville marine alga (red) marine algal (surge algae)  
Leucobryum glaucum (L.) Schimp. Leucobryum sp. moss brackish marsh (drier peaty soil)  
Sphagnum spp.   moss brackish marsh (two species)  

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Wolsak, S., Wingate, D. & Cronk, Q. Environmental change in the terrestrial vegetation of Bermuda: Revisiting Harshberger (1905). Brittonia 70, 257–275 (2018). https://doi.org/10.1007/s12228-017-9509-x

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Keywords

  • Bermuda
  • Ecology
  • Historical ecology
  • Plant invasion
  • Conservation treadmill
  • Environmental change