Research Article |
Corresponding author: Warren L. Wagner ( wagnerw@si.edu ) Academic editor: Gian Pietro Giusso del Galdo
© 2024 Warren L. Wagner, Stephen G. Weller, Ann K. Sakai, Ben Nyberg, Kenneth R. Wood.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wagner WL, Weller SG, Sakai AK, Nyberg B, Wood KR (2024) Schiedea waiahuluensis (Caryophyllaceae), an enigmatic new species from Kaua'i, Hawaiian Islands and the first species discovered by a drone collection system. PhytoKeys 247: 111-121. https://doi.org/10.3897/phytokeys.247.130241
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During a survey by the National Tropical Botanical Garden drone team, an enigmatic Schiedea was observed in December 2021on steep, rocky cliff faces of the Waiahulu Valley in the Waimea Canyon of Kaua'i. Subsequently, another survey was conducted in March 2022 and, by use of a remotely controlled cutting device suspended below the drone, the first herbarium specimen was collected, as well as a seed collection of an undescribed cliff-dwelling species of Schiedea. Detailed study of the collections and plants grown at the University of California, Irvine greenhouse showed that it had enlarged, somewhat whitish sepals similar to those of cliff-dwelling S. attenuata (the sole species in sect. Leucocalyx), yet differed significantly from all other species in the genus. It also shares with S. attenuata a woody habit, hermaphroditic flowers, coloured nectar and styles 5 to 7 or 8. We describe it here as S. waiahuluensis given the only known localities are on the cliffs of this valley and place it in an enlarged sect. Leucocalyx. With the discovery of this new species, there are 36 species in this Hawaiian endemic genus.
Caryophyllaceae, conservation, drone exploration, Hawaiian Islands, Kaua'i, Schiedea
Drone technology is advancing quickly and has become an effective tool for botanical surveys of cliff environments (
While monitoring the walls of Waimea Canyon, Kaua'i for rare plant taxa and using a drone to photograph selected study sites, staff at the National Tropical Botanical Garden (NTBG) documented an enigmatic Schiedea in December 2021. On first examination of our images, we speculated that the unique shrub could possibly represent the extinct Kaua'i species, Schiedea amplexicaulis H.Mann, last observed in 1855, or perhaps a new species. Subsequently, we returned in March 2022 and used a remotely controlled cutting device suspended below the drone to make the first herbarium specimen and seed collections of the Schiedea. On close examination of our herbarium specimen, it became immediately clear that we had discovered a new Schiedea species, bringing the total number of this Hawaiian endemic genus to 36 species (
For the survey portion of this work, we deployed a DJI Phantom 4 pro quadcopter drone. This is a consumer-grade platform that was chosen for its portability and high-resolution image sensor (20 megapixels). Still photographic images were collected approximately 5 m from the cliff surface to allow adequate resolution for the location and identification of small plants.
Photographs taken by the drone were reviewed and classified by species in Adobe Lightroom software. Image classification is a manual process that relies on visual acuity and expert field knowledge. The high-resolution imagery is examined for species identification and phenology, to track plant distribution and abundance and also to guide field operations. GPS location information is embedded into each image file which assists in locating access points and re-finding specific individual plants.
Once the collection target was selected, we deployed an Outreach Robotics Mamba, suspended under a DJI Matrice 300 drone. The Mamba is a robotic sampling manipulator designed to grab, cut and collect plant material in vertical cliff habitats using propellers to advance the device towards an adjacent cliff in a swinging motion, while keeping the carrying drone clear of obstacles.
USA • Hawaiian Islands, Kaua'i: Cultivated at University of California, Irvine from seed collected at Waimea District, north-facing dry cliffs above Waiahulu, 767 m alt., 29 March 2022, B. Nyberg et al. BN023 (PTBG), 12 April 2024, S. G. Weller & A. K. Sakai 1172 (holotype: PTBG 1000097349!; isotypes: BISH!, G!, K!, MO!, UC1, US!).
Schiedea waiahuluensis A habit, stem with leaves and inflorescence B branch of inflorescence, showing open flower with 6 sepals C hair types from petiole D flower, lateral view showing anthers at male stage E flower with 6 sepals and showing rarely dissected nectar shafts, face view F flower with 5 sepals, single petal, face view G flower with 5 sepals, female stage H fruit cut open to reveal seeds I dry fruit J seed. Drawn from living material from cultivated individual at University of California, Irvine (UCI) greenhouse (A, C, F, G, H, J), from photographs of cultivated material at UCI (B, D, I) and photograph taken by drone (E). Illustration by Alice Tangerini.
Erect to spreading shrub to 40 cm long; stems terete, green, glabrous in young plants, but becoming conspicuously viscid, glandular pubescent throughout as plants age, ascending, sprawling, becoming pendent as they elongate in the wild, much branched, with side branches elongating to length of main axis and flowering at the same time as main axis. Leaves opposite, 4.5–7 (–11.9 in cultivation) cm long, 1.5–2.1 (–3.3 in cultivation) cm wide, thin, green, narrowly oblanceolate, mid-rib prominent and 2–5 conspicuous veins branching from the mid-vein near the base of the leaf, glandular pubescent on mature plants, margin entire, slightly thickened, apex acute, base gradually tapering, sessile. Inflorescence terminal, erect, with (4–)9–27(–45) flowers, pseudo-axillary or with a main axis 20.5–36.5 cm long; bracts green, densely glandular pubescent, the lowermost 13–29 mm long, 7–13 mm wide, those of the distal branches and flowers 5–12 mm long, 3–8 mm wide; pedicels spreading at anthesis, densely glandular pubescent (3–) 5–30 (–40) mm long at anthesis. Flowers hermaphroditic, strongly protandrous. Sepals (4–) 5–6, usually subequal in size, 4.5–11.9 mm in length and 2.5–5 mm in width, occasionally two narrower sepals positioned on opposite sides of the flower, green or whitish-green on adaxial face, occasionally with distinctive whitish margins, concave, ovate, orientated at ca. 90° angle to pedicel, adaxial side glabrous, abaxial side glandular pubescent, margins entire, scarious, apex apiculate. Nectary shaft tubular and notched at apex, rarely dissected in up to 3 parts, flap-like to tubular or partially tubular, 2.0–5.0 mm in length, recurved and appressed at the tip to the opposed sepal, small quantities of pale brown or rarely black nectar produced and, in the greenhouse, released on to subtending sepal as stigmas become receptive. Stamens 8–10(–12), 5.3–10 mm long; anthers ca. 0.9 mm long, yellow, dehiscing after flower opens, but before stigmas become receptive. Styles 5–8(–11), 3.6–10.0 mm in length when receptive, stigmatic papillae on distal half of style. Ovary 1.5–3.0 mm in width, 1.0–2.0 mm in height. Capsules 2.6– 3.2 mm long, subglobose, apparently tardily dehiscent. Seeds 13–82, viable following self-pollinations in greenhouse, black, 0.4–0.6 (–0.7) mm long, orbicular-reniform, compressed, the faces transversely rugose, the margins papillose. Chromosome number unknown.
Specific epithet refers to the Waiahulu cliff region of Waimea Canyon, Kaua'i, the only known location where the new species is found.
United States. Hawaiian Islands, Kaua'i: Waimea District, north-facing dry cliffs above Waiahulu, 767 m alt., 29 Mar 2022, B. Nyberg et al. BN023 (PTBG) • Ridge below the Waimea Canyon Lookout, Waiahulu region of Waimea Canyon Complex, Pu'u Ka Pele Forest Reserve, 809 m alt., 12 Sep 2023, Williams et al. AMW818–AMW825 (PTBG), Wood et al. 19397 (PTBG) • NTBG Horticultural Center, 12 Jul 2024, K.R. Wood et al. 19557 (PTBG).
Schiedea waiahuluensis is endemic to the Hawaiian Islands where it is narrowly restricted to the western side of Kaua'i and occurs in open dry to mesic cliff habitat (1000–1500 mm rain/year) above Waiahulu Stream, which occurs in a small tributary of the greater Waimea Canyon located in the Pu'u Ka Pele Forest Reserve (Fig.
Although the native vegetation in Waimea Canyon has been seriously degraded by goats (Capra hircus L.) since their introduction in the late 1790s, there are still rich patches of endemic plant species found in the more inaccessible sections of the forest reserve, especially the vertical dry cliffs. Relictual native shrubs and trees associated in the region where S. waiahuluensis occurs include Nototrichium sandwicense (A.Gray) Hillebr. (Amaranthaceae); Peucedanum sandwicense Hillebr. (Apiaceae); Artemisia australis Less., Bidens sandvicensis ssp. confusa Nagata & Ganders, Wollastonia fauriei (H.Lév.) Orchard, Wilkesia gymnoxiphium A.Gray (Asteraceae); Lobelia niihauensis H.St.John (Campanulaceae); Euphorbia celastroides var. hanapepensis Sherff (Euphorbiaceae); Argemone glauca (Nutt. ex Prain) Pope (Papaveraceae); Dodonaea viscosa Jacq. (Sapindaceae); and Neraudia melastomifolia Gaudich. (Urticaceae). Two native grasses commonly found in this habitat include Eragrostis variabilis (Gaudich.) Steud. and Panicum lineale H.St.John (Poaceae), along with the fern Doryopteris decora Brack. (Pteridaceae). Isodendrion pyrifolium A.Gray (Violaceae), a small federally endangered shrub previously unrecorded from Kaua'i, was also discovered growing with S. waiahuluensis (
Schiedea waiahuluensis A drone collected specimen Nyberg BN 023 with endemic Mirid on upper right bud B habit of plant in native habitat Williams AMW820, photo by Ben Nyberg C seed William AMW 821, photo by seedsofhawaii.org D flower of drone collected specimen, Nyberg BN 023, photo by KR Wood.
Although at least a portion of the sheer vertical cliff habitat of Schiedea waiahuluensis has apparently escaped degradation by feral goats, the former distribution of this species may have been more extensive prior to the introduction of goats. In addition to their immediate negative effects on native plant species, goats in this region aid in the dispersal of invasive non-native plant species, most notably Pluchea carolinensis (Jacq.) G.Don (Asteraceae); Hyptis pectinata (L.) Poit. (Lamiaceae); Festuca bromoides L. (Poaceae), Pentapogon micranthus (Cav.) P.M.Peterson, Romasch. & Soreng (Poaceae); Grevillea robusta A.Cunn. ex R.Br. (Proteaceae); and Lantana camara L. (Verbenaceae).
Critically Endangered - B1ab(i,ii,iii,v)
Schiedea waiahuluensis was assessed for endangerment using the IUCN criteria. Its limited geographic range in Extent of Occurrence (EOO, 2.5 km2) along with its single location and inferred decline of habitat quality led to its classification as Critically Endangered (CR). Further surveys of adjacent cliff habitats are necessary to better understand the distribution and abundance of S. waiahuluensis. Establishment and replication of ex-situ collections is underway; additional collections from unrepresented sub-populations would add to better understanding of the genetic diversity of this species.
The distinctive morphology of this remarkable new species places Schiedea waiahuluensis into sect. Leucocalyx W.L.Wagner & Weller, previously represented by a single cliff-dwelling species, S. attenuata W.L.Wagner, Weller & A.K.Sakai. Besides both being cliff-dwellers with a woody habit, they are also both hermaphroditic, with enlarged greenish-white sepals, coloured nectar and 5 to 7 or 8 styles. Schiedea attenuata occurs in Kalalau Valley, around 7 km to the north of S. waiahuluensis and is restricted to dry/mesic vertical cliffs. Morphological characters of these species and S. viscosa are compared in Table
Comparison of morphological characters of the subclade of species of Schiedea sect. Leucocalyx and S. viscosa (sect. Nothoschiedea).
Character | S. waiahuluensis | S. attenuata | S. viscosa |
---|---|---|---|
Stem shape | Terete | Compressed-terete | Terete |
Pubescence | Glabrous in young plant, becoming densely viscid, glandular pubescent | Glabrous to very sparsely puberulent with minute hairs | Moderately viscid glandular puberulent |
Leaf length and width (cm) | 4.5–7(–11.9) × 1.5–2.1 (–3.3) cm | 5.3–7(–12) × 0.5–0.7(–1.1) cm | 2.5–5 × 0.8–1.8 cm |
Sepal number and aspect | (4–)5–6, usually subequal in size, slightly concave | 5, equal, navicular towards apex | 4–5, flat |
Sepal length | 6–12 mm | 4–5 mm | 6.5–12 mm |
Nectary | Tubular and notched at apex, rarely dissected in up to 3 parts, flap-like to tubular or partially tubular | Tubular and notched at apex | Flap-like |
Nectar colour | Brown or occasionally black | Clear | Black |
Style number | 5–8(–11) | 5–6(–7) | 5–7(–8) |
Capsule length | 2.6–3.2 mm | 3.3–4.5 mm | 8–12 mm |
In contrast to the facultatively self-pollinating species in sect. Nothoschiedea, S. waiahuluensis appears to be outcrossing, based on strong protandry and production of ca. 16,000 pollen grains per flower, a value typical of outcrossing species (
Increasingly, drones are being used to assess and inventory cliff-dwelling plant taxa (
We are grateful to Alice Tangerini for her excellent illustrations of this new species, constructed from limited collections and photographs and for assembling the photo plates of the habitat and features of this unique species. This work was supported by National Science Foundation grants DEB-1753659, DEB-1753664, DEB-1750373 and DEB-1752785: Collaborative Research: Unlocking the evolutionary history of Schiedea (carnation family, Caryophyllaceae): Rapid radiation of an endemic plant genus in the Hawaiian Islands (PIs: A. Sakai, S. Weller, N. Wickett, M. Moore, W. Wagner) and the National Tropical Botanical Garden Drone programme. We thank Outreach Robotics for development and deployment of the Mamba tool that made the drone collections possible. We appreciate the review of the manuscript by Matt Knope. We thank Javier Rodarte for care of plants at the University of California, Irvine greenhouse. For their constant support of fieldwork on Kaua'i, we thank the State of Hawai – Division of Forestry and Wildlife and the Plant Extinction Prevention Program, especially staff that assisted with this study: Adam Williams, Scott Heintzman and Susan Deans.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was supported by National Science Foundation grants DEB-1753659, DEB-1753664, DEB- 1750373 and DEB-1752785: Collaborative Research: Unlocking the evolutionary history of Schiedea (carnation family, Caryophyllaceae): Rapid radiation of an endemic plant genus in the Hawaiian Islands (PIs: A. Sakai, S. Weller, N. Wickett, M. Moore, W. Wagner) and the National Tropical Botanical Garden Drone Program.
Warren L. Wagner / lead author and coordination of other authors’ contributions; Stephen G. Weller / co-author and oversaw greenhouse cultivation and gathering of measurements and descriptive data; Ann K. Sakai / co-author; Ben Nyberg / co-author, leader of drone team and field research; Kenneth R. Wood / co-author, drone team and field research member.
Warren L. Wagner https://orcid.org/0000-0001-5012-8422
Ben Nyberg https://orcid.org/0000-0002-5894-5088
Kenneth R. Wood https://orcid.org/0000-0001-6446-1154
All of the data that support the findings of this study are available in the main text.