Research Article |
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Corresponding author: Isaac H. Lichter Marck ( ilichtermarck@gmail.com ) Academic editor: Oscar Vargas
© 2025 Debra L. Manley, Isaac H. Lichter Marck, Keily Peralta, Arturo Castro Castro, Kelsey A. Wogan, Carolyn V. Whiting, A. Michael Powell.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Manley DL, Lichter Marck IH, Peralta K, Castro Castro A, Wogan KA, Whiting CV, Powell AM (2025) Ovicula biradiata, a new genus of Compositae from Big Bend National Park in Trans-Pecos Texas. PhytoKeys 252: 141-162. https://doi.org/10.3897/phytokeys.252.137624
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Here, we describe and illustrate a new monospecific genus of Compositae, Ovicula biradiata gen. et sp. nov., from the Chihuahuan Desert in Big Bend National Park, Texas. Ovicula biradiata is a very locally abundant, yet range-limited, spring annual herb found in coarse calcareous alluvium. Based on its pistillate ray florets, pappus of hyaline, aristate scales, tomentose foliage and slightly saucer-shaped to flat, epaleate receptacle, we determine that the new species has affinities with the Helenioid subtribe Tetraneurinae in the Heliantheae alliance. Molecular phylogenetic analysis of nrDNA (ITS) sequence data supports the phylogenetic position of Ovicula biradiata within the subtribe Tetranuerinae, where it is resolved as the sister lineage to the genus Psilostrophe. We also present detailed habitat information, high-resolution images captured using a dissecting microscope and scanning electron micrographs of vegetative and reproductive characters of Ovicula biradiata and related taxa, as well as provide an updated key to the genera of Tetraneurinae. Finally, we discuss the significance of this remarkable discovery for community science, biodiversity exploration and plant conservation in the Chihuahuan Desert.
Se ilustra y describe un nuevo género monoespecífico de Compositae, Ovicula biradiata gen. et sp. nov., del Desierto Chihuahuense en el Parque Nacional Big Bend, Texas. Ovicula biradiata es una hierba annual muy localmente abundante, pero con un rango limitado, florece en la primavera y se encuentra en aluviones calcáreos gruesos. Con base en sus flores pistiladas con corola radiada, vilano de escamas aristadas y hialinas, follaje tomentoso y receptáculo ligeramente en forma de platillo a plano y epaleado, determinamos que la nueva especie tiene afinidades con Helenieae subtribu Tetraneurinae, en la alianza Heliantheae. Análisis cladísticos moleculares de secuencias del ADN ribosomal nuclear (ITS) apoyan la posición filogenética de O. biradiata dentro de la subtribu Tetraneurinae, donde resulta el lineage hermano del género Psilostrophe. También presentamos información detallada sobre el hábitat, imágenes de alta resolución usando un microscopio de disección y micrografías electrónicas de barrido de caracteres vegetativos y reproductivos de O. biradiata y taxa relacionados, así como una clave actualizada para los géneros de Tetranuerinae. Finalmente, discutimos la significancia del descubrimiento en relación a la ciencia ciudadana, exploración de biodiversidad, y la conservación de plantas en el Desierto Chihuahuense.
Asteraceae, biodiversity, calciphile, Chihuahuan Desert, Helenieae, taxonomy, Tetraneurinae
Asteraceae, biodiversidad, calcifilo, Desierto Chihuahuense, Helenieae, taxonomía, Tetraneurinae
The Chihuahuan Desert is the largest and most biologically diverse warm desert in North America (
Previous floristic and wildflower studies centred in and near Big Bend National Park, include
On 2 March 2024, while traversing cross country in search of rare plant populations, the first author photographed an anomalous composite and posted images on iNaturalist. These diminutive plants, observed during the peak of their growing season, were inconspicuous annuals, from less than one centimetre to 3–7 centimetres across, prostrate and densely white-woolly, matching the whitish colour of their calcareous gravel substrate (Figs
After the National Park Service granted a research permit for this study (BIBE-2024-SCI-0015), plants were collected from two field locations, briefly dried and deposited in the A. Michael Powell (SRSC) and California Academy of Sciences (CAS) Herbaria for mounting and further study. To our knowledge, besides these collections, this previously unknown species has not been deposited in herbaria before. Dried vegetative and reproductive material of putative close relatives was obtained for detailed morphological study and DNA sequencing from herbarium specimens at SRSU and CAS. Sampling included representatives of genera in subtribe Tetraneurinae, i.e. Amblyolepis DC., Baileya Harv. & A. Gray ex Torr., Hymenoxys Cass., Psilostrophe DC. and Tetraneuris Greene. A complete list of specimens sampled and GenBank accession numbers is presented in Table
Specimen voucher data and GenBank accession numbers for herbarium material used in molecular phylogenetic analyses and scanning electron microscopy.
| Taxon | Purpose | Accession number | Collector | Collector number | Date | GenBank accession number |
|---|---|---|---|---|---|---|
| Psilostrophe bakeri Greene | DNA/SEM | CAS 818281 | A. Cronsquist | 11645 | 13 June 1980 | PQ144335 |
| Baileya pauciradiata Harv. & A. Gray | SEM | CAS 288097 | H.S. Gentry | 25 Feb 1933 | ||
| Tetraneuris acaulis var. arizonica | SEM | CAS 731062 | J. Henrickson | 10576 | 4 June 1973 | |
| Psilostrophe sparsiflora (A.Gray) Nelson | DNA | CAS 5935742 | M. Butterwick | 7526 | 19 May 1981 | PQ144336 |
| Tetraneuris scaposa (DC.) Greene | DNA/SEM | CAS 700109 | B. Turner | 15128 | 5 June 1983 | PQ144339 |
| Baileya multiradiata harv. & A. Gray | SEM | CAS 713832 | P. Munz | 13688 | 3 May 1935 | |
| Amblyolepis setigera DC. | SEM | CAS 765096 | B. Ertter | 5598 | 13 March 1985 | |
| Psilostrophe villosa Rydb. ex Britton | DNA | CAS 507562 | P. Raven | 19297 | 7 June 1964 | PQ144334 |
| Hymenoxys cooperi var. cooperi | SEM | CAS 1005608 | J. Henrickson | 10521 | 4 June 1973 | |
| Psilostrophe mexicana R.C. Br. | DNA | CAS 720959 | J.L. Villaseñor | 1591 | 23 September 1982 | PQ144337 |
| Psilostrophe gnaphalodes DC. | DNA | CAS 701425 | S. Sunderberg | 1214 | 15 August 1981 | PQ144338 |
| Ovicula biradiata Manley | DNA/SEM | SRSC 00058752 | D. Manley | 2 | 17 April 2024 | PQ144333 |
We examined morphological characters from field collections of the new species and exsiccate of putative close relatives using dissecting microscopy. Images of microscopic features were captured using a Leica M60 stereomicroscope (Leica Camera, Wetztlar, Germany) outfitted with a digital camera. Morphology was compared with representatives of all recognised genera of tribe Helenieae (Table
Following removal of the woolly indumentum under a dissecting microscope, fresh, field-collected leaves were dried for one week using silica gel and pulverised in a Qiagen tissue lyser (Qiagen, inc., Valencia, California) with a mixture of zircon beads and autoclaved sand. Genomic DNA was extracted using the DNEasy plant mini-kit (Qiagen, inc., Valencia, California) in the Center for Comparative Genomics at the California Academy of Sciences. We followed the provided protocol with a modified incubation in a cell-lysis buffer extended to 16 hours. A Polymerase Chain Reaction (PCR) master mix containing 9.1 ul H20, 0.3 ul DnTPs, 0.15 ul Taq polymerase, 0.75 ul MgCl2, 1.5 ul 10× PCR buffer and 0.6 ul Bovine serum Albumin (BVA) was combined with two primers for amplifying the Internal Transcribed Spacer region (ITS), ITS4 and LEU (
Following an initial search of the NCBI BLAST database to confirm a close match between our ITS sequence and putative closely-related taxa, we visually aligned the ITS sequence for the new species with the
Preliminary searches of the NCBI nucleotide BLAST database showed a significant match between ITS sequences of the new species and core members of subtribe Tetraneurinae, including Psilostrophe cooperi (A. Gray) Greene (88.69%), Baileya multiradiata Harv. & A. Gray (88.89%), Tetraneuris acaulis Greene (88.48%) and Hymenoxys lemmonii Cockerell (88.68%). The ML phylogenetic tree, based on the ITS alignment from
Micro-morphological features targeted using SEM for their value in evaluating phylogenetic relationships in the Heliantheae alliance included the surface texture of cypselae, pappus elements, trichomes, style trichomes, stigmatic surface, pollen shape and glands of vegetative and reproductive structures. A comparative table (Table
| Character/taxon | Hymenoxys | Tetraneuris | Amblyolepis | Psilostrophe | Ovicula | Baileya |
|---|---|---|---|---|---|---|
| Life span | Annual, biennial or perennial | Annual or perennial | Annual | Biennial, perennial or shrubby | Ephemeral annual | Annual, biennial or perennial |
| Stems | 5–120+ long; erect, often branched; glabrous or pilose | 5–50 cm long; erect, or plants, acaulescent; sparsely to densely pilose | Usually 12–50 cm long; erect to decumbent; sparsely to moderately pilose | 8–50+ cm long; spreading to erect; often densely woolly | 1–4 cm long; prostrate; densely woolly | Usually 15–75 cm long; mostly erect; woolly |
| Leaves/blades | Basal and cauline; simple or 1–2-pinnately lobed; glabrous or pilose | Basal or basal and cauline; linear to lanceolate; glabrous or pilose | Cauline; linear to spatulate; pilose | Basal and cauline; linear to spatulate; sparsely to densely woolly | Basal ovate, involute to nearly folded; densely woolly | Basal and cauline; linear to ovate, often pinnately lobed; often densely woolly |
| Heads/peduncles | Single or several; peduncles 0.4–20+ cm long | Single or several; peduncles 0.5–40+ cm long | Usually single; peduncles to 20 cm long | Single or in clusters; peduncles 0.5–60+ cm long | Single; sessile or peduncles 1 mm long | Single or several; peduncles 2–12 cm long |
| Involucres | 2.5–30 mm wide | 6–20 mm wide | 12–20 mm wide | 2–7 mm wide | To 4–6 mm wide | 5–25 mm wide |
| Phyllaries | 2–3-seriate; sparsely to moderately pilose | 3-seriate; sparsely to densely pilose | 2-seriate; sparsely to moderately pilose | 1–2-seriate; densely woolly | 3-seriate; densely woolly | 2-seriate; moderately to densely woolly |
| Ray florets | (3–)8–13+; corollas yellow to orange, corollas yellow to orange, nerves colourless or greenish; ray floret corollas 0.7–12 mm wide | None or 7–27; corollas yellow, nerves colourless, greenish, sometimes reddish-brown to maroon; ray floret corollas 2.5–6 mm wide | Usually 8–13; corollas yellow, nerves colourless or greenish, sometimes darker than background of laminae; ray floret corollas 4.5–10 mm wide | 1–8; corollas yellow, nerves greenish, sometimes darker than background; ray floret corollas 3–20 mm wide | 2 (-3); corollas whitish, nerves maroon; ray floret corollas 0.6–1 mm wide | 5–7 or 20–55; corollas yellow, nerves colourless to greenish; ray floret corollas 4–7 mm wide |
| Disc florets | Usually 25–50+; corollas yellow to brownish-yellow, 1.5–7.4 mm long; pubescent distally, trichomes to 0.2 mm long | 20–250+; corollas yellow, purplish distally, 1.6–3 mm long; pubescent mainly distally, trichomes to 0.1–0.2 mm long | 20–-50; corollas yellow, 5–7 mm long; essentially glabrous distally | 5–25+; corollas yellow to orange, 3.5–5.5 mm long; pubescent distally, trichomes 0.1–-0.2 mm long | 10–12; corollas pale yellow, ca. 2–-3 mm long; tomentose distally, trichomes 0.3–0.5 mm long | 10–20, or 40–100+; corollas yellow, 2.5–4 mm long; densely pubescent distally, trichomes to ca. 0.2 mm long |
| Cypselae | Obconic or obpyramidal, 1.4–4.7 mm long; glabrous or pilose | Obconic or obpyramidal, 1.5–4 mm long; moderately to densely tomentose | Obconic, 3–4.5 mm long, prominently 10-ribbed; ribs densely tan-tomentose | Cylindrical to clavate or obpyramidal, striate-ribbed, 2.5–4 mm long; glabrous, gland-dotted, or villous | Obconic-obpyramidal, faintly ribbed, 1.5–2 mm long; densely tomentose | Narrowly obpyramidal, 3–4 mm long, weakly ribbed or striate; glandular |
| Pappus | None or of 2–11(–15) usually aristate, obovate to lanceolate, scales, 0.8–4.3 mm long | Usually 4–8 aristate lanceolate, obovate, to oblanceolate, scales, 1–4.5 mm long | 5–6 ovate scales 2–3.5 mm long | 4–8 oblong, elliptic or lanceolate, scales, 1.5–3.2 mm long | 5 aristate, oval scales, to 2 mm long | Usually absent, rarely scales |
| Base chromosome number | x = 15 | x = 15 | x = 19 | x = 16 | x = ? | x = 16 |
SEM images of representatives of related genera revealed similarities in the paleaceous and finely pleated structure of the pappus (Figs
Annuals
, small, flowering plants usually 1–2(–3) cm tall, from less than 1 cm wide to 3–7 cm across, whole plants densely white-tomentose. Root single, thread-like, 0.5–1 mm wide at the plant base. Stems unbranched, erect or branches, if evident, lateral, prostrate, spreading 1–4 cm in one or more directions, internodes ca. 1 cm long. Leaves basal, mostly in tight clusters or at nodes on short stems, proximal leaves spreading, distal ascending, petioles 1–4 mm long, blades ovate, 4–7 × 2.5–5 mm, entire, planar, gently involute or nearly conduplicate. Heads heterogamous, borne singly, essentially sessile (peduncles to ca. 1 mm long), obscured by woolly leaves. Involucres 5–7 × 4–6 mm, broadly funnelform to campanulate or subglobose. Phyllaries in 3 series, ca. 1–2 in outer series, ca. 1–2 in second series, outer slightly spreading, those in outer 2 series 3–4 × 2–3 mm, ovate, inner series ca. 7, linear, ca. 2 mm wide, with scarious margins ca. 0.5 mm wide, densely white-tomentose. Receptacles ca. 1 mm across, slightly saucer-shaped to flat, sometimes with a very small conic enation from near centre, otherwise basically smooth or with faint floret scars, epaleate. Ray florets 2(–3) per head, 3–6 × 0.6–1 mm long, positioned on opposing sides, pistillate and fertile, strap-like; corolla tube 2–3 mm long, densely pilose distally, with wavy trichomes 0.3–1 mm long, laminae 3–6 × 0.6–1 mm, 3-lobed, whitish, markedly 4-nerved proximally, 6-nerved distally, nerves maroon, proximal portion of the abaxial ray laminae densely covered with sessile or short stipitate glandular trichomes. Disc florets 10–12 per head, perfect and fertile; corolla pale yellow, ca. 2–3 mm long, tube 0.6–0.9 mm long, throat 1.6–1.8 mm long, lobes 5, 0.1–0.3 mm long, distalmost throat and lobes densely pilose with wavy trichomes 0.3–0.5 mm long; anthers yellow, distal anther appendage narrowly obovate to subsagittate; style tip appendage truncate, apex papillate. Cypselae of ray and disc florets similar, 1.5–2 mm long, obconic-obpyramidal, slightly compressed or obscurely 4–5-angled (prismatic), ribs 4–5, densely pubescent with straight, ascending-appressed, silvery trichomes 0.5–0.9 mm long, minutely forked at tip, partially obscuring the bases of pappus scales. Pappus of ray and disc florets similar, scales 5, ca. 1–3 × 0.8–1 mm, ovate, hyaline, with an apical arista ca. 1 mm long; the scales spreading when dry (Figs
Similar to members of tribe Helenieae (sensu
USA • Texas: Brewster Co.; Big Bend National Park, low gravelly limestone exposure, eroded alluvial flats, NE of Dagger Mt.; elev. 800 m, 20 Apr 2024, Debra Manley 2, with C. Whiting, C. Hoyt, P. Manning, and S. Menzies; holotype: SRSC 00058752 (BIBE 61799); isotype: CAS 1352777 (BIBE 61820).
Paratypes. USA • Texas; Brewster Co.: Big Bend National Park, low gravelly limestone exposure, eroded alluvial flats, NE of Dagger Mt.; elev. 792.5 m, 20 Apr 2024, Debra Manley 3, with C. Whiting, C. Hoyt, P. Manning, and S. Menzies; BIBE 61800 (SRSC 00058751).
Known habitat of Ovicula biradiata. A, B Habitat with evident pediment slopes C slight habitat variation with iron-bearing rocks present in calcareous cobbles D overview of population locality with individual plants circled in red E close-up of individual plants in habitat illustrating cryptic appearance amongst calcareous surficial deposits. Photos by James Bailey (A) in April 2024 and Deb Manley on 20 April 2024 (B–E).
The generic name from Latin Ovis “sheep” and -cula (diminutive ending) references the dense woolly indumentum of this new plant. The name honours the desert bighorn sheep (Ovis canadensis nelsonii), an iconic, but threatened desert animal that is currently rebounding in this part of the Chihuahuan Desert, providing hope for other rare species like O. biradiata. The specific epithet biradiata references the typically two conspicuous ray florets, occasionally three per head, positioned on opposing margins of the capitulum (Figs
Ovicula biradiata is known from limestone pediments of eastern Big Bend National Park where only three small populations have been found. Within these subpopulations, individual plants were abundant, but short-lived, indicating an ephemeral life history. The species was discovered on 2 March 2024 when plants were in full flower (Fig.
The general area of the three known locations for the new taxon, as so far observed, consists of a broad floodplain composed of fine sand and clay sediments and braided with drainage. This alluvial basin terrain is fringed with low, gravel-capped pediments which then extend into foothills and steeper slopes of a flanking limestone mountain range. The locations are within 625 m of each other and occur where a shallow layer of mixed alluvial gravel and stones overlie bedrock of the Boquillas Formation. This composite substrate occurs on both the Ernst and San Vicente members of the formation and the observed habitat exposures consist of thinly-bedded limestone, carbonate shale and siltstone overlain by Quaternary gravel, which is a heterogeneous mix of surrounding geologic substrates. One site includes a significant presence of iron-bearing rocks. The known locations receive full sun throughout the day with very little shade provided by the sparse vegetation or the flat topography (Figs
Widely-distributed species noted in the habitat include Vachellia vernicosa (Britton & Rose) Seigler & Ebinger, Larrea tridentata (DC.) Coville, Tiquilia greggii (Torr. & A. Gray) A.T. Richardson, T. hispidissima (Torr. & A. Gray) A.T. Richardson, Agave lechuguilla Torr., Thymophylla acerosa (DC.) Strother, Plantago sp. L., Oenothera sp. L., Physaria sp. (Nutt.) A. Gray, Nerisyrenia camporum Greene, Krameria sp. Loefl., Bouteloua sp. Lag., Aristida sp. L., Dasyochloa pulchella (Kunth) Willd. ex Rydb., Ariocarpus fissuratus K. Schum., Echinocactus horizonthalonius Lem., Opuntia sp. (L.) Mill. and Grusonia aggeria (Ralston & Hilsenb.) E.F. Anderson. Cryptobiotic soil is present in the habitat as well. See Figs
Ovicula biradiata is, so far, known only from within a small area in a seldom accessed part of Big Bend National Park. Nevertheless, the extremely narrow range and ephemerality of the species suggests that it is highly sensitive to variable weather patterns. Recently, this part of the Chihuahuan Desert has been under severe drought conditions and aridity is predicted to increase in this region due to climate change (
Approximate range map of Ovicula biradiata. Geographical location of the known range of O. biradiata in Big Bend National Park in Brewster County, Texas. The boundaries of the park are illustrated in dark green. The purple dot marks the approximate area where three sub populations, each with abundant individuals of O. biradiata, were located. An inset illustrates the location of the study site within the Chihuahuan Desert.
Phylogenetic relationships of Ovicula biradiata and representative photographs of genera of Helenieae. A Maximum Likelihood (ML) phylogenetic tree of Helenieae, based on an aligned matrix of nrDNA sequence data from the Internal Transcribed Spacer region. ITS sequences generated as part of this study have collector numbers indicated to the right. Subtribes are indicated with text. Ovicula biradiata is in bold B Tetraneuris scaposa C Hymenoxys cooperi D Ovicula biradiata E Psilostrophe tagetina F Amblyolepis setigera G Baileya pleniradiata H Gaillardia pulchella I Helenium amphibolum. Photographs by Peri Lee Pipkin (C) and James Bailey (B, D–I).
The discovery of Ovicula biradiata underscores that the task of documenting and describing plant diversity is far from finished in the Chihuahuan Desert. Furthermore, that discoveries are not limited to unexplored or unpopulated regions and that interest and purposeful attention may still reveal novelties in places such as National Parks that might be considered “well-trodden” or fully understood. Encounters with novel plant species sufficiently different from their relatives to warrant description at generic rank are very uncommon in North America, but when they do occur, these often tend to be rare species associated with arid or edaphic micro-habitats where selection for unique growth forms is most pronounced (
With the addition of Ovicula biradiata, subtribe Tetraneurinae contains six genera and 46 minimum rank taxa, making it the most diverse subtribe of Helenieae (
Scanning electron micrographs (SEM) of Ovicula biradiata. A Cypsela trichomes appear stiff, linear and end in a bifurcate (forked) tip B trichomes on leaf surface with a flexible, helical structure C short-stalked capitate glands on abaxial surface of ray corolla D pollen E ray floret without cypsela F disc corolla apex G anther column and exserted stigma H pappus palea tip with fine pleated serrations I style branch apex, with papillate trichomes sweeping pollen grains.
Morphological features of Ovicula biradiata initially appeared to suggest a close link between the new genus and Tetraneuris, including maroon linear markings on the ray floret corollas (typically only visible on the abaxial face of the ray lamina in Tetraneuris), cypselae with a dense indument of fork-tipped trichomes and pappus of 4–6 hyaline, aristate scales. Some combination of these traits is present in other genera of subtribe Tetraneurinae, however, suggesting they may be shared ancestral characteristics. Molecular phylogenetic (ITS) data support a more distant relationship between O. biradiata and Tetraneuris than was expected from morphology and resolves the new genus as the sister lineage to the paper flowers (Psilostrophe). Ovicula biradiata bears resemblance to Psilostrophe in terms of its dense tomentose trichomes, leaves that are both basal and cauline and typically non-scapiform heads. ITS is a relatively easy-to-sequence DNA region that has been used for decades to resolve relationships at a variety of scales in Compositae, yet it represents only one line of genetic evidence. The possibility that conflicting relationships amongst genera of Tetraneurinae may be supported by alternative DNA regions or potentially reveal a role for other processes such as hybridisation in producing enigmatic evolutionary lineages like O. biradiata, are hypotheses that are worth exploring in future studies.
Scanning electron micrographs (SEM) of various genera of Tetraneurinae. A Pappus of Amblyolepis setigera B pollen of A. setigera C short-stalked capitate glands on disc corolla of Baileya pauciradiata D stiff, twin hairs on cypsela of Hymenoxys cooperi E hyaline, aristate palea-like pappus of Hymenoxys cooperi F vesicular trichomes on abaxial surface of disc corolla lobes in Psilostrophe bakeri G ridges on the surface of a cypsela in P. bakeri H sweeping papillate trichomes on style branch apices in Tetraneuris scaposa I paleaceous pappus of T. scaposa with terminal, antrorsely setose bristle.
Micro-anatomical observations enabled by SEM revealed several characteristic features noted as diagnostic for the tribe (e.g. as Gaillardiinae in
Finally, to encourage further study of this fascinating group, we present an updated key to the genera of Tetraneurinae, including Ovicula, based on information compiled from floras and observations of herbarium specimens in SRSC and CAS:
| 1 | Phyllaries in 2 series, outer herbaceous, inner very short, hyaline, scale-like; cypselae 10-ribbed, the ribs densely tan-pubescent; herbage notably pleasant-scented; annuals; proximal leaves usually oblanceolate to broadly spatulate, semi-clasping, blades with long brownish trichomes, mostly on the margins | Amblyolepis |
| – | Phyllaries in 1–3 series, all herbaceous, inner not short, sometimes with scarious margins; cypselae 2–5-ribbed or angled, often weakly so, faintly striate in some taxa, the ribs or angles gland-dotted, naked or pubescent with long or short trichomes, these whitish or silvery; herbage lacking notable scent; annuals, biennials or perennials; proximal leaves of various shapes, not clasping, trichomes if present not long and brownish, instead white or colourless, often densely tomentose | 2 |
| 2 | Foliage glabrous or tomentellous (sparsely hairy), at the base often woolly, densely silky in H. subintegra; outer phyllaries usually partially connate; ray corollas ultimately withering and falling | Hymenoxys |
| – | Foliage densely woolly to tomentellous, sometimes glabrous; outer phyllaries distinct; ray corollas usually persistent in fruit, except readily dislodged in Ovicula | 3 |
| 3 | Ray florets 2(–3), corollas white with maroon nerves, readily dislodged from developing cypselae, laminae 0.6–1 mm wide; plants minute annuals, usually 1–2(–3) cm tall, 1–7 cm wide, branches if present lateral, prostrate, whole plants densely woolly; leaves crowded basally, entire, 4–7 mm long, 2.5–5 mm wide | Ovicula |
| – | Ray florets 1–55, rarely 0, corollas yellow or orange, with yellow or maroon nerves, usually persistent in fruit, laminae 0.7–20 mm wide; plants annuals, biennials or perennials, 5–100 cm tall (except 2–40 cm in several species of Tetraneuris), stems ascending to erect, scapiform in most Tetraneuris, glabrous or tomentellous to densely wololy; leaves basal or cauline, densely woolly or not, entire to pinnately lobed, 20–120 mm long, 5–30 mm wide | 4 |
| 4 | Pappus absent; leaf blades woolly, mostly 3-lobed or pinnate | Baileya |
| – | Pappus of 4–8 hyaline scales; leaf blades tomentellous to woolly, margins mostly entire, sometimes toothed or lobed | 5 |
| 5 | Ray florets 7–27, except none in T. verdiensis; disc florets 20–200+; plants tomentellous to somewhat woolly, scapiform, except stems erect in the annual T. linearifolia; heads mostly single | Tetraneuris |
| – | Ray florets usually 1–6; disc florets usually 5–17; plants woolly, not scapiform; heads single (in P. cooperi) or in clusters with peduncles 0.5–2.5 cm or more long | Psilostrophe |
The research in Big Bend National Park was carried out under Scientific Research and Collecting Permit BIBE-2024-SCI-0015 (Study: BIBE-00694). This study was carried out with the support of the Center for Comparative Genomics and Scanning Electron Microscopy (SEM) Lab at the California Academy of Sciences. Keily Peralta was supported by the Summer Systematics Institute, a programme of NSF (DBI 2243994) and the Robert T. Wallace endowment for undergraduate education. Isaac Lichter Marck was supported by NSF DBI 2209393. The botanical illustration was funded by the Babe Turner Herbarium Endowment Fund at SRSC. For productive suggestions that improved the manuscript, we thank David Keil, Mauricio Bonifacino de Leon, Oscar Vargas and one anonymous reviewer. The authors would also like to thank Cathy Hoyt (co-discoverer), Shirley Powell, Dana Sloan, Ellen Ruggia, Patty Manning, Jimmy Duke, Stephen Menzies, Jim Henrickson, Jesse Kelsch, Shea Cadrin, James Bailey, Joey Santore, Ricardo Kreibel, Emily Magnaghi, Sarah Jacobs, Gary Williams, Athena Lam, Grace Kim, Tom Daniels, Bruce Baldwin and Sophia Winitsky. Key technical assistance was provided by Wren and Alice Marck.
The authors have declared that no competing interests exist.
No ethical statement was reported.
No funding was reported.
Conceptualization: AMP, KAW, DLM, IHLM. Data curation: KAW, AMP, DLM, ACC, KP, IHLM. Formal analysis: IHLM, KP. Funding acquisition: IHLM. Investigation: AMP, KAW, DLM, CVW. Methodology: IHLM. Project administration: CVW, AMP, DLM, KAW. Supervision: AMP. Visualization: KAW, KP. Writing – original draft: IHLM, AMP. Writing – review and editing: DLM, IHLM, AMP, ACC, KP, KAW, CVW.
Debra L. Manley https://orcid.org/0009-0003-9345-4166
Isaac H. Lichter Marck https://orcid.org/0000-0003-3575-6003
Arturo Castro Castro https://orcid.org/0000-0002-2864-5180
Carolyn V. Whiting https://orcid.org/0000-0002-7665-1577
All of the data that support the findings of this study are available in the main text.