$¢PhytoKeys PhytoKeys 252: 141-162 (2025) DOI: 10.3897/phytokeys.252.137624 Research Article Ovicula biradiata, a new genus of Compositae from Big Bend National Park in Trans-Pecos Texas Debra L. Manley’™®, Isaac H. Lichter Marck®, Keily Peralta?, Arturo Castro Castro?®, Kelsey A. Wogan’, Carolyn V. Whiting'’®, A. Michael Powell* 1 Science and Resource Management, Big Bend National Park, PO Box 129, Big Bend National Park, TX 79834-0129, USA 2 Department of Botany, Institute for Biodiversity Science and Sustainability, California Academy of Sciences, 55 Music Concourse Dr., San Francisco, CA 94118, USA 3 Centro Interdisciplinario de Investigacion para el Desarrollo Integral Regional, Sigma 119, Fracc. 20 de Noviembre II, Durango, Mexico 4 Department of Natural Sciences, Sul Ross State University, East Highway 90, Alpine, TX 79832, USA Corresponding author: Isaac H. Lichter Marck (ilichtermarck@gmail.com) OPEN Qaceess Academic editor: Oscar Vargas Received: 22 September 2024 Accepted: 20 December 2024 Published: 18 February 2025 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 Copyright: This is an open access article distributed under the terms of the CCO Public Domain Dedication. Abstract 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 Tetraneurinae, where itis 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. Resumen Se ilustra y describe un nuevo género monoespecifico 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, flo- rece en la primavera y se encuentra en aluviones calcareos gruesos. Con base en sus flores pistiladas con corola radiada, vilano de escamas aristadas y hialinas, follaje tomentoso y receptaculo ligeramente en forma de platillo a plano y epaleado, determinamos que la nue- va especie tiene afinidades con Helenieae subtribu Tetraneurinae, en la alianza Heliantheae. Analisis cladisticos moleculares de secuencias del ADN ribosomal nuclear (ITS) apoyan la posicion filogenética de O. biradiata dentro de la subtribu Tetraneurinae, donde resulta el lineage hermano del género Psilostrophe. También presentamos informacion detallada sobre el habitat, imagenes de alta resolucion usando un microscopio de diseccion y micro- grafias electronicas de barrido de caracteres vegetativos y reproductivos de O. biradiata y taxa relacionados, asi como una clave actualizada para los géneros de Tetraneurinae. Final- mente, discutimos la significancia del descubrimiento en relacion a la ciencia ciudadana, exploracion de biodiversidad, y la conservacion de plantas en el Desierto Chihuahuense. 141 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Key words: Asteraceae, biodiversity, calciphile, Chihuahuan Desert, Helenieae, taxono- my, Tetraneurinae Palabras clave: Asteraceae, biodiversidad, calcifilo, Desierto Chihuahuense, Helenieae, taxonomia, Tetraneurinae Introduction The Chihuahuan Desert is the largest and most biologically diverse warm des- ert in North America (Bell et al. 2014). Big Bend National Park is in southern Brewster County, Texas, bordered to the south by the Rio Grande. Its 801,165 acres (324,220 hectares) include some of the best representation of Chihua- huan Desert microhabitat diversity in the United States, including within the Chisos Mountain range (highest elevation ca. 7825 ft. (2385 m)) and numer- ous smaller peaks separated by low desert bajadas. Eighty-nine plant species of conservation concern are found in the park (Louie 1996; Poole et al. 2007, Texas Parks and Wildlife Department 2024). Many of these species of concern have limited distributions that extend into adjacent areas in Mexico or into Chi- huahuan Desert habitats to the immediate north and east of the park boundary (Powell and Worthington 2018). Previous floristic and wildflower studies centred in and near Big Bend Na- tional Park, include McDougall and Sperry (1951), Warnock (1970, 1974, 1977), Fenstermacher et al. (2008), Morey (2008, 2024), Hardy (2009), Weckesser and Terry (2014) and Powell and Worthington (2018). Wauer (1973, 1997) explored many off-the-trail areas searching for plants and animals of natural history in- terest. Potentially relevant floristic treatises of wider range include Correll and Johnston (1970), Henrickson et al. (1997), Turner et al. (2003), Turner (2013), Eason (2018) and Allred et al. (2020). Even though the park has been rather thoroughly botanised in the vicinity of most accessible areas, additional new plant discoveries are possible because of its extensive habitat diversity in stretches of remote terrain. On 2 March 2024, while traversing cross country in search of rare plant popula- tions, 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 cen- timetres across, prostrate and densely white-woolly, matching the whitish colour of their calcareous gravel substrate (Figs 1-9). Following a review, Park author- ities granted us permission to collect a few individuals for further study. From the limited material on hand and the photos we had taken, we were able to dis- cern characters suggestive of relationships to the tribe Helenieae Lindl. (sensu Baldwin et al. (2002)), especially Tetraneuris Greene. These characters included obconic fruits with five paleaceous, aristate scales and pistillate ray florets with maroon linear markings (Bierner and Turner 2006; Funk et al. 2009; Spellenberg and Zucker 2019). To test the hypothesised relationship to Helenieae, we carried out a more detailed study of inflorescence and fruit characters using scanning electron microscopy and sequenced nrDNA sequence data for one DNA gene region, the Internal Transcribed Spacer (ITS). Here, we present morphological, mi- cro-anatomical and molecular phylogenetic evidence that supports description of this plant as a new genus and species of subtribe Tetraneurinae Rydb. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 142 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Methods Field and herbarium collections 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 Scienc- es (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 rel- atives 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 1. Morphological study We examined morphological characters from field collections of the new spe- cies and exsiccate of putative close relatives using dissecting microscopy. Images of microscopic features were captured using a Leica M60 stereomi- croscope (Leica Camera, Wetztlar, Germany) outfitted with a digital camera. Table 1. Specimen voucher data and GenBank accession numbers for herbarium material used in molecular phylogenet- ic analyses and scanning electron microscopy. Taxon Psilostrophe bakeri Greene Baileya pauciradiata Harv. & A. Gray Tetraneuris acaulis (Greene) K.F. Parker var. arizonica Psilostrophe sparsiflora (A.Gray) Nelson Tetraneuris scaposa (DC.) Greene Baileya multiradiata harv. & A. Gray Amblyolepis setigera DC. Psilostrophe villosa Rydb. ex Britton Hymenoxys cooperi Cockerell var. cooperi Psilostrophe mexicana R.C. Br. Psilostrophe gnaphalodes DC. Ovicula biradiata Manley PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 : GenBank Accession Collector : Purpose Collector Date accession number number number DNA/SEM CAS 818281 A. Cronsquist 11645 13 June 1980 PQ144335 SEM CAS 288097 H.S. Gentry 25 Feb 1933 SEM CAS 731062 J. Henrickson 10576 A June 1973 DNA CAS 5935742 M. Butterwick 7526 19 May 1981 PQ144336 DNA/SEM CAS 700109 B. Turner 15128 5 June 1983 PQ144339 SEM CAS 713832 P. Munz 13688 3 May 1935 SEM CAS 765096 B. Ertter 5598 13 March 1985 DNA CAS 507562 P. Raven 19297 7 June 1964 PQ144334 SEM CAS 1005608 J. Henrickson 10521 4 June 1973 DNA CAS 720959 J.L. Villasefior 1591 23 September 1982 PQ144337 DNA CAS 701425 S. Sunderberg 1214 15 August 1981 PQ144338 DNA/SEM | SRSC 00058752 D. Manley 2 17 April 2024 PQ144333 143 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Table 2. Comparison of morphology amongst genera of subtribe Tetraneurinae. Character/taxon Life span Stems Leaves/blades Heads/ peduncles Involucres Phyllaries Ray florets Disc florets Cypselae Pappus Base chromosome number Hymenoxys Annual, biennial or perennial 5-120+ long; erect, often branched; glabrous or pilose Basal and cauline; simple or 1—2-pinnately lobed; glabrous or pilose Single or several; peduncles 0.4- 20+ cm long 2.5-30 mm wide 2-3-seriate; sparsely to moderately pilose (3-)8-13+; corollas yellow to orange, corollas yellow to orange, nerves colourless or greenish; ray floret corollas 0.7-12 mm wide Usually 25-50+; corollas yellow to brownish-yellow, 1.5-7.4 mm long; pubescent distally, trichomes to 0.2 mm long Obconic or obpyramidal, 1.4-4.7 mm long; glabrous or pilose None or of ZT) usually aristate, obovate to lanceolate, scales, 0.8- 4.3 mm long x=15 Tetraneuris Annual or perennial 5-50 cm long; erect, or plants, acaulescent; sparsely to densely pilose Basal or basal and cauline; linear to lanceolate; glabrous or pilose Single or several; peduncles 0.5- 40+ cm long 6-20 mm wide 3-seriate; sparsely to densely pilose None or 7-27; corollas yellow, nerves colourless, greenish, sometimes reddish-brown to maroon; ray floret corollas 2.5-6 mm wide 20-250+; corollas yellow, purplish distally, 1.6-3 mm long; pubescent mainly distally, trichomes to 0.1-0.2 mm long Obconic or obpyramidal, 1.5-4 mm long; moderately to densely tomentose Usually 4-8 aristate lanceolate, obovate, to oblanceolate, scales, 1-4.5 mm long x=15 Amblyolepis Annual Usually 12- 50 cm long; erect to decumbent; sparsely to moderately pilose Cauline; linear to spatulate; pilose Usually single; peduncles to 20 cm long 12-20 mm wide 2-seriate; sparsely to moderately pilose Usually 8-13; corollas yellow, nerves colourless or greenish, sometimes darker than background of laminae; ray floret corollas 4.5-10 mm wide 20--50; corollas yellow, 5-7 mm long; essentially glabrous distally Obconic, 3-4.5 mm long, prominently 10-ribbed; ribs densely tan- tomentose 5-6 ovate scales 2-3.5 mm long PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 Psilostrophe Biennial, perennial or shrubby 8-50+ cm long; spreading to erect; often densely woolly Basal and cauline; linear to spatulate; sparsely to densely woolly Single or in clusters; peduncles 0.5- 60+ cm long 2-7 mm wide 1-2-seriate; densely woolly 1-8; corollas yellow, nerves greenish, sometimes darker than background; ray floret corollas 3-20 mm wide 5-25+; corollas yellow to orange, 3.9-5.5.mmM long; pubescent distally, trichomes 0.1--0.2 mm long Cylindrical to clavate or obpyramidal, striate-ribbed, 2.5-4 mm long; glabrous, gland- dotted, or villous 4-8 oblong, elliptic or lanceolate, scales, 1.5-3.2 mm long Ovicula Ephemeral annual 1-4 cm long; prostrate; densely woolly Basal ovate, involute to nearly folded; densely woolly Single; sessile or peduncles 1 mm long To 4-6 mm wide 3-seriate; densely woolly 2 (-3); corollas whitish, nerves maroon; ray floret corollas 0.6-1 mm wide 10-12; corollas pale yellow, ca. 2--3 mm long; tomentose distally, trichomes 0.3-0.5 mm long Obconic- obpyramidal, faintly ribbed, 1.5-2 mm long; densely tomentose 5 aristate, oval scales, to 2 mm long Baileya Annual, biennial or perennial Usually 15- 75 cm long; mostly erect; woolly Basal and cauline; linear to ovate, often pinnately lobed; often densely woolly Single or several; peduncles 2-12 cm long 5-25 mm wide 2-seriate; moderately to densely woolly 5=70r 20-55; corollas yellow, nerves colourless to greenish; ray floret corollas 4-7 mm wide 10-20, or 40- 100+; corollas yellow, 2.5-4mm long; densely pubescent distally, trichomes to ca. 0.2 mm long Narrowly obpyramidal, 3-4 mm long, weakly ribbed or striate; glandular Usually absent, rarely scales 144 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Morphology was compared with representatives of all recognised genera of tribe Helenieae (Table 2). In addition, surface morphology of floral and vege- tative structures was analysed and imaged using a Hitachi SU3500 Scanning Electron Microscope (SEM; Hitachi, Tokyo, Japan) at the California Academy of Sciences. Initially, inflorescence and fruit structures were disassembled un- der a dissecting microscope and loaded on to an 18 mm pin-mounted SEM stub using double sticky tape. To enhance the electron conductivity of sam- ples, we then used a Cressington Sputter Coater 108 (Cressington, Watford, UK) at a vacuum pressure of 0.8 Pa to apply a 5 nm layer of gold-palladium to the sample for 50 seconds. We observed traits of potential phylogenetic informativeness following Robinson (1981) and King and Robinson (1970) at 15 kV and a working distance of 7 mm, under automated controls for focus, contrast and stigmation. DNA extraction, amplification and sequencing 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 extract- ed 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 H,0, 0.3 ul DnTPs, 0.15 ul Taq polymerase, 0.75 ul MgCl, 1.5 ul 10x PCR buffer and 0.6 ul Bovine serum Albumin (BVA) was com- bined with two primers for amplifying the Internal Transcribed Spacer region (ITS), ITS4 and LEU (White et al. 1990). Two ul of undiluted genomic DNA was combined with the PCR master mix and transferred to a thermal cycler programmed to the following conditions: 97 degrees for 1 min; 40 cycles of 97 degrees for 10 sec, 48 degrees for 30 sec, 72 degrees C for 20 seconds; and 72 degrees C for 7 minutes. Post-PCR products were checked for suc- cessful amplification using gel electrophoresis and unpurified PCR-product was forward and reverse Sanger sequenced by Genewiz (Azenta US Inc., Burlington, MA). 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 Baldwin et al. (2002) published data matrix for epaleate tribes of the Heliantheae alliance. Once we recovered strong evidence for the sister relationship of the new species with Psilostrophe in tribe Helenieae, we generated additional sequence data for all recognised minimum-rank taxa of Psilostrophe and Tetraneuris scaposa (DC.) Greene using leaf tissues sampled from herbarium specimens. For these additional samples, we followed an extraction and amplification procedure identical to that described above. Selection of a model of molecular substitu- tion and Maximum Likelihood (ML) inference of a phylogenetic tree, based on aligned data matrices of ITS, was inferred using IQTREE2 (Minh et al. 2020) and bootstrap support for nodes was calculated, based on 1000 iterations using fast-bootstrapping. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 145 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Results Phylogenetic relationships Preliminary searches of the NCBI nucleotide BLAST database showed a sig- nificant 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 phylo- genetic tree, based on the ITS alignment from Baldwin et al. (2002), resolved the new species with very high (98 bs) support as nested in tribe Helenieae, where it was more closely related to Psilostrophe than other members of subtribe Tetraneurinae (Fig. 10). The new species + Psilostrophe, in turn, form the sister lineage to the clade containing Amblyolepis setigera DC., Tetraneuris acaulis, Tetraneuris scaposa (DC.) Greene, Hymenoxys ambigens var. floribunda (A.Gray) W.L. Wagner, Hymenoxys hoopesii (A. Gray) Bierne, and Hymenoxys lemmonii Cockerell. Addition of DNA sequence data (ITS) for five previously unsampled taxa of Psilostrophe resolves all currently recognised minimum-rank taxa in this genus as a monophyletic group separate from and sister to the new species. Amongst taxa of Psilostrophe, the narrowly endemic P. bakeri is resolved as sister to the rest of the genus, with P. sparsiflora next to diverge, followed by P cooperi. Relationships amongst the highly-nested taxa P mexicana, P. gnaph- alodes, P. tagetina and P villosa were not well supported with ITS data alone. Micro-anatomy Micro-morphological features targeted using SEM for their value in evaluating phy- logenetic 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 2) of these characteristics for genera of Tetraneurinae is given along with plates of SEM images (Figs 11, 12). Micro-anatomical features of the new species revealed by SEM include the dentate margins and pleated structure of hyaline pap- pus scales, style branch apices with terminal papillae and short stipitate glands that are present on the abaxial surface of ray and disc corolla lobes. Two types of trichomes were observed. Cypselae trichomes appear stiff, linear and end in a bifurcate tip. These conform with the cypselar trichomes observed in many other Compositae, also called twin hairs (“Zwillingshaare”) by Hess (1938). Trichomes on leaf tissues have a dilatated base (foot) that is notably wider than the rest of the structure, which has an elongated, flagellate body, an apex that ends in a simple, unbranched tip and a flexible, convoluted, helical structure, presumably giving the plant its characteristic woolly appearance. These trichomes conform to the oblique septate flagellate trichome type identified by Ramayya (1962), which occur in many groups of Compositae and often render the plant surface tomentose. Pollen grains of the new species measure approximately 20 micrometres in diameter and are oblate spheroidal in shape, with evenly spaced, symmetrical echinate projections. SEM images of representatives of related genera revealed similarities in the paleaceous and finely pleated structure of the pappus (Figs 11H, 12A, E, I), pol- len (Figs 11D, 12D), short-stalked stipitate glands (Figs 11C, 12C), stiff cypse- lae trichomes with forked tips (Figs 11A, 12D), presence of the flexible helical PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 146 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) trichomes that give the new species its woolly appearance (Fig. 11B) and style branch apices with sweeping papillae (Figs 111, 12H). Some consistent differ- ences that were noted between the new species and its sister genus Psilostro- phe included the vestiture of disc and ray floret corolla lobes, which consist of papillae in Psilostrophe, whereas the new species possesses helical trichomes along ray throats and on abaxial surfaces of disc lobes (Figs 11F, 12F). Finer variability was evident in the size and shape of the apex in paleaceous pappus elements at shallower taxonomic scales (Figs 11H, 12A, E, |). Taxonomic treatment Ovicula biradiata Manley, gen. et. sp. nov. urn:lsid:ipni.org:names:77356807-1 Figs 1-8 Description. 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 di- rections, 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 x 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 x 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 spread- ing, those in outer 2 series 3-4 x 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 x 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, lam- inae 3-6 x 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 ap- pendage truncate, apex papillate. Cypselae of ray and disc florets similar, 1.5- 2mm 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 x 0.8-1 mm, ovate, hyaline, with an apical arista ca. 1 mm long; the scales spreading when dry (Figs 1-7). Chromosome number unknown. Similar to members of tribe Helenieae (sensu Baldwin et al. (2002)), espe- cially Tetraneuris, with its annual habit, radiate heads, phyllaries in 3 series, PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 147 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Figure 1. First photograph of Ovicula biradiata taken by Deb Manley on 2 March 2024. convex, epaleate receptacles, ray florets pistillate and fertile, strap-like 3-lobed ray floret laminae with prominent, coloured veins, cypselae obconical, faintly ribbed and pappus of hyaline aristate scales, disc florets perfect and fertile, corollas yellow, 5-lobed; differs from other Helenieae genera by its smaller size, shorter stems, tightly clustered small leaves, greater tomentum density and smaller, sessile heads with only 2(-3) ray florets. Type. USA * Texas: Brewster Co.; Big Bend National Park, low gravelly lime- stone exposure, eroded alluvial flats, NE of Dagger Mt.; elev. 800 m, 20 Apr 2024, Debra Manley 2, with C. Whiting, C. Hoyt, R Manning, and S. Menzies; holo- type: 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). PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 148 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Figure 2. Researchers examining individuals in habitat. A Patty Manning scanning the ground in appropriate habitat for individuals of Ovicula biradiata B NPS botanist Carolyn Whiting photographing O. biradiata (circled in red). Photos by Cathy Hoyt on 20 April 2024. Figure 3. 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). PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 149 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Etymology. The generic name from Latin Ovis “sheep” and -cula (diminu- tive 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 1, 4). A recommended common name for O. biradiata is “woolly devil”, in reference to the woolly indumentum, the proximity of populations to the locality known as Devil's Den and the tendency for the ray florets to resemble horns. Distribution and phenology. Ovicula biradiata is known from limestone ped- iments of eastern Big Bend National Park where only three small populations have been found. Within these subpopulations, individual plants were abun- dant, but short-lived, indicating an ephemeral life history. The species was discovered on 2 March 2024 when plants were in full flower (Fig. 1). It is not presently known how early the plants may produce flowers, but, in the same general area, there are other species in several families that may bloom in early February or even earlier. By late May, after a period of warm and dry weather, the delicate annual plants had ceased vegetative growth and only desiccated inflorescences could be found (Fig. 4). Habitat and associated taxa. The general area of the three known locations for the new taxon, as so far observed, consists of a broad floodplain composed Figure 4. Images of Ovicula biradiata individuals representing the “small” growth habit that occurs most frequently in all three known locations. Photographs by James Bailey in April 2024 (A), Kelsey Wogan on 27 April 2024 (B, E), Cathy Hoyt on 2 March 2024 (C), Dana Sloan on 27 April 2024 (D). PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 150 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Figure 5. Examples of moderately sized individuals found occasionally throughout the known locations. Photographs by Cathy Hoyt (A) & Deb Manley (B) on 2 March 2024 and James Bailey in April 2024 (C). Figure 6. Largest individuals of Ovicula biradiata encountered by researchers in known localities thus far. Photographs by Deb Manley (A, B) on 20 April 2024 & A. Michael Powell (C) on 27 April 2024. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 15] Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) SRSC INU 00058752 Recession _ BIBE-O1 702 Figure 7. Scan of the Holotype of Ovicula biradiata. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 N ase E 61799 Sul Ross State Univ. Herbarium (SRSC) Ovicula biradiata D. L. Manley Holotype A.M. Powell Nov 2024 PLANTS OF TRANS-PECOS TEXAS Brewster County Gen. et sp. nov. Asteraceae Big Bend Natl. Park, low gravelly to rocky limestone exposure, eroded alluvial flats, Chihuahuan Desert scrub, NE of Dagger ML.; 2610 ft. (795.53 m); associated plants, Agave lechuguilla, Larrea, Krameria, Plantago, Oenothera, Nerisyrenia camporum, Bouteloua, Grusonia aggeria, Aristida, Dasyochloa pulchella, Ariocarpus fissuratus. Plants annual, locally abundant on the limestone exposure; woolly, tiny, mostly inconspicuous, larger plants to ca. 5 em across, with 1-few short, prostrate stems, spreading; disc corollas pale yellow, ray floret ligules whitish with maroon nerves. Deb Manley 2 20 Apr 2024 with C. Whiting, C. Hoyt, P. Manning, and S. Menzies 152 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) ee Beg ea i Nee “ha | | | | Figure 8. Line drawing of Ovicula biradiata gen. et. sp. nov. A Ray floret without cypsela B disc floret without cypsela C cypsela D habit with close up of leaf surface illustrating nature of indumentum. Illustration by Ellen Ruggia, based on material from the paratype (Manley 3). of fine sand and clay sediments and braided with drainage. This alluvial ba- sin terrain is fringed with low, gravel-capped pediments which then extend into foothills and steeper slopes of a flanking limestone mountain range. The lo- cations 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 3, 4). 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. Richard- son, 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 horizon- thalonius Lem., Opuntia sp. (L.) Mill. and Grusonia aggeria (Ralston & Hilsenb.) E.F. Anderson. Cryptobiotic soil is present in the habitat as well. See Figs 2, 3 for habitat photos and Fig. 9 for a distribution map. Conservation. Ovicula biradiata is, so far, Known only from within a small area ina seldom accessed part of Big Bend National Park. Nevertheless, the extreme- ly narrow range and ephemerality of the species suggests that it is highly sensi- tive to variable weather patterns. Recently, this part of the Chihuahuan Desert has PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 153 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Ovicula biradiata Mexico 72, S ) aa EA qo Ranch State Park { ida Area Proteg| Big Bend § National Park Maderas del Carmen 7 Canon de Santa Elena Area Heat toko Ocampo Area Protegida Figure 9. 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. been under severe drought conditions and aridity is predicted to increase in this region due to climate change (Climate Change Response Program 2024). Under current IUCN guidelines for assessment of conservation status (IUCN Standards and Petition Committee 2022), O. biradiata would, therefore, preliminarily quali- fy as being vulnerable (VU) and under a high threat of extinction. More study is needed on the reproductive biology and population structure of O. biradiata, as well as potential threats to its habitat, to determine if the species should be listed by the U.S. Fish and Wildlife Service under the Federal Endangered Species Act. Due to the extreme sensitivity of the known collection sites the geocoordinates of the locality have been withheld and the locality is obscured on the map (Fig. 9). PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 154 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) 6 6r Psilostrophe mexicana Villasefor 159 66 Psilostrophe gnaphalodes Sunderberg 12 Psilostrophe tagetina Psilostrophe villosa raven 19297 Psilostrophe cooperi Psilostrophe sparsiflora Butterwick 7526 Psilostrophe bakeri Cronquist 11645 Tetraneurinae Ovicula biradiata Manley 2 Tetraneuris scaposa Turner 15128 Tetraneuris acaulis Amblyolepis setigera Hymenoxys ambigens Hymenoxys hoopesii Hymenoxys lemmonii Baileya multiradiata Baileya pleniradiata Baileya pauciflora Balduina uniflora . . Gaillardia pul. Gaillardinae Helenium bigelovii : : Plateilemapalmeri | Plateileminae Marshallia obo. Marshallia caes. oD) © we c xv ) - Pelucha trifida Psathyrodes annua Marshallinae Psathyrodes ramosissima Trichoptilium incisum 0.07 subst. Figure 10. Phylogenetic relationships of Ovicula biradiata and representative photographs of genera of Helenieae. A Max- imum 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 | Helenium amphibolum. Photographs by Peri Lee Pipkin (C) and James Bailey (B, D-I). Discussion 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 re- gions 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 (Stebbins 1952). Past examples of unique monospecific genera discovered in deserts or unique soils include Apacheria chiricahuensis C.T. Mason, Dedeckera eurekensis Reveal & J.T. Howell, Megacorax gracielanus S. Gonzalez & W. Wagner and Yermo xanthocephalus Dorn. Conservation man- agement of O. biradiata will depend on gathering more detailed observations of its habitat specialisation, population size, reproductive biology, geographic range and life cycle and these are data that should be gathered with urgency. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 155 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) As drought conditions continue to increase in frequency and severity, opportu- nities to observe annual plants, including O. biradiata, may occur less frequently. Evolutionary implications With the addition of Ovicula biradiata, subtribe Tetraneurinae contains six gen- era and 46 minimum rank taxa, making it the most diverse subtribe of Hele- nieae (Baldwin 2009). Extant diversity in this group is concentrated in west- ern North America where they are distributed across a broad range of habitats from high mountains to low deserts (Baldwin and Wessa 2000). An ephemeral, annual life history has evidently evolved multiple times in this group apart from O. biradiata, as in Baileya pauciradiata Harvey & A. Gray, Tetraneuris linearifolia Greene and Amblyolepis setigera DC., O. biradiata stands out amongst other members of Tetraneurinae, however, for its minute stature, sessile heads and densely woolly foliage that effectively camouflages the plant into a background of coarse calcareous gravel. A salient, visually conspicuous characteristic of O. biradiata is its ephemeral ray florets, which usually appear in pairs (Figs 1-6). i aN r ae rd 500m _Manleytypert Le 20k 06/13/2024 Figure 11. 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 stig- ma H pappus palea tip with fine pleated serrations I style branch apex, with papillate trichomes sweeping pollen grains. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 156 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Phylogenetic relationships 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 combi- nation 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. bira- diata 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 Figure 12. 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 | paleaceous pappus of T: scaposa with terminal, antrorsely setose bristle. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 157 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) 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 alter- native 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. Micro-anatomy Micro-anatomical observations enabled by SEM revealed several characteris- tic features noted as diagnostic for the tribe (e.g. as Gaillardiinae in Robinson (1981)). These include a style apex with sweeping papillae, stigma with two receptive lines and oblate spheroidal pollen with regular echinate spines. Some characters revealed by SEM images for O. biradiata, include the pleated, serrate structural anatomy of the aristate pappus scales and foliar trichomes with a flagellate, helical body, which make up the plants woolly-tomentose indumen- tum. Short-stipitate glands present on the ray and disc corollas of O. biradi- ata resemble those found in many Compositae, which are often associated with sesquiterpene lactone synthesis (Robinson 2009). Phytochemical stud- ies of Compositae, including members of tribe Helenieae, have yielded unique chemical compounds (e.g. Helenolins) with potential for anti-inflammatory and anti-cancer activity. The presence of short-stipitate glands in O. biradiata sug- gests this new species might contain secondary metabolites worthy of study for their potential medicinal value. Finally, to encourage further study of this fascinating group, we present an updated key to the genera of Tetraneurinae, including Ovicula, based on in- formation compiled from floras and observations of herbarium specimens in SRSC and CAS: Key to the genera of Tetraneurinae 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 broad- ly spatulate, semi-clasping, blades with long brownish trichomes, mostly OMACMESIALOIAS LR. # Beceem, 8S AO enh ers eae 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 no- table 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 ..............cccccccscccesseeeeessseeeees 2 2 Foliage glabrous or tomentellous (sparsely hairy), at the base often wool- ly, densely silky in H. subintegra; outer phyllaries usually partially connate; ray corollas ultimately withering and falling...................ccceee Hymenoxys PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 158 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) - Foliage densely woolly to tomentellous, sometimes glabrous; outer phyl- laries distinct; ray corollas usually persistent in fruit, except readily dis- [GCS CITT VIC Ciao ets 8 ORS ren eee aA aa nlts aa cainachan anit abana tecatees tins cadec 3 3. Ray florets 2(-3), corollas white with maroon nerves, readily dislodged from developing cypselae, laminae 0.6-1 mm wide; plants minute an- nuals, 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 pF: LONG 25S WAG e ec teeta. dos cect eee es eteree sae come dee | Ovicula - Ray florets 1-55, rarely 0, corollas yellow or orange, with yellow or ma- roon 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 sev- eral 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, BBO MMM WIGS. sh teed tels cil hs wos dears tonne tubadernctsne ode cz tiohe watts nehton aaentalivnes ccnates eae 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............c cc ccecccceeeesceeeceseseeees 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 ..............:ccceceeseeeees Tetraneuris - Ray florets usually 1-6; disc florets usually 5-17; plants woolly, not scapi- form; heads single (in P. cooperi) or in clusters with peduncles 0.5-2.5 cm OMIMIOIE, LONG sazcch wscves & Actes ck et vec. dopeesccel beetes eats trast or ReE a eneter ne Psilostrophe Acknowledgements The research in Big Bend National Park was carried out under Scientific Re- search and Collecting Permit BIBE-2024-SCI-0015 (Study: BIBE-00694). This study was carried out with the support of the Center for Comparative Genom- ics 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 manu- script, we thank David Keil, Mauricio Bonifacino de Leon, Oscar Vargas and one anonymous reviewer. The authors would also like to thank Cathy Hoyt (co-dis- coverer), 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. Additional information Conflict of interest The authors have declared that no competing interests exist. PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 159 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Ethical statement No ethical statement was reported. Funding No funding was reported. Author contributions 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. Supervi- sion: AMP. Visualization: KAW, KP. Writing — original draft: IHLM, AMP. Writing — review and editing: DLM, IHLM, AMP, ACC, KP KAW, CVW. Author ORCIDs Debra L. Manley © https://orcid.org/0009-0003-9345-41 66 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 Data availability All of the data that support the findings of this study are available in the main text. References Allred KW, Jercinovic EM, Ivey RD (2020) Flora Neomexicana III: An illustrated Identifica- tion Manual. Part 2: Dicotyledonous Plants. Allred and Jercinovic, 799 pp. Baldwin BG (2009) Heliantheae Alliance. In: Funk VA, Susanna A, Stuessy TF, Bayer RJ (Eds) Systematics, evolution, and biogeography of Compositae. International Associ- ation for Plant Taxonomy, Vienna. Baldwin BG, Wessa BL (2000) Phylogenetic placement of Pelucha and new subtribes in Helenieae sensu stricto (Compositae). Systematic Botany 25(3): 522-538. https:// doi.org/10.2307/2666694 Baldwin BG, Wessa BL, Panero JL (2002) Nuclear rDNA evidence for major lineages of helenioid Heliantheae (Compositae). Systematic Botany 27(1): 161-198. Bell GP, Yanoff S, Karges J, Montoya JA, Najera S, Arango AM, Sada AG (2014) Con- servation blueprint for the Chihuahuan Desert ecoregion. In: Hoyt CA, Karges J (Eds) Proceedings of the Sixth Symposium on the Natural Resources of the Chi- huahuan Desert Region, October 14-17. Chihuahuan Desert Research Institute, Fort Davis, 1-36. Bierner MW, Turner BL (2006) Tetraneuris. Flora of North America north of Mexico. Vol. 21. Oxford University Press, New York, 447-453. Climate Change Response Program (2024) Big Bend National Park climate futures sum- mary. National Park Service, Fort Collins. Correll DS, Johnston MC (1970) Manual of the vascular plants of Texas. Texas Research Foundation, Renner. Eason M (2018) Wildflowers of Texas. Timber Press, Inc., Portland. Fenstermacher J, Powell AM, Sirotnak J, Terry M (2008) Annotated vascular flora of the Dead Horse Mountains, Big Bend National Park, Texas, with notes on local vegetation PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 160 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) communities and regional floristic relationships. Journal of the Botanical Research Institute of Texas 2(1): 685-730. Funk VA, Susanna A, Stuessy TF, Bayer RJ [Eds] (2009) Systematics, evolution, and bio- geography of Compositae. International Association for Plant Taxonomy, Vienna. Hardy JE (2009) Flora and vegetation of the Solitario Dome: Brewster and Presidio Counties, Texas. Iron Mountain Press, Houston. Henrickson J, Johnston MC, et al. (1997) A flora of the Chihuahuan Desert Region. Edi- tion 1.2 as ms. Hess R (1938) Vergleichende Untersuchungen Uber die Zwillingshaare der Compositen. Botanische Jahrbucher fur Systematik. 68: 435-496. IUCN Standards and Petitions Committee (2022) Guidelines for Using the IUCN Red List Categories and Criteria. Version 15.1. Prepared by the Standards and Petitions Com- mittee. https://www.iucnredlist.org/documents/RedListGuidelines.pdf [accessed: 15 June 2024] King RM, Robinson H (1970) The new synantherology. Taxon 19(1): 6-11. https://doi. org/10.2307/1217907 Louie DA (1996) The Rare and Threatened Plant Species of Big Bend National Park, Tex- as. MS Thesis, Sul Ross State University, Alpine. McDougall WB, Sperry OE (1951) Plants of Big Bend National Park. U.S. Government Printing Office, Washington, DC. Minh BQ, Schmidt HA, Chernomor O, Schrempf D, Woodhams MD, Von Haeseler A, Lan- fear R (2020) IQ-TREE 2: New models and efficient methods for phylogenetic infer- ence in the genomic era. Molecular Biology and Evolution 37(5): 1530-1534. https:// doi.org/10.1093/molbev/msaa015 Morey R (2008) Little Big Bend: Common, uncommon, and rare plants of Big Bend Na- tional Park. Texas Tech University Press, Lubbock. Morey R (2024) The other side of nowhere; Exploring Big Bend Ranch State Park and its flora. Texas A&M University Press, College Station. Poole JM, Carr WR, Price DM, Singhurst JR (2007) Rare plants of Texas. Texas A&M University Press, College Station. Powell AM, Worthington RD (2018) Flowering Plants of Trans-Pecos Texas and adjacent areas. Sida, Bot. Misc. 49. Botanical Research Institute of Texas, Fort Worth. Ramayya N (1962) Studies on the trichomes of some Compositae |. General structure. Botanical Survey of India 4(1-4): 177-188. https://doi.org/10.20324/nelumbo/ v4/1962/76550 Robinson HE (1981) A revision of the tribal and subtribal limits of the Heliantheae (As- teraceae). Smithsonian Contributions to Botany 51: 1-102. https://doi.org/10.5479/ si.0081024X.51 Robinson H (2009) An introduction to micro-characters of Compositae. In: Funk VA, Susanna A, Stuessy TF, Bayer RJ (Eds) Systematics, evolution, and biogeography of Compositae. International Association for Plant Taxonomy, Vienna. Spellenberg R, Zucker N (2019) The sunflower family: A guide to the family Asteraceae of the contiguous United States. Sida, Bot. Misc. 52. Botanical Research Institute of Texas, Fort Worth. Stebbins Jr GL (1952) Aridity as a stimulus to plant evolution. American Naturalist 86(826): 33-44. https://doi.org/10.1086/281699 Texas Parks and Wildlife Department (2024) Diversity and habitat assessment species of greatest conservation need. https://tpwd.texas.gov/wildlife/wildlife-diversity/ swap/sgcn/ [accessed 11.1.2024] PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 161 Debra L. Manley et al.: Ovicula biradiata gen. et. sp. nov. (Compositae) Turner BL (2013) The comps of Mexico, vols. 16-18. Phytologia Memoirs. Texensis Publishing, Gruver. Turner BL, Nichols H, Denny G, Doron O (2003) Atlas of the vascular plants of Texas. Sida Botanical Miscellany, Number 24, Volume 1. Botanical Research of Texas, Fort Worth. Warnock BH (1970) Wildflowers of the Big Bend Country, Texas. Sul Ross State Univer- sity Press, Alpine. Warnock BH (1974) Wildflowers of the Guadalupe Mountains and the Sand Dune Coun- try, Texas. Sul Ross State University Press, Alpine. Warnock BH (1977) Wildflowers of the Davis Mountains and Marathon Basin, Texas. Sul Ross State University Press, Alpine. Wauer RH (1973) Naturalist’s Big Bend. Texas A&M University Press, College Station. Wauer RH (1997) For all seasons: A Big Bend journal. University of Texas Press, Austin. Weckesser W, Terry M (2014) An annotated flora of Reed Plateau and adjacent areas, Brewster County, Texas, USA. Journal of the Botanical Research Institute of Texas 8(1): 353-379. White TJ, Bruns T, Lee SUW, Taylor J (1990) Amplification and direct sequencing of fun- gal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White WJ (Eds) PCR protocols: a guide to methods and applications. Academic Press inc., San Diego, 315-322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1 PhytoKeys 252: 141-162 (2025), DOI: 10.3897/phytokeys.252.137624 162