﻿Taxonomic novelties in Haplopappus (Asteraceae, Astereae) from Chile

﻿Abstract Two new species of Haplopappus (Asteraceae) from central Chile are described in this article. Haplopappuscolliguayensissp. nov. is restricted to La Chapa hill, Colliguay, Valparaíso Region, and is most similar to H.undulatus but differs from the latter in its stem indumentum, leaf shape and margin, and synflorescence arrangement. Haplopappusteillierisp. nov. has been recorded from four high-Andean localities in the Choapa, Petorca, Rocín and Aconcagua river basins, and is most similar to H.punctatus but differs from the latter in its leaf length and margin, number of peduncles per twig, width of outer phyllaries, number of ray florets per capitulum, and achene dimensions. Additionally, we propose the reinstatement of H.kingii in agreement with an exhaustive revision of type material and protologues, as well as the study of herbarium specimens. Haplopappuskingii is restricted to mountainous areas in the southern portion of the Atacama Region, and resembles H.parvifolius and H.retinervius but differs from both by its leaf margin and apex, besides additional differences from each. We provide morphological descriptions, field images, distributional maps, conservation assessments, and taxonomic notes for the three species treated, as well as illustrations for the novel taxa.


Introduction
Haplopappus Cassini is a strictly endemic South American genus in the tribe Astereae Cassini (Nesom and Robinson 2007;Funk et al. 2009), composed of 67 specific and intraspecific taxa (Klingenberg 2007;García et al. 2018), distributed in southern South America.Most of the genus diversity is restricted to Chile with 65 taxa, of which 48 (74%) are endemic to this country (Rodríguez et al. 2018).
Recent floristic studies in the Valparaiso Region of Central Chile led to the discovery of two new Haplopappus species.In addition, we also propose the reinstatement of H. kingii (Phil.)Reiche, a name that was placed under the synonymy of H. remyanus Wedd.by Klingenberg (2007).
Herein, we describe H. colliguayensis and H. teillieri, two new species from Central Chile, and provide information to support the taxonomic status of H. kingii.In addition, we provide illustrations and/or photographs, distribution maps, conservation status assessments, and taxonomic notes for all the species treated here.

Methods
At first, recently collected material was checked against taxonomic keys and descriptions of Haplopappus, which were provided by Gay (1849), Philippi (1858), Reiche (1902), Hall (1928), Brown and Clark (1982) and Klingenberg (2007).Then, the specimens were compared to general and type material held at Chilean herbaria (CONC, EIF, SGO, ULS).Digital images of specimens available on the websites of the following herbaria were also examined: BAA, BM, CAS, E, F, GH, K, LIL, M, MSB, NY, P, PH, S, SI, US, WU.Botanical terminology of the descriptions follows Beentje (2012).Leaf widths were measured over the widest portion of the lamina not considering the teeth, which were described separately.Two capitula per sample were rehydrated in 70% ethanol for 24 hours and were subsequently dissected.The different parts of the capitula (e.g., phyllaries, florets, gynoecium) were mounted on a white cardboard and scanned; measurements of capitulum parts were obtained with the free software ImageJ (Schneider et al. 2012).Distribution maps and the estimation of the area of occupancy and the extent of occurrence (sensu IUCN 2022) were generated using the GIS software ArcGis version 10.4 (ESRI 2015).Diagnosis.Haplopappus colliguayensis is similar to H. undulatus Klingenb., but differs from the latter by its stems with capitate glands and multicellular flagelliform trichomes (vs.sessile glands), leaves oblong to lanceolate (vs.obovate to oblanceolate), leaf margins flat, entire to shortly dentate with up to 5 teeth per side (vs.margins undulate, conspicuously dentate to serrate with 5-9 teeth per side), and paniculiform synflorescences with two or more capitula (vs.solitary capitulum).
Distribution and habitat.Haplopappus colliguayensis has been recorded only in the La Chapa hill, Colliguay (~33.1°S;Fig. 1A), which is part of the coastal mountain range (cordillera de la Costa) between the Aconcagua and Maipo rivers.It inhabits rocky outcrops in south-to southwest-facing positions from the base of the hill (680 m a.s.l.) towards its summit (~1680 m a.s.l.).The surrounding zonal vegetation corresponds to sclerophyllous coastal forest; however, the vegetation associated with the rocky outcrops corresponds to a xerophilous scrub with predominance of Adesmia pirionii I.M.Johnst., Gochnatia foliolosa (D.Don) D.Don ex Hook.& Arn., Puya coerulea Lindl., and Chusquea cumingii Nees.
Etymology.The specific epithet refers to Colliguay, a locality situated to the south of the city of Quilpué in the Valparaíso Region of Chile.
Phenology.This species has been recorded flowering in February, but the period probably extends between January and March.Fruits have been recorded between February and April.Conservation status.According to IUCN (2022), Haplopappus colliguayensis can be considered as Critically Endangered (CR) according to criteria B2ab(iii, v), because its Area of Occupancy (AOO) is <10 km 2 (4 km 2 ).The criterion "a" was selected because it is known from a single locality.Although it is estimated that the only known population maintains around 1,000 individuals, all of them grow concentrated in a reduced area, which is susceptible to be affected by natural catastrophes or anthropogenic interventions (e.g., fires, droughts).The criterion "b(iii)" was selected because there is an inferred and projected decline in the extent and the quality of the habitat, given projected replacement of the vegetation due to climate change and the high probability of fire occurrence in the area where the species inhabits.According to MMA (2023b), the climate scenarios for 2050 predict an increase in temperature (of 1-2 °C) and a decrease in precipitation (~60-80 mm) in the county of Quilpué, where the species has been recorded.The predicted changes in precipitation allowed to classify this area under high risk of loss of flora (MMA 2023a).Moreover, the region of Valparaíso is one of the Chilean regions that has experienced one of the highest numbers of fires and its burned area was extensive between 1985-2018 (González et al. 2020).As a proof of this risk, a major fire that consumed 10 km 2 in December of 2022, affected a valley contiguous to Colliguay and reached only 2 km away from the location of H. colliguayensis.The persistence of this trend with the probability of such future events in La Chapa hill would generate a decrease in the number of mature individuals, affecting the persistence of the species (criterion "b(v)").
Distribution and habitat.Haplopappus teillieri inhabits the Mediterranean Andean low scrub of Chuquiraga oppositifolia D.Don and Nardophyllum lanatum (Meyen) Cabrera, between 1500-2600 m a.s.l.(Luebert and Pliscoff 2017).Only four populations have been recorded in the Andes, along the valleys of rivers Choapa, Petorca, Rocín and Aconcagua (Fig. 1B).It seems to be a very rare species but it can become locally dominant.
Etymology.The specific epithet teillieri honours the Chilean botanist Sebastián Teillier Arredondo (1956-), who has made significant contributions to the knowledge of the vascular flora of Chile.
Phenology.Haplopappus teillieri starts flowering in January, probably extending its bloom until early March.Fruits from February onwards.
Conservation status.Haplopappus teillieri was rarely collected since 1924, within a very restricted area in the Andes mountains of the Choapa and San Felipe de Aconcagua provinces (Fig. 1B), which suggests that it is a rare species.Here, we propose the species conservation status as Endangered (EN), considering the criteria B1+B2ab(iii).It has been assessed under the criterion "B1" as its Extent of Occurrence (EOO) is <5,000 km 2 (266 km 2 ), while the criterion "B2" corresponds to the Area of Occupancy (AOO) <500 km 2 (16 km 2 ).Criterion "a" is invoked given the species presence in less than five localities (4).Its populations are potentially threatened since all these mountainous areas are located within a zone of high interest for mining development and consequently is fully covered by mining petitions (SONAMI 2023).The criterion "b(iii)" corresponds to the inferred and projected decrease in the quality of the habitat due to the presence of bovine and caprine livestock and mining activities (e.g., opening of roads, prospecting, excavations, removal of soil and vegetation due to installation of facilities).The habitat will also be affected by climate change the consequences of which are a decline in precipitation (35-50 mm) and temperature increase (~2 °C) (MMA 2023a).In this sense, the species inhabits an area that is projected to suffer a moderate to high risk of loss of the flora because of precipitation decrease (MMA 2023a).
Distribution and habitat.This species is endemic to the Atacama Region in Chile (28°25'-29°05'S), mostly occurring in the middle portions of the Carrizal and Huasco river basins (Fig. 1C).It has been registered growing mostly on roadsides in mountainous areas, between 1500-3200 m a.s.l.There is a single record in the coast south of Huasco, which we consider may be an accidental occurrence.
Etymology.The specific epithet honours Thomas King, English citizen who collected several specimens in the Atacama Desert during the late 19 th century.
Phenology.Flowering from November to January and fruiting from January to March.
Conservation status.Haplopappus kingii is only known from few herbarium specimens and field photographs (P.Dandois, personal communication, 7 July 2023).In accordance with the IUCN (2022), the species is known from 12 localities (Fig. 1C), presenting an estimated Extent of Occurrence (EOO) of 7,087 km 2 and Area of Occupancy (AOO) of 56 km 2 .Although the estimations of EOO and AOO reach the values of threatened categories (Vulnerable and Endangered, respectively), there is not much information about the current state of the populations.The lack of these data does not allow us to classify the species under any threatened category.However, it is known that the species inhabits an area affected by the development of mining activities (SONAMI 2023) and the severe drought in Central Chile, which has produced a shortfall on normal precipitation of about 20-40% between 2010-2014 in the study area (CR2 2015).Moreover, the projections of climate change to 2050 estimate a decrease in precipitation (2-8 mm) and increase in temperature (1.8-2.5 °C) (MMA 2023b).Consequently, the area where the species is distributed will face a moderate to high risk of the loss of flora given the changes in precipitation (MMA 2023a).Considering all of the above, we inferred a change in the quality of the habitat of H. kingii but as the number of known localities exceeds thresholds for threatened categories, we propose the species conservation status as Near Threatened (NT).
Additional specimens examined.Chile.Taxonomic notes.Haplopappus kingii had been considered a distinct species in treatments of Haplopappus by Reiche (1902) andHall (1928).However, Klingenberg (2007) reduced H. kingii into the synonymy of H. remyanus in H. sect.Leiachaenium DC., a decision that was followed by the latest catalogue of the vascular flora of Chile (Rodríguez et al. 2018).Haplopappus kingii can be differentiated from the latter species by its hispid indumentum (vs.glabrous and glutinous), leaves evenly distributed throughout the stem up to the synflorescence (vs.leaves distinctly clustered towards the base of the plant, flowering branches sparsely foliate below the capitula), and outer series of phyllaries 1.7-2.5 mm wide (vs.2.5-3.0 mm wide).However, a close inspection of descriptions and herbarium specimens suggest that H. kingii better fits within Klingenberg's (2007) H. sect.Chromochaeta DC., where it most closely resembles H. parvifolius (DC.) A. Gray and H. retinervius (Kuntze) Klingenb.Haplopappus kingii differs from both species by its leaves with mostly flat margin (vs.margin undulate) and leaf apex acute (vs.obtuse to rounded).More specifically, H. kingii differs from H. parvifolius by its hispid indumentum (vs.glabrous plants), villous achenes (vs.glabrous achenes) and green leaves (vs.glaucous leaves), and from H. retinervius by its leaves oblong to oblanceolate (vs.broadly obovate to nearly orbicular) and more than 35 florets (vs.less than 30 florets) per capitulum (Klingenberg 2007).
In the citation of the type material, we recognize Thomas King as the collector of the sample, which differs from the name mentioned on the protologue, "Georgius King" (Philippi 1894: 615).The holotype at SGO (Fig. 7A) is accompanied by four pieces of paper, each of them with the following information: name of the species ("Pyrrhocoma kingii Ph."), the name of the collector and the year of collection ("George King 1885"), the locality ("Carrizal") and what we interpreted as the collection number ("No 62").On the other hand, the specimen at E (Fig. 7B) is attached to a piece of paper in which it is possible to read "No 62" and "Carrizal".Additionally, there is a printed label saying "Chili.Coll.: Mr. Thos.King.Presented July 1900.",that suggests the sample was sent to Edinburgh by Thomas King himself.The labels with the name of the locality and the collection number seem to have the same handwriting, which suggests they were written by the collector.Thomas (or Tomas) King is a well-known collaborator of R.A. Philippi, who sent him several samples of plants from the Carrizal valley in the Atacama Region and some of these samples were used to describe new species (Philippi 1892; e.g.Leucocoryne narcissoides Phil., Alstroemeria kingii Phil., Valeriana senecioides Phil.).Apart from the description of this species, the name Georgius King has not appeared in other publications.Therefore, we assume that Philippi made a mistake when writing the name of the collector on the label of the holotype specimen of Haplodiscus kingii at SGO and in the species protologue.

Figure 2 .
Figure 2. Haplopappus colliguayensis M.A.Villalobos, V.Morales & Nic.García A habit B capitulum C flowering branch, inset shows glandular pubescence on stem D style and asymmetrical stigmatic branches of true ray florets E true ray floret F disk floret G style and stigmatic branches of disk floret H achene I series of phyllaries J leaves ("bracts") subtending capitula K leaf.Drawn by Daniel Martinez Piña from N.García et al. 6783, 6785.

Figure 4 .
Figure 4. Haplopappus teillieri A.Cádiz-Véliz, V.Morales & Nic.García A habit B capitulum C immature capitulum D style and asymmetrical stigmatic branches of true ray floret E true ray floret F flowering branch G style and stigmatic branches of disk floret H disk floret I series of phyllaries J fascicle of leaves K leaf.Drawn by Daniel Martinez Piña from A. Cádiz-Véliz et al. 991.

Figure 5 .
Figure 5. Haplopappus teillieri A.Cádiz-Véliz, V.Morales & Nic.García A general view of the Rocín valley B habitat in Andean scrub-grassland of Chuquiraga oppositifolia, Festuca acanthophylla and H. teillieri C habit D detail of branches and leaves E flowering branches F immature capitulum G homogamous mature capitulum H, I heterogamous mature capitula.All photographs by Arón Cádiz-Véliz.

Figure 6 .
Figure 6.Haplopappus kingii (Phil.)Reiche A, B habit C flowering branch D, E detail of the abaxial and adaxial face of leaves, respectively, showing the characteristic hispid pubescence F branch showing several sessile capitula, fascicles of leaves and hispid stem.All photographs by Philippe Dandois.