New relevant chorological and conservation data on Carex (Cyperaceae) and Hypericum (Hypericaceae) from Ecuador

Abstract Background Knowledge of Carex L. (true sedges) and Hypericum L. (St. John's wort) in the Neotropics is fragmentary. New information As a result of a fieldwork campaign in Ecuador and revision of herbarium collections (K, QCA and QCNE), we present here relevant records of twelve Carex (Cyperaceae) and four Hypericum (Hypericaceae) species. Regarding Carex, we present the novel report for South America of C.aztecica, as well as the first Ecuadorian records for C.brehmeri, C.collumanthus, C.fecunda, C.melanocystis and C.punicola. The three later records have additional biogeographical significance, as they represent the new northern limit of these species. We also include observations for another five species included in the Ecuadorian Red List of Endemic Plants. As a result, the list of native Carex reported for Ecuador would now include 52 taxa. With regard to Hypericum, we include the new report of H.sprucei for the province of Bolívar, and the confirmation of the presence of three rare species (H.acostanum, H.matangense, H.prietoi) in their type localities, although with extremely low population sizes. We discuss their conservation status and implications.


Introduction
Carex L. (true sedges), is one of the three most diverse plant genera in the World, with more than 2000 species (Roalson et al. 2021).South America harbors about 200 species (Jiménez-Mejías et al. 2018), most of which are endemic to the continent.Some species are dominant in herbaceous habitats, such as bogs or páramos (high altitude Andean grasslands).However and despite its diversity and ecological importance, there is no comprehensive monographic systematic work on South American Carex.As a result, as new fieldwork and taxonomic research progresses, new findings, such as new species and chorological records, continue to be published (e.g.Jiménez-Mejías and Escudero (2016), Jiménez-Mejías and Roalson (2016), Poindexter et al. (2017), Jiménez-Mejías and Reznicek (2018), Jiménez-Mejías et al. (2020), Jiménez-Mejías et al. (2021a).Further work is therefore needed to achieve a complete understanding of the taxonomy of these plants in the Neotropics.Ecuador is a diversity hotspot for Neotropical Carex (46 spp., of which at least four are endemic to the country: only Argentina and Chile have more Carex diversity than Ecuador (Jiménez-Mejías et al. 2018).Ecuadorian Carex species are mostly distributed at high altitudes in páramo vegetation, with a few species reaching montane forests at lower altitudes.
With over 600 species, Hypericum L. is one of the 100 largest angiosperm genera (Carine and Christenhusz 2013), including economically important species in pharmacology (e.g.H. perforatum L., common St. John's wort) and numerous ornamentals.Overall, Hypericum has its main centre of species richness in the temperate regions of the Old World (Meseguer et al. 2013), but Hypericum species are distributed on almost all continents and radiated in the Neotropics.In South America, Hypericum is represented by 102 species (Nürk et al. 2013), most of them endemic and is a prominent component of the páramo shrub flora, but there are also herbaceous species in Brazil and in lowland regions of temperate South America (Robson 1977).Robson published the most complete taxonomic treatment of Hypericum to date (Robson 1977, Robson 1981, Robson 1985, Robson 1987, Robson 1990, Robson 1996, Robson 2001, Robson 2002, Robson 2006, Robson 2010a, Robson 2010b, Robson 2012).Since then, new species have being described regularly and chorological information has been updated.However, new fieldwork campaigns are needed to improve our understanding of the diversity and distribution of Hypericum.
Ecuador is home to 19 species of Hypericum, nine of which are endemic to the country (Robson 1987, Robson 1990).The Ecuadorian Hypericum species are mainly distributed at high altitudes in páramo vegetation, with few species entering open grassy areas at lower altitudes.
As a result of an international collaboration between the Universidad Autónoma de Madrid (UAM, Madrid, Spain), the Universidad Pablo de Olavide (UPO, Seville, Spain), the Real Jardín Botánico (RJB-CSIC, Madrid, Spain) and the Universidad Tecnológica Indoamérica (UTI, Quito, Ecuador), a fieldwork campaign focusing on Carex and Hypericum was carried out in the Ecuadorian Andes from Loja to Carchi provinces during the summer of 2022.In addition, the complete Carex and Hypericum collections of the two main national Ecuadorian herbaria, QCA and QCNE (acronyms according to Thiers (2022)) were studied in situ.The present study summarises the main chorological results of this work.

Materials and methods
The complete Carex and Hypericum collections of QCA (Herbario de la Pontificia Universidad Católica del Ecuador) and QCNE (Herbario Nacional del Ecuador (QCNE) del Instituto Nacional de Biodiversidad (INABIO), as well as the complete South American Carex collection of K (Royal Botanic Gardens, Kew) were studied in situ.Herbarium specimens from field collections were deposited at HUTI (Herbario de la Universidad Tecnológica Indoamérica), MA (Herbario del Real Jardín Botánico de Madrid, CSIC) and UPOS (Herbario de la Universidad Pablo de Olavide).
Carex specimens were determined using the specialised literature cited for taxon.Species are presented in alphabetical order according to the accepted names provided by Govaerts et al. (2007).The general distribution is also based on Govaerts et al. (2007) and supplemented by the relevant literature cited in each epigraph.Terminology of the inflorescence prophylls (utricles and cladoprophylls) follows suggestions in Jiménez-Mejías et al. (2016).Infrageneric placement follows Roalson et al. (2021) unless otherwise indicated.
Hypericum species are listed in alphabetical order according to the accepted names provided by Robson in Studies in the genus Hypericum ( Robson 1987, Robson 1990).Species distributions follow Robson (Robson 1987, Robson 1990).
Comments for species included in the Red List of the Endemic Plants of Ecuador (León Yánez et al. 2011) are given where relevant.When the Red List includes a name that is no longer in use, it is listed as a synonym under the accepted name.

Taxon discussion
Carex acutata is a member of the Hirta Clade (subg.Carex), a lineage with its highest diversity in the Holarctic, but with regional diversification in South America (P.Jiménez-Mejías, pers.obs.).It is distributed in montane and high elevation páramo habitats of the Tropical Andes, from Bolivia to Venezuela.
The name Carex tessellata was included in the Ecuadorian Red List as a "mysterious species known from a single collection of uncertain precedence".The name was synonymised by Jiménez-Mejías et al. (2020) to C. acutata after examining C. tessellata type (K-000584703).Carex acutata is a species widely distributed in South America and recorded from quite a few locations within Ecuador.Accordingly, the name should be removed from the Red List and the conservation status of C. acutata in Ecuador should be re-evaluated.

Taxon discussion
Carex aztecica was hitherto considered a Mesoamerican endemic species, known from southern Mexico and Guatemala.It belongs to the problematic Decora Clade (subg. Carex), where it was included as part of the variation of sect.Indicae Tuckerman sensu lato, a set of species with lax paniculate inflorescences and utriculiform cladoprophylls.
The Decora clade is one of the two groups of Carex that are exclusively distributed in tropical-subtropical areas, along with sect.Fecundae Kük.(see below).
The records that we present here are the first for Ecuador, but also for South America.
In addition, they constitute the new known southernmost limit of the species.The taxon can be distinguished from the closely related C. polystachya Wahlenb.by its darker female glumes (purple-brown to dark-brown in C. aztecica vs. orange, pale reddishbrown, stramineous or hyaline in C. polystachya) with blunter apex (at least some glumes obtuse or acute vs. all glumes acuminate to awned) and the utricles with the beak reddish-tinged (vs.not tinged) (Hermann (1974), Chater (1980).These localities in Ecuador become the new known southernmost limit of the species.

Taxon discussion
Carex collumanthus is endemic from the northern half of the cordillera, where it has been reported from Venezuela, Colombia, Peru and Bolivia (Wheeler 2002).It belongs to sect.Abditispicae G. A. Wheeler, a group of species with Andean-Patagonian distribution that is nested within the Flacca Clade (subg.Carex).
Here, we report the species for the first time in Ecuador, which was somehow expected since the plant is known from Colombia and Peru.

Taxon discussion
An Ecuadorian endemic that belongs to sect.Uncinia (Pers.)Baill.(subg.Uncinia (Pers.)Peterm.), a mainly Southern Hemisphere group disjunctly distributed between the Neotropic and the SW Pacific.Formerly recognized as its own genus (Uncinia Pers.; Global Carex Group ( 2015)), it is one of the few Carex groups with an unequivocal epizoochorus adaptation: the utricles bear a hooked appendix that allows them to attach to fur or feathers (Fig. 1 A,a).
Carex ecuadorensis was reported as threatened in the Ecuadorian Red List (as Uncinia ecuadorensis) under the category vulnerable (VU).Known from only two localities collected in the 80s, we confirm its persistence in at least one of these (the type location) and provide an exact location with coordinates.The reported population was found in a natural drainage channel of the pajonal.It was healthy, with about twenty individuals in good conditions.The only direct threat we noticed is the erosion of the slopes next to the road, which was more developed in other sectors.Probably, Carex ecuadorensis is growing in similar habitats in additional places of more difficult access within the same area.In any case, the location of this population in a National Park is an important safeguard for the future persistence of this species.Nomenclature =Uncinia tenuifolia G.A. Wheeler & Goetgh., Aliso 14: 144 (1995).

Taxon discussion
Another Ecuadorian endemic that belongs to sect.Uncinia (see C. ecuadorensis).
Carex goetghebeurii was reported as threatened (VU category) in the Ecuadorian Red List (as Uncinia tenuifolia).This was based on several collections of the same single population extending along a slope next to a road in Zamora-Chinchipe.We visited one of them ("13 km E of the pass, just before junction with old road", -3.9764109958906992 S, -79.10201083174987W) and failed to find it.The location has been transformed into a quarry.This unfortunate finding depicts that this plant is already negatively affected by human activities and reveals the need to confirm the persistence of this population and, if positive, to apply urgent conservation measures.

Taxon discussion
This species belongs to sect.Uncinia (see C. ecuadorensis), which remarkably includes three of the four Ecuadorian endemic Carex species (Fig. 1B,b,C,c).
Carex madida was reported as vulnerable (VU) in the Ecuadorian Red List (as Uncinia lacustris), based on the known existence of only three localities, all in protected areas (Cayambe-Coca, Cotacachi-Cayapas and Cotopaxi National Parks) in the provinces of Pichincha, Napo and Imbabura, respectively.Here, we report its presence in two additional provinces (Carchi and Chimborazo) and one addition to a protected area (Reserva Ecológica El Ángel).We confirm its persistence in two of the formerly known localities, where it seems abundant.Since the plants occur in Espeletia formations very close to the Colombian border, it is very likely that this species also occurs in the neighbouring country, from which it has not been reported yet.

Taxon discussion
Carex phalaroides is a Neotropical endemic widely distributed from northern Patagonia to Colombia and Venezuela, with an isolated occurrence known in Guatemala (Jiménez-Mejías et al. 2018).It belongs to sect.Junciformes Kük.(subg.Psyllophorae (Degl.)Peterm.), one of the few Carex groups with a remarkably relatively high species diversity in South America (Benítez-Benítez et al. 2021a).Carex phalaroides is an anomalous member of sect.Junciformes from a morphological point of view as its inflorescence is constituted by several spikes (instead of a solitary one in the remainder of the section) (Benítez-Benítez et al. 2021a, Morales-Alonso et al., in prep.).
Previously known from just a handful locations in Ecuador, our field campaign has resulted in the collection of several additional populations; thus, Carex phalaroides seems to be more widely distributed along Ecuadorian high Andean grasslands.However, to date, it has been collected just a few times (see comments under General considerations at the end of the manuscript), as depicted by its scarce representation in herbaria (just one additional specimen from Ecuador: MO-5869195, Imbabura, Cotacachi-Cayapas, after revising the full collections of K, NY, MO and US).

Taxon discussion
Carex punicola was so far known as a Central Andean element, reported from Peru, Bolivia and Argentina (Wheeler 1988, Poindexter et al. 2017).This species is the only Southern Hemisphere taxon from section Acrocystis Dumort., a mostly Holarctic group greatly diversified in North America.Carex punicola is a diminutive acaulescent sedge from dry soils (Fig. 1D,d).Interestingly, there is an unusually high number of acaulescent Carex species in South America, which suggests a possible evolutionary trend (Jiménez-Mejías et al. 2021a).
Here, we present the first records for Ecuador, which constitutes the new northernmost limit for the species.

Taxon discussion
A recently-described species endemic from the northern Andes in Ecuador and Peru (Jiménez-Mejías and Escudero 2016, Jiménez-Mejías et al., under review).As C. lepida (see comments above), it belongs to sect.Wheelerianae.
In Ecuador, it was only known from the type locality also in Azuay Province, at Sevilla de Oro Municipality (Jiménez-Mejías and Escudero 2016), on the Eastern Cordillera.This constitutes the second record of the species in Ecuador and the first in the

Taxon discussion
Carex sodiroi is a poorly known Ecuadorian endemic species.It belongs to the problematic Decora Clade (subg.Carex) as part of sect.Indicae sensu lato (see comments under C. aztecica) (Fig. 1E,e) .
Carex sodiroi was reported in the Ecuadorian Red List as critically endangered (CR) because of the lack of recent collections.However, this is probably not due to a true rarity of the species, but to the poor taxonomic circumscription of C. sodiroi.Section Indicae in Tropical America has an obscure taxonomy.A number of taxa have been described, based on little material and within a local scale (e.g. C. culmenicola Steyerm.Steyermark (1951)), C. regnelliana Boeck.( Boeckeler (1888)).The whole variation of the species in the field remains understudied and, thus, the taxonomic limits are blurred by specimens that does not fit the characteristics and measurements that are presented in the available descriptions.Carex sodiroi is clearly affected by this lack of understanding.The only description associated with the name is Kükenthal (1904), which is in turn based just on the original collection.To date, no additional study has accounted for this taxon. In

Taxon discussion
The last of the sect.Uncinia species in our account (see C. ecuadorensis).
Carex subsacculata was reported in the Ecuadorian Red List (as Uncinia subsacculata) as vulnerable (VU), despite being known from a single locality, it was located in a protected area (Yanacocha Biological Reserve) in the Province of Pichincha.We confirm the persistence and healthy conservation status of this population despite the recent eruptions of the nearby Pichincha Volcano (last eruption in 2002).Carex subsacculata forms an abundant population in Polylepis forest understorey, at the very top of the Andean Snipe track in the Reserve (Fig. 1 F1,F2,f) .

Taxon discussion
Hypericum acostanum (Fig. 2A,a) is an endemic species of southern Ecuador.It belongs to the New World section Brathys (Mutis ex L.f.) Choisy, which has radiated in the Páramos (Nürk et al. 2013).
This species is listed as vulnerable (VU) in the Ecuadorian Red List.It is known from only two localities, collected in the 50s and 80s in the province of Loja, on the road between San Pedro and Portovelo and on the road between Celica-Guachanamá, respectively.We confirm its persistence in the second locality and provide an exact location with coordinates.In addition, after studying the QCNE herbarium material, we have re-assigned a specimen (voucher nº 5612) from the rovince of Loja to this species.Therefore, we have added a new locality to the distribution of H. acostanum in this rovince.
This species is listed as vulnerable (VU) in the Ecuadorian Red List.It is known onlyfrom the type specimen collected in 1980 in the province of Morona-Santiago.The size and status of this population, and its specific threats, are unknown.Our observation is the second known collection of this species, with coordinates from the only specimen found.
This species is listed as endangered (EN) in the Ecuadorian Red List.Threats associated with its conservation status include overgrazing and burning.Hypericum prietoi has only been collected once at the type locality, in 1945.Our observation is the second known collection of the species.We provide the coordinates of the only specimen found.

Taxon discussion
Hypericum sprucei (Fig. 2D,d) has been reported from a handful of provinces in Ecuador.It belongs to section Brathys (Mutis ex L.f.) Choisy.
Here, we present an additional record for a new province (Bolívar).This species is not included in the Ecuadorian Red List.

General considerations about Ecuadorian Carex
Our work reveals a number of common problems in the knowledge of Carex in South America in general and in Ecuador in particular.Although Ecuador is one of the countries in tropical America with the most developed description of its flora, it is not free from the lack of taxonomic understanding of such a difficult group of plants as sedges.
On the one hand, despite the abundant collections in the Páramos promoted by the Ecuadorian government through research projects and vegetation inventories, the relative amount of sedges in the two main national herbaria (QCA, QCNE) is still low.Our field studies have shown that Carex can be among the most abundant plants in páramos, which highlights their ecological importance.However, only a few of the largest species from sect.Fecundae (i.e. C. jamesonii Boott, C. lemanniana Boott and C. pichinchensis Kunth) and Uncinia (C.phleoides Cav.) have been abundantly collected and are available in the QCA and QCNE collections.In particular, we found a large number of populations of C. phalaroides scattered throughout the cordillera, but only a handful of specimens in the herbaria we visited.Another example, and perhaps the most surprising for us, is the discovery of C. punicola for the first time in Ecuador.This species has been completely overlooked to date despite the fact that it can be a dominant part of the vegetation.In fact, in the population collected in the province of Pichincha, the plant was an abundant element of the sparse vegetation, extending over the lower part of the Ovejería puna and even occurring on such accessible places as the road shoulders of one of the main roads in Cotopaxi National Park.This neglect of Carex diversity is probably due to the combined effect of the small size of some of the species involved and the low interest of most naturalists in graminoids.
On the other hand, some species, such as C. collumanthus or C. via-incaica, were relatively well-represented in the herbaria studied despite their small size.This suggests that such species were probably abundant enough in a number of locations to have been overlooked during field collections.However, most of these specimens were missidentified or simply stored as indeterminate.This illustrates the problem of the lack of a comprehensive treatment for Carex in South America in general, and in Ecuador in particular.The lack of an accurate taxonomic treatment is probably what has prevented the formal description of C. via-incaica by other botanists before 2016, as there were already a number of vouchers in QCA or QCNE.
In conclussion, a greater effort in terms of taxonomic and systematic resources is needed before an accurate knowledge of the genus Carex can be achieved.As a result from this work, future changes in the number of native and endemic Carex species reported for Ecuador can be expected as progress is made towards a complete national checklist.This is of paramount importance since species of this genus, as already mentioned, can be abundant, or even dominant, elements of the páramo flora, and thus play a critical role in the ecological functioning of the habitats in which they thrive.

General considerations about Ecuadorian Hypericum
Two species of Hypericum in Ecuador, H. matangense and H. prietoi, were described in 1990 and 1987, respectively and are only known from their type locality.No other records or additional collections are known to us since then.Our expedition seems to be the first discoverty of these species again.During our field campaign and despite our efforts to find other populations, we only found these two species again in their type localities.Apparently, both species have extremely small populations: we found only one specimen of each species.The distribution of these species should be carefully assessed.Similarly, H. acostanum has only been found in three localities since 1987 and we found it in low abundance in one of them.
Hypericum matangense and H. acostanum were listed as VU in the Ecuadorian Red List, while H. prietoi was listed as EN.Our observations suggest that these species may be critically endangered.If this is confirmed, the Red List status of H. matangense and H. prietoi will need to be updated and urgent conservation plans including in situ and ex situ measurements, will need to be implemented.

fecunda Steud., Syn. Pl. Glumac. 2: 194 (1855) Material
Wheeler (1998)chya is endemic from the Tropical Andes from Colombia to Bolivia.As C. brehmeri, it belongs to sect.Phacocystis Dumort.(seeabove).Carex enneastachya was reported only from a handful locations in Cajas National Park at Azuay Province.Here we present additional records for another three provinces.Originally conceived as a narrowly distributed taxon, this species was listed as EN in the Ecuadorian Red List of Endemic plants under the name C. azuayae.However, C. azuayae was already synonymized byWheeler (1998)to C. enneastachya, which effectively expands its range to other South American countries.As a consequence, this plant should not be listed as an Ecuadorian endemic and its threatened status needs revision.and stems when crushed that persisted after drying.This aromatic property is known from other sedge groups (e.g.Cyperus odoratus L., C. sesquiflorus (Torr.)Mattf.& Kük.), but it has not been previously reported in Carex, including other fresh or dry C. fecunda specimens (P.Jiménez-Mejías, pers.obs.).This may indicate some differences with respect to other more southern C. fecunda populations and deserves further study.

Jim.Mejías & M.Escudero, Phytotaxa 260: 186 (2016)
our fieldwork campaign across the wUPOS)n slopes of the Western Cordillera in Pichincha, in forest remnants near to where the original collection of C. sodiroi was performed, we found plants that clearly match the diagnostic characteristics of C. sodiroi's type, allowing its distinction from the syntopic and closely-related C. polystachya: terminal clusters of spikes densely packed with many flowers, concealing the branch axis vs. with less flowers and lax, with the branch axis exposed; and glumes dark orange-brown vs. glumes lighter, pale reddish-brown, orange, stramineous or hyaline.Moreover, in Pululahua Geobotanic Reserve, we also identified individuals morphologically intermediate(124ECUAMA22 and 130ECUAMA22; HUTI,UPOS)between the co-occurring C. polystachya and C. porrecta Reznicek and Camelb., another species of sect.Indicae, whose morphology also approached the type of C. sodiroi.This makes us think that, perhaps, C. sodiroi is a local established hybrid between these two species.Further biosystematics studies are needed to figure out the taxonomic status of C. sodiroi and its systematic relationships with the other South American sect.Indicae species.