Twelve new and exciting Annonaceae from the Neotropics

Abstract As a result of concerted efforts of the community of Annonaceae taxonomists, increasingly detailed knowledge of the diversity of the Neotropical genera has been documented. With the exception of just two large genera, Annona and Xylopia, all Neotropical Annonaceae have been revised within the last 25 years. Subsequent to these publications, many new specimens have been collected and sent to us in Leiden for identification. These included a number that, despite the advanced state of taxonomic knowledge, proved to represent rarely collected, undescribed species. Here we describe 12 new species of Annona, Guatteria, Klarobelia, Tetrameranthus, and Xylopia. These species serve to illustrate the still underestimated diversity of the Neotropical flora, even in well studied plant groups like Annonaceae.


Introduction
The Neotropics are rich in plant diversity but how rich they are is still a matter of serious debate. For instance, estimating how many tree species exist in the Amazon basin is a nontrivial exercise (e.g. ter Steege et al. 2016, Cardoso et al. 2017, ter Steege et al. 2019). Yet, a proper estimate of this diversity is essential in order to answer questions on the ecology, evolution and origins of the Neotropical flora and fauna and the processes that are responsible for creating and maintaining its hyperdiverse communities (Cardoso et al. 2017).
Annonaceae is a family of ca. 2450 species of trees and lianas that is well represented in the Neotropical tree flora with ca. 950 species (Rainer andChatrou 2006, Maas et al. 2011). Understanding the species delineations in this family therefore contributes noticeably to the aim of understanding Neotropical tree diversity. Ongoing revisional work on Neotropical Annonaceae has already led to the revision of almost all genera during the last 25 years (for an overview see Erkens et al. 2017). Still, in this article new species in several genera of Annonaceae are described, and some notes are added on congeneric species of which the circumscription is affected by the newly described species.
The genus Annona L. is distributed in tropical America and tropical Africa. It is the second-largest genus of Annonaceae in the Neotropics with a total number of ca. 160 species, four of which inhabit Africa. Nowadays, it is taken in its original concept including two genera which were treated for a long time as segregates. This concerns Raimondia Saff. and Rollinia A.St.-Hil.: see the comments under the new species Annona caput-medusae and A. oleifolia.
The new species of Klarobelia Chatrou is an addition to the revision of Chatrou (1998). The Neotropical distribution of this genus is similar to that of Mosannona and Cremastosperma (Pirie et al. 2018). As in Mosannona, species of Klarobelia have small, non-overlapping distributions. Species have been discovered before (e.g. K. megalocarpa, Chatrou 1998) when new areas, in between known distribution areas, were first disclosed by plant collectors. The new species described has become known through collecting efforts, especially in the Peruvian province of Oxapampa.
Guatteria Ruiz & Pav. is the largest Neotropical genus of Annonaceae with more than 175 species. It is distributed from Mexico to south-eastern Brazil and was recently revised by Maas et al. (2015). Both because of very recently received specimens and a reinterpretation of some species complexes, several new species of Guatteria have to be described in the present paper.
The very small and poorly collected genus Tetrameranthus R.E.Fr., quite aberrant from all other genera of Annonaceae because of its spirally arranged (instead of distichous) leaves, was treated twice by Westra (Westra 1985;Westra and Maas 2012).
Tetrameranthus is a small genus with 8 species, occurring in the Amazon Region, neighbouring French Guiana and the Colombian state of Chocó. Recently we received very rich flowering and fruiting material, accompanied by nice field photographs of an undescribed species from Amazonian Peru which is herewith described.
The genus Xylopia L., the only genus of Annonaceae occurring in three continents, has recently been revised for Africa (Johnson and Murray 2018), but treatments for the Asian and American species are needed. The last revision of the Neotropical species of Xylopia dates back to Fries (Fries 1930, various supplements). A recent estimation by David Johnson of the total number of species of Xylopia in the tropics is 160, whereas ca. 50 species inhabit the Neotropics (DM Johnson pers. comm.). In the present paper a new Colombian species with very distinctive leaf features is described.

Materials and methods
All IUCN Redlist assessments were done on data from herbarium collections and following the IUCN guidelines (IUCN 2012(IUCN , 2017. Only criterion B could be used for the assessments since data on species' populations (Criteria A, C and D) and extinction probability (Criterion E) were lacking. The area of occupancy (AOO) was calculated by overlaying the occurrence data points with a 2×2 km grid and adding the area of all occupied cells. The extent of occurrence (EOO) was determined by calculating the area of the minimum convex polygon that was drawn around the outer occurrence points. Both AOO and EOO were calculated in R using the ConR package (Dauby et al. 2017). For several species only one to a few data points were available and these were considered Data Deficient. Although assessments can still be carried out for species with such low numbers of collections (Rivers et al 2011) it was unclear whether a lack of data caused the apparent rarity of these species or if they were actually rare, since no other data was available to the authors. It is important to note that a species with such small amounts of data can be endangered and thus a reassessment is needed when more data becomes available.
For those species that were not considered Data Deficient (i.e. had more than 3 collections) data on forest cover loss (Hansen et al. 2013) was investigated to infer if habitat loss was a threat for those species. This was the case if the species occurred in regions where forest cover loss had been observed in the past years. For this assessment, it was assumed that forest cover loss was regulated differently outside and across different protected areas, and thus every occurrence point within a particular protected area was considered as one location. For occurrence points that were not situated in a protected area, a 10 × 10 km grid was used to estimate separate locations.
It must be noted that no extensive survey on the occurrence of these species was undertaken; the AOO and number of locations are therefore a conservative estimate. Description. Tree 5-7 m tall, cauliflorous; young twigs rather densely covered with appressed brown hairs < 0.5 mm long, soon glabrous. Leaves: petioles 16-18 by 2 mm;  lamina narrowly elliptic, 28-30 by 9-12 cm (leaf index 2.5-3.1), membranous, greenish grey above in sicco, somewhat lighter so below, glabrous above except for the large veins sparsely covered with erect, brown hairs, sparsely covered with appressed hairs to glabrous below, base obtuse, extreme base very shortly attenuate, apex acuminate (acumen 10-15 mm long), primary vein impressed to flat above, secondary veins ca. 15, not loop-forming or loop-forming close to the apex (shortest distance between loops and margin ca. 2 mm), tertiary veins mostly percurrent, domatia present in axils of part of the secondary veins; plants androdioecious, probably: only bisexual flowers seen. Inflorescence borne on the stem on older branches, much-branched thyrsoids bearing many flowers in succession; pedicels 7-11 by 1-2 mm, gradually widening from base to flower, densely covered with appressed, brown hairs to 0.2 mm long; bracts triangular-ovate or broadly triangular-ovate, outer side densely covered with hairs 0.1-0.2 mm long, more or less persistent, upper bract 0.3-0.4 mm from base of pedicel; flower buds narrowly conical; sepals free or connate at the base, broadly ovate to triangular-ovate, ca. 1 mm long, appressed, later spreading to reflexed, apex acuminate, outer side densely covered with brown hairs; outer petals connate at the base, narrowly triangular, ca. 20 by 5 mm, outer side densely covered with brown hairs, inner petals ca. 0.4 the length of the outer ones, torus ca. 3 mm long, the lower third beset with stamens, the apical twothirds beset with carpels; stamens ca. 150, ca. 1.5 mm long, anther oblong, ca. 1 mm long, no apical prolongation of connective; carpels 150-200. Fruit ellipsoid, ca. 6.5 by 3.5 cm, densely covered with brown hairs ca. 0.2 mm long in young stage, becoming glabrous, areoles not or weakly protruding, not apiculate. Seeds ca. 6 by 5 mm, brown.
Habitat and ecology. On industrial ground, in secondary forest. At elevations of 350-450 m. Flowering: March; fruiting: June.
Notes. A domatium here is a small thin membrane in the axil spanning the distance between primary vein and secondary vein. It conforms to the Annona muricata type ( Van den Bos et al. 1989).
Annona caput-medusae clearly falls within a distinct group formerly known as the segregate genus Raimondia (Safford 1913;Westra 1995), but (re-)united later with Annona (Rainer 2007). When using Westra's key A. caput-medusae comes closest to A. quinduensis Kunth which generally is not cauliflorous. Whereas A. quinduensis normally is found at higher elevations up to 2500 m, A. caput-medusae, as known from the scanty material collected thus far, occurs at elevations below 500 m. The flowers we examined appear to be bisexual. However, given the obvious similarity with other former Raimondia species, which are all androdioecious, staminate flowers might be expected in A. caput-medusae as well.
Etymology. Caput (L) = head. Medusa, an ancient Greek goddess whose head was covered with snakes. Referring to the shape of the inflorescence.
Preliminary IUCN conservation status. DD. This species is only known from two localities. Although the collections are not made near each other, more data are needed to determine the AOO and EOO. Also, the current population size and population trend of this species are unknown. Habitat loss because of forest cover loss is a Diagnosis. When using the key of the Flora Neotropica Monograph of Rollinia (Maas, Westra et al. 1992) Annona oleifolia keys out to the SE Brazilian Annona neosericea H.Rainer by an indument of appressed hairs on the lower side of the lamina, non-gibbous sepals, and narrow wings, but it is very distinct from that species by the very low number of carpels (≤25 vs. ≥100, respectively), very narrow leaves (leaf index ≥5 vs. 2.5-3) and the slightly recurved instead of horizontal to erect wings in Annona neosericea.
Habitat and ecology. In forest. At elevations of 100-300 m; flowering: May, June; fruiting: August and September.
Notes. The flower had to be described from photographs. Annona oleifolia falls within the former concept of Rollinia because of the characteristic shape of the corolla, with the outer petals wing-like, and the whole flower suggesting a miniature propellor (see Maas et al. 1992).
Using the key to the species in the monograph of Rollinia (Maas et al. 1992), Annona oleifolia ends near Rollina sericea = Annona neosericea, but it is immediately distinct from that species by the very low number of carpels (≤25 vs. ≥100, respectively).
Preliminary IUCN conservation status. EN B2ab(iii). The EOO (126.356 km2) was too large to classify as threatened, but AOO (24 km2) would classify as Endangered. It was determined that this species occurs in 5 locations. Although the species occurs within national parks in Ecuador, it is also found in heavily fragmented forest regions. Since the current population size and population trend of this species are unknown, we have classified it as Endangered.
Other specimens examined. ECUADOR. Napo: La  Diagnosis. Guatteria aliciae is similar to Guatteria tenera R.E.Fr. in terms of its very small and narrow leaves that are not verruculose, straight young twigs, and secondary veins that are impressed to raised on the upper side of the lamina, but it is distinct from that species by long-pedicellate flowers (20-45 vs. 10-20 mm long) and longer petioles (5-10 vs. 2-5 mm long) and almost smooth (to slightly pitted) seeds.
Distribution. Panama (Veraguas) (Fig. 9). Habitat and ecology. In low forest of 6-8 m tall, with the palm species Colpothrinax aphanopetala R.Evans. At an elevation of ca. 1000 m. Flowering and fruiting: July.
Notes. Guatteria aliciae is named in honour of its collector Alicia Ibañez, who assisted us in all kinds of ways in 2006, during our visit to Panama. This species is only known from one locality in the Panamanian province of Veraguas.
Preliminary IUCN conservation status. DD. This species is only known from one locality with three collections. Therefore AOO and EOO could not be calculated. Given that the species occurs in Santa Fé National Park we assume that currently there are no major threats. However, since the current population size and population trend of this species are unknown, it was assessed as Data Deficient.  Diagnosis. Resembling the Amazonian species G. modesta Diels by the long-attenuate leaf base, but differing by a shorter petiole (2-5 vs. 5-10 mm long), and distinct secondary and tertiary veins which are strongly raised above vs. inconspicuous and flat to slightly raised above.
Habitat and ecology. In non-inundated, Atlantic rain forest. At an elevation of ca. 300 m. Flowering: March; fruiting: not recorded.
Notes. Guatteria attenuata, only known from the type collection and a second sterile collection from the same tree, is according to the label a tree of 35 m high, a size rarely seen in the genus. Also the long-attenuate leaf base is a rare feature in the Guatteria, although in G. modesta from the Amazon region, the base is attenuate.
Preliminary IUCN conservation status. DD. This species is only known from one locality and the current population size and population trend of this species are unknown. Guatteria attenuata was collected in an area that shows habitat loss due to forest cover loss and this is therefore a possible threat for this species. Nonetheless, this species was assessed as Data Deficient given the overall lack of data of this species.
Habitat and ecology. In rich rain forest with Licania, Ebenaceae, tree ferns, and palms. At an elevation of ca. 1330 m. Flowering: June; fruiting: unknown.
Notes. Guatteria kamakusensis was found at a fairly high elevation as compared to most of the Guatteria species in the Guianas, in a poorly collected area. It is similar to G. schomburgkiana Mart. in leaf shape and the short-pedicellate flowers, but the connective shield is papillate with few intermixed hairs, vs. a densely hairy connective shield in G. schomburgkiana. Its place remains unclear though, especially because the fruits are still lacking, and requires further research.
Preliminary IUCN conservation status. DD. This species is only known from one locality in a poorly collected area and the current population size and population trend of this species are unknown. There seem to be no immediate threats to this species in terms of habitat loss. However, since no AOO and EOO could be determined or any other assessment criterium could be used, this species was assessed as Data Deficient. Diagnosis. Guatteria pseudoferruginea superficially resembles the SE Brazilian G. ferruginea A.St.-Hil. by having young twigs densely covered with erect, brown hairs, nonverruculose leaves, and ellipsoid seeds, but it is different from that species by being not cauliflorous, and having smaller leaves (14-20 vs. 17-40 cm long), petals hairy on both sides, shorter pedicels (7-15 vs. 15-70 mm long) and seeds that are smooth vs. pitted.  Description. Tree or shrub 5-12 m tall, 5-15 cm diam.; young twigs densely covered with erect, brown hairs, soon glabrous. Leaves: petioles 4-10 by 1-2 mm; lamina narrowly elliptic, 14-20 by 4-6.5 cm (leaf index 2.8-3.5), chartaceous, not verruculose, dull, greyish above, brown below, densely covered with appressed hairs to glabrous above, sparsely covered with appressed hairs to glabrous below, base acute to slightly attenuate, apex acuminate (acumen 5-15 mm long), primary vein impressed above, secondary veins distinct, 15-17 on either side of primary vein, slightly impressed above, smallest distance between loops and margin ca. 2 mm, tertiary veins flat above, reticulate. Inflorescence axillary, 1-2-flowered; pedicels 7-15 by 1 mm, to 15-20 by 1-2 mm in fruit, densely covered with appressed and erect, brown hairs, articulated at ca. 0.3 from the base; bracts not seen; flower buds subglobose; sepals free, broadly ovatetriangular, 7-8 by 6 mm, reflexed, outer and inner side densely covered with appressed and erect, brown hairs, inner base glabrous; petals green in vivo, ovate-elliptic, 7-12 by 5-11 mm, both sides densely covered with appressed and erect, brown hairs, except for the inner glabrous base; stamens ca. 1.5 mm long, connective shield papillate. Monocarps 10-50, colour in vivo not recorded, black in sicco, ellipsoid to narrowly ellipsoid, 10-20 by 5 mm, glabrous, apex apiculate (apiculum < 0.5 mm long), wall ca. 0.2 mm thick, stipes 15-20 by 5 mm. Seed ellipsoid, dark, shiny brown, ca. 8 by 5 mm, smooth.
Habitat and ecology. In non-inundated forest or gallery forest. At elevations of 100-1000 m. Flowering: January to April; fruiting: March, June.
Notes. Specimens of Guatteria pseudoferruginea have previously been identified as G. ferruginea A.St.-Hil. from E and SE Brazil, which differs from G. pseudoferruginea in being cauliflorous. Both species are characterised by a dense indument of brownish, erect hairs on the twigs. However the new species differs from G. ferruginea by characters as given in the diagnosis, notably by petals covered with brown hairs on both sides (vs. on the outer side only in G. ferruginea), and by the absence of cauliflory .The description of G. pseudoferruginea, formerly named Guatteria sp. 2 in the monograph of Guatteria (Maas et al. 2015), could be completed by the first author during a recent visit to Colombia and the COL Herbarium in Bogotá.
Preliminary IUCN conservation status. EN B2ab(iii). EOO (378.742 km 2 ) was too large to classify as threatened, but AOO (20 km 2 ) would classify as Endangered. It was determined that this species has 5 locations almost all in heavily deforested areas outside national parks. Furthermore, because no information is available on the current population size and population trend of this species, we have classified it as Endangered.
Distribution. Panama (Comarca Ngabe-Buglé) (Fig. 9).  included the appropriate illustration as Pl. 7a in our work. G. rotundata, it should be pointed out, is unique among Central American species of Guatteria on account of its leaves having a rounded apex. Recently we received the corresponding herbarium material and it became clear that the photographed plant did not represent G. rotundata at all, but an undescribed species instead. G. pseudorotundata differs from G. rotundata by characters as given in the diagnosis, but notably the lack of verruculae in the lamina. Although more or less hidden from view in the photograph just mentioned, the leaf apex in G. pseudorotundata tends to be acute rather than obtuse or rounded (excl. the extreme tip) as in G. rotundata.
Preliminary IUCN conservation status. DD. This species is only known from three nearby collections and therefore no AOO and EOO was calculated (that would constitute one location) in a region that is partially deforested. More continuous forest is, however, available nearby but it is unclear whether this species occurs there. Habitat loss because of forest cover loss is therefore a possible threat for Guatteria pseudorotundata. Unfortunately, no other assessment criterium could be used for this species since no information is available on the current population size and population trend of this species. Hence, this species was assessed as Data Deficient.
Notes. Guatteria turrialbana can be distinguished by its essentially glabrous, dark green, shiny and conspicuously reticulate leaves on both sides, especially upon drying, lamina commonly narrowly obovate with a flat to slightly raised primary vein above. The rostrate to apiculate apex of the monocarps is also a characteristic feature. Material of this species has previously been identified as G. lucens Standl. However, the lamina is not verruculose in G. turrialbana vs. up to densely verruculose in G. lucens. Other differences can be found in the monocarps: G. turrialbana has rostrate to apiculate ones vs. ellipsoid ones in G. lucens. Lastly, G. lucens is a lowland (0-900 m) species whereas G. turrialbana has mainly been recorded at higher elevations (Maas et al. 2015). Preliminary IUCN conservation status. EN B1ab(iii)+2ab(iii). Both EOO (10 km2) and AOO (8 km2) would classify as Endangered. It was determined that this species has 3 locations,all of them in heavily deforested areas. Deforestation is therefore a major threat to this species and habitat quality is expected to decline in the near future. No information is available on the current population size and population trend of this species. Given all this we assessed this species as Endangered.
Distribution. Peru, only known from a small area in the department of Pasco, districts of Palcazú and Villa Rica (Fig. 3).
Habitat and ecology. In primary forest. At elevations of ca 500-1400 m. Flowering: October and November; fruiting: between February and August (mature fruits collected in July and August).
Notes. Klarobelia rocioae is easy to recognise through its combination of large leaves, hairy flowers, and relatively large and ellipsoid monocarps. Of the Amazonian species of Klarobelia Chatrou, K. napoensis Chatrou has comparably large leaves, but differs in the flowers that are glabrous on the outer side, and in the globose monocarps. K. pandoensis Chatrou and K. pumila Chatrou are two other Amazonian species that share a small habit with K. rocioae (Chatrou 1998;Chatrou and Pirie 2003). With K. rocioae they share outer petals that are hairy on the outer side, giving the flowers a brown appearance rather than glabrous flowers in other species of Klarobelia that appear black when dried. K. pumila Chatrou, can easily be distinguished from K. rocioae by the smaller leaves (12-16 by 4-6 cm vs. 17-35 by 6.5-11.5 cm in K. rocioae) and smaller monocarps (9-15 by 6-8 mm vs. 20-27 by 10-12 mm). K. pandoensis Chatrou can be distinguished from K. rocioae by the smaller leaves (8.5-12.5 by 3-4.5 cm) and smaller monocarps (15-18 by 6-10 mm) too. Additionally, K. pandoensis can easily be distinguished from the two other short stature species by the dense indument on petioles, young twigs and inflorescences axes (glabrous to sparsely hairy in K. rocioae and K. pumila). The three species have non-overlapping areas of distribution in the Diagnosis. Tetrameranthus trichocarpus resembles T. globuliferus Westra from Ecuador in leaf shape and in the young twigs covered with brown, stellate hairs, but differs by 5-merous (vs. 6-merous) flowers and hairy (vs. glabrous) monocarps, and also by smaller leaves (16-28 vs. 27-37 cm long).   Description. Tree ca. 10 m tall. Young twigs and petioles densely covered with stiff, brown, mostly stellate hairs to 1-2 mm long. Leaves: petioles 8-10 by 3-4 mm; lamina narrowly obovate, 16-28 by 5-8 mm (leaf index 2.5-4), bright shiny green above and pale green below in vivo, dark greenish grey above and greenish brown below in in sicco, densely covered with brown hairs ≥1 mm long on primary vein and less densely so on secondary veins above, elsewhere rather densely to sparsely covered with stellate and simple hairs, to at last glabrous above, densely to rather densely covered on primary vein and secondary veins below, elsewhere sparsely covered with stellate and simple hairs mainly on lesser veins below, base narrowly acute, apex acuminate (acumen ca. 5 mm long), primary vein slightly prominent to almost flat above, becoming canaliculate in sicco, secondary veins 12-20 on either side of primary vein, mostly loop-forming, shortest distance between loops and margin 1-2 mm, tertiary veins percurrent. Flowers solitary in axils of leaves; peduncles 7-10 by 2 mm; pedicels 30-35 by 2{-3} mm, to 4 mm diam. in fruit, peduncles and pedicels densely covered with hairs as on twigs; bracts not seen; perianth in 5-merous whorls, petals pale greenish creamy suffused with purple, inner base of inner petals yellowish white, sepals (±) free, narrowly trangular, 4-5 by 10-12 mm, outer side densely covered with hairs as on pedicels to 1 mm long, the inner side same but less densely; outer petals narrowly elliptic-ovate or elliptic-oblong, 30-37 by 10-12 mm, inner side with basal callus to ca. 2/5 of the length and triangular in shape, inner petals narrowly obtriangular-elliptic, about as long as outer petals, slightly narrower than outer petals, markedly recurved about the middle, basal callus on inner side ca. 2/3 of the length and almost touching the side, all petals densely covered with similar though somewhat smaller hairs as on sepals, except for callose parts sparsely so; stamens ∞, apical prolongation of connective shield-like, ca. 1 mm in diam., glabrous; carpels ca. 8, ca. 4 mm long, densely covered with erect hairs to 0.5 mm long on the abaxial side. Monocarps 4-6, pinkish green and somewhat shiny in vivo, brown and with shriveled wall in sicco, ovoid to globose, 4-4.5{-5} by 3-3.5{-4.5} cm, with a conical, obtuse apicule ca. 3-4 mm long, with an oblique constriction (2-seeded forms, only visible in sicco), densely to rather densely covered with stiffly, erect, whitish, stellate and simple hairs. Seeds (1-)2 per monocarp.
Notes. Tetrameranthus trichocarpus is very similar to T. globuliferus Westra, from Ecuador (Maas et al. 1988), and also a narrow endemic. Apart from being 5-merous in T. trichocarpus vs. 6-merous in T. globuliferus, the flowers of the two species resemble each other very much. Both these species share two features with the far-remote T. guianensis Westra & Maas, namely a thick fruit wall that shrivels with drying, and an indument of coarse, stellate and simple hairs on vegetative parts. To our knowledge, this is the only species of Tetrameranthus with permanently hairy fruits.
Preliminary IUCN conservation status. DD. This species is only known from one collection and therefore no AOO and EOO could be calculated. Also, no other assessment criterium could be used for this species since no information is available on the current population size and population trend of this species. The species seems to occur in a large, pristine forest area and habitat loss does not seem to be an immediate threat to Tetrameranthus trichocarpus. However, given the overall lack of data, it was assessed as Data Deficient.
Habitat and ecology. In high rain forest or high caatinga forest. At elevations of 250-500 m. Flowering and fruiting: April.
Notes. Xylopia longicaudata is easily distinguished from other species by the caudate leaf apex. It slightly resembles X. uniflora R.E.Fr., described from caatinga forests in Amazonian Brazil (Ducke RB 29017, Brazil, Amazonas, Rio Curicuriary, Cajú cataracts, 29 February 1936), but the leaf venation in the latter species is much less raised than in X. longicaudata and the apex acuminate instead of caudate.
Preliminary IUCN conservation status. NT. EOO (35.750 km 2 ) was too large to classify as threatened, but AOO (20 km 2 ) would classify as Endangered. It was determined that this species has 5 locations, none of them in national parks and some in slightly fragmented areas of which the habitat is expected to decline in the near future. Overall, however, this species occurs in large stretches of pristine forest and although no information is available on the current population size and population trend of this species, we expect the population size to be rather large. Therefore this species was classified as Near Threatened. Other