Eriosyce Phil.

Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, Phytotaxa 550 (2), pp. 79-98 : 91-94

Eriosyce Phil. (see a proposed key below):

—The results of the molecular phylogeny by Guerrero et al. (2019b) show that Eriosyce s.l. (see also Kattermann 1994) species from Chile, Perú, and Argentina form a strongly supported monophyletic clade, but only with the exclusion of Rimacactus laui (Lüty) Mottram , a species that had been included within Eriosyce section Neoporteria subsection Chileosyce by Kattermann (1994, as Eriosyce laui Lüthy ) and within the “ Islaya -group” by Hunt et al. (2006, 2013, as Eriosyce laui ).

Phylogenetic analyses retrieved seven major clades in the genus ( Guerrero et al. 2019b). The first branching clade (Section Eriosyce Katt. ) comprises two species from Chile, one from Argentina, and two from Perú. The next branching clade (Section Campanulatae P.C. Guerrero & Helmut Walter) comprised two taxa from southern-central Chile [ Eriosyce marksiana (F.Ritter) , Eriosyce marksiana var. lissocarpa (F.Ritter) Katt. ]. Subsequently branching is a clade (Section Pyrrhocactus (A.Berger) P.C. Guerrero & Helmut Walter) harbouring four species endemic to Argentina. Then, a group of 12 taxa endemic to south and and northern-central Chile (Sect. Horridocactus (Backeb.) P.C. Guerrero & H.E. Walter). The next branching clade formed by three species endemic to north central Chile (Section Diaguita P.C.Guerrero & Helmut Walter). Subsequently branching is a clade comprising 15 taxa endemic to south and northcentral Chile (Sect. Horridocactus (Backeb.) P.C. Guerrero & H.E. Walter), and finally, an unnamed clade composing 22 taxa from northern Chile. In Kattermann (1994), Wallace proposed an infrageneric classification with two sections [sect. Eriosyce (with subsect. Eriosyce , subsect. Islaya (Backeb.) Katt. , and subsect. Pyrrhocactus (A.Berger) Katt. and sect. Neoporteria (with subsect. Neoporteria (Britton & Rose) Katt. , subsect. Horridocactus (Backeb.) Katt. , and subsect. Chileosyce Katt. ), whereas Hunt et al. (2006, 2013) and Hoffmann & Walter (2004) proposed six “groups” (= subgenera) based on the former genera Islaya Backeb. , Pyrrhocactus A.Berger , Neoporteria Britton & Rose , Horridocactus Backeb. , Thelocephala Ito , and Eriosyce Phil. None of these concepts were corroborated by the results of the molecular phylogeny ( Guerrero et al. 2019b). Moreover, at species level, the large “species complexes” E. napina (Phil.) Katt. , E. odieri (Lem. ex Salm-Dyck) Katt. , E. heinrichiana (Backeb.) Katt. , E. subgibbosa (Haw.) Katt. , (Backeb.) Katt and E. curvispina (Bertero ex Colla) Katt. were not supported by the results of Guerrero et al. (2019b) (see Table 2 ). In addition, the taxonomic delimitation of E. curvispina Bertero ex Colla including several infraspecies did not resist standing upright much longer, by not complying with the principle of monophyly as reported by Guerrero et al (2019b), and by new molecular data of several putative members of the E. curvispina complex (Villalobos-Barrantes et al. 2022).

Concerning Eriosyce kunzei (C.F. Först.) Katt. , a nomenclatural note is necessary.In the protologue of Echinocactus kunzei, Förster gave “ Chile ” as the type locality but mentioned that the plants are sometimes covered by a light layer of snow in winter. F. Ritter (1980) referred Förster’s E. kunzei to the plants from near Copiapó, that perfectly match Förster’s description (see Table 3 F . Ritter, Kakteen in Südamerika, p.p. 955 and 956 and the specimen at SGO 121487, Ritter 220 loc. 2) clearly shows the typically long and narrow areole of this taxon. Kattermann (1994), however, chose material from Guanta, Provincia de Elqui, FK 459 (DBG) for his typification of Eriosyce kunzei and based his decision not to refer the plants from around Copiapó to Förster’s Echinocactus kunzei (as proposed by F. Ritter, 1980) on the single argument that it never snows in this area. Yet, according to meteorological data ( Vergara 2011) this assumption cannot be upheld. Hunt (2003) stated that “… the editorial preference would be to supersede Kattermann’s neotypification and to substitute FR 220 material from Paipote (SGO 121487)”. Moreover, several relevant morphological characters ( Kattermann 1994; stems, areoles and spines, see Table 3 ) of the plants from Guanta and its vicinity do not match Försters original description. We thus follow F. Ritter’s proposal to refer Förster’s Echinocactus kunzei to the plants from the vicinity of Copiapó. This makes E. confinis (F.Ritter) Katt. a heterotypic synonym of E. kunzei (C.F. Först.) Katt. (see list of species).

With regard to Echinocactus jussieui Monv. ex Salm Dyck a note is necessary. E. jussieui was published by Salm-Dyck (1849: 34, 170–171) in his Hortus Dyckensis with a short diagnosis (“ Caule aterrime virente, tuberculis gibberatis in costas 13 subconfluentibus, aculeisque brunneis ad praecedentem valde accedit; sed differt aculei centrali validissimo ”). No specimen referring to the original material could be traced and, therefore, a neotypification is required according to the Art. 9.8 of ICN (Turland et al. 2018). Ritter (1980) used the epiteth jussieui for plants occurring in the middle and upper Elqui Valley ( Pyrrhocactus jussieui (Monv. ex Salm-Dyck) F.Ritter var. jussieui and var. spinosior F.Ritter ), argueing that these plants match Salm-Dyck’s original protologue (see below) based on “body blackish-green”, “ribs 13”, “ribs tuberculate”, “spines brown”, “strong central spine”. Kattermann (1994: 156), however, considered E. jussieui as “a name of doubtful application, possibly referable to Eriosyce heinrichiana (Backeb.) Katt. ”. To avoid the epithet “jussieui”, he chose the name E. heinrichiana subsp. intermedia var. intermedia (F.Ritter) Katt. for the plants in and around the Elqui Valley. Yet, Ritter’s locality for his P. setosiflora var. intermedia was “ 60 km south of the Elqui Valley low coastal hills”. The morphology of the plants occurring in this region differs in several character states from the plants in the middle- and upper Elqui Valley (plants from the Elqui Valley in brackets): stems subglobose to flat on the ground (globose to somewhat elongating), stem diameter 2–5 cm (6–9 cm), stem colour often reddish-brown (dark-green); spines thin (thick), mostly straight (mostly curved upward), 1.5–4.0 cm long (3–6 cm); flowers 4–6.5 (3–4.5); seeds small, 0.9 × 0.7 mm (1.2 × 0.9 mm). Moreover, Kattermann’s “subspecies intermedia ” is not at all related to E. heinrichiana subsp. heinrichiana , but a species in its own right ( Guerrero et al. 2019b), thus the name E. heinrichiana subsp. intermedia can not be uphold and, consequently, a new name for the plants from the middle- and upper Elqui Valley must be found. Following Ritter’s (1980) arguments, we here propose a new combination for these plants under the genus Eriosyce as well as a neotypification of Echinocactus jussieui based on a Ritter’s collection:

1. Stem diameters large; ribs many; areoles large; roots always fascicular ...........................................................................................2

- Stem diameters from very small to medium-sized; ribs less numerous; areoles smaller; roots various............................................5

2. Seed testa cell appendages lacking or inconspicuous, interstices pitted or sunken; flowers funnel-form (Sect. ERIOSYCE ) .........3

- Seed testa cell appendages present, interstices not pitted nor sunken; flowers campanulate (Sect. CAMPANULATAE ) .................. ................................................................................................................................................................................. 31. E. marksiana

3. Flowers and fruits with erect spine-like bristles; fruits not elongating ..............................................................................................4

- Flowers and fruits not as above; fruits much elongating, balloon-like ..................................................................... 25. E. islayensis

4. Stems very thick; fruits completely covered by wool; loculus pulpless; dehiscent by a basal pore ................................ 5. E. aurata

- Stems thinner; fruits not completely covered by wool; loculus with mucilagineous pulp; indehiscent ............. 39. E. rodentiophila

5. Stems subglobose, globose to somewhat elongating; flowers always diurnal, always funnel-form, usually not fuchsia-colour, interior perianth segments directed outward; nectary small; seed notched below hilum or not ........................................................6

- Stems elongating; flowers mostly tubular, fuchsia-colour, interior perianth segments inclining inward to erect; flowers usually remaining open during the night; nectar chamber usually very large; seeds never notched below hilum (sect. NEOPORTERIA ) ... ..........................................................................................................................................................................................................37

6. Nectary always tubular; ovary always elongate to isodiametric; roots various .................................................................................7

- Nectary widened at base; ovary compressed (with a few exceptions); roots never facicular (unnamed Section)...........................18

7. Pericarpel, hypanthium and fruits with inconspicuous white wool and without bristles; tubercles arranged in parastichies (Sect. HORRIDOCACTUS ).........................................................................................................................................................................8

- Flowers and fruits always covered by long dense wool and numerous long porrect bristles; ribs not arranged in parastichies (Sect. DIAGUITA )......................................................................................................................................................................................16

8. Stems green, never pruinose, medium; fruits short, ovoid to barrel-shape, perianth remnant attachment area large; roots various... ............................................................................................................................................................................................................9

- Stems grey-green to grey brown, often ±pruinose, small, fruits elongating, perianth remnant attachment area small; taproots always present...............................................................................................................................................................................................14

9. Stems with a tendency towards basal branching ..............................................................................................................................10

- Stems not branching .........................................................................................................................................................................11

10. Fruit dry, dehiscing by partial circumscissile splitting; spines finely acicular; ribs low............................................ 3. E. aspillagae

- Fruit-wall fleshy when mature, dehiscing by a complete circumscissile slit; spines thicker; ribs steep......................... 2. E. armata

11. Stems ±elongating, spines numerous, often obscuring stem............................................................................................................12

- Stems subglobose to globose or somewhat elongating, spines less numerous.................................................................................13

12. Spines long, white to yellowish, tipped dark; floral bristles scant, only near hypanthium rim ................................... 15. E. engleri

- Spines shorter, yellow, not turning grey; bristles numerous, distributed all over hypanthium .............................. 21. E. garaventae

13. Ribs numerous; ovary much elongated; rootstock a long taproot; spines thickly acicular ..................................... 28. E. limariensis

- Ribs fewer; ovary ±isodiametric; roots fascicular; spines thinner ........................................................................... 13. E. curvispina

14. Ribs dissolved into tubercles arranged in parastichies; stems very small; spines very short...........................................................15

- Ribs arranged in orthostichies; stems medium-sized; spines much longer .................................................................. 44. E. jussieui

15. Stems never elongating, tubercles large; spines black; flowers with brownish wool and black bristles ..................... 32. E. napina

- Stems elongating with age; tubercles small, spines horn-coloured to white; flowers with white wool and white bristles.................. ................................................................................................................................................................................... 14. E. duripulpa

16. Stems to 6 cm; ribs to 21 discernible, tubercles rhomboid...............................................................................................................17

- Stems 2-4 cm; ribs 12, tubercles rounded ...................................................................................................................... 38. E. riparia

17. Stems not branching; roots neckless; adult plants spineless; style yellow; seed large, hilum pyriform ............... 19. E. fankhauseri

- Stems branching; roots with a narrow neck; spines present; style red; seed smaller, hilum oval .............................. 47. E. tenebrica

18. Stems medium-sized, ±globose sometimes elongated; spine length various, centrals always present ............................................19

- Stems very small, flattened to subglobose; spines ≤ 1.5 cm, centrals often absent ..........................................................................28

19. Spines long, hair-like, flexible, often contorted; stem epidermis grey-brown, ±pruinose .............................................. 12. E. crispa

- Spines shorter, not hair-like or flexible; epidermis usually green to grey-green, blackish-green or glaucous, not pruinose............... ..........................................................................................................................................................................................................20

20. Spines numerous, occulting stems....................................................................................................................................................21

- Spines less numerous, not occulting stems.......................................................................................................................................24

21. Flowers purple, narrowly funnel-form; exterior perianth segments ±bent downwards ...................................................................22

- Flowers never purple; funnel-form; exterior perianth segments not as above .................................................................................23

22. Perianth segments concolourous; interior ones nearly erect; ribs numerous ............................................................. 42. E. sociabilis

- Perianth segments bicolourous; interior ones directed outward; ribs fewer............................................................. 46. E. taltalensis

23. Stems elongating; areoles round-oval; spines yellow, not turning grey; fruit red and juicy when ripe ................ 17. E. eriosyzoides

- Stems globose; areoles narrow; spines brownish, soon turning grey; fruit dry when ripe............................................. 27. E. kunzei

24. Fruits clavate; spines short; flowers very short ................................................................................................................................25

- Fruits not clavate; spines longer; flowers longer..............................................................................................................................26

25. Stems globose to somewhat elongating; flowers yellow; ribs to16 ....................................................................... 24. E. iquiquensis

- Stems small, subglobose; flowers reddish; ribs fewer............................................................................................... 7. E. caligophila

26. Stems green or glaucous; ribs few; flowers whitish-yellowish ........................................................................................................27

- Stems blackish grey-green; ribs more numerous; flowers reddish............................................................................. 4. E. atroviridis

27. Stems (sub)globose, green, areoles not woolly; style whitish; fruits yellowish.......................................................... 6. E. calderana

- Stems ±elongating, glaucous, areoles woolly; style red; fruits red ...................................................................... 36. E. paucicostata

28. Ribs dissolved into tubercles arranged in parastichies ....................................................................................................................29

- Ribs arranged in orthostichies ..........................................................................................................................................................35

29. Hypanthium and pericarpel with long porrect bristles .....................................................................................................................30

- Hypanthium and pericarpel not as above .........................................................................................................................................33

30. Stems branching, not elongating; spines often completely lacking, not interwoven, short .............................................................31

- Stems often elongating with age, not branching; spines interwoven, to 1 cm, always present................................... 1. E. aerocarpa

31. Tubercles, areoles and spines tiny ................................................................................................................................. 26. E. krausii

- Tubercles and areoles larger, spines longer ......................................................................................................................................32

32. Stem apex with long white wool; fruit areoles on top of large nipple-shaped tubercles; seed testa surface shiny, smooth, cells lacking appendages................................................................................................................................................... 30. E. malleolata

- Stem apex not woolly; fruit etuberculate; seed testa opaque, rugose, cell appendages present; flowers whitish................................ .............................................................................................................................................................................. 18. E. esmeraldana

33. Stems to 6 cm diam.; spines radiating, straight; roots conical without a neck................................................................ 35. E. odieri

- Stems <4 cm ¸ spines appressed, recurved; roots a long taproot with a neck-like constricton .........................................................34

34. Stems flat to the ground, not elongating; tubercles large, obtuse; spines few, often lacking; fruits with long white wool ................. ................................................................................................................................................................................ 22. E. glabrescens

- Stems elongating with age; tubercles smaller, pronounced; spines numerous; fruits with short wool ............................. 20. E. fulva

35. Fruits short barrel-shape; perianth remnant area and attach basal pore large; fruit-wall thick at maturity.............. 43. E. spectabilis

- Fruits ellipsoid, elongating; perianth remnant area small, basal pore small; fruit-wall thin ….. 36

36. Stems hard, bluish grey-green; fruits with black bristles .......................................................................................... 37. E. recondita

- Stems soft, grey-brown; fruits with white bristles......................................................................................................... 34. E. occulta

37. Flowers tubular, always fuchsia, hypanthium long in relation to pericarpal length.........................................................................40

- Flowers (narrowly)funnel-form, fuchsia or not hypanthium short in relation to pericarpal length .................................................38

38. Stems much elongating; flowers fuchsia; roots fibrous ................................................................................................ 9. E. chilensis

- Stems only moderately or not elongating; flowers not fuchsia; taproots .........................................................................................39

39. Stems somewhat elongating, apex woolly; flowers with plenty of white hairs and bristles ...................................... 41. E. simulans

- Stems (sub)globose, apex not woolly; flowers not as above................................................................................ 23. E. heinrichiana

40. Roots tuberous or (short)taproots; perianth segments narrowly lanceolate with long tips ..............................................................41

- Roots fascicular; perianth segments broader, apex often rounded ...................................................................................................49

41. Flowers not exceeding spines; fruits medium-sized to small, ellipsoid or slender ..........................................................................42

- Flowers exceeding spines; fruits ovoid to globose, larger................................................................................................................43

42. Flowers medium-sized, carmine-fuchsia; fruits, medium-sized, ellipsoid.......................................................................................44

- Flowers small, pale fuchsia; fruits small, slender.............................................................................................................................45

43. Ribs up to16; spines numerous, acicular, often hair-like, contorted .............................................................................. 49. E. villosa

- Ribs fewer; spines less numerous, thicker, never hair-like nor contorted ........................................................... 48. E. vallenarensis

44. Spines numerous, completely obscuring stems; these small; areoles small................................................................. 29. E. litoralis

- Spines less numerous; stems thicker; areoles larger.......................................................................................... 50. E. wagenknechtii

45. Stems much elongating, clavate; ribs few; spines few, short ........................................................................................ 10. E. clavata

- Stems only moderately elongating; ribs more numerous; spines more numerous, longer...............................................................46

46. Spines flexible, soft, completely hiding stems .................................................................................................................................47

- Spines not as above ..........................................................................................................................................................................48

47. Stems elongating; flowers very large, bristles inconspicuous......................................................................................... 40. E. senilis

- Stems smaller; flowers smaller, bristles long and numerous..................................................................................... 16. E. elquiensis

48. Flowers very large, bristles numerous; stems green.............................................................................................. 11. E. coimasensis

- Flowers smaller, bristles few; stems dark grey-green ........................................................................................... 33. E. nigrihorrida

49. Stems much elongating; stigma lobes not exserted beyond perianth segments ...................................................... 45. E. subgibbosa

- Stems only moderately elongating; stigma lobes exserted............................................................................................ 8. E. castanea