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Oliveira, Anabela Silveira De, Pontes, Rodrigo Corrêa (2023): Parodia hofackeriana (Cactaceae-Notocacteae), a new species from southwest of Rio Grande do Sul, Brazil. Phytotaxa 598 (4): 283-292, DOI: 10.11646/phytotaxa.598.4.2, URL: http://dx.doi.org/10.11646/phytotaxa.598.4.2

Parodia hofackeriana A. S. Oliveira & R. Pontes sp. nov. (Figs. 1A–D, 2A–D and 3A–D). Type:— Brazil. Rio Grande do Sul: Santana do Livramento, 265 m, 12 October 2015, Oliveira & Deble 84b (holotype PACA!, isotype PACA!). Diagnosis (latin):— Parodia hofackeriana similis est Parodia mueller-melchersii subsp. mueller-melchersii a quo magnitudine minore differt (9 × 10 vs. 20 × 15 cm), figura caulis (globoso-cylindraceo vel brevissimo vs. rotundato vel elongato); costarum numero minus (18–20 vs. 20–25); aculeis centralibus (3–4 vs. 1); plures aculei radiales (13–19 vs. 15–18), flores maiores (usque ad 6 vs. usque ad 5 cm). Diagnosis (english):— Parodia hofackeriana is similar to Parodia mueller-melchersii subsp. melchersii from which differs by its smaller size (9 × 10 vs. 20 × 15 cm), shape of the stem (globose to short-cylindric vs. rounded to elongated); less number of ribs (18–20 vs. 20–25); number of central spines (3–4 vs. 1); number of radial spines (13–19 vs. 15–18), larger flowers (up to 6 vs. up to 5 cm). Description:— Saxicolous plants with succulent habit, globose to short-cylindrical, 9–10 cm high × 7–9 cm wide, single, not producing offsets naturally (excepted when the areoles are damaged), epidermis dark-green, opaque, slightly squamose, with umbilicate apex; with 18–20 ribs, straight, well-defined but slightly tuberculate, with little sinuses between successive tubercles, 0.1–0.2 cm tall × 0.9–1.1 cm thick, with almost with no proeminent edges below the areoles; areoles located at the slightly sinuses between consecutive tubercles, rounded, 0.3–0.5 cm diameter, densely set, 0.7–0.8 cm apart, with whitish wool when juvenile, later glabrescent; spines: prominent, acicular, straight, pungent; radial spines 11–15–(20), 0.4–0.9 × 0.03–0.04 cm; reddish to grey (when older); central spines 1–2, 1.0–1.5 × 0.07–0.10 cm dark redish to grey (generally when older), easly distinguished from the radials, longer and thicker; flowering areoles producing longer and thicker spines, up to 2 cm, like observed in wild; flower buds with a dense cover of white to pale brown woll and bristles; flowers subapical, diurnal, opening along several days, flowering 8 hours after the sunrise and closing circa four hours later, self-fertile; flowers infundibuliform, to 4.0– 4.5 cm, 5–6 cm in diameter (Fig. 3); floral tube to ca. 1 cm; internal walls pale yellow and external walls yellowish green, with triangular bract scales subtending areoles that bear white to pale brownish hairs and long reddish brown bristles to 1.5–2.0 cm; tepals pale-yellow, perianth multiseriate, segments pale-yellow oblanceolate; stamens numerous, spreading, arranged along the inner walls of the floral tube and around the style; filaments with 1.6–1.9 cm, style pale - yellow, ca. 1.6–2.0 cm; stigma lobes exserted, 9–10, light-purple; fruit 2.0–2.5 × 1.0–2.0 cm, green to dark purple, pericarpel with areoles bearing long brown hairs and bristles, dehiscing at the base and later falling off, pulp white; dry perianth remnants either persistent on the fruit or falling off; seeds broadly oval, 0.9–1.2 mm broad, 1.1–1.4 mm long, 1.1–1.3 mm high, dark brown, matt, border little expanded around hilum, testacells uniform, elongated, antilclinal boundaries cannelled, cells on the bottom black, on top light brown, hilum basal, superficial, hilum-microphylar region oval, microphyle conjunct (Fig. 4). Etymology:— The specific epithet honors to Andreas Hofacker, researcher of Cactaceae of Brazil. Phenology:— Flowering season is September–October, whereas mature fruits can be found in November– December. The flowers, in cultivation, open only on very hot days, around 11 a.m., closing about 4 hours after the beginning of the anthesis. Distribution and habitat:— The species was found in three isolated locations in the area between the municipalities of Alegrete, Rosário do Sul and Santana do Livramento (Fig. 5), possibly endemic to the southwestern region of Rio Grande do Sul, Brazil. Parodia hofackeriana grows on rocky outcrops of volcanic origin (basalt) on hills, among fragments of weathered rock, with the presence of humus and organic matter. It is sympatric with Frailea perumbilicata F. Ritter (1970:125) and Gymnocalycium uruguayense (Arechavaletae 1905: 218) Britton & Rose (1922: 172). Nearby, F. pumila (Lemaire 1838: 21) Britton & Rose (1922: 209), Parodia glaucina, P. mammulosa, P. mueller-melchersii subsp. mueller-melchersii, P. orthacantha, P. tenuicylindrica, and P. erinacea are growing, in the same environmental conditions. Conservation status:— Firstly, Parodia hofackeriana was located by the authors in 2015. With complementary observations, carried out during 2019, 2020, 2021 and 2022, it was concluded that the taxon is an independent species. No morphological deviations were observed in its population, showing that it is a species with peculiar characteristics. So far, it is recognized for three localities, ca. 80 km apart, found in a rocky region, between fragments of basaltic rock with grasses. This area is inserted in the Pampa, a biome internationally known for biotic endemism (Andrade et al. 2023). In the area, pastures with cattle and sheep are frequent, which can be potential threats to the survival of these populations. With these findings, it is concluded that this new taxon is endangered in their locations. Following the criteria of IUCN (2022), we propose that Parodia hofackeriana has to be considered Critically Endangered (CR), based on geographic distribution, occupation area, and population size [CR B1+B2b (i,ii,v) + c (iv) +C2a(i),b]. Taxonomical notes: — Parodia hofackeriana is closely related to the P. mueller-melchersii subsp. mueller-melchersii due to the presence of tubercular ribs, larger pale-yellow flowers, and hemispherical berry-like fruits. P. hofackeriana is distinguished from P. mueller-melchersii by the number and shape of ribs (18–20 vs. 20–25); shape of central spines (hemispheric vs. flat); color of central spines (reddish-brown to grey vs. whitish-horn-colored) and color of radial spines (reddish-brown to grey-brown vs. yellowish white). From Parodia mammulosa, it is distinguished by the greater number of ribs (18–20 vs. 14–20), shape of central spines (hemispheric vs. flat); color of central spines (reddish-brown to grey vs. whitish-horn colored, brown-red to purplish-brown, often lighter at the tip); shape of radial spines; color of radial spines (reddish-brown to grey-brown vs. yellowish to reddish) brown. It should be noted that the species in question differs from related species also by flowering season, being earlier than P. mammulosa and P. mueller-melchersii subsp. mueller-melchersii. It blooms at the beginning of spring (September–October) bearing fruits in November–December. The Table 1 includes additional differences between P. hofackeriana and its related species. P. mammulosa and P. mueller-melchersii subsp. mueller-melchersii were also found in the immediate vicinity of the type location (circa 1 km away). No transitions and hybrids were observed and all taxa are clearly distinguishable.An important feature for differentiating individual species is the issue of hybridization of taxa that are sympatric or occur close together. If hybrids are not found, they can be considered independent taxa (Hunt et al. 2006: 4). Thus, P. hofackeriana should be recognized as a separate taxon. Investigations were also carried out in nearby locations, with the aim of finding new habitats for P. hofackeriana, but such actions were unsuccessful. Due to accessibility, sometimes difficult, the possibility of the existence of the species in other areas not yet investigated can not be ruled out. Morphological differences (Tab. 1 and Fig. 6) and proximity to populations of Parodia mammulosa and P. mueller-melchersii subsp. mueller-melchersii make it necessary to recognize the taxon as an independent species. Additional materials examined:— Parodia mammulosa. BRAZIL. Rio Grande do Sul: Quaraí, BR 293, close to Sarandi II, 15.XI.1016, Oliveira & Deble ex. cult. 127az (PACA); Santana do Livramento, Cerros Verdes, 02 October 2018, Oliveira & Deble ex. cult. 128az (PACA). Parodia mueller-melchersii. BRAZIL. Rio Grande do Sul: Alegrete, Caverá, 31 March 2018, Oliveira & Deble ex cult. 70b (PACA); Bagé, Passo do Cação, out 2018, Oliveira & Deble ex cult. 96b (PACA); Quaraí, Cerro Chato, 22 March 2017, Oliveira & Deble ex cult. 56b (PACA); between Sarandi II e Sarandi III, 12 November 2017, Oliveira & Deble ex cult. 51b e 52b (PACA). URUGUAY. Artigas, Masoller, east of Cuchilla de Belén, 21 March 2017, Oliveira & Deble ex. cult. 82b (PACA); Tacuarembó, Valle Eden, 12 October 2016, Oliveira & Deble ex cult. 100 (PACA)., Published as part of Oliveira, Anabela Silveira De & Pontes, Rodrigo Corrêa, 2023, Parodia hofackeriana (Cactaceae-Notocacteae), a new species from southwest of Rio Grande do Sul, Brazil, pp. 283-292 in Phytotaxa 598 (4) on pages 284-290, DOI: 10.11646/phytotaxa.598.4.2, http://zenodo.org/record/7983774, {"references":["Ritter, F. (1970) Nieuwe cactussen uit Zuid-Amerika I. Succulenta 49 (7): 108 - 109.","Britton, N. L. & Rose, J. N. (1922) The Cactaceae. Descriptions and illustratiopns of plants of the cactus familiy, vol. III. Carnegie Institution, Washington, 255 pp.","Lemaire, C. (1838) Echinocactus pumilus. Cactearum aliqot nomarum ac insetuarium in horto Monvilliana cultarum accurata desciptio. F. G. Levrault, Lutetiae-Parisiorum, 44 pp.","Andrade, B. O., Dr ˆ se, W., Aguiar, C. A., Aires, E. T., Alvares, D. J., Barbieri, R. L., Carvalho, C. J. B., Bartz, M., Becker, F. G., Bencke, G. A., Beneduzi, A., Silva, J. B., Blochtein, B., Boldrini, I. I., Boll, P. K., Bordin, J., Silveira, R. M. B., Martins, M. B., Bosenbecker, C., Braccini, J., Braun, B., Brito, R., Brown, G. G., B ¸ neker, H. M., Buzatto, C. R., Cavalleri, A., Cechin, S. Z., Colombo, P., Constantino, R., Costa, C. F., Dalzochio, M. S., Oliveira, M. G., Dias, R. A., Santos, L. A., Duarte, A. F., Duarte, J. L. P., Durigon, J., da Silva, M. E., Ferreira, P. P. A., Ferreira, T., Ferrer, J., Ferro, V. G., Fontana, C. S., Freire, M. D., Freitas, T. R. O., Galiano, D., Garcia, M., dos Santos, T. G., Heydrich, I., Iop, S., Jarenkow, J. A., Jungbluth, P., Kaffer, M. I., Kaminski, L. A., Kenne, D. C., Kirst, F. D., Krolow, T. K., Kr ¸ ger, R. F., Kubiak, B. B., Leal-Zanchet, A. M., Loebmann, D., Lucas, D. B., Lucas, E. M., Luza, A. L., Machado, I. F., Madalozzo, B., Maestri, R., Malabarba, L. R., Maneyro, R., Marinho, M. A. T., Marques, R., Marta, K. S., Martins, D. S., Martins, G. S., Martins, T. R., Mello, A. S., Mello, R. L., Mendonca Junior, M. S., Morais, A. B. B., Moreira, F. F. F., Moreira, L. F. B., Moura, L. A., Nervo, M. H., Ott, R., Paludo, P., Passaglia, L. M. P., Perico, E., Petzhold, E. S., Pires, M. M., Poppe, J. L., Quintela, F. M., Raguse-Quadros, M., Pereira, M. J. R., Renner, S., Ribeiro, F. B., Ribeiro, J. R. I., Romanowski, H. P., Ruschel, T. P., Saccol, S. S. A., Savaris, M., Schmitz, H. J., Silva Filho, P. J. S., Soares, A. G., Somavilla, A., Sperotto, P., Spies, M. R., Tirelli, F. P., Tozetti, A. M., Verrastro, L., Vogel Ely, C., da Silva, A. Z., Zank, C., Zefa, E. & Overbeck, G. E. (2023) 12,500 + and counting: biodiversity of the Brazilian Pampa. Frontiers of Biogeography. https: // doi. org / 10.21425 / F 5 FBG 59288","IUCN (2022) The IUCN Red List of Threatened Species. Version 2022 - 2. Available from: https: // www. iucnredlist. org (accessed 28 March 2023).","Hunt, D. R., Taylor, N. P. & Charles, G. (eds.) (2006) The new cactus lexicon. dh Books, Milborne Port, 373 pp."]}

Key to the species of Deamia 1. Stems 0.6–2 cm in diameter, the ribs (6–)7 or 8, acute or rounded; flowers longistylous.................................................................2 - Stems 2.5–15 cm in diameter, the ribs 3–10, acute; flowers longistylous or brevistylous.................................................................3 2. Spines up to 1.4 cm long; stem ribs rounded; flowers 8–10 cm long............................................................................ Deamia funis - Spines up to 2.6 cm long; stem ribs acute; flowers (23–) 27–30 cm long........................................................... Deamia montalvoae 3. Flowers 6–9 cm long, brevistylous with the stamens 0.4–0.5 cm long and the style 2–2.1 cm long, the stigma positioned well below the level of anthers; stems 2.5–3.5 cm in diameter............................................................................................ Deamia chontalensis - Flowers 22–28 cm long, longistylous with the stamens 9–20 cm long and the style 22–24 cm long, the stigma positioned among or just beyond the anthers; stems 3–15 cm in diameter.............................................................................................. Deamia testudo

Deamia funis Hammel & S. Arias sp. nov. (Figs 1, 2) Type:— NICARAGUA. Matagalpa: 9.1 km N of Matiguás (highway) along road through Tierra Blanca, 2.9 km N of Tierra Blanca, deciduous forest, rocky and heavily grazed, 12°53’53”N, 85°30’15”W, 423 m, 15 March 2017 (fl), Stevens & Montiel 39242 (holotype, MO!; isotype HULE). Diagnosis:— In its flowers Deamia funis is outwardly similar to D. chontalensis, but has stamens (1.5–3 vs. 0.4–0.7 cm long) and styles (6–8 vs. 2–2.2 cm long) much longer, and with the stigma exerted among or slightly beyond the anthers (vs. the brevistylous flowers of the latter), furthermore, its stems are less than half the diameter of those of the latter, and with more ribs; the stems are similar to those of D. montalvoae, but the flowers are smaller, less than 1/2 (mostly 1/3) as long as in that species. Description:—Root-climbing plants or trailing epiphytes, with straight spines; stems ± straight and usually pendulous, irregularly branched, the internodes from ca. 0.12 up to ca. 10 m long, 0.6–1.1 cm in diameter, ± cylindrical with ca. (6–)8 relatively rounded ribs, with a brittle, transparent, varnish like epidermal coating and abundant adventitious roots, monomorphic; areoles 1–1.5 cm apart, with 4–18 orange to black, persistent spines, to ca. 1.4 cm long, the trichome-spines lacking or a few persistent. Leaves scalelike, to ca. 0.04 cm long, deltate, acute, soon deciduous. Flowers mostly restricted to the distal, new growth of pendulous stems, nocturnal, 8–10 cm long, actinomorphic, funnel-shaped, the pericarpel ca. 1 cm long, with ca. 8 series of indistinct pustules (podaria) bearing the areoles with the bracteole dark reddish brown, ca. 0.5 mm long, deltate, with dense and conspicuous felt and with few or no spines but abundant trichome-spines (hairs); receptacular tube pale green to greenish yellow, sometimes tinged pink, 5–7 cm long, funnel shaped (the narrow portion of the tube up to ca. 3 × 0.7 cm, the wider portion up to ca. 3.5 × 3 cm), with numerous scattered bracteoles (sometimes intergrading with or positioned upon the outer tepals), up to ca. 2 mm long, the areoles sometimes with a few spines (in the proximal 1/3 to 1/2 of the tube) and with abundant, white to orange trichome-spines to ca. 1.5 cm long; perianth with numerous, slightly differentiated tepals, up to 1.5–2 cm long, the outer tepals pale yellow green to white, sometimes apically pale brown, ascending, ± oblong, obtuse to acute, the inner tepals white, ascending, oblong, acute; stamens numerous, 1.5–3 cm long, attached in two bands, one in the distal 1/3 of the narrow portion of the tube, to ca. 1.5 cm wide, the other uniseriate and born at the mouth of the tube (at its widest point), the filaments white, included (within the cup of the limb), the anthers 1–1.4 × ca. 0.8 mm; style to ca. 6–8 cm long; stigma white to cream, with ca. 8 simple lobes. Submature fruit yellowish green, ca. 2 × 2.5 cm, ovoid, smooth, the areoles with ca. 20 spines, to ca. 0.7 cm long, not completely covering the fruit surface; pulp and seeds not seen. Additional specimens examined:— NICARAGUA. Chontales: 2 km NE of Cuapa along road to Cerro Oluma, dry forest patch on hillside, 12°16’51”N, 85°22’27”W, 355 m, 20 January 2009 (fl.), Stevens & Montiel 27909 (HULE, MO). Matagalpa: 11.9 km N of Esquipulas (at Puente Kaulapa) along road to San Dionysio; dry forest, 12°44’23”N, 85°47’16”W, 322 m, 3 April 2018 (fl., immature fruit.), Stevens 40324 (HULE, MO); ibd., 9.1 km N of Matiguás (highway) along road through Tierra Blanca, 2.9 km N of Tierra Blanca, deciduous forest, rocky and heavily grazed, 12°53’53”N, 85°30’15”W, 423 m, 4 January 2013 (fl.), Stevens & Montiel 33938 (HULE, MO); ibd., 25 February 2022 (flower bud, immature fruits), Stevens & Calero 44070 (HULE, MO). Distribution and habitat:— Deamia funis is known only from low to mid elevations (322–423 m) of fog-zones of dry, seasonal forests on the Atlantic watershed, from central Nicaragua. Although presently endemic to Nicaragua, the species should be sought in similar habitat just to the north, in Honduras. Deamia testudo, the only other species in the genus known from Nicaragua is more widespread, and is known to occur sympatrically, at least as close as ca. 25 km (Fig. 3). Several sterile collections from limestone outcrops in a mixture of moist forest and savana of the Atlántico Norte region (northeastern Nicaragua), may belong to this species, but lacking fertile collections from that area, we have not included them. Phenology:—Flowering plants have been found only in the months of January to April, during the dry season. Etymology:—The epithet refers to the long, dangling, rope-like stems common in this species when found growing on horizontal branches of trees. Preliminary conservation status:—The new species is not known for sure to occur within any protected area, but the type and two other collections were found within a few km of the Reserva Natural Sierra Quirragua and one other collection from very near the Reserva Natural Sierra Amerrisque. The species has been collected from only three different localities and we estimate that a polygon encompassing all three of them covers an area of no more than 5000 km 2. We consider that the area where this plant is known to occur, being subject to cattle farming, suffers from continuing habitat destruction. Thus, by the guidelines for using the IUCN Red List categories and criteria (2022), table 2.1, B1 a and b, this species ought to be considered endangered (EN). In any case, all species of plants everywhere, except common weeds and cultivated plants should be considered endangered. Human caused habitat destruction and climate change are not making things any better. Discussion:—By its stems 0.6–1.1 cm in diameter with ca. (6–)8 ribs (vs. stems 1–2 cm in diameter with 7 or 8 ribs) this new species overlaps somewhat with those of Deamia montalvoae, but in floral characters it differs markedly from that species by its much shorter flowers [8–10 vs. (23–) 27–30 cm long in D. montalvoae]. However, in size and overall form the funnel-shaped flowers of the present species are almost identical to those of D. chontalensis, which however, has stems 2.5–3.5 cm in diameter with only 5 or 6(–7) acute ribs. The fruits of D. funis differ from both latter two species by the surface being hardly at all covered by the areole spines, and in that respect are more similar to the fruits of D. testudo. The flowers of Deamia chontalensis itself are unique among species of the genus, and at least uncommon among all Cactaceae.An illustration showing the three main floral types in Cactaceae in the review by Mandujano et al. (2010) shows only styles with the stigma exerted among or slightly beyond the anthers (i.e., longistylous). They mention only one study of one species (Opuntia robusta H.L. Wendl. ex Pfeiffer) with a brevistylous morph. The flowers of D. chontalensis, however, were originally described as unusual for having an “extremely short style” (Alexander, 1950), and were illustrated (Bravo-Hollis 1978) with the stigma as being positioned well below the level of the anthers, in fact, below the insertion of the short stamens (i.e., brevistylous). Furthermore, the protologue includes a curious note attributed to the original collector “Mr. MacDougall notes that the style extends 7 mm beyond the stamens at anthesis but is back in place below the stamens when the flower closes.” This juxtaposition, so contrary to the usual—where the stigma is exerted slightly beyond or among the anthers—could appear to be some sort of error in the description (the original collection having been lost and the lectotype, a photo that reveals nothing in this respect). Nonetheless, the short style (2–2.2 cm long) with the stigma positioned well below the anthers, has been confirmed with recent collections of D. chontalensis (Fig. 4). No species of Deamia with more than one flower morph is known. The suggestion that the style stretches out and then shrinks back has not been confirmed. The stamens of D. chontalensis are also unusually short for a flower of this size, ranging from only 0.4 to 0.5 cm long; stamen length in the other species of Deamia varies from 1.5–20 cm long. These characters in the floral morphology of D. chontalensis are strongly suggestive of a particular pollination syndrome worthy of investigation. Thus, although the flowers of D. chontalensis and D. funis are outwardly similar, the latter is further distinguished by its longer style (6–8 cm long) and stamens (1.5–3 cm long), with the stigma among or slightly exceeding the anthers. The submature fruits are similar to those of other species in the genus, in having a smooth surface with spiny areoles, the spines (as in the fruits of Deamia testudo) not completely covering the surface; we are confident that all characters of the new species, especially that of the unique epidermal wax of the stems, often with a sort of cob-web pattern of cracking, are congruent with its placement in Deamia. To facilitate identification of the four known species of Deamia we include a key., Published as part of Hammel, Barry & Arias, Salvador, 2022, A new species of Deamia (Cactaceae) from Nicaragua, pp. 220-226 in Phytotaxa 576 (2) on pages 221-225, DOI: 10.11646/phytotaxa.576.2.8, http://zenodo.org/record/7461375, {"references":["Mandujano, M. d. C., Carrillo-Angeles, I., Martinez-Peralta, C. & Golubov, J. (2010) Reproductive biology of Cactaceae. In: Ramawat, K. (Ed.) Desert plants: biology and biotechnology. Springer, Berlin, Heidelberg, pp. 197 - 230. https: // doi. org / 10.1007 / 978 - 3 - 642 - 02550 - 1 _ 10","Alexander, E. J. (1950) A new Nyctocereus from southern Mexico. Cactus and Succulent Journal (Los Angeles) 22: 131 - 133.","Bravo-Hollis, H. (1978) Las cactaceas de Mexico. 2 nd ed., vol. 1. Universidad Nacional Autonoma de Mexico, Mexico City, 743 pp."]}

Deamia funis sp. nov. (Cactaceae) from Nicaragua, in the Mesoamerican region, is described and illustrated based on morphological data. The new species is characterized by its often long, dangling (ropey), and relatively narrow, multiribbed stems with relatively small flowers like those of D. chontalensis, but with the stigma exerted among or slightly beyond the anthers as occurs in D. montalvoae and D. testudo.

Key to the species of Deamia 1. Stems 0.6–2 cm in diameter, the ribs (6–)7 or 8, acute or rounded; flowers longistylous.................................................................2 - Stems 2.5–15 cm in diameter, the ribs 3–10, acute; flowers longistylous or brevistylous.................................................................3 2. Spines up to 1.4 cm long; stem ribs rounded; flowers 8–10 cm long............................................................................ Deamia funis - Spines up to 2.6 cm long; stem ribs acute; flowers (23–) 27–30 cm long........................................................... Deamia montalvoae 3. Flowers 6–9 cm long, brevistylous with the stamens 0.4–0.5 cm long and the style 2–2.1 cm long, the stigma positioned well below the level of anthers; stems 2.5–3.5 cm in diameter............................................................................................ Deamia chontalensis - Flowers 22–28 cm long, longistylous with the stamens 9–20 cm long and the style 22–24 cm long, the stigma positioned among or just beyond the anthers; stems 3–15 cm in diameter.............................................................................................. Deamia testudo, Published as part of Hammel, Barry & Arias, Salvador, 2022, A new species of Deamia (Cactaceae) from Nicaragua, pp. 220-226 in Phytotaxa 576 (2) on pages 225-226, DOI: 10.11646/phytotaxa.576.2.8, http://zenodo.org/record/7461375

A detailed morphological and molecular study of the four Opuntia species belonging to the ser. Streptacanthae is carried out. No published phylogenetic study of Opuntia includes these four species. Our data support the recognition of the four taxa as separated species but related by the characteristic of having a glabrous epidermis, yellow flowers, red and sweet fruits, in the present study it is recovered forming a monophyletic group within Opuntia s.str.

Opuntia lasiacantha Pfeiffer (1837: 160) Neotype (designated by Scheinvar et al. 2010: 280): — MEXICO. Guanajuato, San Luis de la Paz, 25 May 1999, Reyes-Agüero & Carlín 1857 (MEXU!). Description:— Arborescent habit, 2.0–3.0 m high (Fig. 7A). Defined trunk of 50 cm in diameter, slightly grayish scaly bark with light brown tones (Fig. 7B). Glabrous epidermis (Fig. 10C). Oblanceolate to obovate cladodes, 45–62 cm long x 24–29 cm wide, green in color, covered with a layer of wax (Fig. 8A). Areolas arranged in 11–12 series, 3.8 cm spacing between series, 3.1 cm spacing between elliptical areolas, 0.4 × 0.3 cm, short black trichomes (Fig. 8B). Short yellow glochids (0.2 cm), arranged at the top of the areola. 1–4 yellowish white, rigid, cylindrical spines, some twisted, erect, and diffuse, unequal 0.4–3.3 cm long (Fig. 8C) with yellow apex. Juvenile cladodes with very prominent tubers, areolas with short yellowish trichomes, subulate, erect, green leaves with slightly uncinate reddish apex (Fig. 7C). Acute flower buds, reddish perianth segments with acuminate apex, slightly cylindrical pericarpel, very marked and elevated tubers, areolas with short brown to black trichomes, yellow glochids and areolas with generally yellow bristles and small brown shades (Fig. 8D and 8E). Yellow flowers (Fig. 8F), 9–10.5 cm long, cylindrical pericarp, ca. 6.0 × 2.5 cm, areolas arranged in 6–7 series, 0.42 cm spacing between them, with crass basal scale, outer obovate segments of perianth, acuminate apex, smooth yellow edges, with medium reddish striation, reddish apex, inner obovate segments, emarginated apex, upper yellow edges generally smooth, stamens 1/3 of the length of the perianth, white filaments, white-yellowish anthers, slightly cylindrical style with white with pink tones, ca. 2 cm long, usually eleven green papillary lobes of the stigma, ca. 0.6 cm long (Fig. 9B). Pyriform to elliptical red fruits (Figure 9C), 8.0– 9.3 cm long, slightly sunken floral striated scar, large areolas with short, black, semicircular to circular trichomes (Fig. 9E), arranged in 6–7 series, 1.2 cm spacing from each other and 1.4 cm between series, absent spines, yellow glochids, orange-red juicy and sweet funicles (Fig. 9D). Kidney shaped seed with irregular lateral aril, lateral thread-micropillar region, microphile and funiculus included (Fig. 9F). Micromorphology:— Thornswith lignified epidermal cells 12.72 ± 1.4 × 3.95 ± 2.3 μm in the distal part (Fig. 10B), 75.11 ± 1.10 × 8.89 ± 1.7 μm, continuous epidermal cells and rough texture, in the middle part (Fig. 10A). Glabrous epidermis, irregular epidermal cells (Fig. 10C) and paracytic stomas (Fig. 10D). Polyhedron-shaped pollen grain, pantoaperturate (Fig. 10E), 105.10 ± 1.50 μm in diameter, 8327.73 μm ² in area, 15.30 ± 1.1 μm distance between each distal opening (pores), with a diameter of 20.15 ± 1.50 μm, 16–18 pores with walls of 2.46 ± 0.04 μm width and wart (Fig. 10F). Vernacular names:— Nopal de tuna colorada, Nopal de espinas lacias and Nopal de cerro (see e.g., Anderson 2001, Scheinvar 2004). Phenology:— Opuntia lasiacantha blooms from March–April, whereas fruiting time if June–July. Habitat and distribution:— Endemic to Mexico, it is known in the states of Aguascalientes, Chiapas, Ciudad de México, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México, Michoacán, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tamaulipas, Tlaxcala, Veracruz and Zacatecas (see e.g., Arias et al. 1997, Guzmán et al. 2003, Scheinvar 2004, Scheinvar et al. 2010, Arias et al. 2012, Hernández et al. 2014) (Fig. 2). It grows in xerophytic scrub (Rzedowski 1978)., Published as part of Martínez-González, César Ramiro, Gallegos-Vázquez, Clemente, Mascorro- Gallardo, José O. & Barrientos-Priego, Alejandro F., 2022, Molecular and morphological notes on Opuntia ser. Streptacanthae (Cactaceae), pp. 1-28 in Phytotaxa 576 (1) on pages 9-12, DOI: 10.11646/phytotaxa.576.1.1, http://zenodo.org/record/7440876, {"references":["Pfeiffer, L. G. K. (1837) Enumeratio diagnostica Cactearum Hucusque Cognitarum. Sumtibus Ludovici Oehmigke, Berlin, 192 pp. https: // doi. org / 10.5962 / bhl. title. 147375","Scheinvar, L., Gallegos-Vazquez, C., Olalde-Parra, G. & Rodriguez-Fuentes, A. (2010) Opuntia ser, Streptacanthae (Cactaceae) - neotypifications and taxonomic notes for four species. Schumannia 6: 277 - 296.","Anderson, E. F. (2001) The Cactus family. Timber Press, 776 pp.","Scheinvar, L. (2004) Flora cactologica del estado de Queretaro. Diversidad y riqueza. Fondo de Cultura Economica, Mexico, 390 pp.","Arias, S., Gama S. & Guzman, U. (1997) Cactaceae. Flora del Valle de Tehuacan-Cuicatlan, Fasc. 14. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico City, 146 pp.","Guzman, U. L., Arias, S. & Davila, P. (2003) Catalogo de cactaceas mexicanas. Universidad Nacional Autonoma de Mexico. Mexico City, 315 pp.","Arias, S., Gama, S., Guzman, U. & Vazquez, B. (2012) Cactaceae. Flora del Valle de Tehuacan-Cuicatlan, Fasc. 95. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico City, 235 pp.","Hernandez, H. M., Gomez, H. C., Barcenas, R. T., Puente, P. & Reyes, A. J. (2014) A checklist of the subfamily Opuntioideae (Cactaceae) from North and Central America. In: Hunt, D. R. (Ed.) Further Studies in Opuntioideae (Cactaceae). Succulent Plant Research 8: 185 - 200.","Rzedowski, J. (1978) Vegetacion de Mexico. Limusa, Mexico City, 432 pp."]}

Opuntia hyptiacantha F.A.C. Weber (1898: 894) Neotype (designated by Scheinvar et al. 2010: 287): — MEXICO. San Luis Potosí, 13 Km northwest of San Luis Potosí, 01 May 1973, Sánchez-Mejorada 1734 (MEXU!). Description:— Plant 2.0– 2.5 m high tree (Fig. 15A). Defined trunk, 35 cm in diameter, slightly grayish scaly bark with light brown tones (Fig. 15B). Glabrous epidermis (Fig. 18C). Elliptical to oval cladodes, 32–40 cm long x 14–24 cm wide and 2–3 cm thick, green in color, covered with a light layer of wax (Fig. 16A). Areolas arranged in 12–13 series, 3.6 cm spacing between series, 3.4 cm spacing between elliptical areolas, 0.4 × 0.3 cm, short black trichomes, (Fig. 16B). Very small and scarce yellow glochids, arranged at the top of the areola. 0–9 white spines, yellowish apex, rigid, generally erect and diffuse, some of which are generally adpressed at the edge, unequal in length, 0.3–2.1 cm, the lower part of the cladode has no spines (Fig. 16C). Juvenile cladodes with prominent tubers, areolas with short brown to black trichomes, some reddish white bristle hairs, erect subulate green leaves, with reddish apex (Fig. 15C). Acute flower buds, reddish perianth segments with accumulated apex, elliptical pericarpel, slightly elevated tubers, areolas with short brown trichomes, brown glochids and areolas with brown bristles (Fig. 16D and 16E). Yellow flowers (Fig. 16F), 6.0–7.0 cm long, elliptical pericarp, ca. 3.4 × 2.3 cm, areolas arranged in 5–6 series, 0.66 cm spacing between them, with crass basal scale, outer segments of the perianth with obovate form, mucroned apex, smooth yellow edges, with medium greenish striation, reddish apex, oblanceolated inner segments, emarginated apex, yellow with reddish apical part, upper edges slightly dentate, stamens 1/3 of the length of the perianth, reddish and yellow filaments, yellow anthers, cuneiform style, ca. 1.9 cm long, light green, usually eight green papillary lobes of stigma, ca. 0.5 cm long, with medium white pink striation (Fig. 17B). Red elliptical fruits (Fig. 17C), 5.0– 6.5 cm long, sunken floral striated scar, large and circular to elliptical areolas, (Fig. 17E), arranged in 4–5 series, 1 cm spacing from each other and 0.7 cm between series, long black trichomes, some spines, short yellow glochids, juicy and sweet pink red funicles (Fig. 17D). Lenticular seed with irregular lateral aril, lateral trhead-micropilar region, micropilum and funiculus included (Fig. 17F). Micromorphology:— Thornswith lignified epidermal cells 98.35 ± 1.40 × 23.64 ± 1.10 μm in the distal part (Fig. 18B), 61.22 ± 1.30 × 12.04 ± 1.40 μm, continuous epidermal cells and rough texture, in the middle part (Fig. 18A). Glabrous epidermis, irregular epidermal cells (Fig. 18C) and paracytic stomas (Fig. 18D). Polyhedron-shaped pollen grain hexagonal, pantoaperturate (Fig. 18E), 111.16 ± 1.40 μm in diameter, 8432.28 μm ² in area, 10.10 ± 1.90 μm distance between each distal opening (pores), with a diameter of 22.50 ± 1.50 μm, 15–17 pores with walls of 1.85 ± 0.14 μm width and wart (Fig. 18F). Vernacular names:— Nopal cascarón, tuna mansa, tuna corriente and Tuna chaveña (see e.g., Bravo-Hollis 1978, Anderson 2001, Scheinvar 2004). Phenology:— Opuntia hyptiacantha blooms from March–April, whereas fruiting time if June–July. Habitat and distribution:— Endemic to Mexico, it is known in the states of Aguascalientes, Ciudad de México, Guanajuato, Hidalgo, Jalisco, México, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tlaxcala and Zacatecas (see e.g., Arias et al. 1997, Guzmán et al. 2003, Scheinvar 2004, Scheinvar et al. 2010, Arias et al. 2012, Hernández et al. 2014) (Fig. 2). It grows in xerophytic scrub (Rzedowski 1978)., Published as part of Martínez-González, César Ramiro, Gallegos-Vázquez, Clemente, Mascorro- Gallardo, José O. & Barrientos-Priego, Alejandro F., 2022, Molecular and morphological notes on Opuntia ser. Streptacanthae (Cactaceae), pp. 1-28 in Phytotaxa 576 (1) on pages 20-23, DOI: 10.11646/phytotaxa.576.1.1, http://zenodo.org/record/7440876, {"references":["Weber, F. A. C. (1898) Opuntia Tournef. In: Bois, D. (Ed.) Dictionnaire d´Horticulture. P. Klincksieck, Paris, pp. 892 - 899.","Scheinvar, L., Gallegos-Vazquez, C., Olalde-Parra, G. & Rodriguez-Fuentes, A. (2010) Opuntia ser, Streptacanthae (Cactaceae) - neotypifications and taxonomic notes for four species. Schumannia 6: 277 - 296.","Bravo-Hollis, H. (1978) Las Cactaceas de Mexico, vol. 1. Universidad Nacional Autonoma de Mexico, Mexico City, 351 pp.","Anderson, E. F. (2001) The Cactus family. Timber Press, 776 pp.","Scheinvar, L. (2004) Flora cactologica del estado de Queretaro. Diversidad y riqueza. Fondo de Cultura Economica, Mexico, 390 pp.","Arias, S., Gama S. & Guzman, U. (1997) Cactaceae. Flora del Valle de Tehuacan-Cuicatlan, Fasc. 14. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico City, 146 pp.","Guzman, U. L., Arias, S. & Davila, P. (2003) Catalogo de cactaceas mexicanas. Universidad Nacional Autonoma de Mexico. Mexico City, 315 pp.","Arias, S., Gama, S., Guzman, U. & Vazquez, B. (2012) Cactaceae. Flora del Valle de Tehuacan-Cuicatlan, Fasc. 95. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico City, 235 pp.","Hernandez, H. M., Gomez, H. C., Barcenas, R. T., Puente, P. & Reyes, A. J. (2014) A checklist of the subfamily Opuntioideae (Cactaceae) from North and Central America. In: Hunt, D. R. (Ed.) Further Studies in Opuntioideae (Cactaceae). Succulent Plant Research 8: 185 - 200.","Rzedowski, J. (1978) Vegetacion de Mexico. Limusa, Mexico City, 432 pp."]}

Opuntia streptacantha Lemaire (1839: 62) Neotype (designated by Scheinvar et al. 2010: 283): — MEXICO. Hidalgo, Zempoala, east of Téllez basis of El Cerrito, 23 March 1973, Scheinvar & Ahuatzin 2024 (MEXU!). = Opuntia streptacantha subsp. aguirriana Scheinvar & Rodríguez (2003: 304). Description:— Plant 2.0–3.0 m high tree (Fig. 11A). Defined trunk of 60 cm in diameter, grayish scaly bark with black shades (Fig. 11B). Glabrous epidermis (Fig. 14C). Oval cladodes, 32–40 cm long × 23–35 cm wide and 2–3 cm thick, green in color, covered with a very light layer of wax (Fig. 12A). Areolas arranged in 10–11 series, 3 cm spacing between series, 2.1 cm spaing between elliptical areolas of 0.4 × 0.3 cm, short black trichomes (Fig. 12B). Very small and scarce yellow gloquids, arranged at the top of the areola. 0–8 white spines with gray and black tones, flexible, acicular, some slightly twisted, adpressed and diffuse, very few erect, unequal of 0.2–2.2 cm long, yellowish apex, in the lower part of the cladode some areolas do not have spines (Fig. 12C). Juvenile cladodes with prominent tubers, areolas with short brown to black trichomes, some white bristel hairs with reddish tones, subulate, erect, green leaves, with uncinate slightly reddish apex, white spines with reddish apex (Fig. 11C). Acute flower buds, green perianth segments with reddish acuminate apex, elliptic pericarpel, slightly raised tubers, areolas with short black trichomes, brown glochids and brown bristle areolas (Fig. 12D and 12E). Yellow flowers (Fig. 12F), 5.5–6.1 cm long, elliptical pericarp, ca. 3.2 × 2.0 cm, areolas arranged in 4–5 series, 0.62 cm spacing between them, outer segments of perianth with spatulate shape, rounded apex, smooth yellow edges, with medium greenish striation, reddish apex, oblanceolated inner segments, emarginated apex, yellow, smooth upper edges, stamens 1/3 of the length of the perianth, yellow filaments, yellow anthers, white, cuneiform style, ca. 2.3 cm long, usually nine green papillary lobes of stigma, ca. 0.4 cm long, with medium white striation (Fig. 13B). Red elliptical to obovate fruits (Fig. 13C), 4–6 cm long, slightly sunken striated floral scar, circular to elliptical large areolas (Fig. 13E), arranged in 6–7 series, 1.1 cm spacing from each other and 1 cm between series, long black trichomes, some spines, yellow glochids, red juicy and sweet funicles (Fig. 13D). Discoid seed with irregular lateral aril, lateral thread-micropillar region, microphile and funiculus included (Fig. 13F). Micromorphology:— Thornswith lignified epidermal cells 13.38 ± 1.90 × 3.59 ± 1.3 μm in the distal part (Fig. 14B), 114.01 ± 1.10 × 12.79 ± 1.20 μm, non-continuous epidermal cells with fissures between cells and smooth texture, in the middle part (Fig. 14A). Glabrous epidermis, irregular epidermal cells (Fig. 14C) and cyclocytic stomas (Fig. 14D). Polyhedron-shaped pollen grain, pantoaperturate (Fig. 14E), 114.64 ± 1.90 μm in diameter, 9219.1 μm ² in area, 13.89 ± 2.10 μm distance between each distal opening (pores), with a diameter of 21.49 ± 1.50 μm, 22–24 pores with walls of 2.43 ± 0.04 μm width and wart (Fig. 14F). Vernacular names:— Nopal cardón, Tuna cardona and Nopal hartón (see e.g., Bravo-Hollis 1978, Scheinvar 2004). Phenology:— Opuntia streptacantha blooms from March–April, whereas fruiting time if June–July. Habitat and distribution:— Endemic to Mexico, it is known in the states of Aguascalientes, Chihuahua, Ciudad de México, Coahuila, Durango, Guanajuato, Hidalgo, Jalisco, México, Michoacán, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tlaxcala, Veracruz and Zacatecas (see e.g., Arias et al. 1997, Guzmán et al. 2003, Scheinvar 2004, Scheinvar et al. 2010, Arias et al. 2012, Hernández et al. 2014) (Fig. 2). It grows in xerophytic scrub and tropical deciduous forest (Rzedowski 1978)., Published as part of Martínez-González, César Ramiro, Gallegos-Vázquez, Clemente, Mascorro- Gallardo, José O. & Barrientos-Priego, Alejandro F., 2022, Molecular and morphological notes on Opuntia ser. Streptacanthae (Cactaceae), pp. 1-28 in Phytotaxa 576 (1) on pages 12-19, DOI: 10.11646/phytotaxa.576.1.1, http://zenodo.org/record/7440876, {"references":["Lemaire, C. (1839) Cactearum genera nova speciesque novae et omnium in horto Monvilliano cultarum ex affinitatibus naturalibus ordinatio nova indexque methodicus. Lutetiis, Paris, 115 pp.","Scheinvar, L., Gallegos-Vazquez, C., Olalde-Parra, G. & Rodriguez-Fuentes, A. (2010) Opuntia ser, Streptacanthae (Cactaceae) - neotypifications and taxonomic notes for four species. Schumannia 6: 277 - 296.","Bravo-Hollis, H. (1978) Las Cactaceas de Mexico, vol. 1. Universidad Nacional Autonoma de Mexico, Mexico City, 351 pp.","Scheinvar, L. (2004) Flora cactologica del estado de Queretaro. Diversidad y riqueza. Fondo de Cultura Economica, Mexico, 390 pp.","Arias, S., Gama S. & Guzman, U. (1997) Cactaceae. Flora del Valle de Tehuacan-Cuicatlan, Fasc. 14. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico City, 146 pp.","Guzman, U. L., Arias, S. & Davila, P. (2003) Catalogo de cactaceas mexicanas. Universidad Nacional Autonoma de Mexico. Mexico City, 315 pp.","Arias, S., Gama, S., Guzman, U. & Vazquez, B. (2012) Cactaceae. Flora del Valle de Tehuacan-Cuicatlan, Fasc. 95. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. Mexico City, 235 pp.","Hernandez, H. M., Gomez, H. C., Barcenas, R. T., Puente, P. & Reyes, A. J. (2014) A checklist of the subfamily Opuntioideae (Cactaceae) from North and Central America. In: Hunt, D. R. (Ed.) Further Studies in Opuntioideae (Cactaceae). Succulent Plant Research 8: 185 - 200.","Rzedowski, J. (1978) Vegetacion de Mexico. Limusa, Mexico City, 432 pp."]}

Opuntia megacantha Salm-Dyck (1834: 363) Neotype (designated by Scheinvar et al. 2010: 278): — MEXICO. Hidalgo, Villa de Tezontepec, 25 February 2009, Martínez-González & Scheinvar 7632 (MEXU!). Description:— Arborescent habit, 2.0– 2.2 m high (Fig. 3A). Defined trunk of 30 cm in diameter, grayish scaly bark (Fig. 3B). Glabrous epidermis (Fig. 6C). Elliptical to oval cladodes, 28–35 cm long × 18–28 cm wide and 2–3 cm thick, green in color, covered with a light layer of wax (Fig. 4A). Areolas arranged in 8–9 series, 2.4 cm spacing between series, 2.2 cm spacing between areolas, elliptical shape, 0.3 × 0.2 cm, short black trichomes (Fig. 4B). Very short brown glochids (0.1 cm). 1–7 flexible, radial and diffuse white spines, yellowish apex, conical rigid central spines, some twisted, erect, diffuse and radial, unequal of 0.4–2.6 cm long (Fig. 4C). Juvenile cladodes with prominent tubers, areolas with short brown to black trichomes, reddish bristel hairs, green erect subulate leaves, with reddish uncinate apex, white spines with reddish apex (Fig. 3C). Acute flower buds, green perianth segments with reddish acuminate apex, obovate to elliptical pericarpel, slightly elevated tubers, areolas with short light brown trichomes, brown glochids and yellow bristle areolas (Fig. 4D and 4E). Yellow flowers (Fig. 4F), 6.0–7.0 cm long, turbinate pericarp, ca. 3.2 × 2.6 cm, areolas arranged in 4–5 series, 0. 6 cm spacing between each other, with crass basal scale, oblanceolated outer segments of perianth, mucroned apex, smooth, yellow edges, with medium green striation, oblanceolated inner segments, emarginated apex, yellow, slightly toothed upper edges, stamens 1/3 of the length of the perianth, white filaments with yellowish tones, yellow anthers, white cuneiform style, ca. 1.9 cm long, usually ten green papillary lobes of the stigma, ca. 0.6 cm long, with medium white striation (Fig. 3D). Elliptical to obovate (Fig. 5B), red fruits, 5.1–7.0 cm long, striated and slightly sunken floral scar, large, circular areolas (Fig. 5C), arranged in 6–7 series, 1 cm spacing from each other and 1 cm between series, long black trichomes, absent spines, brown glochids, sweet and juicy red funicles (Fig. 5D). Kidney shaped seed with irregular lateral aril, lateral yellow-micropillar region, micropiles and funicles included (Fig. 5F). Micromorphology:— Thornswith lignified epidermal cells 140.51 ± 3.1 × 36.32 ± 2.3 μm in the distal part (Fig. 6B), 94.97 ± 1.50 × 18.20 ± 2.40 μm, non-continuous epidermal cells with fissures between cells and smooth texture, in the middle part (Fig. 6B). Glabrous epidermis, irregular epidermal cells (Fig. 6C) and tetracytic stomas (Fig. 6D). Polyhedron-shaped pollen grain, pantoaperturate (Fig. 6E), 102.08 ± 1.90 μm in diameter, 8050.69 μm ² in area, 16.27 ± 1.80 μm distance between each distal opening (pores), with a diameter of 22.21 ± 1.50 μm, 20–22 pores with walls of 1.15 ± 2.00 μm width and wart (Fig. 6F). Vernacular names:— Sangrita (e.g., Reyes-Aguëro et al. 2009). Phenology:— Opuntia megacantha blooms from March–April, whereas fruiting time if June–July. Habitat and distribution:— Opuntia megacantha is species endemic to Mexico, it is known in the states of Aguascalientes, Coahuila, Durango, Guanajuato, Hidalgo, Jalisco, Oaxaca, Querétaro, San Luis Potosí, Sonora, Tamaulipas, Tlaxcala and Zacatecas (see e.g., Guzmán et al. 2003, Scheinvar et al. 2010, Hernández et al. 2014) (Fig. 2). It grows in xerophytic scrub (Rzedowski 1978)., Published as part of Martínez-González, César Ramiro, Gallegos-Vázquez, Clemente, Mascorro- Gallardo, José O. & Barrientos-Priego, Alejandro F., 2022, Molecular and morphological notes on Opuntia ser. Streptacanthae (Cactaceae), pp. 1-28 in Phytotaxa 576 (1) on pages 7-9, DOI: 10.11646/phytotaxa.576.1.1, http://zenodo.org/record/7440876, {"references":["Salm-Dyck, J. (1834) Hortus Dyckensis oder Verzeichniss der in dem botanischen Garten zu Dyck wachsenden Pflanzen. Dusseldorf, Arnz.","Scheinvar, L., Gallegos-Vazquez, C., Olalde-Parra, G. & Rodriguez-Fuentes, A. (2010) Opuntia ser, Streptacanthae (Cactaceae) - neotypifications and taxonomic notes for four species. Schumannia 6: 277 - 296.","Reyes-Aguero, J. A., Aguirre, J. R., Carlin, F. & Gonzalez A. (2009) Catalogo de las principales variantes silvestres y cultivadas de Opuntia en la Altiplanicie Meridional de Mexico. UASLP., SAGARPA and CONACYT, San Luis Potosi, Mexico, 350 pp.","Guzman, U. L., Arias, S. & Davila, P. (2003) Catalogo de cactaceas mexicanas. Universidad Nacional Autonoma de Mexico. Mexico City, 315 pp.","Hernandez, H. M., Gomez, H. C., Barcenas, R. T., Puente, P. & Reyes, A. J. (2014) A checklist of the subfamily Opuntioideae (Cactaceae) from North and Central America. In: Hunt, D. R. (Ed.) Further Studies in Opuntioideae (Cactaceae). Succulent Plant Research 8: 185 - 200.","Rzedowski, J. (1978) Vegetacion de Mexico. Limusa, Mexico City, 432 pp."]}

The recently debated identity and incertus status of Cereus serruliflorus, Opuntia taylorii, O. tuna. and of its presumed close relatives, O. dillenii and O. jamaicensis from the Caribbean region, are here investigated to clarify the correct application of the names through an analysis of the protologues, original material, and examination of other specimens. One new endemic species to Jamaica (O. kingstoniana) was described; O. repens subsp. taylorii comb. nov. is proposed; an epitipe was designated for Cactus tuna (≡ O. tuna) and Cereus serruliflorus (≡ Serrulatocereus serrulatus); O. jamaicensis is here accepted as a valid species, distinct from O. tuna sensu auct.

3. Opuntia repens Bello (1881: 277) subsp. taylorii (Britton & Rose) Guiggi, comb. et stat. nov. ≡ Opuntia taylorii Britton & Rose (1908: 520, as “ Opuntia taylori ”). Type: ― HAITI. Between Gonaives and La Hotte Rochée: on road to Terre Neuve, 12 August 1905, Nash 1587 (holotype NY386061, non videmus; isotype US 00117058!, corpus, areolae, spinae). = Opuntia hattoniana Britton & Rose (1914: 97). Lectotype (designated here): ― DOMINICAN REPUBLIC. Azua, March 1913, Rose 3835 [US00180245! (two sheets), corpus, areolae, spinae; isolectotypes: NY1496093! (sub Opuntia taylori), corpus, areolae, spinae, HMGBH!, corpus, areolae, spinae (Fig. 6)]. Etymology: ―Dedicated to the British botanist, explorer and co-collector Norman Taylor (1883–1967) (Crook & Mottram 2004, Eggli & Newton 2004). Tyfication of Opuntia hattoniana: ―No type or herbarium were cited by Britton & Rose (1914: 97) in the protologue. The species was later synonymized by the same authors under Opuntia taylori (see Britton & Rose 1919: 103). Guiggi & Mariotti (2019: 194) traced three type collections with the same number and date deposited at NY, US and HMGBH, but being the type necessarily a single specimen conserved in one herbarium (see Art. 8.1 of ICN), a lectotype is designated here (herbarium US) because includes larger and easier identifiable specimens; isolectotypes at NY and HMGBH. Taxonomical notes: ―We here propose to consider the taxon taylorii at subspecies rank of Opuntia repens. In fact, the ecology of Britton & Rose’s Opuntia taylorii is different from Bello’s O. repens [desert on calcareous rocks vs. saline soils (see Majure & Clase 2020: 107)]; the distribution areas are: region of Hispaniola for O. taylorii (Britton & Rose 1919: 103) and, concerning O. repens, Puerto Rico (including Mona Island), Virgin Islands and in northern part of Dominican Republic (Majure & Clase 2020: 107) and in Haiti, recognised here for the first time (see Majure & Clase 2020: 107), vicinity of Fond Parisien (see Leonard 4124 sub O. taylori, 1496092 NY!, corp, ar, sp, icon). Morphologycally, taxon taylorii is, characterised by cylindrical stem-segments, that reduce the exposed surface compared with the more frequent lateral compressed segments of the typical species, probably in response to the colonisation of drier habitats, whereas the typical O. repens has flattened stem-segments. Illustrations examined: ― Britton & Rose (1919: plate 17 fig. 2) represents a joint collected by Rose, Fitch and Russel at Azua, Dominican Rep., in 1913). Chorology: ―Endemic to Hispaniola (Majure & Clase 2020: 107). Relevant literature: ―Britton (1908), Britton & Rose (1919). Specimen examined:― HAITI. Dept. du Nord-Ouest. Presqu’il du Nord-Ouest, Port-de-Paix, road to Jean Rabel, in Morne Palmiste, 16 May 1925, Ekman H4037 sub Opuntia taylori Britton & Rose (1496088 NY!, corp, ar, sp; 00180237 US! corp, ar, sp, fl); vicinity of La Vallée, Tortue Island, 3–10 May 1929, Leonard 15312 sub O. taylori (1496086 NY!, corp, ar, sp, fl). Dept. du Ouest. vicinity of Etang Saumatre, 4–12 April 1920, Leonard 3501 sub O. taylori (1496091 NY!, corp, ar, sp, fl); vicinity of Etang, Etang Saumatre, arid wooded plain of the Cul de Sac, 13–17 May 1920, Leonard 4250 sub O. taylori (1496085 NY!, corp, ar, sp, fl). DOMINICAN REP. Pedernales Prov., 10 km E del cruce de Cabo-Rojo Pedernales en el camino a Oviedo, sobre Loma El Guano, 7 May 1982, Zanoni 20527 sub Opuntia caribaea Britton & Rose (01496083 NY!, corp, ar. sp, fl, fr).

2. Opuntia kingstoniana Guiggi sp. nov. (Figs. 3–4) Type: ― JAMAICA. Kingston, June 1914, Rose 18503 sub Opuntia tuna (L.) Mill. (holotype US00180263!, corpus, areolae, spinae; isotype NY1496104!, corpus, areolae, spinae). – Opuntia tuna sensu Britton & Rose (1919: 113–114), non (L.) Mill. Diagnosis: ―It differs from Opuntia tuna by its lower habit to 0.9 m tall (vs. to 3 m); smaller cladodes to 16 cm long (vs. to 40 cm), light green (vs. bluish green), with entire margins (vs. ± undulate); spines acicular, straight, greyish in age (vs. subulate, curved or twisted, yellow ± brown banded), to 6 in number (vs. to 10); fruit 3 cm long (vs. 5.0– 7.5 cm). Description: ―Shrubby habit, 60–90 cm high, with horizontal branching pattern; cladodes light green, darker around the areoles, normally obovate, to 16 cm long; leaves small, early deciduous; areoles large, brownish later grayish; spines initially brownish-yellow, later greyish darker at apex, normally straight and reflexed, 2–6 in number, 3.5–5.0 cm long; glochids yellowish; flower yellowish slightly tinged of red, 5 cm in diameter, inner tepals oblong, rounded at apex, outer tepals orbicular, filaments greenish below, style and stigma-lobes cream or yellowish; pericarpel obovoid, intense green, with areoles and tufts of yellow glochids; fruit reddish, obovoid, ca. 3 cm long; seeds 3–4 mm in diameter. Etymology: ―The name refers to Kingston, the capital city of Jamaica, where the species grows. Taxonomical notes: ―This new species was reported by Britton & Rose (1919: 113–114) as Opuntia tuna, a name with priority referred over Opuntia dillenii (see the discussion below about O. tuna), but it can easily distinguished by its lower habit, smaller cladodes not glaucous, acicular spines and smaller fruits (see the above diagnosis). O. kingstoniana with its yellow flowers is also easily distinct from another Jamaican endemism O. sanguinea Proctor (1982: 239) which is characterized by its oxblood ” sanguineis ” flowers. The synonyms included in Britton & Rose (1919: 113) sub Opuntia tuna of uncertain application are not referable to O. kingstoniana. Chorology: ―Endemic to Jamaica (Britton & Rose 1919: 114). Its disjointed distribution in Cayman Islands (see Adams 1972: 74) has not been confirmed by Proctor (1984: 320). Illustrations examined: ― Britton & Rose (1919: 114 figs. 141–142) represent a plant and two cladodes collected by W. Harris near Kingston in 1913 (Fig. 5); Howard & Touw (1982: 176–177 figs. 8–9). Relevant literature: ― Britton & Rose (1919), Adams (1972)., Published as part of Guiggi, Alessandro & Mariotti, Mauro, 2022, Taxonomic and nomenclatural novelties in some Cactaceae of Greater Antilles, pp. 215-230 in Phytotaxa 573 (2) on page 219, DOI: 10.11646/phytotaxa.573.2.3, http://zenodo.org/record/7349827, {"references":["Britton, N. L. & Rose, J. N. (1919) The Cactaceae, vol. 1. Carnegie Institution, Washington, 236 pp.","Proctor, G. R. (1982) More additions to the Flora of Jamaica. Journal of the Arnold Arboretum 63 (3): 199 - 315. https: // doi. org / 10.5962 / p. 37029","Adams, C. D. (1972) Flowering plants of Jamaica. University of West Indies, Jamaica, 848 pp. Areces, A. M. (2018) Neohaiticereus a new subgenus for the rediscovered Cereeae depicted in Plumier's Plate 26 of the Botanicon","Proctor, G. R. (1984) Flora of the Cayman Islands. Royal Botanic Gardens, Kew, 724 pp.","Howard, R. A. & Touw, M. (1982) Opuntia species in the Lesser Antilles. Cactus & Succulent Journal 54: 170 - 179."]}

3. Opuntia repens Bello (1881: 277) subsp. taylorii (Britton & Rose) Guiggi, comb. et stat. nov. ≡ Opuntia taylorii Britton & Rose (1908: 520, as “ Opuntia taylori ”). Type: ― HAITI. Between Gonaives and La Hotte Rochée: on road to Terre Neuve, 12 August 1905, Nash 1587 (holotype NY386061, non videmus; isotype US 00117058!, corpus, areolae, spinae). = Opuntia hattoniana Britton & Rose (1914: 97). Lectotype (designated here): ― DOMINICAN REPUBLIC. Azua, March 1913, Rose 3835 [US00180245! (two sheets), corpus, areolae, spinae; isolectotypes: NY1496093! (sub Opuntia taylori), corpus, areolae, spinae, HMGBH!, corpus, areolae, spinae (Fig. 6)]. Etymology: ―Dedicated to the British botanist, explorer and co-collector Norman Taylor (1883–1967) (Crook & Mottram 2004, Eggli & Newton 2004). Tyfication of Opuntia hattoniana: ―No type or herbarium were cited by Britton & Rose (1914: 97) in the protologue. The species was later synonymized by the same authors under Opuntia taylori (see Britton & Rose 1919: 103). Guiggi & Mariotti (2019: 194) traced three type collections with the same number and date deposited at NY, US and HMGBH, but being the type necessarily a single specimen conserved in one herbarium (see Art. 8.1 of ICN), a lectotype is designated here (herbarium US) because includes larger and easier identifiable specimens; isolectotypes at NY and HMGBH. Taxonomical notes: ―We here propose to consider the taxon taylorii at subspecies rank of Opuntia repens. In fact, the ecology of Britton & Rose’s Opuntia taylorii is different from Bello’s O. repens [desert on calcareous rocks vs. saline soils (see Majure & Clase 2020: 107)]; the distribution areas are: region of Hispaniola for O. taylorii (Britton & Rose 1919: 103) and, concerning O. repens, Puerto Rico (including Mona Island), Virgin Islands and in northern part of Dominican Republic (Majure & Clase 2020: 107) and in Haiti, recognised here for the first time (see Majure & Clase 2020: 107), vicinity of Fond Parisien (see Leonard 4124 sub O. taylori, 1496092 NY!, corp, ar, sp, icon). Morphologycally, taxon taylorii is, characterised by cylindrical stem-segments, that reduce the exposed surface compared with the more frequent lateral compressed segments of the typical species, probably in response to the colonisation of drier habitats, whereas the typical O. repens has flattened stem-segments. Illustrations examined: ― Britton & Rose (1919: plate 17 fig. 2) represents a joint collected by Rose, Fitch and Russel at Azua, Dominican Rep., in 1913). Chorology: ―Endemic to Hispaniola (Majure & Clase 2020: 107). Relevant literature: ―Britton (1908), Britton & Rose (1919). Specimen examined:― HAITI. Dept. du Nord-Ouest. Presqu’il du Nord-Ouest, Port-de-Paix, road to Jean Rabel, in Morne Palmiste, 16 May 1925, Ekman H4037 sub Opuntia taylori Britton & Rose (1496088 NY!, corp, ar, sp; 00180237 US! corp, ar, sp, fl); vicinity of La Vallée, Tortue Island, 3–10 May 1929, Leonard 15312 sub O. taylori (1496086 NY!, corp, ar, sp, fl). Dept. du Ouest. vicinity of Etang Saumatre, 4–12 April 1920, Leonard 3501 sub O. taylori (1496091 NY!, corp, ar, sp, fl); vicinity of Etang, Etang Saumatre, arid wooded plain of the Cul de Sac, 13–17 May 1920, Leonard 4250 sub O. taylori (1496085 NY!, corp, ar, sp, fl). DOMINICAN REP. Pedernales Prov., 10 km E del cruce de Cabo-Rojo Pedernales en el camino a Oviedo, sobre Loma El Guano, 7 May 1982, Zanoni 20527 sub Opuntia caribaea Britton & Rose (01496083 NY!, corp, ar. sp, fl, fr)., Published as part of Guiggi, Alessandro & Mariotti, Mauro, 2022, Taxonomic and nomenclatural novelties in some Cactaceae of Greater Antilles, pp. 215-230 in Phytotaxa 573 (2) on page 222, DOI: 10.11646/phytotaxa.573.2.3, http://zenodo.org/record/7349827, {"references":["Britton, N. L. & Rose, J. N. (1908) A preliminary treatment of the Opuntioideae of North America. Smithsonian miscellaneous collections 50: 503 - 539.","Britton, N. L. & Rose, J. N. (1914) In the West Indies. In: Johnston & Tryon. Report of the Prickly-pear Travelling Commission. 1 st November, 1912 - 30 th April, 1914. pp. 90 - 101","Crook, R. & Mottram, R. (2004) Opuntia Index Part 10: T - V. Bradleya 22: 53 - 76. https: // doi. org / 10.25223 / brad. n 22.2004. a 7","Eggli, U. & Newton, L. E. (2004) Etymological dictionary of succulent plant names. Springer-Verlag, Berlin, 266 pp. https: // doi. org / 10.1007 / 978 - 3 - 662 - 07125 - 0 _ 1","Britton, N. L. & Rose, J. N. (1919) The Cactaceae, vol. 1. Carnegie Institution, Washington, 236 pp.","Majure, L. C & Clase, T. (2020) Opuntia repens en Republica Dominicana y notas sobre la taxonomia del complejo Curassavica. Moscosoa 20: 105 - 113."]}

4. Opuntia tuna (L.) Miller (1768: OPUNTIA n° 3) ≡ Cactus tuna Linnaeus (1753: 468). Lectotype (designated by Crook & Mottram (2004: 61):―[Icon] pl. 295 fig. 380 in Dillenius (1732) (Fig. 7). Epitype (designated here): ― JAMAICA. St. Andrew: Hope River, about 1 mi E of Mona, ca. 152 m, 12 December 1957, Yuncker 17727 (NY1496096!, corpus, areolae, spinae, flos, fructus, Fig. 8). = Opuntia dillenii (Ker Gawler 1818: pl. 255) Haworth (1819: 79). Lectotype (designated by Benson 1969: 126):―[Icon] pl. 255 in Ker Gawler (1818) (reproduced in Crook & Mottram 1996: 118, fig. 13). Etymology: ―From Spanish tuna, a vernacular name referring to the fruit of Opuntia (Crook & Mottram 2004, Eggli & Newton 2004). Typification of Cactus tuna: ―Doubts and questions over the iconographic lectotype proposed by Hunt et al. (2006: 213) can be resolved choosing an epitype from preserved material to better define the name (Art. 9.9 of ICN). The descriptions and the images reported in the protologue of Cactus tuna (see Linnaeus 1753: 468), as well as the lectotype [Dillenius’ (1732) pl. 295 fig. 380], designated by Crook & Mottram (2004: 61) and the epitype [Yuncker 17727 (NY1496096!), here designated] of the species morphologycally match the Opuntia dillenii species concept (Proctor 1984: 320–321 fig. 90; Howard 1989: 412; Liogier 1994: 322–325 fig. 95–8) and its type [lectotype, i.e. the plate no 255 in Ker Gawler (1818), designated by Benson 1969: 126]; important characters are: cladodes from ovate to oblong, spines subulate, brownish yellow, yellow flower and fruit normally long pyriform with a narrowed base. All these characters are different from those characterizing O. kingstoniana (– Opuntia tuna sensu Britton & Rose; here newly described, see above) which can be distinguished by its shrubby habit, with cladodes obovate, spines acicular, greyish and fruit short-obovoid. Furthermore, note that the provenance of Cactus tuna reported by Linnaeus (1753: 468), i.e. “ Jamaica & America calidiore ”, refers to O. dillenii for Jamaica, being this species the only one occurring in the island with this cited distribution. According to Schumann (1899: 724), O. tuna is the valid name with priority for the plant known nowadays as O. dillenii and it differs from the Jamaican species here described and collected in Kingston by Rose (O. kingstoniana). The collectors of the specimens here studied have correctly applied the name O. tuna as conceptually defined here. Chorology: ―Caribbean, SE-United States, E-Mexico, N-South America (Britton & Rose 1919: 162). Illustrations examined: ― Dillenius (1732: pl. 295 figs. 380–381, pl. 296 fig. 382, reproduced in Crook & Mottram 1996: 123 fig. 16, 126 fig. 17), Sloane (1725: pl. 224 fig. 1, deposited in Sloane’s Herbarium at BM, Fig. 9), Edwards (1818: pl. 255, reproduced in Crook & Mottram 1996: 118 fig. 13). Relevant literature: ― Schumann (1899), Britton & Rose (1919). Specimen examined:― CAYMAN ISLANDS. East End, 1 July 1938, 3–4.5 m, Kings 226 (1496098 NY!, corp, ar, sp, fl)., Published as part of Guiggi, Alessandro & Mariotti, Mauro, 2022, Taxonomic and nomenclatural novelties in some Cactaceae of Greater Antilles, pp. 215-230 in Phytotaxa 573 (2) on pages 222-224, DOI: 10.11646/phytotaxa.573.2.3, http://zenodo.org/record/7349827, {"references":["Miller, P. (1768) The gardeners dictionary. 8 th edition. Published by the author, London, unpaged.","Linnaeus, C. (1753) Species plantarum, vol. 1. Laurentii Salvii, Holmiae, 560 pp.","Crook, R. & Mottram, R. (2004) Opuntia Index Part 10: T - V. Bradleya 22: 53 - 76. https: // doi. org / 10.25223 / brad. n 22.2004. a 7","Dillenius, J. J .. (1732) Hortus Elthamensis. Sumptibus auctoris, Londini, 204 pp.","Haworth, A. H. (1819) Supplementum Plantarum Succulentarum. Impensis J. Harding, Londini, 160 pp.","Crook, R. & Mottram, R. (1996) Opuntia Index Part 2: Nomenclatural note and C - E. Bradleya 14: 99 - 144. https: // doi. org / 10.25223 / brad. n 14.1996. a 15","Eggli, U. & Newton, L. E. (2004) Etymological dictionary of succulent plant names. Springer-Verlag, Berlin, 266 pp. https: // doi. org / 10.1007 / 978 - 3 - 662 - 07125 - 0 _ 1","Hunt, D., Taylor, N. & Charles, G. (Eds.) (2006) New cactus lexicon. David Hunt Books, Milborne Port, 371 pp.","Proctor, G. R. (1984) Flora of the Cayman Islands. Royal Botanic Gardens, Kew, 724 pp.","Howard, R. A. (1989) Flora of the Lesser Antilles, vol. 5, part. 2. Harvard University Jamaica Plain, Massachusetts, 604 pp.","Liogier, H. A. (1994) Descriptive Flora of Puerto Rico and Adjacent Islands, vol. 3. Editorial de la Universidad de Puerto Rico, 462 pp.","Schumann, K. (1899) Gesamtbeschreibung der Kakteen (Monographia Cactacearum). J. Neudamm Verlag, 832 pp. https: // doi. org / 10.5962 / bhl. title. 10394","Britton, N. L. & Rose, J. N. (1919) The Cactaceae, vol. 1. Carnegie Institution, Washington, 236 pp.","Sloane, H. (1725) A voyage to the islands Madera, Barbados, Nieves, S. Christophers and Jamaica. The British Museum, for the author. Cacti 2: 148 - 161."]}

1. Opuntia jamaicensis Britton & Harris in Britton (1911: 130). Lectotype (designated here): ― JAMAICA. St. Catharine: roadside plains, near Salt Ponds, 31 August 1908, Britton & Harris 10887 [NY386051!, corpus, areolae, spinae, flos, fructus, semen (Fig. 1); isolectotypes US00115891!, corpus, areolae, spinae, flos, fructus, semen, UCWI (ex JM), non videmus]. Etymology: ―The epiteth derives from Jamaica with the Latin suffix - ensis, referred to its Jamaican geographic origin (Crook & Mottram 1999, Eggli & Newton 2004). Typification of Opuntia jamaicensis: ― Britton (1911: 130) reported “ Typus ” for collection n. 10887 by Britton & Harris (or Harris & Britton as cited in the label of the existent herbarium samples). However, no herbarium was cited in the protologue. Moreover, there are three type collections with the same number and date deposited at NY, US, and UCWI (ex JM). As consequence, according to the Art. 8.1 of ICN, a lectotype is designated here among the syntypes known (Art. 9.6 of ICN), opting for NY specimen, being the most complete with all vegetative and reproductive elements. Taxonomical notes: ―The comparison of the illustrations and descriptions by Britton (1911: 130) and Britton & Rose (1919: 161), as well as by examining herbaria specimens, supported this taxon as a distinct species from Opuntia tuna sensu auct. (see next paragraphs), differing from the other Jamaican taxon (here described as Opuntia kingstoniana), for its cladodes more sphatulate with a narrow base (vs. usually obovate), areoles oval (vs. circular), fewer spines (usually 1–2 per areoles (vs. usually 2–5), up to 2.5 cm long (vs. up to 5.0 cm long), spreading (vs. reflexed), whitish (vs. brownish-yellow), usually arranged only along the apical margins (vs. the entire surface), fruit pyriform with the lower part very narrow and elongated, 3.5–4.0 cm long (vs. short-obovoid, about 3 cm long). Also habitat and distribution areas are different between these two taxa: O. jamaicensis grows on saline soils (vs. rocky or sandy soils for O. kingstoniana), at 0–8 m a.s.l. (vs. 0–120 m), in St. Catherine and Manchester Parishes (vs. St. Andrew and St. Thomas Parishes) [see also Britton & Rose 1919: 113–114, Adams 1972: 74, and Britton & Harris 10887 sub Opuntia jamaicensis (NY, US), Rose 18503 sub O. tuna (US, NY)]. Notes on flowering time: ―The plant start to flower at 11 a.m. and begin to close after 4 p. m. (Britton 1911: 130). Chorology: ―Endemic to Jamaica (Britton & Rose 1919: 113). Illustrations examined: ― Britton & Rose’s (1919: plate 18 figs. 4–5) figures are a fruiting cladode with a fruit section; plate 19 figs. 1–7 are a flowering plant, two flowers, one flower section, stamens and style from the plant type (Fig. 2). Relevant literature: ― Britton (1911), Britton & Rose (1919), Adams (1972)., Published as part of Guiggi, Alessandro & Mariotti, Mauro, 2022, Taxonomic and nomenclatural novelties in some Cactaceae of Greater Antilles, pp. 215-230 in Phytotaxa 573 (2) on pages 216-217, DOI: 10.11646/phytotaxa.573.2.3, http://zenodo.org/record/7349827, {"references":["Britton, N. L. (1911) An undescribed Opuntia from Jamaica. Torreya 11: 130.","Crook, R. & Mottram, R. (1999) Opuntia Index Part 5: I - L. Bradleya 17: 109 - 131. https: // doi. org / 10.25223 / brad. n 17.1999. a 8","Eggli, U. & Newton, L. E. (2004) Etymological dictionary of succulent plant names. Springer-Verlag, Berlin, 266 pp. https: // doi. org / 10.1007 / 978 - 3 - 662 - 07125 - 0 _ 1","Britton, N. L. & Rose, J. N. (1919) The Cactaceae, vol. 1. Carnegie Institution, Washington, 236 pp.","Adams, C. D. (1972) Flowering plants of Jamaica. University of West Indies, Jamaica, 848 pp. Areces, A. M. (2018) Neohaiticereus a new subgenus for the rediscovered Cereeae depicted in Plumier's Plate 26 of the Botanicon"]}

5. Serrulatocereus serruliflorus (Haworth 1830: 113) Guiggi (2018: 1). Basionym: Cereus serruliflorus Haworth (1830: 113). ≡ Harrisia serruliflora (Haworth 1830: 113) Lourteig (1991: 408). Lectotype (designated by Lourteig 1991: 408):― HAITI. Grand Cul de Sac: between Port-au-Prince and Léogane, in hot, dry woods; plant portrayed in September 1689 –1690, or 1693, by Charles Plumier (1689 –1697: 26, pl. 3: 26) (Fig. 10). Epitype (designated here):― HAITI. Ouest Dept.: Plaine Cul-de-Sac, Croix-des-Bouquets, 18 December 1925, Ekman H-5377, sub Cereus repandus (S15-7962!, two sheets, corpus, areolae, spinae, flos, fructus, Fig. 11; isoepitype: US 00171443!, corpus, areolae, spinae). = Cereus haitiensis A.R.Franck & Peguero in Franck et al. (2017: 2), non (Schumann 1903: 183) Schelle (1926: 120), nom. illeg. (Arts. 53.1 of ICN). Type: ― HAITI. Nord-Ouest Dept.: arid thickets, W of Môle gorge, vicinity of Môle-St-Nicolas, 16 February 1929, Leonard & Leonard 13311 sub Cephalocereus nobilis (Haw.) Britton & Rose (holotype US 00171001!, corpus, areolae, spinae, flos, fructus; isotype NY1495818!, corpus, areolae, spinae, flos, fructus). = Cereus ayisyen Van der Meer (2019: 14), nom. illeg. (Art. 52.1 of ICN). Etymology: ―From the Latin serrulatus referring to the serrulate apical margin of the perianth segments (Eggli & Newton 2004, Guiggi, 2018). Typification of Cereus serruliflorus: ―Given the considerable uncertainty associated with this taxon (see discussion below) and the designation of an iconographic lectotype, an epitype represented by a herbarium sample referred to and compatible with all elements included in the Plumier’s drawing is here chosen to establish the application of the name (see Turlan et al. 2018, Art. 9.9 of ICN). Charles Plumier in his unpublished Botanicon Americanum (1689–1697) included 29 plates describing 24 taxa of Cactaceae discovered during his trips in the Greater and Lesser Antilles. However, his drawings in many cases are not accurate, containing errors or out-of-scale elements (see Areces 2018: 108, 112), probably because they were illustrated from memory (Mottram 2002: 86, Areces 2018: 112). Some examples are Consolea moniliformis (Linnaeus 1753: 468) Berger (1926: 94), based on Plumier’s plate 11, Harrisia divaricata based on plate 23, and Neogriseocereus fimbriatus (Lam. in Lamarck et al. 1785: 539) Guiggi (2013: 1), based on plate 25 (Britton & Rose 1919: 207, Mottram 2002: 88, 2020: 2, 63). Their identification and validity have never been questioned, unlike plate 26 (Fig. 10), used by Haworth (1830: 113) to describe his Cereus serruliflorus. Franck et al. (2017: 5) suspected that C. serruliflorus was a mix with H. divaricata and interpreted their narrowed lectotype in the sense of the latter species. We are not in agreement with this position (Cereus serruliflorus was a mix with Harrisia divaricata) for several reasons. H. divaricata with its different characters is depicted by Plumier (1689 –1697) on plate 24. Oversized elements including the scales on the flower tube and on the fruits included in pl. 26 occur also in the previously cited cases (i.e. plates nos. 11, 23 and 25). The locality (i.e. Plaine Cul-de-Sac, Croix-des-Bouquets) of Ekman’s collection (H-5377), here chosen as an epitype of Cereus serruliflorus (see Franck et al. 2017: 2–3, Areces 2018: 116), is the same of the plant on which Plumier (1689 –1697: pl. 26) based his drawing. The morphology of the stem in Plumier’s pl. 26, with the visible crenate margins and short and weak spines, are also conspecific with those included in Ekman (H-5377), with the exception of the lower number of ribs (8 vs. 11–13; see Franck et al. 2017: 2), here considered not so decisive, in consideration of artistic interpretations or imprecise memories. In addition, the lectotype interpretation as Harrisia divaricata by Franck et al. (2017: 5) is in conflict with the protologue that described a large plant (to 4–5 m high, see Franck et al. 2017: 2, Areces 2018: 113) with a large fruit, subconical (Haworth 1830: 113; see also the Plumier’s plate 26), that for this reason could be rejected (Art. 9.19 of ICN). In contrast, H. divaricata is a shrubby plant (2–3 m high, see Franck 2016: 30, 115 fig. 66, 116 fig. 67) normally with globose fruit at maturity (Franck, 2016: 118, fig. 69). As support of these arguments, the morphology of the flower occurring in Plumier’s plate 26 is not the same apart the scales already discussed, as that of H. divaricata characterised by non-deflexed and triangular perianth segments and non-exserted stamens and stigma (see Franck 2016: 117, fig. 68). Furthermore, in the English translation of the Plumier’s description associated with the pl. 26 (Mottram 2002: 113, 2020: 64), the characters reported (“ […] the trunk is almost as thick as a man’s body […] flowers, opening wide, monopetalous, truly campanulate […], with numerous narrow acuminate fimbriate segments […]. Its pericarpel and receptacle tube with small green scale-like leaves […] having a funnelform, many-branched stigma. […] an oblong or cucumber-shaped fruit”) matches the Areces’s concept of Cereus serruliflorus (see Areces 2018: 109-113, figs. 2, 6, 9, 10). All things analysed, we do not consider the Plumier’s plate 26 to refer to more the one species (Art. 9.17 of ICN). Therefore, the lectotyfication of Lourteig is valid in our opinion (1991: 408), whereas that of Franck et al. (2017: 5) is unnecessary and the new species Cereus haitiensis A.R.Franck & Peguero in Franck et al. (2017: 2) is also suspected to be a later homonym of C. haitiensis (K.Schum.) Schelle (1926: 120). Its replaced name Cereus ayisyen Van der Meer (2019: 14) see Mottram (2020: 64, 68) needs to be interpreted as illegitimate superfluous name. Taxonomic notes: ―The original inclusion of the taxon serruliflorus under the genus Cereus Miller (1754) by Haworth (1830: 113; see also Areces 2018: 113, Wisnev 2018: 223) appears inappropriate according to Guiggi (2018: 1), who highlighted that 1) the genus Cereus is not native from Caribbean region and 2) C. serruliflorus has vegetative and reproductive characters more related to other Caribbean genera, i.e Harrisia Britton (1908: 561) and Leptocereus (Berger 1905: 79) Britton & Rose (1909: 433) (see Guiggi 2018: 1). C. serruliflorus is distinguishable from Harrisia and Leptocereus by the following characteristics: flower strongly reflexed with oblong perianth segments, apical margins finely serrulate, style and stamens strongly exserted from the perianth, fruit long and subconical and pendulous. This led to the description of a new genus, Serrulatocereus Guiggi (2018: 1); it would belong to the tribe Leptocereae Buxbaum (1958: 178) sensu Guiggi (2020: 1–5), but new molecular data need (see Guiggi 2020: 1). Notes on flowering time: ―This species has a specific flowering time during the dry season from December to March (Areces 2018: 118). Chorology: ―Endemic to Haiti (Areces 2018: 113). Illustrations examined: ― Mottram (2002: 113 Plumier plate’s 26 Fig.10), Areces (2018: 108–116, figs 2–16), Franck et al. (2017: 10–17, figs. 3–15). Relevant literature: ― Mottram (2002, 2020), Franck et al. (2017), Areces (2018), Guiggi (2018), Wisnev (2018). Specimens examined: ― HAITI. Nord-Ouest Dept.: road Jean-Rabel to Môle-St-Nicolas, near Môle-St-Nicolas, 3 July 1925, Ekman H-4446 sub Cereus repandus (L.) Mill. (15-7969 S!, corp, ar, sp; 00171444 US!, corp, ar, sp, ico); vicinity of Jean Rabel, thicket along Mole Road, 3 March 1929, Leonard & Leonard 13682 sub Cephalocereus nobilis (1495817 NY! corp, ar, sp, fl; 00171002 US!, corp, ar, sp, fl); Môle-St-Nicolas, 2 February 1985, Zanoni et al. 33542 sub Harrisia sp. (65076 JBSD!, corp, ar, sp)., Published as part of Guiggi, Alessandro & Mariotti, Mauro, 2022, Taxonomic and nomenclatural novelties in some Cactaceae of Greater Antilles, pp. 215-230 in Phytotaxa 573 (2) on pages 225-228, DOI: 10.11646/phytotaxa.573.2.3, http://zenodo.org/record/7349827, {"references":["Haworth, A. H. (1830) Twelfth decade of new succulent plants. Philosophical Magazine Journal 37: 106 - 118.","Guiggi, A. (2018) Serrulatocereus Guiggi. A new proposed genus for the recent re-evaluated Cereus serruliflorus Haworth from Haiti (Cactaceae). Cactology 5 (suppl. 6): 1 - 3.","Lourteig, A. (1991) Nomenclatura plantarum Americanum. XVI. Cactaceae. Bradea 5: 400 - 411.","Plumier, C. (1689 - 1697) Botanicon Americanum seu historia plantarum in Americanis insulis nascentium (unpublished), vols. 1 - 30. Bibliotheque Centrale, Museum d'Histoire Naturelle, Paris, 1335 pp.","Franck, A. R., Peguero, B., Cinea, W. & Jestrow, B. (2017) A new species of Cereus s. str. (Cactaceae) endemic to Haiti. Phytoneuron 2017 - 29: 1 - 17.","Schumann, K. (1903) Reiseerinnerungen vom Jahre 1903. J. Neumann, Neudamm, for the Deutsche Kakteen-Gesellschaft. Monatsschrift Kakteen-kunde 13 (12): 182 - 189.","Schelle, E. (1926) Kakteen. Kurze Beschreibung nebst Angaben uber die Kultur der gegenwartig im Handel befindlichen Arten und Formen. Alexander Fischer Verlag, Tubingen, 368 pp.","Van Der Meer, M. H. J. (2019) A new name for Cereus haitiensis A. R. Franck & Peguero (Cactaceae). Cactologia Phantastica 4 (2): 13 - 15.","Eggli, U. & Newton, L. E. (2004) Etymological dictionary of succulent plant names. Springer-Verlag, Berlin, 266 pp. https: // doi. org / 10.1007 / 978 - 3 - 662 - 07125 - 0 _ 1","Mottram, R. (2002) Charles Plumier, the King's Botanist - his life and work. With a facsimile of the original cactus plates and text from Botanicon Americanum (1689 - 1697). Bradleya 20: 79 - 120. https: // doi. org / 10.25223 / brad. n 20.2002. a 12","Linnaeus, C. (1753) Species plantarum, vol. 1. Laurentii Salvii, Holmiae, 560 pp.","Berger, A. (1926) Die Entwicklungslinien der Kakteen. Gustav Fischer Verlag, Jena, Germany, 105 pp.","Lamarck, De J. B. (1785) Dictionnaire encyclopedique, col. 1 (2) Hotel de Thou, Paris, 409 pp.","Guiggi, A. (2013) Cactology III addenda et corrigenda III. Cactology 3 (suppl. 4): 1 - 2.","Britton, N. L. & Rose, J. N. (1919) The Cactaceae, vol. 1. Carnegie Institution, Washington, 236 pp.","Mottram, R. & Hoxey, P. (2020) An evaluation of the cacti of Charles Plumier (1646 - 1704). Cactician 13: 1 - 99.","Burmann, J. (1758) Plantarum Americanarum, Fasciculus Octavus. Petrum Schouten, Leiden and Gerard. Potuliet, & Theodor. Haak, 26 pp.","Franck, A. R. (2016) Monograph of Harrisia (Cactaceae). Phytoneuron 2016 - 85 (4): 102 - 134.","Miller, P. (1754) The gardeners dictionary. 4 th edition. Published by the author, London, unpaged.","Wisnev, M. A. (2018) Arecesocereus a new subgenus of Cereus. Cactus & Succulent Journal (United States) 90 (3): 222 - 223. https: // doi. org / 10.2985 / 015.090.0301","Berger, A. (1905) A systematic revision of the genus Cereus Mill. Annual Report Missouri Botanical Garden 16: 57 - 86. https: // doi. org / 10.2307 / 2400072","Britton, N. L. & Rose, J. N. (1909) The genus Cereus and its allies in North America. Contributions from the United States National Herbarium 12 (10): 413 - 437.","Buxbaum, F. (1958) The phylogenetic division of the subfamily Cereoideae, Cactaceae. Madrono 14: 177 - 216.","Guiggi, A. (2020) New circumscription and combinations in the tribe Leptocereae Buxbaum sensu Guiggi (Cactaceae-Cactoideae) from the Caribbean region. Cactology 5 (suppl. 9): 1 - 5."]}

The species of Eilema Hübner, [1819] sensu lato, present in Europe and North Africa, are studied on the basis of morphological characteristics and molecular genetics. Images of adults, genitalia of both sexes, immature stages, as well as phylogenetic trees obtained from the combined analysis of three different genetic markers are presented. Data on life cycles, food plants, and geographic distribution are also included. The morphological differences observed within Eilema sensu lato have led the authors to create the genera Indalia gen. nov. and Pseudokatha gen. nov. New combinations are stablished: Manulea iberica (Mentzer, 1980) st. rest., comb. nov.; Indalia marcida (Mann, 1859) comb. nov.; Indalia predotae (Schawerda, 1927) comb. nov.; Indalia albicosta (Rogenhofer, 1894) comb. nov.; Indalia interposita (Rothschild, 1914) comb. nov.; Indalia uniola (Rambur, [1866]) comb. nov.; Indalia lutarella (Linnaeus, 1758) comb. nov.; Indalia pygmaeola (Doubleday, 1847) comb. nov.; Pseudokatha rungsi (Toulgoët, 1960) comb. nov., and Eilema albicosta witti Kobes, 1993 syn. nov.

Wittia sororcula (Hufnagel, 1766) Original combination: Phalaena Noctua sororcula Hufnagel, 1766 Berlinisches Magazin 3: 398–399. T. L.: Berlín, Germany. Material examined. SPAIN: ARABA: 1 ♂ and 1 ♀, Berganzo, 500 m, 16.v.1998, J. Gastón leg.: 1 ♂, Zárate, 700 m, 29.v.1983, J. Gastón leg.; 1 ♂, Markínez, 790 m, 18.v.2018, J. Gastón leg.; 4 ♂ and 5 ♀, Pto. de Herrera, 1000 m, 13.vi.1981, J. Gastón leg. BARCELONA: 6 ♂ and 5 ♀, Els Munts, Sant Agustí de Lluçanès, 958 m, 31 TDG25, 5.v.2000, 12.v.2001, 15.viii.2009, R. Macià & J. Ylla leg.; 15 ♂ and 10 ♀, El Surreig, St. Bartomeu del Grau, 650 m, 3.v.1997, 30.iv.2007, 24.iv.2009, 1.v.2011, R. Macià & J. Ylla leg.; 6 ♂ and 4 ♀, Ermita de Sant Jaume, St. Bartomeu del Grau, 750 m, 1.vii.2014, R. Macià & J. Ylla leg.; 8 ♂ and 5 ♀, Molí del Llopart, Sta. Eugènia de Berga, 538 m, 14.vi.2003, 18.vi.2005, R. Macià & J. Ylla leg.; 7 ♂ and 4 ♀, St. Julià de Cabrera, 1008 m, 2.vii.2003, 1.vii.2005, R. Macià & J. Ylla leg.; 3 ♂, Urb. La Roca, Taradell, 650 m, 25.v.1991, R. Macià leg.; 5 ♂, 4 ♀, Vespella, Gurb, 650 m, 5.v.2000, R. Macià & J. Ylla leg.; 9 ♂ and 6 ♀, Coll d´Hueres, Collsuspina, 980 m, 31 TDG36, 15.vi.2013, R. Macià leg.; 4 ♂ and 2 ♀, Mas El Bruguer, Sau, 650 m, 20.v.2006, 5.v.2007, R. Macià & J. Ylla leg.; 1 ♂, Urb. Urbisol, Calders, 650 m, 15.v.1994, T. Mira leg.; 14 ♂ and 10 ♀, Vallvidrera, 362 m, 31.vii.1921, 15.v.1922, I. Sagarra leg. in coll. MCNB.; 1 ♂, Santa Susanna, Montseny, 630 m, 9.v.1914, I. Sagarra leg. in coll. MCNB.; 6 ♂ and 5 ♀, Sant Pere de Vilamajor, 305 m, 10.viii.2014, 4.ix.2014, I. Sagarra leg. in coll. MCNB. BIZKAIA: 1 ♀, Lendoño Goikoa, 460 m, 14.vi.1991, J. Gastón leg. ZAMORA: 5 ♂ and 3 ♀, Sierra de la Culebra, Ferreras, 890 m, 23.vi.2001, R. Macià & J. Ylla leg. BURGOS: 5 ♂ and 6 ♀, San Martín de Don, 850 m, 16.vi.1984, J. Gastón leg.; 5 ♂ and 6 ♀, Loma del Rey, Quintanilla Sobresierra, 1050 m, 30 TVN41, 28.v.2008, R. Macià leg. CASTELLÓN: 3 ♂ and 2 ♀, Serra del Turmell, Vallibona, 1080 m, 2.viii.2007, R. Macià & J.Ylla leg.; 2 ♂, Corral de Gracià, Barracas, 1029 m, 16.vi.2009, R. Macià & J. Ylla leg.; GIRONA: 3 ♂, Tamariu, 63 m, 19.viii.2006, R. Macià & J. Ylla leg.; 6 ♂ and 4 ♀, Urb. Vall Repós, Romanyà de la Selva, 325 m, 29.iv.2006, R. Macià & J. Ylla leg.; 4 ♂ and 1 ♀, Coll de Jou, Ogassa, 1650 m, 24.v.1996, R. Macià leg.; 1 ♂ and 2 ♀, Sant Pere Pescador, 15 m, 9.viii.1996, R. Macià leg.; 3 ♂, Fontclara, Palau-Sator, 70 m, 30.iv.2006, R. Macià & J. Ylla leg.; 3 ♂, Montgrony, Gombren, 1350 m, 29.v.1993, R. Macià & J. Ylla leg. FRANCE: 8 ♂ and 5 ♀, Nohèdes, Prades, Pirineus Orientals, 1080 m, 18.v.2002, R. Macià & J. Ylla leg.; 7 ♂ and 9 ♀, Mont Ventoux, Provenza, 1020 m, 29.vii.1994, R. Macià & J. Ylla leg. HUNGARY: 5 ♂ and 3 ♀, Örsèg, Kétvölgy, 1020 m, 31.vii.2005, A. Azabó leg. ITALY: 4 ♂ and 1 ♀, Sovicille, Siena, Toscana, 265 m, 23.vii.1996, R. Macià & J. Ylla leg. Diagnosis. The bright orange yellow colour and the shape of the forewing, very wide and rounded, make it easily distinguishable from any other Eilema sensu lato. Re-description. Imago (Figs. 43–44). Average wingspan males 28.0 mm (n=10; 25–30 mm); average wingspan females 28.7 mm (n=10; 25–31 mm). Small morphological differences between males and females. Genitalia (Fig. 70). Male genitalia: Uncus very long, thin, tubular, and curved, topped with a small hook; narrow and poorly sclerotised tegumen; valvae oval, elongated with rounded cucullus; the distal process of the sacculus very powerful, large, strongly sclerotised and curved towards the cucullus, ending with a small spine; the rest of the sacculus is strongly sclerotised; juxta trapezoidal, weakly sclerotised and topped on its upper part by two very sclerotised finger-like processes covered with small spines; vinculum short, triangular and bilobed; short cylindrical aedeagus with one apical spine and four cornuti comparable in size. Female genitalia: Anal papillae slightly sclerotised with fine, medium-sized posterior processes; 8th segment narrow with medium-sized, thin anterior processes; bell-shaped ostium with raised edge and strongly sclerotised; lamella antevaginalis large, elliptical with the long axis in a horizontal position, narrow and elongated, provided (generally) with two strongly sclerotised hooks next to the ostium; ductus bursae short, thin arched with a sclerotised base; an elongated pear-shaped corpus bursae with sclerotised plates (signum) inside. In its upper part it has a membranous wart of appreciable proportions from where the ductus seminalis arises. Immature stages (Fig. 91). The last instar larva is 20 to 25 mm long. Cephalic capsule dark, well-developed, rounded, prothoracic and anal shield with undefined verrucae. Body with dark grey verrucae, all with long, translucent setae mixed with black ones; numerous black spots mixed with grey give it a dark appearance; broad black dorsal line touching D1, then a light grey one between D1 and D2, and then a dark one below D2, with an orange or reddish spot attached to D1 and D2 on the lateral border; sides greywith black spots. The pupa with 2 spines curved to the sides at tip of the abdomen. Pupae of both sexes very similar. The cocoon woven among lichens. Molecular data. The Wittia sororcula samples cluster in a well-supported clade (PP=1). The distance from other species under study (average TN-dist = 8.59) is also large enough to justify its inclusion in a separate genus (Wittia). The published samples available in BOLD form two different BIN clusters (BOLD: AAC1077, n=40; BOLD: AAL5773; n=6). The samples we analysed coincide with the main cluster BOLD: AAC1077 which includes samples from 10 different countries including Austria and Italy, being the second one BOLD: AAL5773 comprised of six samples from Austria and Italy. Because of the inclusion of these two BIN clusters, the internal distance for the Wittia sororcula BOLD dataset is considerably large (dist=0.85%) but does not relate to any groups of specimens with morphological, or geographical differences. Biology. Univoltine in the northern European regions and bivoltine in the south, where it is on the wing from April to June, and then from August to October. Frequent and abundant, it is found in very diverse biotopes, but always with a moderate level of humidity, showing a preference for mixed and deciduous forests at elevations between sea level and 1000–1200 m. Adults are active by day and both sexes are attracted to bait and artificial light. The larvae feed on lichens on trunks of conifers and Quercus, as well as on beech leaves, conifer needles and certain species of algae (Ylla et al. 2010). In captivity it accepts artificial diet. W. sororcula is the only species of Eilema sensu lato that overwinters as a pupa, unlike the other species in which the larva is the overwintering stage (Ylla et al. 2010). Distribution (Fig. 116). Mediterranean-Asiatic. Its range extends throughout southern and central Europe to the Near East, northern and central Asia. In the Iberian Peninsula it is found in a northern strip of variable width, which extends from Catalonia to Galicia and northern of Portugal, south to mountainous areas of Castellón and Teruel; present in Mallorca., Published as part of Macià, Ramon, Ylla, Josep, Gastón, Javier, Huertas, Manuel & Bau, Josep, 2022, The species of Eilema Hübner, [1819] sensu lato present in Europe and North Africa (Lepidoptera: Erebidae: Arctiinae: Lithosiini), pp. 1-87 in Zootaxa 5191 (1) on pages 46-47, DOI: 10.11646/zootaxa.5191.1.1, http://zenodo.org/record/7144073, {"references":["Hufnagel, J. S. (1766) Zwote Fortsetzung der vierten Tabelle von den Insecten, besonders von denen so genannten Nachteulen als der zwoten Klasse der Nachtvogel hiesiger Gegend. Berlinisches Magazin, 3 (4), 398 - 399.","Ylla, J., Macia, R. & Gaston, F. J. (2010) Manual de identificacion y guia de campo de los Arctidos de la Peninsula Iberica y Baleares. Arganda Editio, Barcelona, 290 pp."]}

Genus Wittia de Freina, 1980 Wittia de Freina, 1980. Nachrichtenblatt der Bayerischen Entomologen 29 (4):80. Type species: Bombyx aureola Hübner, [1803] 1796; Sammlung Europäischer Schmetterlinge 3: pl. 24, fig. 98, subsequent designation by Moore, 1878; Proceedings of the Zoological Society of London 1878: 18. Diagnosis. Imago. Forewings moderately wide with a curvature of the costal margin, yellow, without pattern. Male genitalia. Uncus very long and narrow; valvae ovoid and weakly sclerotised; saccular process apically sclerotised, long, curved upwards; juxta without apical process; vinculum short, bilobed. Aedeagus with an apical process; vesica short pouch-shaped with indistinct short lobes and several rather similar cornuti. Female genitalia. Very short and narrow ductus bursae, moderately sclerotised and the anterior partially sclerotised appendix, in part helical. The corpus bursae is relatively small, discoidal-globular. Molecular data. A single species of this genus (Wittia sororcula) has been analysed, so we cannot be certain if the results presented for this species would also apply to the whole genus. According Dubatolov (2011), there are many more species included in Wittia, for example Wittia klapperichi (Daniel, 1954) and Wittia yazakii Dubatolov, Kishida & Wang, 2012. None of them have been available to the authors and so it has not been possible to include their study in the present work. Future molecular studies including species placed in the genus Wittia will provide additional insight into this genus. Taxa included. Wittia sororcula (Hufnagel, 1766), Published as part of Macià, Ramon, Ylla, Josep, Gastón, Javier, Huertas, Manuel & Bau, Josep, 2022, The species of Eilema Hübner, [1819] sensu lato present in Europe and North Africa (Lepidoptera: Erebidae: Arctiinae: Lithosiini), pp. 1-87 in Zootaxa 5191 (1) on page 45, DOI: 10.11646/zootaxa.5191.1.1, http://zenodo.org/record/7144073, {"references":["Freina, J. J. (1980) Wittia nom. nov. (Lepidoptera, Arctiidae) Nachrichtenblatt der Bayerischen Entomologen Bd. 29. S. 80.","Moore, F. (1878) A revision of certain Genera of European and Asiatic Lithosiinae, with characters of new Genera and Species. Proceedings of the Scientific Meetings of the Zoological Society of London, 3 - 37 pl. 1 - 3. https: // doi. org / 10.1111 / j. 1469 - 7998.1878. tb 07927. x","Dubatolov, V. V. & Zolotuhin, V. V. (2011) Does Eilema Hubner, [1819] (Lepidoptera, Arctiidae, Lithosiinae) present one or several genera? Eurasian Entomological Journal, 10 (3), 367 - 379.","Hufnagel, J. S. (1766) Zwote Fortsetzung der vierten Tabelle von den Insecten, besonders von denen so genannten Nachteulen als der zwoten Klasse der Nachtvogel hiesiger Gegend. Berlinisches Magazin, 3 (4), 398 - 399."]}

I Opuntia stricta (Haw.) Haw. Bas.: Cactus strictus Haw. Thorny perennial herb or subshrub, in wooded savannah and disturbed sites such as cropland and fallows. C.N.: pitela, cacto (pt). Uses: food, medicinal. Introduced and naturalized species, native to Tropical America., Published as part of Monteiro, Francisca, Costa, Esperança Da, Kissanga, Raquel, Costa, José Carlos & Catarino, Luís, 2022, An annotated checklist of the vascular flora of Quiçama National Park, Angola, pp. 1-67 in Phytotaxa 557 (1) on page 23, DOI: 10.11646/phytotaxa.557.1.1, http://zenodo.org/record/6985699

Subtribe Trichocereinae Britton & Rose (1920): —Most of the genera in Subtribe Trichocereine are also found in the traditional concept “Trichocereae” and the same applies to all the five Chilean genera [Haageocereus Backeb., Leucostele Backeb., Lobivia Britton & Rose, Oreocereus (A.Berger) Riccob., and Soehrensia Backeb.] in the Subtribe Trichocereinae. Based on DNA sequences and a broad sampling, Schlumperger & Renner (2012) showed that the large genus Echinopsis Zucc. is polyphyletic. Consequently, many former segregate genera were reinstalled, and the names of the Chilean taxa needed to be changed from Echinopsis s.l. into Leucostele Backeb., Lobivia Britton & Rose, and Soehrensia Backeb. (Schlumpberger 2012). Hunt (2012, 2016, see Table 2), accepted Schlumperger’s concept as “alternative”, while Rodríguez et al. (2018) still upheld the Echinopsis s.l. concept. Concerning the genus Leucostele, the old name Cactus coquimbanus Mol. has recently been rejected for being ambiguous (Eggli & Walter 2012) and Wilson (2016). Consequently, all combinations using this basionym must be abandoned. Among them is “ Leucostele coquimbana (Mol.) Schlumpb. ”. A new combination of Cereus nigripilis Phil., which represents the shrubby columnar plants occurring along the coast from the Coquimbo- to the Atacama Region was recently proposed [Leucostele nigripilis (Phil.) P. C. Guerrero & Helmut Walter (see Guerrero & Walter 2019 and Table 2)]. Several new taxa in the genus Leucostele were also recently proposed in different journals. As none of them had been included in the sampling of a molecular-based study, we decided not to accept them as long as they are supported by molecular phylogenies: L. faundezii (Albesiano) Schlumpb., L. pectinifera (Albesiano) Schlumpb., and L. undulosa (Albesiano) Schlumpb. (Korotkova et al. 2021).

Eulychnia: —The genus includes two main clades according to Larridon et al. (2018) and Merklinger et al. (2021). The two groups are morphologically and phylogenetically supported and clearly associated with geographical changes in the Atacama Desert (a proposed key is reported below). According to the results of the phylogenetic analyses three former species-complexes were resolved: Eulychnia acida Phil., E. breviflora Phil., and E. iquiquensis (K.Schum.) Britton & Rose. 1. Ribs steep and narrow; flowers and fruits densely covered with wool; perianth segments lanceolate (“ breviflora -group”)............2 - Ribs broad and flattened; flowers and fruits with less and shorter wool; perianth segments spathulate (“ acida -group”)................6 2. Spines numerous, central ones very long; seed large.................................................................................................. 3. E. breviflora - Spines less numerous and shorter; seed smaller.................................................................................................................................3 3. Pulp orange; areoles with long wool; plants low to medium-sized....................................................................................................4 - Pulp whitish; areoles with shorter wool; plants higher......................................................................................................................5 4. Plants shrubby; branches sub-prostrate, thin........................................................................................................... 2. E. barquitensis - Plants arborescent; branches ascending, thicker..................................................................................................... 7. E. saint-pieana 5. Areoles far apart, wool dark brown........................................................................................................................... 8. E. taltalensis - Areoles closely set, wool greyish............................................................................................................................. 6. E. iquiquensis 6. Flower and fruit areoles spiny....................................................................................................................................... 4. E. castanea - Flower and fruit areoles spineless......................................................................................................................................................7 7. Plants shrubby; branches (sub)prostrate with superior portions pointing upwards; ribs 8–12...........................................................8 - Plants arborescent; branches ascending to erect; ribs up to 16.......................................................................................... 1. E. acida 8. Shrubs low; branches thin, grey-green; flowers with short wool; new areoles with abundant white felt................ 5. E. chorosensis - Shrubs higher; branches thicker, yellowish-green; flowers with in conspicuous hairs; areoles with short grey felt.......................................................................................................................................................................................................... 9. E. vallenarensis

Tribe Cylindropuntieae Doweld: —The only species in this tribe occurring in Chile [Cylindropuntia tunicata (Lehmann) Knuth] has been introduced from Northern Mexico and Arizona., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 82, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038

Tribe Cereeae Salm-Dyck: — Nyffeler & Eggli (2010) proposed a broader circumscription of the tribe Cereeae than traditional concepts (see e.g., Anderson 2001, 2005, Hunt et al. 2013). Lendel (2006) and Ritz et al. (2007) showed, in fact, that the traditionally circumscribed Cereeae and Trichocereeae are not monophyletic. Therefore, we follow Nyffeler & Eggli’s (2010) concept of a tribe Cereeae with three subtribes, i.e. Cereinae Britton & Rose, Rebutiinae Donald (incl. Browningieae Buxb.) and Trichocereinae Britton & Rose., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 95, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Nyffeler, R. & Eggli, U. (2010) A farewell to dated ideas and concepts: molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6: 109 - 149.","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Lendel, A. (2006) Phylogenetic relationships in the tribe Trichocereeae (Cacteae) inferred from cpDNA sequence data analysis. IOS Bulletin 14: 11 - 12","Ritz, C., Martins, L., Mecklenburg, R., Goremykin, V. & Hellwig, F. H. (2007) The molecular phylogeny of Rebutia (Cactaceae) and its allies demonstrates the influence of paleography on the evolution of South American mountain cacti. American Journal of Botany 94: 1321 - 1332. https: // doi. org / 10.3732 / ajb. 94.8.1321"]}

Sphaeropuntia Guggi: — Nyffeler & Eggli (2010) remarked that the exclusively West-Andean species Cumulopuntia sphaerica (C.F.Först.) F. Anderson “is unambiguously shown as a separate lineage”, as it appears in a trichotomy with Austrocylindropuntia Backeb. and Cumulopuntia F. Ritter (Wallace & Dickie, 2002). Also, in Griffith and Porter (2009) the two accessions of C. sphaerica were not placed within the C. boliviana clade, but in a strongly supported trichotomy with Austrocylindropuntia. Finally, Ritz et al. (2012) showed that the well-supported C. sphaerica clade is sister to the C. boliviana clade. The morphology of Sphaeropuntia support above mentioned findings. In fact this genus differs from Cumulopuntia in many character states: shrubs low, 10–15 cm high, forming loose groups Sphaeropuntia is distributed between the latitudes of 33° S and 18°S and from sea-level to the Pre-Cordillera (100–3700 m), while members of the genus Cumulopuntia are mainly distributed in the Altiplano regions (3700–4300 m). According to these findings we accept Sphaeropuntia Gucchi., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 87, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Nyffeler, R. & Eggli, U. (2010) A farewell to dated ideas and concepts: molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6: 109 - 149.","Wallace, R. S. & Dickie, S. L. (2002) Systematic implications of chloroplast DNA sequence variation in subfam. Opuntioideae (Cactaceae) Studies in the Opuntieae (Cactaceae). Succulent Plant Research 6: 9 - 24.","Griffith, M. P. & Porter, J. M. (2009) Phylogeny of Opuntioideae (Cactaceae). International Journal of Plant Science 170: 107 - 116. https: // doi. org / 10.1086 / 593048","Ritz, C. M., Reiker, J., Charles, G., Hoxey, P., Hunt, D., Lowry, M., Stuppy, W. & Taylor, N. (2012) Molecular phylogeny and character evolution in terete - stemmed Andean opuntias (Cactaceae - Opuntioideae). Molecular Phylogenetics and Evolution 65: 668 - 681. https: // doi. org / 10.1016 / j. ympev. 2012.07.027","Stuppy, W. (2002) Seed characters and the classification of the Opuntioideae. Studies in the Opuntieae (Cactaceae). Succulent Plant Research 6: 25 - 58."]}

Eriosyce fulva (F.Ritter) P.C. Guerrero & Helmut Walter comb. nov. ≡ Thelocephala fulva F.Ritter, Kakt. Südamer. 3: 1011. 1980. Holotype:— CHILE: Atacama, Totoral, 1956, Ritter 500 (U0249077! https://data.biodiversitydata.nl/naturalis/specimen/ U %20%200249077)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 93, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038

Eriosyce jussieui (Monv. ex Salm-Dyck) P.C. Guerrero & Helmut Walter, comb. nov. ≡ Echinocactus jussieui Monv. ex Salm-Dyck, Cact. Hort. Dyck 1849: 170 (1850). Type (neotype, designated here):— CHILE: Coquimbo, Elqui, 20 km West of Vicuña, Ritter 252a [SGO121545!, corpus, areoles, spines (includes a small rooted seedling)]., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 93, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038

Tribe Tephrocacteae Doweld: —According to phylogenetic analyses (Ritz et al. 2012, Walker et al. 2018, Majure et al. 2019), the monophyletic clade “ Tephrocacteae ” represents a widely distributed group which includes the genera Pterocactus K.Schum., Maihueniopsis Speg., Tephrocactus Lem., Austrocylindropuntia Backeb., and Cumulopuntia F.Ritter. The study by Ritz et al. (2012) retrieved two main clades, one of which comprises Pterocactus and Maihueniopsis and the other one harbours Tephrocactus and the sister pair Cumulopuntia and Austrocylindropuntia. Former taxonomic concepts (e.g. Hunt et al. 2006: p. 12) suggested a close relationship between Maihueniopsis and Cumulopuntia mainly for their shared gross morphology. Wallace & Dickie’s (2002) results suggest a close relationship between Maihueniopsis and Thephrocactus and, furthermore, Pterocactus was treated as a tribe, an opinion shared by Nyffeler & Eggli (2010). Yet, in Griffith & Porter (2009), Pterocactus was not isolated but was placed together with Maihueniopsis in a basal grade to the rest of the terete-stemmed Opuntioideae. Ritz et al. (2012) showed that Pterocactus is not isolated, appearing as sister to Maihueniopsis. This finding supports Stuppy’s (2001) assumption that the genus might be closely related to Maihueniopsis. Although some morphological characters of Pterocactus are unique in the tribe and the Opuntioideae (the base of the glochids is rounded; the flowers are immersed into the apex of the segment; at an early oncological stage the fruit is not clearly separated from the surrounding tissue of the segment; the funicular girdle of the seed forms a broad papery and undulating wing), it shares some important morphological characters with Maihueniopsis. According to all these findings our classification follows the results of Ritz et al. (2012). According to the results of Ritz et al. one large species-complex was split: Cumulopuntia boliviana (Salm-Dyck) F.Ritter (Table 2)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 82, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Ritz, C. M., Reiker, J., Charles, G., Hoxey, P., Hunt, D., Lowry, M., Stuppy, W. & Taylor, N. (2012) Molecular phylogeny and character evolution in terete - stemmed Andean opuntias (Cactaceae - Opuntioideae). Molecular Phylogenetics and Evolution 65: 668 - 681. https: // doi. org / 10.1016 / j. ympev. 2012.07.027","Walker, J. F, Yang, Y., Feng, T., Timoneda, A., Mikenas, J., Hutchison, V., Edwards, C., Wang, N., Ahluwalia, S., Olivieri, J., Walker-Hale, N., Majure, L. C., Puente, R., Kadereit, G., Lauterbach, M., Eggli, U., Flores-Olvera, H., Ochoterena, H., Brockington, S. F., Moore, M. J. & Smith, S. A. (2018) From cacti to carnivores: improved phylotranscriptomic sampling and hierarchical homology inference provide further insight into the evolution of Caryophyllales. American Journal of Botany 105: 446 - 462. https: // doi. org / 10.1002 / ajb 2.1069","Majure, L. C., Baker, M. A., Cloud-Hughes, M., Salywon, A. & Neubig, K. M. (2019) Phylogenomics in Cactaceae: A case study using the chollas sensu lato (Cylindropuntieae, Opuntioideae) reveals a common pattern out of the Chihuahuan and Sonoran deserts. American Journal of Botany 106: 1327 - 1345. https: // doi. org / 10.1002 / ajb 2.1364","Hunt, D., Taylor, N. & Charles, G. (2006) The New Cactus Lexicon. DH Books, Milborn Port, 526 pp.","Wallace, R. S. & Dickie, S. L. (2002) Systematic implications of chloroplast DNA sequence variation in subfam. Opuntioideae (Cactaceae) Studies in the Opuntieae (Cactaceae). Succulent Plant Research 6: 9 - 24.","Nyffeler, R. & Eggli, U. (2010) A farewell to dated ideas and concepts: molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6: 109 - 149.","Griffith, M. P. & Porter, J. M. (2009) Phylogeny of Opuntioideae (Cactaceae). International Journal of Plant Science 170: 107 - 116. https: // doi. org / 10.1086 / 593048"]}

Copiapoa Britton & Rose: —This endemic genus has formerly been regarded as a member of the tribe Notacacteae Buxb., e.g. by Barthlott & Hunt (1993),Anderson (2001), Hoffmann & Walter (2004), and Hunt et al. (2006). Molecular studies by Nyffeler (2002), Korotkova et al. (2010), Nyffeler & Eggli (2010), Arakaki et al. (2011), and Hernández-Hernández et al. (2011) suggested that Copiapoa is not a member of this tribe as it appears isolated on its own clade. Nyffeler & Eggli (2010) treated Copiapoa as a genus of uncertain relationship (incertae sedis), whereas Korotkova et al. (2010) suggested a close relation between Copiapoa and Calymmanthium F.Ritter, although morphology, ecology, and distribution of the two genera are very different. In Nyffeler (2002) and Hernández-Hernández et al. (2011), both genera appear together in a polytomy. Finally, Hunt et al. (2013) accepted Doweld`s (2002) recognition of a “tribe Copiapoae ”. In consideration of the complex situation and the lack of data, we here consider Nyffeler & Eggli’s (2010) concept (incertae sedis). Several infrageneric classifications of Copiapoa based on morphological characters had been proposed by Ritter (1980) [two subgenera (subgenus Pilocopiapoa F.Ritter and subgenus Copiapoa F.Ritter) and five unnamed sections]. Doweld (2002) proposed three sections (sect. Pilocopiapoa (F.Ritter) Doweld (one serie), sect. Echinopoa Doweld (two series: Echinoidei Doweld and Cinerei Doweld), and sect. Copiapoa (two series: Humilis Doweld and Copiapoa); mainly based on general morphological data (Hunt 2001) and stem mucilage and/or root types. Taylor (2001) proposed two subgenera (subgen. Pilocopiapoa (F.Ritter) F.Ritter) and subgen. Copiapoa (five unformal “groups”: “ marginata ”, “ cinerea ”, “ hypogaea ”, “ cinerascens ”, and “ humilis ”). None of these concepts were corroborated by the molecular phylogeny presented by Larridon et al. (2015) whose data retrieved the four well supported sections: sect. Pilocopiapoa (with 1 species), sect. Mammilopoa Helmut Walter & Larridon (1 species), sect. Copiopoa [with two subsections: subsect. Cinerei (Doweld) Helmut Walter & Larridon (2 species), subsect. Copiapoa (22 species)], sect. Echinopoa (Doweld) Helmut Walter & Larridon (5 species) and two basal unnamed monotypic clades (“ Copiapoa australis ” and “ Copiapoa laui ”) (see the proposed key below). According to the results of the phylogenetic analyses five speciescomplexes were split: C. humilis (Phil.) Hutchison, C. cinerea (Phil.) Britton & Rose, C. taltalensis (Werderm.) Looser, C. montana F.Ritter, and C. coquimbana (Rümpler) Britton & Rose. 1. Bract scales numerous distributed all over pericarpel and hypanthium, axils very woolly; ribs to 3.5 cm high (Sect. PILOCOPIAPOA).......................................................................................................................................................... 31. C. solaris - Bract scales few, mainly near hypanthium rim, axils only rarely with fine tiny hairs; ribs ≤ 2 cm high............................................2 2. Ribs in mature plants dissolved into ±conical tubercles; stems soft to the touch..............................................................................3 - Ribs not dissolved into conical tubercles (except for C. longispina), stems soft or not so................................................................4 3. Tubercles obtuse; epidermis ±pruinose; ribs 8-10 (unnamed Section) ......................................................................... 5. C. australis - Tubercles pronounced; epidermis not pruinose; ribs more numerous................................................................................................5 4. Stems dwarf (1–2 cm diam.); tubercles and spines minute (Unnamed Section.................................................................. 21. C. laui - Stems to 8 cm diam.; tubercles large, spines much longer (Sect. HUMILIS)............................................................... 19. C. humilis 5. Plants usually mound-forming; ribs to 2 cm high; fruits large, 1.5–2.0 cm (Sect. ECHINOPOA)...................................................6 - Plants mound-forming or not; ribs lower; fruits ................................................................................ 10 6. Tubercles inconspicuous; roots fascicular..........................................................................................................................................7 - Tubercles well pronounced; taproots of different lengths..................................................................................................................8 7. Mounds up to 2 × 1.5 m; stems often pruinose; ribs to 30; areoles far apart.............................................................. 11. C. dealbata - Mounds much smaller, ribs to 15; stems never pruinose; areoles closely set......................................................... 14. C. echinoides 8. Stems 5–8 cm diam.; tubercles with large chins.................................................................................................... 16. C. fiedleriana - Stems 8–18 cm diam., usually chinless..............................................................................................................................................9 9. Stems 8–12 cm diam.; new spines black, to 4 cm................................................................................................. 10. C. coquimbana - Stems to 18 cm diam.; new spines brownish, later golden yellow, to 6 cm.................................................................... 3. C. armata 10. Rib number up to 40; roots always fascicular; stem tissue very hard (Subsect. CINAREI.............................................................11 - Rib number 8. C. cinerea - Plants forming large dense mounds; apical wool (orange)-brown.............................................................................. 17. C. gigantea 12. Ribs dissolved into conical tubercles....................................................................................................................... 23. C. longispina - Ribs not so........................................................................................................................................................................................13 13. Stem tissue somewhat hard (except for C. hypogaea)......................................................................................................................14 - Stem tissue ±soft-fleshy...................................................................................................................................................................29 14. Bract-scales on hypanthium and fruits large, broad and fleshy; hilum very large................................................... 26. C. megarhiza - Bract scales and hilum not as above.................................................................................................................................................15 15. Rib number usually 15–25................................................................................................................................................................24 - rib number usually low (8–15).........................................................................................................................................................16 16. Roots fascicular................................................................................................................................................................................17 - Roots usually tuberous or long to short taproots..............................................................................................................................18 17. Stem diam. 4–7 cm............................................................................................................................................... 9. C. conglomerata - Stem diam. larger, to 15 cm................................................................................................................................ 23. C. longistaminea 18. Interior perianth segments yellow, without red mid-stripes; style whitish-yellow..........................................................................19 - Interior perianth segments yellowish with red mid-stripes of different widths; style pink to red....................................................22 19. Plants mound-forming; stems elongating; spines short (1.5–2.5 cm), straight................................................................................20 - Plants solitary to few-headed; stems only slightly elongating; spines long (to 5 cm), partly curved...................... 32. C. taltalensis 20. Rib tubercles much pronounced, furrows undulate.......................................................................................... 30. C. serpentisulcata - Ribs only somewhat tuberculate, furrows not undulate...................................................................................................................21 21. Plants forming loose mounds; stems ca. 8 cm diam.; longer ones prostrate; fruit 1cm......................................... 12. C. decorticans - Plants forming compact mounds; stems apically flattened, to 15 cm diam.; stems not prostrate; fruit 1.5 cm..................................................................................................................................................................................................................... 7. C. cinerascens 22. Plants mound-forming; stems green, not pruinose, to12 × 50 cm; flowers 3.0– 4.5 cm, funnel-form; spines to 5 cm, thick........................................................................................................................................................................................................................23 - Plants solitary to few-headed; stems grey-green, ±pruinose, to 8 × 20 cm; flowers small, (2.5 cm), campanulate; spines short, to 2.5 cm, thin.................................................................................................................................................................... 2. C. aphanes 23. Mounds usually loose, spines to 10, (red)-brown, partly curved................................................................................ 29. C. rupestris - Mounds compact, spines to 20, completely hiding stem, black, straight............................................................... 13. C. desertorum 24. Plants large, forming loose or dense mounds; rib tubercles inconspicuous, areoles closely set in older plants..............................25 - Plants small to medium-sized, not much elongating, few-headed, rib tubercles pronounced; areoles not crowded.....................................................................................................................................................................................................................................27 25. Plants forming large, dense mounds; stems to 15 cm diam., pruinose.................................................................... 4. C. atacamensis - Mounds smaller, loose; stems to 10 cm diam., only sometimes pruinose........................................................................................26 26. Stems light grey-green, sometimes pruinose, spines to 3 cm...................................................................................... 6. C. calderana - Stems green, never pruinose, spines to 5 cm............................................................................................................ 25. C. marginata 27. Plants subglobose, temporarily geophytic; epidermis grey-brown........................................................................... 20. C. hypogaea - Plants globose to somewhat elongate; epidermis grey-green...........................................................................................................28 28. Stems to 10 cm diam., not pruinose; areoles large..................................................................................................... 28. C. montana - Stems 4–7 cm diam.; ± pruinose; areoles smaller...................................................................................................... 22. C. leonensis 29. Plants mound-forming; stems to 10 cm diam.; tubercles not chinned................................................................... 18. C. grandiflora - Plants solitary to few-headed; stems 3–7 cm diam.; tubercles ±chinned.........................................................................................30 30. Ribs broad and flattened............................................................................................................................................ 27. C. mollicula - Ribs narrower and deeper.................................................................................................................................................................31 31. Flowers large, broadly campanulate; stems to 7 cm diam. green, not pruinose................................................... 15. C. esmeraldana - Flowers small, narrowly funnel-form; stems to 4 cm, grey-brown, somewhat pruinose........................................ 1. C. angustiflora Several new taxa in the genus Copiapoa were recently proposed in different journals. As none of them had been included in the sampling of a molecular-based study, we decided not to accept them as long as they are not corroborated by molecular phylogenies: Copiapoa coquimbana subsp. rubrispina Piombetti in Xerophilia 4(3): 76. 2015; Copiapoa longispinea subsp. imperialis Piombetti in Xerophilia 4(3): 78. 2015; Copiapoa corralensis Schaub & Keim in Cactus Explorer 16: 48. 2016; Copiapoa fusca Schaub, Cactus Explorer 16: 42. 2016; Copiapoa humilis subsp. matancillensis Schaub & Keim in Cactus & Co 20(1): 15. 2016. Concerning Copiapoa gigantea Backeb., Hunt et al. (2006) use the epithet “ haseltoniana ” (instead of “ gigantea ”) in the combination C. cinerea subsp. haseltoniana (Backeb.) N.P.Taylor (see species list). The results by Larridon et al. (2015, 2018b) suggest that this taxon is not closely related to C. cinerea and should thus be considered as a valid species. Also, C. gigantea has priority over C. haseltoniana at species level (ICN, Art. 11.2) as below reported., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on pages 88-90, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Hoffmann, A. E. & Walter, H. E. (2004) Cactaceas en la flora de Chile, 2 nd ed. Fundacion Claudio Gay, Santiago de Chile, 307 pp.","Hunt, D., Taylor, N. & Charles, G. (2006) The New Cactus Lexicon. DH Books, Milborn Port, 526 pp.","Nyffeler, R. (2002) Phylogenetic relationships in the Cactus family. American Journal of Botany 89: 312 - 326. https: // doi. org / 10.3732 / ajb. 89.2.312","Korotkova, N., Zabel. I., T., Quandt, D. & Bartlott, W. (2010) A phylogenetic analysis of Pfeiffera and the reistatement of Lymanbensonia as an independently evolved lineage of epiphytic Cactaceae within a new tribe Lymanbensonieae. Willdenowia 40: 151 - 172. https: // doi. org / 10.3372 / wi. 40.40201","Nyffeler, R. & Eggli, U. (2010) A farewell to dated ideas and concepts: molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6: 109 - 149.","Hernandez-Hernandez, T., Hernandez, H. M., De-Nova, J. A., Puente, R., Eguiarte, L. E. & Magallon, S. (2011) Phylogenetic relationships and evolution of growth form in Cactaceae (Caryophyllales, Eudicotyledoneae). American Journal of Botany 98: 44 - 61. https: // doi. org / 10.1002 / tax. 12066","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Ritter, F. (1980) Kakteen in Su ¨ damerika, vol. 3. Friedrich Ritter Selbstverlag, Spangenberg, 381 pp.","Doweld, A. B. (2002) On the phylogeny and systematics of the genus Copiapoa Britton & Rose. Sukkulenty 2001, series I - II 4: 46 - 56.","Hunt, D. (2001) Coping with Copiapoa - continued. Cactaceae Systematics Initiatives 12: 15 - 17.","Taylor, N. P. (2001) Roots and mucilage in Copiapoa. Cactaceae Systematics Initiative 12: 18."]}

Tribe Opuntieae: —The membership of the two Chilean genera (Miqueliopuntia Frič ex F.Ritter and Tunilla D.Hunt & Iliff) in the Opuntieae is corroborated by various studies (e.g. Wallace & Dickie 2002, Griffith & Porter 2009, Majure et al. 2012)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 82, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Wallace, R. S. & Dickie, S. L. (2002) Systematic implications of chloroplast DNA sequence variation in subfam. Opuntioideae (Cactaceae) Studies in the Opuntieae (Cactaceae). Succulent Plant Research 6: 9 - 24.","Griffith, M. P. & Porter, J. M. (2009) Phylogeny of Opuntioideae (Cactaceae). International Journal of Plant Science 170: 107 - 116. https: // doi. org / 10.1086 / 593048","Majure, L. C., Puente, R., Griffith, M. P., Judd, W. S., Soltis, P. S. & Soltis, D. E. (2012) Phylogeny of Opuntia s. s. (Cactaceae): clade delineation, geographic origins, and reticulate evolution. American Journal of Botany 99: 847 - 864. https: // doi. org / 10.3732 / ajb. 1100375"]}

Tephrocactus Lem.: — Kiesling (1984) and Rodríguez et al. (2018) placed Tephrocactus nigrispinus (K.Schum.) Backeb. within the genus Maihueniopsis. This assumption was not supported by molecular studies (Wallace & Dickie 2002, Griffith & Porter 2009, Ritz et al. 2012)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 87, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Kiesling, R. (1984) Estudios en Cactaceae de Argentina: Maihueniopsis, Tephrocactus y generosafines (Opuntioideae). Darwiniana 25: 171 - 215.","Rodriguez, R., Marticorena, C., Alarcon, D., Baeza, C., Cavieres, V., Finot, L., Fuentes, N., Kiessling, A., Mihoc, M., Pauchard, A., Ruiz, E., Sanchez, P. & Marticorena, A. (2018) Catalogo de las plantas vasculares de Chile. Gayana Botanica 75: 1 - 430. http: // dx. doi. org / 10.4067 / S 0717 - 66432018000100001","Wallace, R. S. & Dickie, S. L. (2002) Systematic implications of chloroplast DNA sequence variation in subfam. Opuntioideae (Cactaceae) Studies in the Opuntieae (Cactaceae). Succulent Plant Research 6: 9 - 24.","Griffith, M. P. & Porter, J. M. (2009) Phylogeny of Opuntioideae (Cactaceae). International Journal of Plant Science 170: 107 - 116. https: // doi. org / 10.1086 / 593048","Ritz, C. M., Reiker, J., Charles, G., Hoxey, P., Hunt, D., Lowry, M., Stuppy, W. & Taylor, N. (2012) Molecular phylogeny and character evolution in terete - stemmed Andean opuntias (Cactaceae - Opuntioideae). Molecular Phylogenetics and Evolution 65: 668 - 681. https: // doi. org / 10.1016 / j. ympev. 2012.07.027"]}

Austrocactus: —The Austrocactus species from around the town of Chile Chico was shown to be A. coxii (K.Schum.) Backeb. (see Walter 2019) and not A. patagonicus Hosseus according to Rodríguez et al. (2018) and Hunt et al. (2013, 2016). A. coxii can easily be distinguished from A. patagonicus (= A. bertinii Britton & Rose) by large multi-headed cushions (vs. simple, rarely few-headed in A. patagonicus), branches 10–15 cm long (vs. 50–60 cm) and 5 cm thick (vs. to 15 cm), all spines straight (vs. hooked), centrals 2.0– 2.5 cm (vs. 2–4 cm), flowers yellow to orange (vs. pinkish to white)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 88, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Walter, H. E. (2019) The cacti of the Chilean Patagonia. Cactus World 37: 49 - 56.","Rodriguez, R., Marticorena, C., Alarcon, D., Baeza, C., Cavieres, V., Finot, L., Fuentes, N., Kiessling, A., Mihoc, M., Pauchard, A., Ruiz, E., Sanchez, P. & Marticorena, A. (2018) Catalogo de las plantas vasculares de Chile. Gayana Botanica 75: 1 - 430. http: // dx. doi. org / 10.4067 / S 0717 - 66432018000100001","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Hunt, D. (2016) CITES Cactaceae Checklist Third Edition. Milborne Port, Remous Limited, 175 pp."]}

Copiapoa gigantea Backeb., Jahrb. Deutsche Kakt. Ges. 1: 104. 1936. Type:—Not designated. = Copiapoa eremophila F. Ritter, Kakt. Südam. 3: 1104–1105. 1980. Type:— CHILE, östl. Paposo, s.d. [1956], Ritter 208b [holotype SGO124843, (corpus), areoles, spines]. Notes: —Backeberg (1936: 104) gave a vague type locality (“ Chile: Provinz Antofagasta, auf nachts oft nebelfeuchten Berggipfeln” = “ Chile: Province Antofagasta, on mountain peaks that are often damp at night”) in the protologue of his Copiapoa gigantea. Later, the same author (Backeberg 1977: 107) specified the locality as “above Paposo”. Ritter (1980: 1100) accepted Backeberg’s species for plants occurring east of Paposo area (“Gegen die Wüste des Hinterlandes wächst sie aber nur gegen Süden, während sie im gleichen Klima und in gleicher Höhenlage etwas nördlicher, nämlich östlich von PAPOSO...” = “Towards the desert of the hinterland, however, it [Copiapoa gigantea var. gigantea] only grows towards the south, while in the same climate and at the same altitude it grows a little further north, namely east of PAPOSO...”) and proposed C. haseltoniana Backeb. at variety rank of C. gigantea (Ritter 1980: 1101) for plant occurrring in “... nördlich Paposo an der Küste...” (= “... north of Paposo on the coast...”). Concerning Copiapoa eremophila, Ritter (1980: 1105) reported “Typusort. östlich von PAPOSO am Rand der Vollwüste als einzige noch wachsende Kakteenart; nur von hier bekannt. Von mir entdeckt 1956. Nr. FR 476 (=208a)”, the “Typusort” can be considered as “ holotype ”. We traced this specimen at SGO. Its morphology corresponds to the current concept of C. gigantea (see e.g., Larridon et al. 2015). The “current” concept of Copiapoa gigantea (based on molecular evidence) is Larridon 2015. There are no other works occupying with the status of C. gigantea. Former concepts treated C. gigantea as a synonym of C. cinerea subsp. haseltonia (Hunt et al., Hoffmann & Walter (2004), Anderson (2001) as a synonym of Copiapoa haseltonia and Slaba, (in Kaktussy (special) 33: 3. 1997) as C. cinerea subsp. gigantea for sharing the following characters: Apical wool orange-brown; large dense multi-headed mounds; stem diameter 10–25 cm, farina-covered; spines honey-coloured, later turning greyish-blackish, more numerous at higher elevations; young areoles orange brown.Accordingly, and since the type locality of C. eremophila is nearly the same of that of C. gigantea (Paposo), we here propose to synonymize the two names., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 90, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Ritter, F. (1980) Kakteen in Su ¨ damerika, vol. 3. Friedrich Ritter Selbstverlag, Spangenberg, 381 pp.","Hoffmann, A. E. & Walter, H. E. (2004) Cactaceas en la flora de Chile, 2 nd ed. Fundacion Claudio Gay, Santiago de Chile, 307 pp."]}

Maihueniopsis Speg.: — Hunt’s et al. (2016) and Hunt’s (2013) circumscription of the genus Maihueniopsis includes only four species occurring in Chile, i.e. the two endemics M. archiconoidea F.Ritter and M. conoidea F.Ritter, and two species occurring also in Argentina (M. darwinii Hnsl. and M. ovata Pfeiff.). These authors merged the rest of the Chilean species described by Ritter (1980) and Espinosa (1933) into the widespread Argentinean species M. glomerata (Haw.) R.Kiesling without any evidence (i.e. detailed descriptions, morphological keys, and/or molecular phylogenies). They did not realize, however, that the Chilean Maihueniopsis are clearly divided into two main groups (the “ glomerata - group” and the “ domeykoensis- group”) when all members of the domeykoensis group [M. camachoi (Espinosa) F.Ritter, M. colorea F.Ritter, M. crassispina F.Ritter, M. domeykoensis F.Ritter, M. grandiflora F.Ritter, and M. wagenknechtii F.Ritter] are included in the sampling. All members of the domeykoensis group are endemic to the western slopes of the Chilean Andes. Populations of M. glomerata s.str. had not yet been documented in Chile. The molecular-based phylogeny presented by Ritz et al. (2012) shows that M. domeykoensis [a species that was lumped into M. glomerata by Hunt (2011)] is sister to the rest of the nine taxa (all of them being members of the glomerata group). Ritz et al. (2012) found it “interesting” that M. domeykoensis was not placed within the “ glomerata ” group. Their sampling was unbalanced, as they included nine species assigned to the “ glomerata ” group (see above) but only one assigned to the “ domeykoensis ” group (i.e. M. domeykoensis). Maihueniopsis domeykoensis is likely a separate lineage. So, we conclude that a) as M. domeykoensis did not cluster with M. glomerata (or other members of the glomerata -clade), it cannot be considered a synonym of M. glomerata, and thus b) the rest of the members of the domeykoensis group, who share the same morphological characters with M. domeykoensis, cannot be considered synonyms of M. glomerata. This conclusion is corroborated by a broadly sampled (i.e. all the members of the domeykoensis group) molecular phylogeny (Guerrero, in preparation) that showed that Maihueniopsis is split into two groups congruent with the above presented morphylogy-based groups (a morphologybased key is reported below). According to the results of the phylogenetic analyses one large species-complex was split: Maihueniopsis glomerata. However, Hunt (2016) withdrew his opinion and reinstalled all of the formerly not accepted species of the domeykoensis-clade. A nomenclatural note on the no validly published name “ Maihueniopsis leoncito (Werderm.) F.Ritter ” is necessary. According to Art. 41.5 ICN (Turland et al. 2018), Ritter’s “ Maihueniopsis leoncito ” is invalid because it does not include the place of publication of the basyonym. The correct name Maihueniopsis leoncito (Werdermann ex F.Ritter) P.C. Guerrero & Helmut Walter was recently published by Guerrero & Helmut Walter (2019); 1. Branch segments ≤ 5 cm; spines ± flattened; mature fruits unspined, ±ovoid or cylindric; seeds pale yellow, with short trichomes (“ glomerata -group”)...........................................................................................................................................................................2 - Branch segments 5–10 cm, spines terete in cross-section; fruits spiny, obconical urn-shaped; seeds brown, with long trichomes (“ domeykoensis -group”).....................................................................................................................................................................7 2. Spines strongly flattened and broad at base, often deflexed..............................................................................................................3 - Spines less flattened, not broad at base, thinner, not deflexed...........................................................................................................4 3. Compact, large cushions 20–60 cm high; segments to 4 cm, broadly conical, apex acute; spines to 4 cm, pointed downwards to porrect......................................................................................................................................................................... 10. M. leoncito - Small, loose cushions up to 20 cm diam. and 10 cm high; segments 1. M. archiconoidea 4. Branch segments M. darwinii - Cushions loose, to 10 cm high; areoles few, 1–2 mm, only upper third to upper half spiniferous; stigma-lobes red.................................................................................................................................................................................................................. 11. M. ovata 6. Small, loose cushions; spines directed sideward to ±downwards, new ones blackish-violet; fruits cylindric, areoles wooly................................................................................................................................................................................................ 5. M. conoidea - Large compact cushions; spines porrect to erect, new ones yellowish; fruits ovoid, areoles not woolly............. 2. M. atacamensis 7. Areole number M. wagenknechtii 8. Spines red-brown, base whitish; seeds small.....................................................................................................................................9 - Spines white or light honey-brown; seeds large...............................................................................................................................10 9. Stigma lobes red; spines to 12 cm.............................................................................................................................. 3. M. camachoi - Stigma lobes green; spines to 6 cm................................................................................................................................ 4. M. colorea 10. Shrubs up to 2 m diam.; spines white, those on flowers and fruits contorted........................................................ 9. M. grandiflora - Shrubs to 50 cm diam.; spines honey-brown, stiff...........................................................................................................................11 11. Segments light yellowish-green; most spines directed upwards; stigma green.................................................. 8. M. domeykoensis - Segments bluish grey-green; most spines directed; sideward or downwards; stigma ruby-red............................... 6. M. crassispina, Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on pages 86-87, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Hunt, D. (2016) CITES Cactaceae Checklist Third Edition. Milborne Port, Remous Limited, 175 pp.","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Ritter, F. (1980) Kakteen in Su ¨ damerika, vol. 3. Friedrich Ritter Selbstverlag, Spangenberg, 381 pp.","Espinosa, M. R. (1933) Una Opuntia nueva chilena. Revista Chilena de Historia Natural 37: 126 - 130.","Ritz, C. M., Reiker, J., Charles, G., Hoxey, P., Hunt, D., Lowry, M., Stuppy, W. & Taylor, N. (2012) Molecular phylogeny and character evolution in terete - stemmed Andean opuntias (Cactaceae - Opuntioideae). Molecular Phylogenetics and Evolution 65: 668 - 681. https: // doi. org / 10.1016 / j. ympev. 2012.07.027","Hunt, D. (2011) Classification of the \" cyndroid \" opuntias of South America. Cactaceae Systematics Initiatives 25: 5 - 29.","Walter, H. E. (2019) The cacti of the Chilean Patagonia. Cactus World 37: 49 - 56."]}

The relevance of classifying the Cactaceae family into different subfamilies helps to better understand the subdivision into clades that differ morphologically (see key below). On the other hand, the delimitation of some subfamilies has shown to been difficult (for example the paraphyly of the Pereskioideae K.Schum. and the ambiguous relationships of the Maihuenioideae P.Fearn with other subfamilies). Also, the tribes within the Opuntioideae K.Schum. and Cactoideae Eaton have undergone several changes, as molecular-based phylogenies provided new information about evolutionary relationships of its members. Most of the proposed changes at these taxonomic levels have an impact on the classification of Chilean cacti because the cactus flora in the Atacama Desert, the Altiplano, the Mediterranean area, and the Chilean Patagonian steppe harbours three of the four subfamilies and seven of the ten tribes. 1. Stems with functional leaves, at least on immature shoots; a 739 bp non-coding intron region in the chloroplast-encoded gene rpo C 1 present.....................................................................................................................................................................................2 - Stems without functional leaves; a 739 bp non-coding intron region in the chloroplast-encoded gene rpo C 1 is lost (subfam. Cactoideae).........................................................................................................................................................................................3 - Stems without functional leaves; a 739 bp non-coding intron region in the chloroplast-encoded gene rpo C 1 is lost (Subfamily CACTOIDEAE)................................................................................................................................................................................10 2. Leaves persistent, terete; glochids on areoles absent; seeds without funicular envelope; stems astomatous, barky... (subfam. Maihuenioideae).......................................................................................................................................................... 12. Maihuenia - Leaves ephemeral, flat; areoles with spines and glochids; seeds encased in a funicular envelope; stems stomatous, bark formation delayed (subfam. Opuntioideae).........................................................................................................................................................4 4. Branch segments flat to subterete; shrubs to 20 cm high.................................................................................................... 22. Tunilla - Segments cylindric; shrubs to 1.5 m high........................................................................................................... 14. Miqueliopuntia 3. Branches flattened to subterete or cylindric.........................................................................................................Tribe OPUNTIEAE - Branch segments never flattened, terete in cross-section...................................................................................................................5 5. Shrubs to 60 cm high, then mound-forming; segments to 10 cm, with determinate growth; spines without sheath..................................................................................................................................................................................... TEPHROCACTEAE (pro parte) - Shrubs to 1.5 m; branch segments to 30 cm, with indeterminate growth; spines with a papery sheath (CYLINDROPUNTIEAE)................................................................................................................................................................................. 6. Cylindropuntia 6. Flowers dark red; petaloids few, 21. Tephrocactus (nigrispinus) - Flowers not dark red; petaloids>10; fruits 2–5 cm, greenish yellow, not glabrous, pulp never red.................................................7 7. Seeds laterally compressed; perisperm large......................................................................................................................................8 - Seeds ± globose; perisperm small......................................................................................................................................................9 8. Flowers immersed into apex; funicular girdle of seed formed into a papery wing.................................................... 17. Pterocactus - Flowers not immersed; seeds not winged.............................................................................................................. 13. Maihueniopsis 9. Segments ±globose, etuberculate, easily detaching; spines on segments and fruits 20. Sphaeropuntia - Segments ±conical, tuberculate; not detaching; spines much longer; areoles crowded towards apex; fruits cylindric; lateral ridges present......................................................................................................................................................................5. Cumulopuntia 10. Hypanthium shorter than pericarpel; stems always cylindric (PHYLLOCACTEAE subtribe CORRYOCACTINAE).................11 - Hypanthium longer than pericarpel; stems cylindric or not.............................................................................................................13 11. Plants low, branches 10–50 cm; central spines 1. Austrocactus - Plants tall, 0.8–5 m; central spines longer, to 15 cm; flowers densely covered with large bracts; fruits 5–10 cm..........................12 12. Flowers to 7 cm, whitish, narrow funnelform................................................................................................................ 8. Eulychnia - Flowers 8–11 cm, yellow, campanulate..................................................................................................................... 4. Corryocactus 13. Stems ±globose to elongating, small to medium-sized; flowers 1.5-6 cm; fruits never with pulp..................................................14 - Stems cylindric, (subcylindric in Lobivia ferox), mostly tall; flowers large, 7-18 cm; fruits with pulp or pulpless (CEREEAE)...17 14. Fruit dehiscing apically, funiculi juicy at fruit maturity (seed ant-dispersed) (INCERTAE SEDIS).............................. 3. Copiapoa - Fruit dehiscion not as above; funiculi dry at fruit maturity (seed not ant-dispersed) (Tribe NOTOCACTEAE)............................15 15. Flowers 2–7 cm, bract scales numerous, axils woolly and bristly; fruit always dehiscing basally (except for Eriosyce rodentiophila).................................................................................................................................................................... 7. Eriosyce - Flowers small, to 2 cm, bract scales few, axils naked; fruit dehiscence lateral or basal..................................................................16 16. Stems 1–3 cm, areoles subtended by tiny persistent leaf; fruits elongating to 3 cm, dehiscing basally.................... 18. Rimacactus - Stems to 15 cm diam., leaf absent; fruits .............................................................15. Neowerdermannia 17. Flower scales large, fleshy, imbricate; axils naked; stems with densely branched crown (subtribe REBUTIINAE).. 2. Browningia - Flower scales small, not fleshy, not imbricate; axils with ± wool (subtribe TRICHOCEREINAE)................................................18 18. Flowers brightly carmine red, zygomorphic; fruits without pulp, dehiscing basally................................................... 6. Oreocereus - Flowers never carmine red, actinomorphic; fruits, with pulp never dehiscing basally....................................................................19 19. Stamens inserted in a single series; fruits indehiscent.............................................................................................. 9. Haageocereus - Stamens inserted in two series; fruits dehiscent...............................................................................................................................20 20. Plants> 0.5 m, branches columnar or thickly cylindric....................................................................................................................21 - Plants 11. Lobivia [ferox] 21. Plants ±branching, shrubby or treelike; flowers narrowly funnelform, whitish........................................................... 10. Leucostele - Plants usually unbranched, thickly cylindric; flowers campanulate, yellow............................................................... 19. Soehrensia The comparison of the taxonomic classifications shows that there are important differences in the number of Chilean genera and species (Table 1). Hunt’s classification (2006, 2013, 2016) includes the lowest number of species (95), whereas Rodríguez et al. (2018) reports the lowest number of genera (17). The percentage of endemic species also varies: 73% by Hunt et al. (2006, 2013, 2016), 74% by Rodríguez et al. (2018), 81% in the present study. These results highlight the importance of updating the taxonomic classifications in the Flora of Chile, because underestimating the endemism and number of species may have major negative effects on conservation (see e.g., Mace 2004, Duarte et al. 2014). The number of species accepted here means that the Cactaceae represent 6% of the Chilean native angiosperms and 5% of endemic species of Chile (Rodríguez et al. 2018)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on pages 80-81, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Rodriguez, R., Marticorena, C., Alarcon, D., Baeza, C., Cavieres, V., Finot, L., Fuentes, N., Kiessling, A., Mihoc, M., Pauchard, A., Ruiz, E., Sanchez, P. & Marticorena, A. (2018) Catalogo de las plantas vasculares de Chile. Gayana Botanica 75: 1 - 430. http: // dx. doi. org / 10.4067 / S 0717 - 66432018000100001","Hunt, D., Taylor, N. & Charles, G. (2006) The New Cactus Lexicon. DH Books, Milborn Port, 526 pp.","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Hunt, D. (2016) CITES Cactaceae Checklist Third Edition. Milborne Port, Remous Limited, 175 pp.","Mace, G. M. (2004) The role of taxonomy in species conservation. Philosophical transactions of the Royal Society of London. Series B, Biological sciences 359: 711 - 719. https: // doi. org / 10.1098 / rstb. 2003.1454","Duarte, M., Guerrero, P. C., Carvallo, G. & Bustamante, R. O. (2014) Conservation network design for endemic cacti under taxonomic uncertainty. Biological Conservation 176: 236 - 242."]}

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 - S, Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on pages 91-94, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Guerrero, P. C., Walter, H. E., Arroyo, M. K. T, Pena, C., Tamburrino, I., De Benedictis, M. & Larridon, I. (2019 b) Molecular phylogeny of the large South American genus Eriosyce (Notocacteae, Cactaceae): Generic delimitation and proposed changes in infrageneric and species ranks. Taxon 68: 557 - 573. https: // doi. org / 10.1002 / tax. 12066","Kattermann, F. (1994) Eriosyce (Cactaceae) The genus revised and amplified. Succulent Plant Research 1: 5 - 176.","Hunt, D., Taylor, N. & Charles, G. (2006) The New Cactus Lexicon. DH Books, Milborn Port, 526 pp.","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Vergara, E. (2011) World's driest desert hits snow rain. NBC News. Available from http: // www. nbcnews. com (accessed 27 January 2022).","Hunt, D. (2003) Subg. Horridocactus. Cactaceae Systematics Initiatives 16: 9 - 10.","Hoffmann, A. E. & Walter, H. E. (2004) Cactaceas en la flora de Chile, 2 nd ed. Fundacion Claudio Gay, Santiago de Chile, 307 pp.","Ritter, F. (1980) Kakteen in Su ¨ damerika, vol. 3. Friedrich Ritter Selbstverlag, Spangenberg, 381 pp.","Salm-Dyck, J. (1849) Cactaceae Hortus Dyckensis 34: 170 - 171."]}

Eriosyce fulva (F.Ritter) P.C. Guerrero & Helmut Walter comb. nov. ≡ Thelocephala fulva F.Ritter, Kakt. Südamer. 3: 1011. 1980. Holotype:— CHILE: Atacama, Totoral, 1956, Ritter 500 (U0249077! https://data.biodiversitydata.nl/naturalis/specimen/ U %20%200249077).

Tribe Opuntieae: —The membership of the two Chilean genera (Miqueliopuntia Frič ex F.Ritter and Tunilla D.Hunt & Iliff) in the Opuntieae is corroborated by various studies (e.g. Wallace & Dickie 2002, Griffith & Porter 2009, Majure et al. 2012).

Tephrocactus Lem.: — Kiesling (1984) and Rodríguez et al. (2018) placed Tephrocactus nigrispinus (K.Schum.) Backeb. within the genus Maihueniopsis. This assumption was not supported by molecular studies (Wallace & Dickie 2002, Griffith & Porter 2009, Ritz et al. 2012).

Tribe Phyllocacteae Salm-Dyck: — Nyffeler & Eggli (2010) stated that the monophyly of the tribe is well supported by Nyffeler (2002) and Crozier (2005). Moreover, for nomenclatural reasons, the name Phyllocacteae (incl. Echinocereeae Buxb.) must be used because it has priority over all the other relevant tribal names, i.e. Hylocereeae Buxb., Leptocereeae Buxb., Pachycereeae Buxb. and Peniocereeae Doweld. Despite of this, Hunt et al. (2013) placed the three Chilean genera (Austrocactus Britton & Rose, Eulychnia Phil., and Corryocactus Britton & Rose) within Echinocereeae Buxb., Nyffeler & Eggli (2010) proposed the following three subtribes, one of which (Subtribe Corryocactinae Buxb.) includes the three Chilean genera. The relationships between Austrocactus, Eulychnia, and Corryocactus was corroborated by various molecular-based studies (e.g. Nyffeler 2002, Hernández-Hernández et al. 2011, Bárcenas et al. 2011)., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 88, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Nyffeler, R. & Eggli, U. (2010) A farewell to dated ideas and concepts: molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6: 109 - 149.","Nyffeler, R. (2002) Phylogenetic relationships in the Cactus family. American Journal of Botany 89: 312 - 326. https: // doi. org / 10.3732 / ajb. 89.2.312","Crozier, B. S. (2005) Systematics of Cactaceae Juss.: phylogeny cpDNA evolution and classification, with emphasis on the genus Mammilaria Haw. Dissertation, The University of Texas at Austin, 154 pp.","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp.","Hernandez-Hernandez, T., Hernandez, H. M., De-Nova, J. A., Puente, R., Eguiarte, L. E. & Magallon, S. (2011) Phylogenetic relationships and evolution of growth form in Cactaceae (Caryophyllales, Eudicotyledoneae). American Journal of Botany 98: 44 - 61. https: // doi. org / 10.1002 / tax. 12066"]}

Eriosyce jussieui (Monv. ex Salm-Dyck) P.C. Guerrero & Helmut Walter, comb. nov. ≡ Echinocactus jussieui Monv. ex Salm-Dyck, Cact. Hort. Dyck 1849: 170 (1850). Type (neotype, designated here):— CHILE: Coquimbo, Elqui, 20 km West of Vicuña, Ritter 252a [SGO121545!, corpus, areoles, spines (includes a small rooted seedling)].

Subtribe Rebutiinae Donald. (incl. Browningieae Buxb.): —Various molecular-based studies (e.g. Lendel et al. 2006, Ritz et al. 2007) showed that Browningia Britton & Rose is closely related to Rebutia K.Schum., while other authors using only morphology-based characters (e.g. Hunt et al. 2013, Anderson 2001, 2005) placed the two genera in different tribes: Hunt et al. (2013) placed Browningia in tribe Cereeae and Rebutia in tribe Trichocereeae, whereas Anderson (2001, 2005) placed Rebutia in Trichocereeae and Browningia in Browningieae (together with Stetsonia Britton & Rose, Jasminocereus Britton & Rose, Armatocereus Backeb. and Neoraimondia Britton & Rose). We here accept to include Browningieae in the tribe Rebutiinae., Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 95, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Lendel, A. (2006) Phylogenetic relationships in the tribe Trichocereeae (Cacteae) inferred from cpDNA sequence data analysis. IOS Bulletin 14: 11 - 12","Ritz, C., Martins, L., Mecklenburg, R., Goremykin, V. & Hellwig, F. H. (2007) The molecular phylogeny of Rebutia (Cactaceae) and its allies demonstrates the influence of paleography on the evolution of South American mountain cacti. American Journal of Botany 94: 1321 - 1332. https: // doi. org / 10.3732 / ajb. 94.8.1321","Hunt, D., Taylor, N. & Charles, G. (2013) The New Cactus Lexicon, Atlas of Illustrations. Dh Books, Milborn Port, 373 pp."]}

Eulychnia: —The genus includes two main clades according to Larridon et al. (2018) and Merklinger et al. (2021). The two groups are morphologically and phylogenetically supported and clearly associated with geographical changes in the Atacama Desert (a proposed key is reported below). According to the results of the phylogenetic analyses three former species-complexes were resolved: Eulychnia acida Phil., E. breviflora Phil., and E. iquiquensis (K.Schum.) Britton & Rose. 1. Ribs steep and narrow; flowers and fruits densely covered with wool; perianth segments lanceolate (“ breviflora -group”)............2 - Ribs broad and flattened; flowers and fruits with less and shorter wool; perianth segments spathulate (“ acida -group”)................6 2. Spines numerous, central ones very long; seed large.................................................................................................. 3. E. breviflora - Spines less numerous and shorter; seed smaller.................................................................................................................................3 3. Pulp orange; areoles with long wool; plants low to medium-sized....................................................................................................4 - Pulp whitish; areoles with shorter wool; plants higher......................................................................................................................5 4. Plants shrubby; branches sub-prostrate, thin........................................................................................................... 2. E. barquitensis - Plants arborescent; branches ascending, thicker..................................................................................................... 7. E. saint-pieana 5. Areoles far apart, wool dark brown........................................................................................................................... 8. E. taltalensis - Areoles closely set, wool greyish............................................................................................................................. 6. E. iquiquensis 6. Flower and fruit areoles spiny....................................................................................................................................... 4. E. castanea - Flower and fruit areoles spineless......................................................................................................................................................7 7. Plants shrubby; branches (sub)prostrate with superior portions pointing upwards; ribs 8–12...........................................................8 - Plants arborescent; branches ascending to erect; ribs up to 16.......................................................................................... 1. E. acida 8. Shrubs low; branches thin, grey-green; flowers with short wool; new areoles with abundant white felt................ 5. E. chorosensis - Shrubs higher; branches thicker, yellowish-green; flowers with in conspicuous hairs; areoles with short grey felt.......................................................................................................................................................................................................... 9. E. vallenarensis, Published as part of Walter, Helmut E. & Guerrero, Pablo C., 2022, Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy, pp. 79-98 in Phytotaxa 550 (2) on page 88, DOI: 10.11646/phytotaxa.550.2.1, http://zenodo.org/record/6641038, {"references":["Merklinger, F. F., Bohnert, T., Arakaki, M., Weigend, M. & Quandt, D. (2021) Luebert F. Quaternary diversification of a columnar cactus in the driest place on earth. American Journal of Botany 108: 184 - 199. https: // doi. org / 10.1002 / ajb 2.1608"]}

Aerangis luteoalba (Kraenzl.) Schltr. var. rhodosticta (Kraenzl.) J.Stewart — Habit: Herb. Habitat: LMWF; up to 2 400 m. Distribution: II. Voucher: East Mount Kenya Forest, Alt. 1 524–1 829 m, Battiscombe K692 (EA, K). References: Blundell (1987), Cribb (1989b), Stewart & Campbell (2003), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 25, DOI: 10.11646/phytotaxa.546.1.1

6. Opuntia Miller (1754: 974) Type (indicated by Linnaeus 1753: 468):— Cactus opuntia L. Species richness:— Opuntia is a genus with wide geographic distribution and comprises approximately 200 species. In Brazil, there are 5 species of which ocurr in Rio Grande do Sul (Carneiro et al. 2016). General distribution:— The genus occurs from southern Canada to Uruguay, with the largest number of species in Mexico (Kiesling 2005). Distribution in Santa Maria river basin:— In SMRB only Opuntia megapotamica Arechavaletae (1905: 42) occurs (Fig. 3A) and it has a wide distribution in fields, rocky outcrops and woods. Morphologic notes:— The species is characterized by its shrub or arboreal size, with articulated branches, acicular thorns and usually diurnal orange or yellow flowers with fleshy fruits. It blooms from October to March, bearing fruit shortly after flowering., Published as part of Oliveira, Anabela Silveira De, Pontes, Rodrigo Corrêa, Robaina, Luis Eduardo De Souza & Trentin, Romário, 2022, The Cactaceae family in the Santa Maria river basin (Rio Grande do Sul, Brazil), pp. 153-166 in Phytotaxa 542 (2) on page 161, DOI: 10.11646/phytotaxa.542.2.3, http://zenodo.org/record/6413545, {"references":["Linnaeus, C. (1753) Species plantarum, vol. 1. Salvius, Stockholm, 560 pp.","Carneiro, A. M., Farias-Singer, R., Ramos, R. A. & Nilson, A. D. (2016) Cactos do Rio Grande do Sul. Fundacao Zoobotanica do Rio Grande do Sul, Porto Alegre, 224 pp.","Kiesling, R. (2005) Cactaceae. In: Burkart, A. & Bacigalupo, N. (Eds.) Flora ilustrada de Entre Rios - Argentina, Tomo IV, Parte IV. Coleccion Cientifica del I. N. T. A, Buenos Aires, pp. 441 - 444."]}

4. Gymnocalycium Pfeiffer ex Mittler (1844:124) Type (designated by Haworth 1816: 137):— Gymnocalycium gibbosum (Haworth 1816: 137) Pfeiffer ex Mittler. Species richness:— Gathers approximately 50 species, four species occur in Rio Grande do Sul (Kiesling 2005). General distribution:— Gymnocalycium is distributed in southern Brazil, Uruguay, Argentina, Paraguay, and Bolivia. Distribution in Santa Maria river basin:— Two species were found, Gymnocalycium denudatum (Link & Otto 1828: 9) Pfeiffer ex Mittler (1844: 124) (Fig. 2K), and G. uruguayense (Arechavaletae 1905: 218) Britton & Rose (1922: 172) (Fig. 2L), they are widespread in the study area and grow on rocky outcrops and the Saxon vegetation, tolerate low light. Morphologic notes:— These are usually solitary or clustered plants, globose of more than 4 cm in diameter, 4–15 sides, bell-shaped, white calyx, pink or yellowish-white flowers. Dried or fleshy fruits up to 3 cm in length. They usually bloom from October to December, bearing fruit soon after., Published as part of Oliveira, Anabela Silveira De, Pontes, Rodrigo Corrêa, Robaina, Luis Eduardo De Souza & Trentin, Romário, 2022, The Cactaceae family in the Santa Maria river basin (Rio Grande do Sul, Brazil), pp. 153-166 in Phytotaxa 542 (2) on page 161, DOI: 10.11646/phytotaxa.542.2.3, http://zenodo.org/record/6413545, {"references":["Mittler, L. (1844) Taschenbch fur Cactusliebliebhaber, Lei Mittler, L. (1844). Taschenbch fur Cactusliebliebhaber, Leipzig 2: 124.","Kiesling, R. (2005) Cactaceae. In: Burkart, A. & Bacigalupo, N. (Eds.) Flora ilustrada de Entre Rios - Argentina, Tomo IV, Parte IV. Coleccion Cientifica del I. N. T. A, Buenos Aires, pp. 441 - 444.","Britton, N. L. & Rose, J. N. (1922) The Cactaceae. Descriptions and Illustrations of Plants of the Cactus Family, vols. 3 - 4 Carnegie Institution, Washington, 322 pp."]}

3. Frailea Britton & Rose (1922: 208) Type (designated by Britton & Rose 1922: 210):— Frailea cataphracta Britton & Rose. Species richness:— Frailea comprises 19 species according to Gerloff (2001). Twelve are endemic, recognized only for Rio Grande do Sul State (Pontes et al. 2018). General distribution:— Frailea has a restricted geographical distribution to southern Brazil, Paraguay, Uruguay, southeastern Bolivia, and northeast Argentina. Distribution in Santa Maria river basin:— In SMRB occur nine species, Frailea albifusca, (Fig. 2C and Fig. 2F), F. castanea Backeberg (1936: 415) (Fig. 2D), F. lepida Buining & Brederoo (1973: 106), (Fig. 2E), F. mammifera, (Fig. 2G), F. pumila (Lemaire 1838: 21) Britton & Rose (1922: 209) (Fig. 2H), F. pygmaea (Spegazzini 1905: 497) Britton & Rose (1922: 210) (Fig. 2I), F. erythracantha R.Pontes, A.S.Oliveira & Deble (2018: 202) (Fig. 2J), in the study area, these species usually grow on rocky outcrops. Morphologic notes:— They are tiny plants less than 4 cm in diameter, sometimes solitary, globular to cylindrical. Undeveloped ribs and very small straight or curved thorns. Flowers very ephemeral, l cleistogamous or casmogamous, yellow or cream-yellow, dense woolly floral bracts and many bristles. Dried fruit with many seeds up to 1.5 cm long. They usually bloom in summer and fruit soon after., Published as part of Oliveira, Anabela Silveira De, Pontes, Rodrigo Corrêa, Robaina, Luis Eduardo De Souza & Trentin, Romário, 2022, The Cactaceae family in the Santa Maria river basin (Rio Grande do Sul, Brazil), pp. 153-166 in Phytotaxa 542 (2) on pages 160-161, DOI: 10.11646/phytotaxa.542.2.3, http://zenodo.org/record/6413545, {"references":["Britton, N. L. & Rose, J. N. (1922) The Cactaceae. Descriptions and Illustrations of Plants of the Cactus Family, vols. 3 - 4 Carnegie Institution, Washington, 322 pp.","Gerloff, N. (2001) Frailea densispina (Hofacker & Herm) N. Gerloff, Comb. & Stat. Nov. Cactus & Co 3 (5): 138 - 147.","Backeberg, C. (1936) Frailea castanea Bckbg. sp. n. In: Backeberg, C. & Knuth, F. M. (Eds.) Kaktus ABC. Gyldendals Forlagstrykkeri, Kobenhavn, p. 415.","Buining, A. F. H. & Brederoo, A. J. (1973) Frailea lepida Buining et Brederoo spec. nov. Kakteen 54: CVIe.","Lemaire, C. (1838) Echinocactus pumilus. Cactearum aliquot novarum acinsuetarum in hortomonvilliano cultarumaccurata descriptio. Lutetiae-Parisiorum, Paris, 40 pp.","Spegazzini, C. L. (1905) Cactacearum Platensium Tentamen. Anales Del Museo Nacional de Buenos Aires 11: 477 - 521. https: // doi. org / 10.5962 / bhl. title. 9303"]}

Parodia ritteriana (Lisal & Kolarik) A. S. Oliveira & R. Pontes comb. nov. ≡ Notocactus ritterianus Lisal & Kolarik (1986: 5) The species grows on vulcanic rocks and it is classified as critically endangered (CR) in its native habitat (IUCN 2022). The populations found are small and disjunct in an area of about 14 km 2. The threats observed were human activities associated with agriculture (removal of soil cover and removal of stones from the field) and livestock (grazing)., Published as part of Oliveira, Anabela Silveira De, Pontes, Rodrigo Corrêa, Robaina, Luis Eduardo De Souza & Trentin, Romário, 2022, The Cactaceae family in the Santa Maria river basin (Rio Grande do Sul, Brazil), pp. 153-166 in Phytotaxa 542 (2) on page 162, DOI: 10.11646/phytotaxa.542.2.3, http://zenodo.org/record/6413545, {"references":["Lisal, K. & Kolarik, J. (1986) Notocactus arnostianus Lisal & Kolarik & Notocactus ritterianus Lisal & Kolarik. Internoto 7 (1): 3 - 19."]}