Lupinus burkartianus C.P.Sm., Sp. Lup.

Planchuelo, Ana María, 2022, New varieties and synonyms of Lupinus species (Fabaceae, Faboideae) of Northwestern Argentina, Phytotaxa 566 (2), pp. 143-170 : 155-158

2. Lupinus burkartianus C.P.Sm., Sp. Lup. : 334 (1943). Tables 1 & 2 (second columns). Figs. 2 & 4 B .

Type:— ARGENTINA: Prov. Tucumán: Dpto. Tafí del Valle: Sierras Calchaquies , Peñas Azules , 3400 m s.n.m., 29 Jan. 1933, A. Burkart 5429 (holotype K! , isotype SI!, UC!) .

= Lupinus campestris-florum C.P.Sm., Sp. Lup. : 334 (1943), syn. nov. Type:— ARGENTINA: Prov. Tucumán: Dpto. Tafí del Valle: San José, de Peñas Azules a campo La Flora, 3000 m s.n.m., Dec. 1931, Schreiter 7011 (holotype US!, isotype A! (now at GH!), LIL! this specimen has a label saying “cotype, Determinavit: Charles Piper Smith”).

= Lupinus hiulcus C.P.Sm., Sp. Lup. : 336 (1943). syn. nov. Type:— ARGENTINA: Prov. Tucumán: Dpto. Chicligasta: Ea. Las Pavas, 3200 m s.n.m., 12 Mar. 1924, S. Venturi 3087 (holotype US! sheets 1546683 and 1546684; isotype A! ex GH! US!, SI!).

= Lupinus intortus C.P.Sm., Sp. Lup. : 339 (1944). syn. nov. Type:— ARGENTINA: Prov. Jujuy: Dpto. El Volcán: Feb. 1920. Castillon 7139 (Holotype DS!; isotype LIL!).

= Lupinus opertospicus C.P.Sm. , syn. nov., Sp. Lup.: 341 (1944). Type: (Data from Smith: “Type. Budin (11793 h 201); h Sm ex 201); h Sm ex 201” (h Sm=Smith; 201=DS); “ Jujuy, Maimara, Huanchie, 3000 m s.n.m., Budin..., 1912 I 17. ” Type data from holotype: Jujuy: Maimará, Huanchín, 3000 m s.n.m., 17 Jan. 1912, Budin 11793. Correct quote: Prov. Jujuy: Dpto. Tilcara: Maimará, Huichaira, 3000 m s.n.m., 17 Jan. 1912, Budin 11793 (Holotype DS! Isotype LIL sheet 46788 only) (no other specimens E. Budin 11793 collected at Huanchie, Huanchín, Hudchin or Hutchin, at LIL; A ex GH, SI, or UC should be considered as type of this taxon). (See comments on this species and on Representative Specimens for L. austrorientalis var. umidicola ).

= Lupinus pavorum C.P.Sm., Sp. Lup. : 342 (1944). Type:— ARGENTINA: Prov. Tucumán: Dpto. Chicligasta: Ea. Las Pavas. 3300 m s.n.m., 6 Dec. 1926. S. Venturi 4687, (holotype: US!. isotypes BA!, F!, A ex GH!, K!, LIL!, MO!, SI!, UC!).

= Lupinus venturianus C.P.Sm., Sp. Lup. : 344 (1944). Type:— ARGENTINA: Prov. Tucumán: Dpto. Chigligasta: Pueblo Viejo, Ea. Santa Rosa, 4000 m s.n.m., S. Venturi 4061, 13 Dec. 1925 (holotype GH!, isotype LIL!).

= Lupinus amnis-churquis C.P.Sm., Sp. Lup. : 348 (1944). Type:— ARGENTINA: Prov. Tucumán: Dpto. Tafí del Valle: El Churqui, 2000 m s.n.m., Dec. 1917, Schreiter 408 (holotype h Sm ex 201” (h Sm=Smith; 201=DS); isotypes A! now GH plants A, B, D, E only, LIL!, plant C only: none SI specimens). (See comments on this species and on L. hieronymi and Representative Specimens of L. austrorientalis var. umidicola ).

= Lupinus solitarius C.P.Sm., Sp. Lup. : 353 (1944). Type:— ARGENTINA: Prov. Catamarca: Dpto. Ambato: El Rodeo, 15 Jan. 1911, Castillon 2097 (holotype: DS! isotype LIL!).

= Lupinus tafiensis C.P.Sm., Sp. Lup. : 354 (1944). syn. nov. Type:— ARGENTINA: Prov. Tucumán; Dpto. Tafí del Valle: Valle de Tafí, Dec. 1911. Castillon 2016 (holotype DS!, LIL!).

Plants (15–) 25–40 cm tall, with a cluster of several stems from a caudex. Indumentum on stems and petioles with semi-appressed, 1.3–1.5 (–2.0) mm long hairs becoming yellowish-golden in herbarium specimens and with an undercoat of curly canescent 0.5 mm long hairs, not totally covering the surface. Stems 9–10 mm diam, sub-fistulous and subangular from the veins of the above petioles, ligneous and hollow below; internodes 2.5–4.5 cm long; Leaves palmately (6) 8–10 foliolate; stipules 8–13 mm long, the narrowly triangular to subulate free tips 4–10 × 1.0– 1.5 mm, pilose to sericeous dorsally; petioles 9–13 (–15) cm long, about (2–) 2½ (–3) times the length of the leaflets, somewhat persistent after the leaflets fall; leaflets oblanceolate, 4–6 × 0.6–1.2 cm, the youngest conduplicate, mostly complanate at maturity, adaxial face papillose and with sparsely appressed hairs all over the surface and abaxial face with semidense indumentums of semi-appressed to appressed long and short hairs, tips obtuse sometime rounded. Peduncles 0.5–6.0 cm. Inflorescence terminal racemes, (9–) 12–16 (–18) cm with verticillate, sometimes with crowded flowers and partially covered by the leaves of the upper nodes; bracts lanceolate to subulate, 5–8 × 1.0– 1.5 mm, deciduous or tardily deciduous, with indumentums similar to stem dorsally; bracteoles triangular to subulate, 0.4–1.0 × 0.1–0.3 mm, attached at 1.0– 1.5 mm below the lips of the lateral sinuses of the calyx; pedicels 3–5 mm long at anthesis, 6–8 mm in fruit with strait and curly hairs. Flowers (9–) 10–11(–12) mm long; calyx with short and curly hairs outside, glabrous within, the base slightly gibbous above, lower-lip lanceolate to lance-oblong, (7.0–) (7.5–8.0 (–8.5) × 1.5–2.5 mm, the tip truncate or rarely dentate, teeth 0.1 mm long; upper-lip (5.0-7.0) mm long, the lobes (2.0–) 2.5–3.5 × 2.0-3.0 x 1.0- 2.5 mm; lips connate 1.0–2.0 mm; banner broad obovate (7.5–) 8–10 × 7.0– 9.5 mm, glabrous; wings (8.0–) 9.0–11.0 × 4.5–6.0 mm, the claw 1.5–2.8 mm long, the lobe above the claw 1–2 mm wide; keel glabrous or with few cilia on the distal part of the upper margin, 2.7-3.0 mm wide in the middle, the angle 100°–120°; ovary with 6–9 ovules. Legumes 3.0–3.2 × 0.7–0.8 cm, covered with dense woolly white indumentum. Seeds with oblong contour 5.0–5.5 × 3.0– 3.5 mm, seed coat light tan with sparsely mottled with browns spots without angle line around the hilar region, reticulate-foveate pattern under 1000 x magnification.

Etymology:— “ burkartianus ” was in homage to the Argentine botanist Arturo E. Burkart who was an expert in Leguminosae taxonomy and also the collector of the type specimen; “ campestris-florum ” is a Latin version of the place where the type was collected; “ hiulcus ” meaning in Latin open, for the open branching of the plants; “ intortus ” is from Latin, meaning twisted, in relation to the twisted branches and petioles of the holotype specimen; “ opertospicus ” is from the Latin combination of “ opertus ” meaning cover and “ apicus ” raceme; “ pavorum ” a Latin version of the place of collection “Estancia Las Pavas”; “ venturianus ” was in homage to the Argentine naturalist Santiago Venturi who collected the type specimen; “ amnis-churquis ” a Latin version “El Churqui” where the type was collected; “ solitarius ” derived by the interpretation by Smith of having only one flower in each verticille in a verticillate inflorescence, instead of considering a raceme with scattered flowers; “ tafiensis ” a Latin version of the place where the type was collected.

Phenology:— Flowering and fruiting time depends on high mountain weather conditions, booming has a range from late spring (December) to late summer (March) and fruiting from mid-summer (February) to mid-autumn (May), following the data collected from herbarium specimens and field observations.

Distribution, Habitat and Red List of Endemic Species Category:— This species is widely distributed through the provinces of NOA especially in Jujuy and Tucumán and less common in Salta, Catamarca and La Rioja which is the southernmost distribution of this taxon ( Fig. 4 B ). It is growing in high valleys and inter-mountain ravines between 2500 and 4000 m s.n.m. The first collection recorded was in Tucumán in 1900, and the last one in Salta in 2003. The major affecting factors are the modifications in normal precipitation patterns caused by the alternation of El Niño (ENSO) that may cause excesses of precipitation and La Niña which produce recurrent drought in the extreme north western areas of its distribution ( Zanvettor, 2019). Another major threatening factor is the expansion of rural communities in Jujuy and Tucumán. This species was assigned in 2010 to the conservation Category 4 (PlanEAr, 2021), and to L. intortus , L. opertospicus and L. tafiensis that are now synonyms, to the Category 5 (because at that time were known only by the holotypes). Taking into consideration the distribution area and the treating factors this author considered that the species should be placed in conservation Category 4 following the scale of PlanEAr and VU (Vulnerable) under the IUCN status scale.

Comments:—The type specimens of L. burkartianus and the other types of eight of its synonyms have a long terminal raceme, with whorl flowers, but the holotype of L. opertospicus has an inflorescence with obsolete peduncle, flowers crowded and partially covered by the branches and leaves of the upper nodes ( Fig. 2 B & D ). The obsolete peduncle was considered by C. P. Smith a key character for L. opertospicus , but the peduncle length in this group of herbaceous therophyte plants has not taxonomic links, since it is a physiological variable related to sensitivity to photo and thermo-periods, as happen in cultivate lupins, mainly in L. mutabilis Sweet (Ravelo & Planchuelo, 1998) .

The two holotype sheets of L. hiulcus ( US 1546683 and 1546684) have an identification label by C. P. Smith with the inscription “ Lupinus hieronymi sp. nov. 1546683” and “ Lupinus hieronymi sp. nov. 1546683” but on the Sp. Lup. page: 335, when C. P. Smith described L. hieronymi does not mention those specimens identified by him, it is probably that the identification labels were pasted before he changed his way of thinking and chose another specimen as holotype of L. hieronymi . The observation of the materials by this author shows that none of those materials have the typical extended hairs that characterize L. hieronymi , consequently the specimens match the description of L. hiulcus treated here as synonym of L. burkartianus . As was explained before, one specimen Lorentz & Hieronymus 696 at NY was cited in Sp. Lup.: 336, 1943 as paratype of L. hiulcus , this specimen differed from the type specimens of L. honoratus that have the same collection number Lorentz & Hieronymus 696 and are at CORD, F, NY and US herbaria.

The structure of the twisted branches present in the holotype of L. intortus could be considered as a structural modification during the drying and pressing process on a plant that was collected on a wet place, this characteristic is not present in other plants representative of the taxon and consequently it is not considered with taxonomic values.

In the label of L. opertospicus holotype specimen at DS (DS 378266) the collection locality is “ Jujuy: Maimará, Huanchín, 3000 m, Leg. Budin 11793, 17 Jan. 1912 ”; which is different from the data “ Huanchie“ cited by C.P. Smith in the protologue (Sp. Lup.: 341, 1944). At LIL herbarium two herbarium sheets have the same collector number (Budin 11793); the sheet LIL 46788, said that was collected in Maimará, Huanchín and is an isotype of the species ( L. opertospicus ); this specimen has an anonymous handwritten note that said “This is the only one that matches Smiths type from (A) Stanford.” The other herbarium sheet LIL 89946, the collection place is Maimará, Hutchín and has four plants mounted, that were assigned by this author with the letters B1, B2, B3 and B4 and identified in 1979 as L. umidicola , now considered L. austrorientalis var. umidicola . At SI one specimen collected in Jujuy, Maimará, Hudchin by E. Budin with the number of the LIL sheet 46788, was identified by A. Burkart as isotype of L. opertospicus C.P.Smith. Two plants are mounted on the herbarium sheet, the leaflets have dense pubescence in the adaxial face, and the banner is orbicular which are not characters of L. opertospicus consequently the specimen is treated here as L. austrorientalis var. jujuyensis . At the herbaria F, GH and UC there are specimens collected by Budin with the LIL number 46788 the plants match the plants of the LIL specimen on sheet LIL 89946 that were identified by the letters B1, B2, B3 & B4, consequently, are not isotype of L. opertospicus and in this treatment are cited as L. austrorientalis var. umidicola . In the “Flora del ConoSur” the picture of the specimen cited as type of L. opertospicus has the following information: Collector Budin, E.; collector number -1 (SI 54595); date 17 Jan. 1912; Argentina, Jujuy, 3000 m s.n.m., Herbaria DS, LIL, SI. As can be see the gather of plants under E. Budin from the original collection in LIL is a mixture of taxa that were sent to other herbaria, with different information about the place of collection, which make those isotypes not a trusted materials to establish the characters to defined the taxon that supposed to represent. A personal communication with the city hall of Maimará the Department of Culture has reported that there was not and are not any locality near Maimará named Huanchie, Huanchin, Hutchin or Hudchin as are written on the specimens’ labels and the near town to Maimará which name start with “H” and is having a “ch” in the middle is Huichaira that is a small village belonging to an ancient aboriginal community which previous name was Yucaira.

The specimen Schreiter 408 collected in “ Tucumán, Dept. Tafí, El Churqui, 2000 m s.n.m., Dec. 1917 ” (Smith Herbarium, now DS) was selected by C. P. Smith as holotype of L. amnis-churquis , and is represented by the upper part of a plant with only one inflorescence that matches the species description in the protologue and also the one of L. burkartianus as treated here. An isotype at A now GH herbarium has mounted on the same cardboard five apical branches with one inflorescence each and a piece of branch with only two leaves. The plants were identified by A. Planchuelo (1978) with the letters A, B, C, D, E, and F. In the specimen is attached a glass slide with one complete and one dissected flower by D. Dunn. A detailed study of the plants designated as A, B, D and E agree with the description on the protologue, and are isotypes of L. amnis churquis , the plant C does not match and the determination is pending and the leaves (F) cannot be identified. At the LIL herbarium there is an isotype that also has several parts of plants from which only the one designated as C by A. Planchuelo (1979) is the one that corresponds to the isotype of the species, the others assigned as A and B, were identified as L. umidicola (now L. austrorientalis var. umidicola ) and the plant assigned as D agrees with the one that was assigned as C of the GH herbarium and was not determined. In the SI herbarium there is another specimen Scheiter 408, collected in the same place and date as the aforementioned and labeled as isotype of L. amnis-churquis , however none of the five plants represents this species. The plants with the letters A and B are equal to plants A and B of LIL (identified as L. umidicola now L. austrorientalis var. umidicola ) and plants C, D, and E are equal to the ones assigned as D (LIL) and C (GH) herbaria and as mentioned above the identification is pending. A pasted note by A.M. Planchuelo (1984) provides the identification of each plant with the letter that was assigned and also has two glass slides with dissected flowers. As was explained on the comments of L. hieronymi , another specimen with collector number Schreiter 408 (collected in Tafí Viejo: Playa del Río de la Hoyada, 22 Nov. 1921), from the DS was chosen by Smith as the holotype of L. schreiteranus , which differs from the other specimens mentioned above for having an indumentum of spreading hairs typically of L. hieronymi (as is treated here) and is not related with the other specimen Schreiter 408 collected in Tafí del Valle.

Representative Specimens:— ARGENTINA: Prov. Catamarca: Dpto. Ambato: El Rodeo, 15 Jan. 1911, Castillon 2097 (DS, LIL, L. solitarius types); idem, 1700 m s.n.m 12 Dec. 1949, J. Arande wout/# LIL374692 (LIL). Dpto. Andalgalá: El Ingenio, 2700 m s.n.m, 5 Dec. 1960, F. Roig 3666 (MRL); idem, Ruiz Leal 21336 ( SI). Prov. Jujuy: Dpto. Cochinoca: Cochinoca, Sierra de Cochinoca, entre Abra Pampa y Rinconada, 3900 m s.n.m., 29 Jan, 1943, A.L. Cabrera 7747 (LP, SI); de ruta 9 a 5 km del desvío a Cochinoca por ruta 71, a orilla de río, 29 Feb. 1996, A.M. Planchuelo 986 (ACOR). Dpto. El Volcán: Estación El Volcán, Feb. 1920. Castillon 7139 (DS, LIL, L. intortus types) Dpto. Humahuaca: 25 km al SE de Tres Cruces, camino a la Cueva del Inca, 27 Jan. 1995, O. Ahumada 7182 (JUJ). Dpto. San Pedro: Miraflores, 17 Jan. 1948, A.L. Cabrera 9288 (LIL). Dpto: Maimará: Huachín (should be Huichaira), 3000 m s.n.m., 17 Jan. 1912, E. Budin 11793 (DS, LIL, sheet 46788 only). Dpto. Santa Catalina: Quebrada San José, 7 Feb. 1982, P. Melchiorre wout/# BAA 18225½ (BAA). Dpto Tilcara: La Laguna 3000 m s.n.m., Jan. 1926, Pereyra 45-MCN5854 (A, LIL). Dpto.Tumbaya: de Purmamarca al Abra de Lipán, 4000 m, 11 Mar. 1982, R. Kiesling et al. 3528 ( SI); camino al Abra de Lipán ca. 3500 m s.n.m., 22 Apr. 1989, R. Kiesling 7061 ( SI); Quebrada de Lipán, 3800, 17 Jan, 1988, F.O. Zuloaga & N.B. Deginani 3575 ( SI). Dpto. Valle Grande: De Río Jordán a Duraznillo, Camino a Alto Calilegua, 29 Dec. 1977, Kiesling et al. 1629 ( SI). Dpto. Yavi: Yavi 29 Jan. 1989, I. Lupo 3 (JUA); Río Sansana 3400 m s.n.m., 10 Feb. T. Meyer et al. 21313 (LIL). Prov. La Rioja: Dpto. Chilecito: Sierra de Famatina , camino a La Mejicana, 3400 m s.n.m, 4 Feb. 1927, L. Parodi 7880 (BAA, GH). Dpto. Famatina : camino a la mina La Mexicana, cueva de Noroña, 3700 m s.n.m, 20 Feb. 1986, Kiesling et al. 6382 (ACOR, SI). Prov. Salta: Dpto. Cachi: Quebrada Sunchales, 9 km al N del ingreso al Valle Encantado, 2003, R. Fortunato 2780 (ACOR, BAB); ruta 33 pasando la Cuesta del Obispo a 3 km del desvío al Valle Encantado, 27 Feb. 1996, A.M. Planchuelo 943 (ACOR); ídem a 5 km del desvío a Valle Encantado 27 Mar. 1996, A.M. Planchuelo 948 (ACOR); ídem a 9 km del desvío a Valle Encantado, 27 Mar. 1996 A.M. Planchuelo 951 (ACOR). Dpto. Cafayate: Río Colorado, pasando la segunda cascada, 28 Dec. 1987, F. Juárez 1657 (MCNS). Dpto. Rosario de Lerma: El Alfarcito, antes de Sta. Rosa de Tastil, 2780 m s.n.m, 1 Dec, 1986, L. Novara 5696 (MCNS). Prov. Tucumán: Dpto. Chicligasta: Estancia Las Pavas, 3200 m s.n.m, 12 Mar. 1924, S. Venturi 3087 (US, sheets 1546683 & 1546684; GH, US, SI, L. hiulcus types); idem, 3300 m s.n.m,, 6 Dec. 1926, S. Venturi 4687. (US. BA, F, GH, K, LIL, MO, UC, L. pavorum types); Pueblo Viejo, Ea. Santa Rosa, 4000 m s.n.m., 13 Dec. 1925, S. Venturi 4061, (GH, LIL, L. venturianus types); del Bolsón a la Cascada, 2500–2700 m s.n.m., 26 Mar. 1953, T. Meyer18190 (LIL). Dpto. Leales: Los Gómez, 3000 m s.n.m, Arenal del río Salí, 19 Nov. 1921, Venturi 630 (BM, LIL, Smith cited this specimen by mistake as L. hilarianus in Sp. Lup.: 352, 1994, whose actual name is L. gibertianus C.P.Sm. , see Dunn & Planchuelo, 1981). Dpto. Tafí del Valle: Tafí del Valle, 27 Jan. 1935, Castellanos 14730 (BA); idem, de Peñas Azules a campo La Flora, 3000 m s.n.m., Dec. 1931, Schreiter 7011 (US, GH, LIL, L. campestris-florum types); Infiernillo, 3000 m s.n.m., 21 Feb. 1949, B. Spane 6004 (LIL); El Churqui, 2000 m s.n.m, Dec. 1917, Schreiter 408 (GH plants A, B, D, E only, DS, LIL plant C only, L. amnis-churquis types); La Banda, 9 Dec. 1945, A. Lourtey 1139 (F, this specimens was cited by mistake as L. tucumanensis in Planchuelo, 1978, K, LIL); Peñas Azules, 3400 m s.n.m, 29 Jan. 1933, A. Burkart 5429 (K, UC, SI, L. burkartianus types); idem, 4000 m s.n.m., 27 Jan. 1933, L. Parodi 10970 (BAA); La Cienaga, 1–17 Jan. 1874, Lorentz & Hieronymus 696 (NY no isotype of L. honoratus ); Quebrada del Barón, 3500–3700 m s.n.m, 28 Mar. 1959, A. Türpe wout/# LIL 187917 (LIL); idem, LIL 458171 (LIL); Cumbres Calchaquíes, Cerro Negrito, 3500 m s.n.m., 27 Feb. 1959, Türpe wout/# LIL 167889 (LIL). Dpto. Tafí Viejo: La Puerta camino de San José a Las Lagunas, 3800 m s.n.m, Jan. 1933, O. Obea, wout/# LIL 68683 (GH, LIL, SI); idem, 4000 m s.n.m, común en faldas pedregosas, 29 Jan. 1933, L. Parodi 10815 (BAA, GH, UMOMO); idem, 30 Jan. 1933, A. Burkart 5568 (US); playa cerca de la Hoyada, 16 Dec. 1900, LIL 46773 (LIL).

Figures

FIGURE 2. Lupinus burkartianus C.P.Sm. A. Isotype UC with typical extended inflorescence. B. Isotype L. opertospicus, LIL with inflorescence covered by leaves. C. Diagram of holotype showing the extended inflorescence D. Diagram of holotype L. opertospicus showing the inflorescence covered by the leaves and petioles that are 2 ½–3 times the length of a leaflet. E. Dissected flowers showing banner broad obovate (Planchuelo 986). F. Plant growing in Jujuy, representative specimen Planchuelo 986 (ACOR) (Photo Planchuelo). G. Detail of inflorescence. H. Microphotograph (150 x) of adaxial face of leaflet papillose with a basal portion of a long hair. I. Photo SEM (100 x) showing the adaxial face of leaflet with esparcelly long hairs (Photos Seisdedos & Planchuelo).

FIGURE 4. Distribution maps of the treated species. The dots indicate the places where representative specimens were collected. Due to the overlap of places, the number of citations is indicated by province in a north-south sequence. A. Lupinus hieronymi C.P.Sm.: Jujuy 4; Salta, 2; Tucumán, 14; Catamarca 4; La Rioja, 1; Córdoba, 9. B. Lupinus burkartianus C.P.Sm.: Jujuy 14; Salta, 6; Tucumán, 16; Catamarca 4; La Rioja, 2. C. Lupinus austrorientalis C.P.Sm. var. austrorientalis: Jujuy 8; Salta, 3; Catamarca, 5; La Rioja, 1. D. Lupinus austrorientalis var. umidicola (C.P.Sm.) Planchuelo: Jujuy, 2; Salta, 13; Tucumán, 4; Catamarca 7; La Rioja, 3; Córdoba, 6. E. Lupinus austrorientalis var. jujuyensis (C.P.Sm.) Planchuelo: Jujuy, 6; Salta, 3.