Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy
Author
Walter, Helmut E.
0000-0001-8614-0159
The EXSIS Project: Cactaceae Ex-Situ & In-Situ Conservation, 31860, Emmerthal, Germany. & hw 582133 @ gmail. com; https: // orcid. org / 0000 - 0001 - 8614 - 0159
hw582133@gmail.com
Author
Guerrero, Pablo C.
0000-0003-1034-1899
Departamento de Botánica, Facultad de Ciencias Naturales & Oceanográficas, Universidad de Concepción, Casilla 160 C, Concepción, Chile. & Institute of Ecology and Biodiversity (IEB), Victoria 631, Barrio Universitario, Concepción, Chile. & Millennium Institute Biodiversity of Antarctic and Sub-Antarctic Ecosystems (BASE), CP 7800003, Santiago, Chile. & pabloguerrero @ udec. cl; https: // orcid. org / 0000 - 0003 - 1034 - 1899
pabloguerrero@udec.cl
text
Phytotaxa
2022
2022-06-14
550
2
79
98
journal article
10.11646/phytotaxa.550.2.1
1179-3163
6641038
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
C1 present .....................................................................................................................................................................................2
- Stems without functional leaves; a 739 bp non-coding intron region in the chloroplast-encoded gene
rpo
C1 is lost (subfam.
Cactoideae
) .........................................................................................................................................................................................3
- Stems without functional leaves; a 739 bp non-coding intron region in the chloroplast-encoded gene
rpo
C1 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, <10; fruits
1cm
, dark red, glabrous, pulp red................................
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 <
5cm
; areoles evenly distributed; fruits subglobose; seeds lack lateral ridges....................................................................................................................
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 <
4 cm
; flowers with few small bracts; fruits
2-5 cm
......................
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 <
1 cm
, dehiscing laterally
.............................................................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 <50(-70) cm, globose, elongating with age .................................................................................................
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
).