Alooideae

'Haworthia' attenuata, copyright Diego Delso.

Belongs within: Asphodelaceae.
Contains: × Gasteraloe, Haworthia, Aloe, Gasteria.

The Alooideae are a group of succulent monocotyledons found primarily in southern Africa (Eggli & Nyffeler 2020).

Synapomorphies (from Eggli & Nyffeler 2020): Leaves succulent; secondary growth often present via secondary thickening meristem; flowers tubular; ovules consistently hemitropous; distinctly bimodal basic karyotype consisting of four long and three short chromosomes; parenchymatous, cap-like inner bundle sheath present at phloem poles; anthrone-C-glycosides present in leaves, 1-methyl-8-hydroxyanthraquinones in roots; steroidal saponins generally absent.

<==Alooideae [Aloaceae, Aloeae, Aloineae]
    |--Aloidendron (Berger) Klopper & Smith 2013 EN20
    |    |--*A. barberae (Dyer) Klopper & Smith 2013 EN20 (see below for synonymy)
    |    `--+--A. pillansii (Guthrie) Klopper & Smith 2013 EN20 (see below for synonymy)
    |       `--A. ramosissimum (Pillans) Klopper & Smith 2013 EN20 (see below for synonymy)
    `--+--Haworthia TS03
       `--+--Aloiampelos Klopper & Smith 2013 EN20
          |    `--*A. ciliaris (Haworth) Klopper & Smith 2013 EN20 [=Aloe ciliaris Haworth 1825 TS03, EN20]
          |         |--A. c. var. ciliaris (see below for synonymy) EN20
          |         |--A. c. var. redacta (Carter) Klopper & Smith 2013 [=Aloe ciliaris var. redacta Carter 1990] EN20
          |         `--A. c. var. tidmarshii (Schönland) Klopper & Smith 2013 (see below for synonymy) EN20
          `--+--Aloe TS03
             `--+--+--Poellnitzia Uitewaal 1940 TS03, EN20
                |  |    `--*P. rubriflora (Bolus) Uitewaal 1940 (see below for synonymy) EN20
                |  `--+--Aristaloe Boatwright & Manning 2014 EN20
                |     |    `--*A. aristata (Haworth) Boatwright & Manning 2014 EN20 (see below for synonymy)
                |     `--Astroloba Uitewaal 1947 TS03, EN20 [=Apicra Haworth 1819 non Willdenow 1811 EN20]
                |          |  i. s.: A. spiralis (Linné) Uitewaal 1947 (see below for synonymy) EN20
                |          |         A. bullulata (Jacquin) Uitewaal 1947 (see below for synonymy) EN20
                |          |         A. cremnophila Van Jaarsveld 2015 EN20
                |          |         A. herrei Uitewaal 1948 (see below for synonymy) EN20
                |          |         A. robusta Roberts Reinecke ex Molteno et al. 2017 [=A. foliolosa ssp. robusta] EN20
                |          |         A. tenax Molteno et al. 2017 EN20
                |          |           |--A. t. var. tenax EN20
                |          |           `--A. t. var. moltenoi Smith & Van Jaarsveld 2017 EN20
                |          |--A. congesta (Salm-Dyck) Uitewaal 1947 TS03, EN20 (see below for synonymy)
                |          `--+--‘Haworthia’ kewensis TS03
                |             `--+--A. foliolosa (Haworth) Uitewaal 1947 TS03, EN20 (see below for synonymy)
                |                `--+--A. corrugata Meyer & Smith 1998 TS03, EN20 (see below for synonymy)
                |                   `--× Astroworthia Rowley 1967 TS03, EN20 (see below for synonymy)
                |                        `--× A. bicarinata (Haworth) Rowley 1973 (see below for synonymy) EN20
                `--+--+--‘Haworthia’ attenuata (Haworth) Haworth 1812 TS03, EN20 (see below for synonymy)
                   |  |    |--H. a. var. attenuata (see below for synonymy) EN20
                   |  |    |--H. ‘radula’ var. asperior Haworth 1821 (n. d.) EN20
                   |  |    |--H. ‘glabrata’ var. concolor (Salm-Dyck) Baker 1880 (n. d.) (see below for synonymy) EN20
                   |  |    |--H. a. var. glabrata (Salm-Dyck) Bayer 2012 (see below for synonymy) EN20
                   |  |    |--H. ‘radula’ var. laevior Haworth 1821 (n. d.) EN20
                   |  |    |--H. ‘radula’ var. magniperlata Haworth 1821 (n. d.) EN20
                   |  |    |--‘Aloe radula’ var. major Salm-Dyck 1817 nec A. margaritifera var. major Aiton 1789 nec A. picta var. major Willdenow 1799 nec A. expansa var. major Haworth 1804 nec A. glauca var. major Haworth 1812 nec A. acuminata var. major Salm-Dyck 1817 nec A. fasciata var. major Salm-Dyck 1817 nec A. nitida var. major Salm-Dyck 1817 nec A. plicatilis var. major Salm-Dyck 1817 nec A. prolifera var. major Salm-Dyck 1817 EN20
                   |  |    |--‘Aloe radula’ var. media Salm-Dyck 1817 non A. margaritifera var. media De Candoll 1799 EN20
                   |  |    |--‘Aloe radula’ var. minor Salm-Dyck 1817 nec A. margaritifera var. minor Aiton 1789 nec A. picta var. minor Willdenow 1799 nec A. glauca var. minor Haworth 1812 nec A. nitida var. minor Salm-Dyck 1817 EN20
                   |  |    |--‘Aloe radula’ var. paulo-major Salm-Dyck 1817 EN20
                   |  |    |--H. ‘glabrata’ var. perviridis (Salm-Dyck) Baker 1880 (n. d.) (see below for synonymy) EN20
                   |  |    `--H. a. var. radula (Jacquin) Bayer 1999 (see below for synonymy) EN20
                   |  `--‘Haworthia’ icosiphylla Baker 1880 TS03, EN20
                   `--+--‘Haworthia’ glauca Baker 1880 TS03, EN20 (see below for synonymy)
                      |    |--H. g. var. glauca [incl. H. carrissoi Resende 1941] EN20
                      |    `--H. g. var. herrei (Von Poellnitz) Bayer 1976 (see below for synonymy) EN20
                      `--Gasteria TS03

Alooideae incertae sedis:
  × Algastoloba Cumming 1999 [incl. Altulisteria Rowley 2014] EN20
  × Aloloba Rowley 1967 [incl. Chamaeloba Cumming 1974, Lomatoloba Cumming 1974] EN20
  × Alworthia Rowley 1973 EN20
  × Bayerara Cumming 1999 EN20
  × Cummingara Rowley 1999 [incl. Smithara Cumming 1999 (nom. illeg.)] EN20
  × Gasteraloe EN20
  × Gasterhaworthia Guillaumin 1931 (see below for synonymy) EN20
  × Gastroloba Cumming 1974 EN20
    `--× G. apicroides [=Gasteria apicroides] EN20
  × Maysara Cumming 1999 [incl. Rowleyara Cumming 1999] EN20
  × Poellneria Rowley 1973 EN20

Nomina invalida: Astroloba hallii Roberts Reinecke 1965 [=Haworthia hallii] EN20
                 A. smutsiana Roberts Reinecke 1965 EN20

*Aloiampelos ciliaris (Haworth) Klopper & Smith 2013 var. ciliaris [incl. Aloe ciliaris var. flanaganii Schönland 1903, Aloe ciliaris f. flanaganii (Schönland) Resende 1943, Aloe ciliaris f. gigas Resende 1938] EN20

Aloiampelos ciliaris var. tidmarshii (Schönland) Klopper & Smith 2013 [=Aloe ciliaris var. tidmarshii Schönland 1903, Aloe ciliaris f. tidmarshii (Schönland) Resende 1943, Aloe tidmarshii (Schönland) Müller ex Dyer 1943] EN20

*Aloidendron barberae (Dyer) Klopper & Smith 2013 EN20 [=Aloe barberae Dyer 1874 TS03, EN20, Aloe bainesii var. barberae (Dyer) Baker 1896 EN20; incl. Aloe bainesii Dyer 1874 EN20, Aloe zeyheri Hort. ex Baker 1880 (nom. inv.) non Salm-Dyck 1836 EN20]

Aloidendron pillansii (Guthrie) Klopper & Smith 2013 EN20 [=Aloe pillansii Guthrie 1928 TS03, EN20, Aloe dichotoma ssp. pillansii (Guthrie) Zonneveld 2002 EN20]

Aloidendron ramosissimum (Pillans) Klopper & Smith 2013 EN20 [=Aloe ramosissima Pillans 1937 TS03, EN20, Aloe dichotoma ssp. ramosissima (Pillans) Zonneveld 2002 EN20, Aloe dichotoma var. ramosissima (Pillans) Glen & Hardy 2000 EN20]

*Aristaloe aristata (Haworth) Boatwright & Manning 2014 EN20 [=Aloe aristata Haworth 1825 TS03, EN20, Tulista aristata (Haworth) Rowley 2013 EN20; incl. Aloe ellenbergeri Guillaumin 1934 EN20, Aloe aristata var. leiophylla Baker 1880 EN20, Aloe longiaristata Schultes & Schultes 1829 EN20, Aloe aristata var. parvifolia Baker 1896 non A. viscosa var. parvifolia (Haworth) Roemer & Schultes 1829 EN20]

Astroloba bullulata (Jacquin) Uitewaal 1947 [=Aloe bullulata Jacquin 1809, Apicra bullulata (Jacquin) Willdenow 1811, Ap. pentagona var. bullulata (Jacquin) Baker 1880, Haworthia bullulata (Jacquin) Parr 1971, Tulista bullulata (Jacquin) Rowley 2013; incl. Astroloba bullata (nom. inv.), Apicra egregia Von Poellnitz 1930, As. egregia (Von Poellnitz) Uitewaal 1947, Haworthia egregia (Von Poellnitz) Parr 1971, As. egregia var. fardeniana Uitewaal 1948, H. egregia var. fardeniana (Uitewaal) Parr 1971] EN20

Astroloba congesta (Salm-Dyck) Uitewaal 1947 TS03, EN20 [=Aloe congesta Salm-Dyck 1854 EN20, Apicra congesta (Salm-Dyck) Baker 1880 EN20, Astroloba foliolosa ssp. congesta (Salm-Dyck) Roberts Reinecke 1965 (nom. inv.) EN20, Haworthia congesta (Salm-Dyck) Parr 1971 EN20, Tulista congesta (Salm-Dyck) Rowley 2013 EN20; incl. Aloe deltoidea Hooker 1873 EN20, Apicra deltoidea (Hooker) Baker 1880 EN20, Astroloba deltoidea (Hooker) Uitewaal 1947 EN20, Haworthia deltoidea (Hooker) Parr 1971 non H. attenuata var. deltoidea Farden 1939 EN20, Apicra deltoidea var. intermedia Berger 1908 EN20, Astroloba deltoidea var. intermedia (Berger) Uitewaal 1947 EN20, Haworthia deltoidea var. intermedia (Berger) Parr 1971 non H. intermedia Von Poellnitz 1937 EN20, H. shielsiana Parr 1971 (nom. inv.) EN20, Apicra turgida Baker 1889 EN20, Ap. deltoidea var. turgida (Baker) Berger 1908 EN20, Astroloba deltoidea var. turgida (Baker) Jacobsen 1954 (nom. inv.) EN20, As. turgida (Baker) Jacobsen 1960 (nom. inv.) EN20, Haworthia deltoidea var. turgida (Berger) Parr 1971 (nom. inv.) nec H. turgida Haworth 1819 nec H. triebneriana var. turgida Triebner 1938 EN20]

Astroloba corrugata Meyer & Smith 1998 TS03, EN20 [=Haworthia corrugata (Meyer & Smith) Hayashi 2000 EN20, Tulista corrugata (Meyer & Smith) Rowley 2013 EN20; incl. Aloe aspera Salm-Dyck 1854 non Haworth 1804 EN20, Haworthia aspera (Salm-Dyck) Parr 1971 non Haworth 1812 EN20, Astroloba muricata Groen 1987 (nom. inv.) EN20, As. rugosa Roberts Reinecke 1965 (nom. inv.) EN20]

Astroloba foliolosa (Haworth) Uitewaal 1947 TS03, EN20 [=Aloe foliolosa Haworth 1804 EN20, Apicra foliolosa (Haworth) Willdenow 1811 EN20, Astroloba spiralis ssp. foliolosa (Haworth) Groen 1987 EN20, Haworthia foliolosa (Haworth) Haworth 1812 EN20, Tulista foliolosa (Haworth) Rowley 2013 EN20]

Astroloba herrei Uitewaal 1948 [=Tulista herrei (Uitewaal) Rowley 2013; incl. A. dodsoniana Uitewaal 1950, A. herrei cv. dodsoniana (Uitewaal) Groen 1987, Haworthia dodsoniana (Uitewaal) Parr 1971; incl. H. harlandiana Parr 1971] EN20

Astroloba spiralis (Linné) Uitewaal 1947 [=Aloe spiralis Linné 1753, Apicra spiralis (Linné) Duval 1809, Haworthia spiralis (Linné) Duval 1809, Tulista spiralis (Linné) Rowley 2013; incl. H. gweneana Parr 1971 (nom. inv.), Aloe spiralis var. imbricata Aiton 1789, Aloe imbricata (Aiton) Haworth 1811, Apicra imbricata (Aiton) Willdenow 1811, Haworthia imbricata (Aiton) Haworth 1812, Aloe spiralis var. pentagona Aiton 1789, Aloe pentagona (Aiton) Haworth 1804, *Apicra pentagona (Aiton) Willdenow 1811, *Astroloba pentagona (Aiton) Uitewaal 1947, As. spiralis cv. pentagona (Aiton) Groen 1987, Haworthia pentagona (Aiton) Haworth 1812, Aloe spiralis Haworth 1804 non Linné 1753, Aloe pentagona var. spiralis (Haworth) Salm-Dyck 1836, Astroloba pentagona var. spiralis (Haworth) Uitewaal 1947, Haworthia pentagona var. spiralis (Salm-Dyck) Parr 1971; incl. H. spirella Haworth 1812, Aloe spirella (Haworth) Salm-Dyck 1817, Apicra pentagona var. spirella (Haworth) Baker 1880, Astroloba pentagona var. spirella (Haworth) Uitewaal 1947, Haworthia pentagona var. spirella (Haworth) Parr 1971, Apicra pentagona var. torulosa Haworth 1821, Astroloba pentagona var. torulosa (Haworth) Uitewaal 1821, Haworthia pentagona var. torulosa (Haworth) Parr 1971] EN20

× Astroworthia Rowley 1967 TS03, EN20 [incl. Apworthia Von Poellnitz 1943 EN20, Astrolista Molteno & Figueiredo 2017 EN20]

× Astroworthia bicarinata (Haworth) Rowley 1973 [=Apicra bicarinata Haworth 1819, Aloe bicarinata (Haworth) Schultes & Schultes 1829, Astrolista bicarinata (Haworth) Molteno & Figueiredo 2017, Astroloba bicarinata (Haworth) Uitewaal 1947, Haworthia bicarinata (Haworth) Parr 1971 non H. altilinea var. bicarinata Triebner 1938, Tulista bicarinata (Haworth) Rowley 2013; Astroloba corrugata × Haworthia margaritifera; incl. Apicra bullulata Uitewaal 1938 non (Jacquin) Willdenow 1811, Haworthia olivettiana Parr 1971, Ap. skinneri Berger 1908, Astroloba skinneri (Berger) Uitewaal 1947, Astroworthia bicarinata nvar. skinneri (Berger) Rowley 1973, Astroworthia skinneri (Berger) Groen 1987, Haworthia skinneri (Berger) Resende 1943] EN20

× Gasterhaworthia Guillaumin 1931 [incl. Gasworthia Gates 1955, Gasworthiopsis Rowley 2014, Haworthio-gasteria Kondo & Megata ex Megata 1943 (nom. inv.)] EN20

‘Haworthia’ attenuata (Haworth) Haworth 1812 TS03, EN20 [=Aloe attenuata Haworth 1804 EN20, Apicra attenuata (Haworth) Willdenow 1811 EN20, Haworthia pumila ssp. attenuata (Haworth) Halda 1997 EN20, Haworthiopsis attenuata (Haworth) Rowley 2015 EN20]

‘Haworthia’ attenuata (Haworth) Haworth 1812 var. attenuata [incl. H. britteniana Von Poellnitz 1937, H. attenuata f. britteniana (Von Poellnitz) Bayer 1982, H. attenuata var. britteniana (Von Poellnitz) Von Poellnitz 1937, H. fasciata var. caespitosa Berger 1908, H. attenuata f. caespitosa (Berger) Pilbean 1983 (nom. inv.), H. attenuata var. caespitosa (Berger) Farden 1939, H. clariperla Haworth 1826, Aloe attenuata var. clariperla (Haworth) Salm-Dyck 1836, A. clariperla (Haworth) Roemer & Schultes 1829, Haworthia attenuata f. clariperla (Haworth) Bayer 1976, H. attenuata var. clariperla (Haworth) Baker 1880, H. attenuata var. deltoidea Farden 1939, H. attenuata var. inusitata Farden 1939 (nom. inv.), H. attenuata var. linearis Farden 1939, H. attenuata var. minissima Farden 1939, H. attenuata var. odonoghueana Farden 1939, H. attenuata var. uitewaaliana Farden 1939 (nom. inv.) non H. uitewaaliana Von Poellnitz 1940] EN20

‘Haworthia’ attenuata var. glabrata (Salm-Dyck) Bayer 2012 [=Aloe glabrata Salm-Dyck 1834, Haworthia glabrata (Salm-Dyck) Bayer 1880, Haworthiopsis attenuata var. glabrata (Salm-Dyck) Rowley 2015] EN20

‘Haworthia’ attenuata var. radula (Jacquin) Bayer 1999 [=Aloe radula Jacquin 1804, Apicra radula (Jacquin) Willdenow 1811, Haworthia radula (Jacquin) Haworth 1812, Haworthia pumila ssp. radula (Jacquin) Halda 1997, Haworthiopsis attenuata var. radula (Jacquin) Rowley 2015; incl. Haworthia radula var. pluriperlata Haworth 1821] EN20

‘Haworthia glabrata’ var. concolor (Salm-Dyck) Baker 1880 (n. d.) [=Aloe glabrata var. concolor Salm-Dyck 1849] EN20

‘Haworthia glabrata’ var. perviridis (Salm-Dyck) Baker 1880 (n. d.) [=Aloe glabrata var. perviridis Salm-Dyck 1849 non A. rugosa var. perviridis Salm-Dyck 1834] EN20

‘Haworthia’ glauca Baker 1880 TS03, EN20 [=Haworthia reinwardtii ssp. glauca (Baker) Halda 1997 EN20, Haworthiopsis glauca (Baker) Rowley 2013 EN20]

‘Haworthia’ glauca var. herrei (Von Poellnitz) Bayer 1976 [=Haworthia herrei Von Poellnitz 1929, Haworthia reinwardtii var. herrei (Von Poellnitz) Halda 1997, Haworthiopsis glauca var. herrei (Von Poellnitz) Rowley 2013; incl. Haworthia armstrongii Von Poellnitz 1937, Haworthia glauca f. armstrongii (Von Poellnitz) Bayer 1976, Haworthia herrei var. depauperata Von Poellnitz 1933, Haworthia eilyae Von Poellnitz 1937, Haworthia jacobseniana Von Poellnitz 1937, Haworthia glauca f. jacobseniana (Von Poellnitz) Pilbean 1983, Haworthia jonesiae Von Poellnitz 1937, Haworthia glauca f. jonesiae (Von Poellnitz) Pilbean 1983, Haworthia eilyae var. poellnitziana Resende 1943 (nom. illeg.) nec H. poellnitziana Uitewaal 1939 nec H. planifolia var. poellnitziana Resende 1940, Haworthia herrei var. poellnitzii Resende 1941 (nom. illeg.), Haworthia eilyae var. zantneriana Resende 1943 non H. zantneriana Von Poellnitz 1935] EN20

*Poellnitzia rubriflora (Bolus) Uitewaal 1940 [=Apicra rubriflora Bolus 1920, Aloe rubriflora (Bolus) Rowley 1981 non A. tenuior var. rubriflora Reynolds 1936, Astroloba rubriflora (Bolus) Smith & Manning 2000, Haworthia rubriflora (Bolus) Parr 1971, Tulista rubriflora (Bolus) Rowley 2013; incl. Apicra jacobseniana Von Poellnitz 1939, Haworthia rubriflora var. jacobseniana (Von Poellnitz) Parr 1972 (nom. inv.) non H. jacobseniana Von Poellnitz 1937, Poellnitzia rubriflora var. jacobseniana (Von Poellnitz) Uitewaal 1955] EN20

*Type species of generic name indicated

REFERENCES

[EN20] Eggli, U., & R. Nyffeler (eds) 2020. Illustrated Handbook of Succulent Plants: Monocotyledons 2^nd ed. Springer.

[TS03] Treutlein, J., G. F. Smith, B.-E. van Wyk & M. Wink. 2003. Phylogenetic relationships in Asphodelaceae (subfamily Aloooideae) inferred from chloroplast DNA sequences (rbcL, matK) and from genomic fingerprinting (ISSR). Taxon 52: 193–207.