Lipochaeta

Seaside nehe Lipochaeta succulenta, copyright David Eickhoff.

Belongs within: Asteraceae.

Lipochaeta, nehe, is a genus of composite-flowered plants endemic to Hawaii.

Lipochaeta
    |--L. lobata F79
    |    |--L. l. var. leptophylla F79
    |    `--+--L. l. var. lobata F79
    |       `--L. l. var. hastulatoides F79
    `--+--+--L. heterophylla F79
       |  `--+--L. degeneri F79
       |     `--L. succulenta F79
       `--+--L. rockii F79
          `--L. connata F79
               |--L. c. var. connata F79
               `--L. c. var. acris F79

*Type species of generic name indicated

REFERENCES

[F79] Farris, J. S. 1979. Review: Cladistics and Plant Systematics. Systematic Zoology 28 (3): 400–411.

Carduoideae

Dicoma tomentosa, copyright D. Wesuls.

Belongs within: Asteraceae.
Contains: Cardueae.

The Carduoideae are a clade of composite-flowered plants supported by molecular data, members of which commonly have spiny leaves and discoid flower heads. Members include the Dicomeae, a group of African herbs, shrubs and small trees with homogamous or heterogamous flower heads that may have bilabiate peripheral florets (Panero & Funk 2002).

<==Carduoideae
    |--+--Cardueae PF02
    |  `--Tarchonantheae PF02
    |       |--Oldenburgia PF02
    |       `--+--Tarchonanthus PF02
    |          `--Brachylaena PF02
    |               |--B. glabra K02
    |               `--B. huillensis R-S02
    `--Dicomeae PF02
         |--Erythrocephalum PF02
         |--Gladiopappus PF02
         |--Macledium PF02
         |--Cloiselia PF02
         |--Pasaccardoa PF02
         |--Pleiotaxis PF02
         `--Dicoma Cassini 1817 PF02
              |--D. cuneneensis CV06
              |--D. dinteri CV06
              |--D. obconica CV06
              `--D. tomentosa PP07

*Type species of generic name indicated

REFERENCES

[CV06] Craven, P., & P. Vorster. 2006. Patterns of plant diversity and endemism in Namibia. Bothalia 36 (2): 175–189.

[K02] Krüger, M. 2002. Revision of the Afrotropical Ennominae of the Drepanogynis group IV: the genus Drepanogynis Guenée (Lepidoptera: Geometridae). Transvaal Museum Monograph 13: 1–220.

[PF02] Panero, J. L., & V. A. Funk. 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae). Proceedings of the Biological Society of Washington 115 (4): 909–922.

[R-S02] Russell-Smith, A. 2002. A comparison of the diversity and composition of ground-active spiders in Mkomazi Game Reserve, Tanzania and Etosha National Park, Namibia. Journal of Arachnology 30 (2): 383–388.

Last updated: 31 December 2021.

Vernonieae

Little ironweed Vernonia cinerea, copyright J. M. Garg.

Belongs within: Cichorioideae.

The Vernonieae are a group of composite-flowered plants, mostly herbs to shrubs, found in tropical and subtropical parts of the world. Species of the type genus Vernonia commonly bear intensely purple flowers and certain species are used in Africa as leaf vegetables or oilseeds.

Characters (from Flora of North America): Annuals, biennials, perennials, or shrubs (rarely trees or lianas) (sap rarely milky). Leaves usually cauline, sometimes basal or basal and cauline; alternate (rarely subopposite distally or opposite); usually petiolate, sometimes sessile (or petioles winged); margins usually more or less dentate, sometimes entire (rarely lobed or dissected). Heads homogamous (discoid, rarely pseudo-radiant or -liguliflorous), usually in corymbiform, paniculiform, or scorpioid arrays, sometimes borne singly or in glomerules (rarely aggregated in second-order heads). Calyculi absent. Phyllaries usually persistent (rarely readily falling), in 2–8+ series, distinct, unequal, herbaceous to chartaceous, margins and/or apices sometimes scarious. Receptacles flat to convex, usually epaleate (often foveolate, sometimes setose). Ray florets absent (rarely corollas of peripheral florets enlarged, zygomorphic, more or less raylike). Disc florets bisexual, fertile; corollas white, ochroleucous, or pink to cyanic (rarely yellow); anther bases more or less sagittate (rarely tailed), apical appendages ovate to lanceolate; styles abaxially hirsutulous (at least distally), branches lance-linear to more or less lanceolate, adaxially continuously stigmatic from bases nearly to apices, apices acute, appendages essentially none. Cypselae more or less monomorphic within heads, columnar to clavate, fusiform, or prismatic, sometimes compressed, not beaked, bodies smooth, nerved, or ribbed (glabrous or hirsutulous to strigillose, sometimes resin-gland-dotted as well); pappi usually persistent, usually in two series (outer series of shorter, stouter bristles or narrow scales, inner of longer, usually barbellate bristles), sometimes in one series (bristles or scales, scales often aristate).

<==Vernonieae
    |--Lessingianthus D03
    |--Hesperomania T00
    |--Pacourina S06
    |--Centratherum S06
    |--Blanchetia S06
    |--Stilpnopappus S06
    |--Lepidaploa D03
    |    `--‘Vernonia’ eriolepis D03
    |--Piptocarpha N10
    |    |--P. angustifolia OB11
    |    `--P. tetrantha SWK87
    |--Mattfeldanthus D03
    |    |--M. andrade-limae (Barroso) Dematteis 2003 (see below for synonymy) D03
    |    `--M. mutisioides D03
    |--Elephantopus N10
    |    |--E. mollis N10
    |    |--E. scaber P03
    |    `--E. tomentosus K03
    `--Vernonia D03
         |--V. auriculifera PB27
         |--V. buxifolia J87
         |--V. cataractarum D03
         |--V. chamaedrys RJ11
         |--V. cinerascens PP07
         |--V. cinerea PP07
         |--V. ekmanii J87
         |--V. gertii Dematteis 2003 D03
         |--V. obionifolia CV06
         |    |--V. o. ssp. obionifolia CV06
         |    `--V. o. ssp. dentata CV06
         |--V. pattens MM96
         |--V. psilophylla D03
         |--V. riedelii H01
         |--V. saepium J87
         `--V. secunda D03

Mattfeldanthus andrade-limae (Barroso) Dematteis 2003 [=Vernonia andrade-limae Barroso 1962; incl. V. nobilis Robinson 1979, Mattfeldanthus nobilis (Robinson) Robinson 1980] D03

*Type species of generic name indicated

REFERENCES

[CV06] Craven, P., & P. Vorster. 2006. Patterns of plant diversity and endemism in Namibia. Bothalia 36 (2): 175–189.

[D03] Dematteis, M. 2003. New species and new combinations in Brazilian Vernonieae (Asteraceae). Taxon 52: 281–286.

[H01] Hempel, A. 1901. Descriptions of Brazilian Coccidae (continued). Annals and Magazine of Natural History, series 7, 7: 556–561.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.

[K03] Kulip, J. 2003. An ethnobotanical survey of medicinal and other useful plants of Muruts in Sabah, Malaysia. Telopea 10 (1): 81–98.

[MM96] Mound, L. A., & R. Marullo. 1996. The thrips of Central and South America: an introduction (Insecta: Thysanoptera). Memoirs on Entomology, International 6: 1–487.

[N10] Norrbom, A. L. 2010. Tephritidae (fruit flies, moscas de frutas). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 909–954. NRC Research Press: Ottawa.

[OB11] Orenstein, R. I., & D. Brewer. 2011. Family Cardinalidae (cardinals). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 330–427. Lynx Edicions: Barcelona.

[PP07] Pandey, R. P., & P. M. Padhye. 2007. Studies on phytodiversity of Arid Machia Safari Park-Kailana in Jodhpur (Rajasthan). Bulletin of the Botanical Survey of India 49: 15–78.

[P03] Paul, T. K. 2003. Botanical observations on the Purulia pumped storage hydropower project area, Bagmundi Hills, Purulia district, West Bengal. Bulletin of the Botanical Survey of India 45: 121–142.

[PB27] Pilsbry, H. A., & J. Bequaert. 1927. The aquatic mollusks of the Belgian Congo, with a geographical and ecological account of Congo malacology. Bulletin of the American Museum of Natural History 53 (2): 69–602, pls 10–77.

[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.

[SWK87] Snyder, N. F. R., J. W. Wiley & C. B. Kepler. 1987. The Parrots of Luquillo: Natural history and conservation of the Puerto Rican parrot. Western Foundation of Vertebrate Zoology: Los Angeles.

[S06] Stuckert, T. 1906. Distribución geográfica de la flora Argentina. Géneros de la familia de las compuestas. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 303–309.

[T00] Thorne, R. F. 2000. The classification and geography of the flowering plants: dicotyledons of the class Angiospermae (subclasses Magnoliidae, Ranunculidae, Caryophyllidae, Dilleniidae, Rosidae, Asteridae, and Lamiidae). The Botanical Review 66: 441–647.

Last updated: 21 December 2019.

Taraxacum

Common dandelion Taraxacum officinale, copyright H. Zell.

Belongs within: Cichorieae.

Taraxacum, the dandelions, is a genus of perennial herbs bearing solitary flower heads, of which the common dandelion T. officinale has been spread widely around the world in association with humans.

Characters (from Hickman 1993): Perennial from taproot; sap milky. Stems naked, hollow. Leaves all basal, toothed or lobed; lobes acute. Inflorescence with heads ligulate, solitary, scapose; phyllaries many, outer ovate to lanceolate, generally reflexed, inner erect, linear; receptacle convex, naked. Florets many; ligules yellow, readily withering. Fruit fusiform; ribs rough; beak slender, >> body; pappus of many, white, slender bristles, not plumose.

<==Taraxacum Wiggers 1780 BR65
    |--T. aleppicum PT98
    |--T. californicum H93
    |--T. eripodum O88
    |--T. hellenicum PT98
    |--T. japonicum MH98
    |--T. magellanicum Comm. ex Sch. Bip. 1855 (see below for synonymy) A61
    |--T. mitalii O88
    |--T. officinale Weber 1787 BR65 [incl. T. dens-leonis Desfontaines 1798 BR65, T. laevigatum H93]
    |--T. palustre C55
    |--T. pseudostenoceras O88
    |--T. sikkimense O88
    |--T. tibetanum O88
    `--T. vulgare LNB03

Taraxacum magellanicum Comm. ex Sch. Bip. 1855 [incl. T. officinale var. glabratus Kirk 1899, Leontodon glabratum, T. officinale var. pygmaea Hooker 1853] A61

*Type species of generic name indicated

REFERENCES

[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[BR65] Black, J. M., & E. L. Robertson. 1965. Flora of South Australia. Part IV. Oleaceae–Compositae. W. L. Hawes, Government Printer: Adelaide.

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 1. Librairie de Victor Masson: Paris.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[LNB03] Ledeganck, P., I. Nijs & L. Beyens. 2003. Plant functional group diversity promotes soil protist diversity. Protist 154 (2): 239–249.

[MH98] Morikawa, H., A. Higaki, M. Nohno, M. Takahashi, M. Kamada, M. Nakata, G. Toyohara, Y. Okamura, K. Matsui, S. Kitani, K. Fujita, K. Irifune & N. Goshima. 1998. More than a 600-fold variation in nitrogen dioxide assimilation among 217 plant taxa. Plant, Cell and Environment 21: 180–190.

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: an introductory note. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

Crepis

Crepis jacquini, copyright Jerzy Opioła.

Belongs within: Cichorieae.

Crepis, hawksbeards, is a primarily Northern Hemisphere genus of herbs with milky sap, bearing heads clustered in cymes (Hickman 1993).

Characters (from Hickman 1993): Annual, biennial, perennial from taproot; sap milky. Stems erect, <8 dm. Leaves basal or cauline, entire to pinnately lobed. Inflorescence with heads ligulate, clustered in cymes; phyllaries in two distinct series; receptacle naked. Florets 5–60; ligules yellow. Fruit tapered at both ends, sometimes beaked; pappus of many soft, hair-like bristles.

<==Crepis [incl. Barkhausia] S06
    |--C. acuminata H93
    |--C. artificialis D37
    |--C. asturica D37
    |--C. bakeri [incl. C. bakeri ssp. cusickii] H93
    |--C. biennis D37
    |--C. bursifolia D51
    |--C. capillaris [incl. C. virens] BR65
    |--C. commutata PT98
    |--C. dioscoridis PT98
    |--C. divaricata D37
    |--C. foetida BR65
    |    |--C. f. ssp. foetida BR65
    |    `--C. f. ssp. vulgaris BR65
    |--C. fuliginosa D51
    |--C. intermedia H93
    |--C. jacquini C55
    |--C. kotschyana D51
    |--C. leontodontoides D51
    |--C. modocensis [incl. C. modocensis ssp. subacaulis] H93
    |--C. monticola H93
    |--C. multiflora PT98
    |--C. nana [incl. C. nana ssp. ramosa] H93
    |--C. neglecta Linnaeus 1767 PL04
    |--C. occidentalis [incl. C. o. ssp. conjuncta, C. o. ssp. costata, C. o. ssp. pumila] H93
    |--C. pleurocarpa H93
    |--C. pygmaea C55
    |--C. rubra D37
    |--C. runcinata H93
    |    |--C. r. ssp. runcinata H93
    |    |--C. r. ssp. andersonii H93
    |    `--C. r. ssp. hallii H93
    |--C. sancta PT98
    |--C. setosa D51
    |--C. taraxacifolia H59
    |--C. tectorum D51
    |--C. tibetica O88
    `--C. vesicaria BR65
         |--C. v. var. vesicaria BR65
         `--C. v. var. taraxacifolia BR65

*Type species of generic name indicated

REFERENCES

[BR65] Black, J. M., & E. L. Robertson. 1965. Flora of South Australia. Part IV. Oleaceae–Compositae. W. L. Hawes, Government Printer: Adelaide.

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 1. Librairie de Victor Masson: Paris.

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

[D51] Dobzhansky, T. 1951. Genetics and the Origin of Species 3^rd ed. Columbia University Press: New York.

[H59] Healy, A. J. 1959. Contributions to a knowledge of the adventive flora of New Zealand, no. 7. Transactions of the Royal Society of New Zealand 86 (1): 113–118.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: an introductory note. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: Flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[PL04] Pohl, G., & I. Lenski. 2004. Zur Verbreitung und Vergesellschaftung von Pennisetum orientale Rich. in Nordeuböa (Griechenland) (Poaceae, Paniceae). Senckenbergiana Biologica 83 (2): 209–223.

[S06] Stuckert, T. 1906. Distribución geográfica de la flora Argentina. Géneros de la familia de las compuestas. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 303–309.

Samooidea

Features of Anaceros pauliani, from Lawrence (1959).

Belongs within: Grassatores.
Contains: Stenostygninae, Hinzuanius, Metabiantes, Biantes, Samoidae, Stygnomma.

The Samooidea are a group of laniatorean harvestmen found in Africa and the Neotropics that commonly possess a scopula and a sexually dimorphic metatarsus III (Sharma & Giribet 2011). Families of the Samooidea include the Biantidae, in which the eyes are not borne on a central eyemound but widely separated on the carapace.

See also: Biantidae: the importance of titillators.

<==Samooidea [Biantoidea]
    |--Biantidae [Biantoidae, Hinzuanidae] SG11
    |    |--Zairebiantes Kauri 1985 S92 [Zairebiantinae KPG07]
    |    |    `--*Z. microphthalmus Kauri 1985 S92
    |    |--Lacurbsinae KSP-G15
    |    |    |--Lacurbs Sørensen 1896 S92 [Lacurbsinae KPG07]
    |    |    |    |--L. nigrimana Roewer 1912 [=L. nigrimanus] S92
    |    |    |    `--L. spinosa Sørensen 1896 S92
    |    |    |--Eulacurbs Roewer 1949 KSP-G15, S92
    |    |    |    `--*E. paradoxa Roewer 1949 S92
    |    |    |--Metalacurbs Roewer 1914 KSP-G15, S92
    |    |    |    |--*M. simoni Roewer 1915 S92, KSP-G15
    |    |    |    |--M. cornipes (Roewer 1958) [=Prolacurbs cornines (l. c.)] S92
    |    |    |    |--M. oedipus (Roewer 1958) [=Prolacurbs oedipus] S92
    |    |    |    `--M. villiersi (Roewer 1953) [=Prolacurbs villiersi] S92
    |    |    `--Prolacurbs Roewer 1949 KSP-G15, S92
    |    |         `--*P. singularis Roewer 1949 S92
    |    |--Stenostygninae P-GAB09
    |    `--Biantinae KPG07
    |         |--Hinzuanius R23
    |         |--Metabiantes KPG07
    |         |--Eubiantes Roewer 1915 R23, S92
    |         |    `--*E. africanus Roewer 1915 S92
    |         |--Acudorsum Loman 1902 R23, S92
    |         |    `--*A. albimanum Loman 1902 S92
    |         |--Hoplobiantes Roewer 1915 R23
    |         |--Cryptobiantes Kauri 1961 K61
    |         |    `--*C. protector Kauri 1961 S92
    |         |--Fageibiantes Roewer 1949 KPG07, S92
    |         |    |--*F. bicornis (Fage 1946) [=Biantes bicornis, Fageibantes (l. c.) bicornis] S92
    |         |    `--F. bispina Lawrence 1959 [=Fageibantes (l. c.) bispina] S92
    |         |--Biantessus Roewer 1949 K61, S92
    |         |    |--*B. nigrotarsus (Lawrence 1933) [=Metabiantes nigrotarsus] S92
    |         |    `--B. vertebralis (Lawrence 1933) S92 [=Metabiantes vertebralis S92, *Biantanius vertebralis L63]
    |         |--Biantes S89
    |         |--Monobiantes Lawrence 1962 L62
    |         |    `--*M. benoiti Lawrence 1962 S92
    |         |--Biantomma Roewer 1942 L62, S92
    |         |    `--*B. nigrospinosum Roewer 1942 S92
    |         |--Clinobiantes Roewer 1927 L62, S92
    |         |    `--*C. paradoxus Roewer 1927 S92
    |         |--Biantella Roewer 1927 L62, S92
    |         |    `--*B. reticulata Roewer 1927 S92
    |         |--Anaceros Lawrence 1959 KSP-G15
    |         |    |--*A. humilis Lawrence 1959 S92
    |         |    |--A. anodonta Lawrence 1959 S92
    |         |    |--A. canidens Lawrence 1959 S92
    |         |    `--A. pauliani Lawrence 1959 S92
    |         |--Hirstienus Roewer 1949 KSP-G15, SPG11
    |         |    `--*H. nanus (Hirst 1913) [=Phalangodes nanus] SPG11
    |         |--Hovanoceros Lawrence 1959 KSP-G15, S92
    |         |    `--*H. bison Lawrence 1959 S92
    |         |--Ivobiantes Lawrence 1965 KSP-G15, S92
    |         |    `--*I. spinipalpis Lawrence 1965 S92
    |         |--Malgaceros Lawrence 1959 KSP-G15, KP-G15
    |         |    `--*M. boviceps Lawrence 1959 S92
    |         |--Probiantes croceus Roewer 1927 KSP-G15
    |         `--Tetebius Roewer 1949 KSP-G15, KP-G15
    |              `--*T. latibunus Roewer 1949 S92
    `--+--Samoidae SG11
       `--Stygnommatidae SG11
            |--Stygnomma SG11
            `--Stygnomimus Roewer 1927 SG11, KP-G15
                 |--*S. conopygus Roewer 1927 [=Stignomimus conopygus] KP-G15
                 `--S. malayensis Suzuki 1969 [=Stignomimus malayensis] KP-G15

Nomen nudum: Lacurbs fernandopoensis Prieto 1999 SP10

*Type species of generic name indicated

REFERENCES

[K61] Kauri, H. 1961. Opiliones. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 9–197. Almqvist & Wiksell: Uppsala.

[KPG07] Kury, A. B., & Pérez González, A. 2007. Biantidae Thorell, 1889. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 176–179. Harvard University Press: Cambridge (Massachusetts).

[KP-G15] Kury, A. B., & A. Pérez-González. 2015. A companion to part 2 of the world checklist of Opiliones species (Arachnida): Laniatores—Samooidea, Zalmoxoidea and Grassatores incertae sedis. Biodiversity Data Journal 3: e6663.

[KSP-G15] Kury, A. B., D. R. Souza & A. Pérez-González. 2015. World checklist of Opiliones species (Arachnida). Part 2: Laniatores—Samooidea, Zalmoxoidea and Grassatores incertae sedis. Biodiversity Data Journal 3: e6482.

[L62] Lawrence, R. F. 1962. Resultats scientifiques des missions zoologiques de l'I.R.S.A.C. en Afrique orientale (P. Basilewsky et N. Leleup, 1957). LXXIV. Opiliones. Annales du Musée Royal de l'Afrique Centrale: Sciences Zoologiques 110: 9–89.

[L63] Lawrence, R. F. 1963. The Opiliones of the Transvaal. Annals of the Transvaal Museum 24 (4): 275–304.

[P-GAB09] Pérez-González, A., & A. Alegre Barroso. 2009. On the enigmatic Heterolacurbs ovalis Roewer, 1912 (Opiliones, Laniatores, Biantidae). Zootaxa 2269: 65–67.

[R23] Roewer, C.-F. 1923. Die Weberknechte der Erde: Systematische Bearbeitung der bisher bekannnten Opiliones. Gustav Fischer: Jena.

[SP10] Santos, R., & C. E. Prieto. 2010. Los Assamiidae (Opiliones: Assamiidae) de Río Muni (Guinea Ecuatorial), con la descripción de ocho nuevas especies. Revista de Biologia Tropical 58 (1): 203–243.

[SG11] Sharma, P. P. & G. Giribet. 2011. The evolutionary and biogeographic history of the armoured harvestmen—Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida). Invertebrate Systematics 25: 106–142.

[SPG11] Sharma, P. P., C. E. Prieto & G. Giribet. 2011. A new family of Laniatores (Arachnida: Opiliones) from the Afrotropics. Invertebrate Systematics 25: 143–154.

[S89] Staręga, W. 1989. Harvestmen (Opiliones) from the Mascarene Islands Indian Ocean and resurrection of the family Zalmoxidae. Annals of the Natal Museum 30: 1–8.

[S92] Staręga, W. 1992. An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33 (2): 271–336.

Grassatores

Microcina edgewoodensis, copyright Friends of Edgewood.

Belongs within: Laniatores.
Contains: Sandokanidae, Epedanoidea, Assamioidea, Zalmoxoidea, Samooidea, Gonyleptoidea, Calicina, Sitalcina, Banksula, Texella.

The Grassatores are a clade of laniatorean harvestmen in which the penis lacks intrinsic muscles, with the glans expanding through hydraulic pressure alone.

Grassatores [Camptonoti] SG14
    |--+--Sandokanidae SG11
    |  `--+--Epedanoidea SG14
    |     `--+--+--Assamioidea SG14
    |        |  `--+--Zalmoxoidea SG11
    |        |     `--Samooidea SG11
    |        `--Gonyleptoidea SG14
    `--Phalangodidae [Lolinae, Paralolidae, Phalangodinae, Phalangodoidea] GV09
         |  i. s.: Ausobskya Martens 1972 UB08
         |           |--A. athos Martens 1972 T96
         |           |--A. brevipes Thaler 1996 T96
         |           |--A. hauseri Silhavy 1976 T96
         |           `--A. mahnerti Silhavy 1976 T96
         |         Idzubius akiyamae TI87
         |         Spalicus oeditarsus Roewer 1949 S01, KSP-G15
         |         Bindoona glauerti Roewer 1929 K54, KSP-G15
         |         Paralola Kratochvil 1951 K18
         |           `--P. buresi Kratochvíl 1951 UD05
         |         Haasus Roewer 1942 U07, M76
         |           `--H. judaeus Roewer 1949 S02
         |         Babrius Thorell 1890 R23
         |           `--B. murcidus Thorell 1891 KSP-G15
         |         Heterobabrius Roewer 1915 R23
         |           |--H. atroluteus Roewer 1915 R23
         |           `--H. granulatus Roewer 1927 KSP-G15
         |         Biconibunus Rower 1915 R23
         |           `--B. fuscus Roewer 1915 R23
         |         Euzalmoxis Roewer 1915 R23
         |           `--E. neoguinensis Roewer 1915 R23
         |         Hoplozalmoxis Roewer 1915 R23
         |           `--H. pallida Roewer 1915 R23
         |         Parazalmoxida Roewer 1916 R23
         |           `--P. solitaria Roewer 1916 R23
         |         Proscotolemon Roewer 1916 U07, S73
         |           `--*P. sauteri Roewer 1916 S73
         |                |--P. s. sauteri S73
         |                `--P. s. latens Suzuki 1973 S73
         |         Metaconomma Pickard-Cambridge 1905 R23, K03
         |           `--*M. femorale Pickard-Cambridge 1905 K03
         |         Belonisculus Roewer 1923 R23
         |           `--*B. jacobsoni Roewer 1923 R23
         |         Isaeolus Roewer 1954 R54, K03
         |           `--*I. zilchi Roewer 1954 K03
         |         Glennhuntia Shear 2001 S01
         |           `--*G. glennhunti Shear 2001 S01
         |         Phalangodinus Roewer 1912 R23, K03
         |           `--*P. surinamensis Roewer 1912 K03
         |         Metapellobunus Roewer 1923 R23, K03
         |           |--*M. unicolor (Roewer 1912) [=Pellobunus unicolor] K03
         |           `--M. lapidosus Roewer 1928 KSP-G15
         |         Neoscotolemon Roewer 1912 R23, K03
         |           |--*N. pictipes (Banks 1908) [=Scotolemon pictipes] K03
         |           `--N. lutzi Goodnight & Goodnight 1942 K03
         |         Pseudomitraceras Roewer 1912 R23, KA-Z11
         |           |--*P. brasiliense Roewer 1912 K03, KA-Z11 (see below for synonymy)
         |           |--P. curvatum Goodnight & Goodnight 1942 KA-Z11 [=P. curvatus K03]
         |           `--P. minutum Goodnight & Goodnight 1942 KA-Z11 [=P. minutus K03]
         |         Paraconomma Roewer 1915 R23, K03
         |           |--*P. argentinum Roewer 1915 [=P. argentina] K03
         |           |--P. ovale Goodnight & Goodnight 1942 [=P. ovala] K03
         |           `--P. spinooculorum Goodnight & Goodnight 1942 K03
         |         Guerrobunus Goodnight & Goodnight 1945 U07, K03 [incl. Caecoa Šilhavý 1974 K03]
         |           |--*G. minutus Goodnight & Goodnight 1945 [=Cynortina minutus, Dapessus minutus] K03
         |           |--G. arganoi (Šilhavý 1974) [=*Caecoa arganoi] K03
         |           `--G. vallensis Vázquez & Cokendolpher 1997 K03
         |         Savoa Forster 1949 F49
         |           `--*S. bonka Forster 1949 F49
         |         Ptychosoma Sørensen 1873 UO05, R23
         |           |--P. catalonicum T96
         |           `--P. vitellinum Sørensen 1873 T96, R23 [=Scotolemon vitellinus R23]
         |         Metagjellerujia Goodnight & Goodnight 1947 GG47
         |           `--*M. jewetti Goodnight & Goodnight 1947 GG47
         |         Foella Goodnight & Goodnight 1948 GG48
         |           `--*F. remingtoni Goodnight & Goodnight 1948 GG48
         |         Spinolatum Goodnight & Goodnight 1942 GG42a
         |           `--*S. mediale Goodnight & Goodnight 1942 GG42a, KA-Z11 [=S. medialis GG42a]
         |         Lola Kratochvíl 1937 non Hamel & Hamel 1929 (ICBN) UO05
         |           `--*L. insularis Kratochvíl 1937 UO05
         |         Nimbadus Roewer 1953 S92
         |           `--*N. femoralis Roewer 1953 SPG11
         |         Epedanestus Roewer 1938 W53
         |           `--E. geniculatus (Pocock 1903) W53 (see below for synonymy)
         |         Tweedielus S72
         |           |--T. brevipes Roewer 1949 S72
         |           `--T. longipes Roewer 1933 KSP-G15
         |--Remyus Roewer 1949 SG11, S92
         |    `--R. gracillimus Roewer 1949 S92
         `--+--Scotolemon Lucas 1860 SG11, M78
            |    |--*S. lespesii Lucas 1860 M78 [=Phalangodes lespesi R23, Ptychosoma lespesi R23]
            |    |--S. balearicus Rambla 1977 T96
            |    |--S. doriae Pavesi 1878 M78
            |    |--S. jaqueti Roewer 1912 R23
            |    |--S. krausi T96
            |    |--S. lucasi Simon 1872 [=Phalangodes lucasi] R23
            |    |--S. roeweri T96
            |    `--S. terricola Simon 1872 [=Phalangodes terricola] R23
            `--+--Calicina UB08
               `--+--Microcinella Ubick & Briggs 2008 UB08
                  |    |--*M. homi (Briggs & Ubick 1989) [=Microcina homi] UB08
                  |    `--M. coensis Ubick & Briggs 2008 UB08
                  `--+--+--+--Sitalcina UB08
                     |  |  `--Enigmina Ubick & Briggs 2008 UB08
                     |  |       |--*E. granita (Briggs 1968) [=Sitalcina granita] UB08
                     |  |       `--E. warrenorum Ubick & Briggs 2008 UB08
                     |  `--Microcina Briggs & Ubick 1989 UB08
                     |       |--*M. tiburona (Briggs & Hom 1966) [=Sitalcina tiburona] UB08
                     |       |--M. edgewoodensis Briggs & Ubick 1989 UB08
                     |       |--M. jungi Briggs & Ubick 1989 UB08
                     |       |--M. leei Briggs & Ubick 1989 UB08
                     |       |--M. lumi Briggs & Ubick 1989 UB08
                     |       |--M. potrero Ubick & Briggs 2008 UB08
                     |       |--M. sanbruno Ubick & Briggs 2008 UB08
                     |       |--M. stanford Ubick & Briggs 2008 UB08
                     |       `--M. tamalpais Ubick & Briggs 2008 UB08
                     `--+--Tularina Ubick & Briggs 2008 UB08
                        |    |--*T. tularensis Ubick & Briggs 2008 UB08
                        |    |--T. plumosa Ubick & Briggs 2008 UB08
                        |    `--T. scopula (Briggs 1968) [=Sitalcina scopula] UB08
                        `--+--Megacina Ubick & Briggs 2008 UB08
                           |    |--*M. cockerelli (Goodnight & Goodnight 1942) [=Sitalcina cockerelli] UB08
                           |    `--+--M. madera (Briggs 1968) [=Sitalcina madera] UB08
                           |       |--M. mayacma Ubick & Briggs 2008 UB08
                           |       `--M. schusteri Ubick & Briggs 2008 UB08
                           `--+--Banksula UB08
                              |--Texella UB08
                              |--Tolus Goodnight & Goodnight 1942 UB08, GG42b
                              |    `--*T. appalachius Goodnight & Goodnight 1942 GG42b [=Phalangodes appalachius K03]
                              |--Undulus Goodnight & Goodnight 1942 UB08, GG42b
                              |    `--*U. formosus Goodnight & Goodnight 1942 GG42b [=Phalangodes formosus K03]
                              |--Wespus Goodnight & Goodnight 1942 UB08, GG42b
                              |    `--*W. arkansasensis Goodnight & Goodnight 1942 GG42b (see below for synonymy)
                              |--Bishopella Roewer 1927 UB08, K03
                              |    |--*B. laciniosa (Crosby & Bishop 1924) K03, GG42b (see below for synonymy)
                              |    `--B. jonesi Goodnight & Goodnight 1942 GG42b
                              |--Phalangodes Tellkampf 1844 UB08, K03 (see below for synonymy)
                              |    |--*P. armata Tellkampf 1844 (see below for synonymy) K03
                              |    `--P. flavipes (Banks 1908) [=Scotolemon flavipes] K03
                              `--Crosbyella Roewer 1927 UB08, K03 (see below for synonymy)
                                   |--*C. spinturnix (Crosby & Bishop 1924) K03, GG42b [=Phalangodes spinturnix GG42b]
                                   |--C. distincta Goodnight & Goodnight 1942 GG42b (see below for synonymy)
                                   |--C. montana Goodnight & Goodnight 1942 GG42b [=Phalangodes montana K03]
                                   |--C. roeweri Goodnight & Goodnight 1942 GG42b (see below for synonymy)
                                   `--C. tuberculata Goodnight & Goodnight 1942 GG42b [=Phalangodes tuberculata K03]

Grassatores incertae sedis:
  Anamota Šilhavý 1979 K03
    `--*A. custodiens Šilhavý 1979 K03
  Belemarua Roewer 1949 K03
    `--*B. nitens (Soares & Soares 1946) [=Paramitraceras nitens, Timoleon nitens] K03
  Bissopius Roewer 1949 K03
    `--*B. rugosus Roewer 1949 K03
  Caecobunus Roewer 1927 K03
    `--*C. termitarum Roewer 1927 K03
  Caribula Šilhavý 1979 K03
    `--*C. longimana Šilhavý 1979 K03
  Cleombrotus Sørensen 1932 K03
    `--*C. minutus Sørensen 1932 K03
  Contuor Roewer 1963 K03
    `--*C. novum Roewer 1963 K03
  Detlefilus Roewer 1949 K03
    `--*D. trispinosus Roewer 1949 K03
  Ignacianulus Roewer 1957 K03
    `--*I. pumilio Roewer 1957 K03
  Jimeneziella Kury & Alonso-Zarazaga 2011 KA-Z11
    |--*J. decui Avram 1970 K03
    `--J. negreai Avram 1970 K03
  Alpazia Özdikmen & Kury 2006 [=Lapazia Roewer 1949 non Ferris 1937] OK06
    `--*A. minima (Roewer 1949) [=*Lapazia minima] OK06
  Manuelangelia Kury & Alonso-Zarazaga 2011 KA-Z11 (see below for synonymy)
    `--*M. tuberosa (González-Sponga 1998) KA-Z11 [=*Limonia tuberosa K03]
  Liomma Roewer 1959 K03
    `--*L. laeve Roewer 1959 K03
  Micrisaeus Roewer 1957 K03
    `--*M. gracillimus Roewer 1957 K03
  Mirda Šilhavý 1973 K03
    `--*M. insulanus (Banks 1901) [=Stygnus insulanus] K03
  Munis Roewer 1963 K03
    `--*M. multivirgatus Roewer 1963 K03
  Octophthalmus Wood 1869 K03
    `--*O. marginatus Wood 1869 (n. d.) K03
  Ortizia Roewer 1952 K03
    `--*O. gracilipes Roewer 1952 K03
  Neoparalus Özdikmen 2006 [=Paralus Roewer 1949 non Rafinesque 1815] O06
    `--*N. granitus (Roewer 1949) [=*Paralus granitus] O06
  Pentos Roewer 1952 K03
    `--*P. pygoplus Roewer 1952 K03
  Pucallpana Avram & Soares 1983 K03
    `--*P. pullex Avram & Soares 1983 K03
  Siryseus Roewer 1949 K03
    `--*S. tibialis Roewer 1949 K03
  Tarmaops Roewer 1956 K03
    `--*T. koepckei Roewer 1956 K03
  Valifema Šilhavý 1979 K03
    `--*V. blanda Šilhavý 1979 K03

*Bishopella laciniosa (Crosby & Bishop 1924) K03, GG42b [=Phalangodes laciniosa GG42b; incl. B. marianna Goodnight & Goodnight 1942 K03]

Crosbyella Roewer 1927 UB08, K03 [incl. Goodnightiella Roewer 1949 non Soares & Soares 1945 K03]

Crosbyella distincta Goodnight & Goodnight 1942 GG42b [=*Goodnightiella distincta K03, Phalangodes distincta K03]

Crosbyella roeweri Goodnight & Goodnight 1942 GG42b [=Crosbiella (l. c.) roeweri K03, Phalangodes roeweri K03]

Epedanestus geniculatus (Pocock 1903) W53 [=Epedanus geniculatus R23, Heterobiantes geniculatus W53, Pseudobiantes geniculatus W53]

Manuelangelia Kury & Alonso-Zarazaga 2011 KA-Z11 [=Limonia González-Sponga 1998 nec Meigen 1803 nec Agassiz 1846 nec Carvalho 1985 K03]

Phalangodes Tellkampf 1844 UB08, K03 [=Acanthocheir Lucas 1861 K03, KA-Z11; incl. Phrixis Cope 1872 K03]

*Phalangodes armata Tellkampf 1844 [=*Acanthocheir armata, A. armatum; incl. *Phrixis longipes Cope 1872, Phryxus (l. c.) longipes] K03

*Pseudomitraceras brasiliense Roewer 1912 K03, KA-Z11 [=Preudomitraceras (l. c.) brasiliensis K03, Ps. brasiliensis K03]

*Wespus arkansasensis Goodnight & Goodnight 1942 GG42b [=Phalangodes arkansasensis K03]

*Type species of generic name indicated

REFERENCES

[F49] Forster, R. R. 1949. Opiliones from the Solomon Islands. Records of the Australian Museum 22 (2): 141–147.

[GV09] Giribet, G., L. Vogt, A. Pérez González, P. Sharma & A. B. Kury. 2009. A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores. Cladistics 25: 1–30.

[GG42a] Goodnight, C. J., & M. L. Goodnight. 1942a. Phalangids from British Guiana. American Museum Novitates 1167: 1–13.

[GG42b] Goodnight, C. J., & M. L. Goodnight. 1942b. New Phalangodidae (Phalangida) from the United States. American Museum Novitates 1188: 1–18.

[GG47] Goodnight, C. J., & M. L. Goodnight. 1947. Report on a collection of phalangids from New Guinea. Transactions of the American Microscopical Society 66 (4): 328–338.

[GG48] Goodnight, C. J., & M. L. Goodnight. 1948. New phalangids from the Southwest Pacific. American Museum Novitates 1371: 1–14.

[K54] Kauri, H. 1954. Report from Professor T. Gislén’s expedition to Australia in 1951–1952. 9. Harvest-spiders from S. W. Australia. Lunds Universitets Ärsskrift. N. F. Avd. 2. 50 (11): 1–10.

[K03] Kury, A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, volumen especial monográfico 1: 1–337.

[K18] Kury, A. B. 2018. Familial nomina in harvestmen (Arachnida, Opiliones). Bionomina 13: 1–27.

[KA-Z11] Kury, A. B. & M. A. Alonso-Zarazaga. 2011. Addenda and corrigenda to the "Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones)". Zootaxa 3034: 47–68.

[KSP-G15] Kury, A. B., D. R. Souza & A. Pérez-González. 2015. World checklist of Opiliones species (Arachnida). Part 2: Laniatores—Samooidea, Zalmoxoidea and Grassatores incertae sedis. Biodiversity Data Journal 3: e6482.

[M76] Martens, J. 1976. Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: Opiliones). Entomologica Germanica 3 (1–2): 51–68.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[O06] Özdikmen, H. 2006. Nomenclatural changes for some Laniatores (Opiliones) genera: New substitute names and new combinations. Mun. Ent. Zool. 1 (1): 63–68.

[OK06] Özdikmen, H., & A. B. Kury. 2006. Three homonymous generic names in Araneae and Opiliones. Journal of Arachnology 34 (1): 279–280.

[R23] Roewer, C.-F. 1923. Die Weberknechte der Erde: Systematische Bearbeitung der bisher bekannnten Opiliones. Gustav Fischer: Jena.

[R54] Roewer, C. F. 1954. Spinnentiere aus El Salvador, I. (Arachnoidea: Pedipalpi, Solifuga, Opiliones-Laniatores). Senckenbergiana Biologica 35: 57–73.

[S02] Schwendinger, P. J. 2002. Penis morphology in Oncopodidae (Opiliones, Laniatores): Evolutionary trends and relationships. Journal of Arachnology 30 (2): 425–434.

[SG11] Sharma, P. P., & G. Giribet. 2011. The evolutionary and biogeographic history of the armoured harvestmen—Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida). Invertebrate Systematics 25: 106–142.

[SG14] Sharma, P. P., & G. Giribet. 2014. A revised dated phylogeny of the arachnid order Opiliones. Frontiers in Genetics 5 (255): 1–13.

[SPG11] Sharma, P. P., C. E. Prieto & G. Giribet. 2011. A new family of Laniatores (Arachnida: Opiliones) from the Afrotropics. Invertebrate Systematics 25: 143–154.

[S01] Shear, W. A. 2001. Two new cave-dwelling harvestmen from Western Australia (Arachnida: Opiliones: Assamiidae and “Phalangodidae”). Records of the Western Australian Museum Supplement 64: 153–158.

[S92] Staręga, W. 1992. An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33 (2): 271–336.

[S72] Suzuki, S. 1972. Opiliones of Semangkok Forest Reserve, Malaysia. Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 24 (1): 1–37.

[S73] Suzuki, S. 1973. Opiliones from the South-west Islands, Japan. Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 24: 205–279.

[T96] Thaler, K. 1996. Neue Funde europäischer Krallenweberknechte (Arachnida, Opiliones: Phalangodidae, Travuniidae). Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 83: 135–148.

[TI87] Tsurusaki, N., & H. Ikeda. 1987. Opiliones of the Hakone district, Japan. Report of the Owakidani Natural History Museum 7: 1–16.

[U07] Ubick, D. 2007. Phalangodidae Simon, 1879. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 217–221. Harvard University Press: Cambridge (Massachusetts).

[UB08] Ubick, D., & T. S. Briggs. 2008. The harvestman family Phalangodidae. 6. Revision of the Sitalcina complex (Opiliones: Laniatores). Proceedings of the California Academy of Sciences, series 4, 59 (1): 1–108.

[UD05] Ubick, D., & J. A. Dunlop. 2005. On the placement of the Baltic amber harvestman Gonyleptes nemastomoides Koch & Berendt, 1854, with notes on the phylogeny of Cladonychiidae (Opiliones, Laniatores, Travunioidea). Mitteilungen Museum für Naturkunde, Geowissenschaftlichen Reihe 8: 75–82.

[UO05] Ubick, D., & R. Ozimec. 2005. On the harvestman genus Lola Kratochvíl (Opiliones: Laniatores). Natura Croatica 14 (3): 161–174.

[W53] Wang F.-C. 1953. A list of Chinese spiders of the order Opiliones. Kunchong Xuebao [Acta Entomologica Sinica] 3: 503–512.