Androsace

Androsace chamaejasme, copyright Denis Barthel.

Belongs within: Primulales.

Androsace is a mostly boreal and alpine genus of plants found in Eurasia and North America.

Characters (from Flora of North America): Herbs usually annual or perennial, rarely biennial, sometimes cushion- or mat-forming, rarely slightly succulent. Rhizomes absent; roots fibrous or a slender taproot. Stems ascending, simple. Leaves in single or multiple basal rosettes, simple; petiole absent or obscure, slightly winged; blade lanceolate to spatulate or cuneate, base attenuate, cuneate, or truncate, margins entire or moderately dentate, apex acute to obtuse (often dentate), surfaces usually hairy, hairs grayish white, simple or forked. Scapes 1-25. Inflorescences umbels, 2-20-flowered, involucrate; bracts 1-10. Pedicels absent or erect to arcuate, elongating with age. Flowers homostylous; sepals 5, green, calyx broadly campanulate to subglobose or hemispheric, ± 5-angled, not keeled, glabrous, pilose, or puberulent, lobes shorter than tube; petals 5, white, sometimes fading to pink in age [pink to shades of red], corolla campanulate to salverform, tube yellow, ± inflated, lobes shorter than tube, apex emarginate to entire; stamens included; filaments indistinct, very short; anthers not connivent. Capsules globose to subglobose, valvate, dehiscent nearly to base. Seeds usually 3-4 in perennial species, 20-50+ in annual species, brown, 4-angled to somewhat trigonous, reticulate to almost smooth. x = 10.

<==Androsace
    |--A. chamaejasme C55
    |--A. delavayi O88
    |--A. globifera O88
    |--A. hookeriana O88
    |--A. imbricata C55
    |--A. lactea C55
    |--A. lehmannii O88
    |--A. muscoidea O88
    |    |--A. m. f. muscoidea O88
    |    `--A. m. f. longiscapa O88
    |--A. nortonii O88
    |--A. obtusifolia C55
    |--A. sarmentosa O88
    |--A. strigillosa O88
    |--A. tapete O88
    |--A. villosa C55
    `--A. zambalensis O88

*Type species of generic name indicated

REFERENCES

[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.

[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.

Tetraoninae

Lesser prairie chicken Tympanuchus pallidicinctus, copyright Greg Kramos.

Belongs within: Phasianidae.
Contains: Lagopus.

The Tetraonidae are the grouse, a group of gamebirds found primarily in colder parts of Eurasia and North America. Grouse are distinguished from other gamebirds by the presence of feathers on the lower leg. Males of the prairie chickens Tympanuchus and the sage grouse Centrocercus have large inflatable sacs on the neck that are used in mating displays. Members of the genus Tetrao are found in cooler parts of the Palaearctic region; males are mostly black birds with red wattles above the eyes.

Synapomorphies (from Dyke et al. 2003): Distal end of condylus ventralis of humerus greatly extended distally; ischium shallow and wide, relative to width of synsacrum; tarsus partially feathered.

<==Tetraoninae [Tetraonidae, Urogallinae]
    |  i. s.: Canachites canadensis FP64
    |         Lyrurus Swainson 1832 CC10, M02 [Lyrurinae]
    |         Tetrastes Keysserling & Blasius 1840 MMJ03, B94 [Tetrastinae]
    |--Bonasa Stephens 1810 JT12, M02
    |    |--B. bonasia (Linnaeus 1758) M02 (see below for synonymy)
    |    `--B. sewerzowi JT12
    `--+--‘Bonasa’ umbellus JT12
       `--+--+--Lagopus BKB15
          |  `--+--Centrocercus BKB15
          |     |    |--C. minimus JT12
          |     |    `--C. urophasianus JT12
          |     `--+--‘Dendragapus’ obscurus JT12
          |        `--Tympanuchus JT12
          |             |  i. s.: T. stirtoni FP64
          |             |--T. cupido BKB15
          |             |    |--T. c. cupido CC10
          |             |    |--T. c. americanus (Reichenbach 1853) L81, CC10 [=Cupidonia americana CC10]
          |             |    |--T. c. attwateri USDI77
          |             |    `--T. c. pinnatus (Brewster 1885) [=Cupidonia pinnata] CC10
          |             `--+--T. pallidicinctus BKB15
          |                `--T. phasianellus (Linnaeus 1758) BKB15, CC10 (see below for synonymy)
          |                     |--T. p. phasianellus CC10
          |                     `--T. p. columbianus (Ord in Guthrie 1815) (see below for synonymy) CC10
          `--+--+--Falcipennis canadensis BKB15
             |  `--Dendragapus BKB15
             |       |--D. canadensis JT12
             |       |--D. falcipennis JT12
             |       `--D. fuliginosus JT12
             `--Tetrao Linnaeus 1758 JT12, M02 [incl. Urogallus Scopoli 1777 B94]
                  |  i. s.: T. canadensis Linnaeus 1758 L58
                  |         T. cupido Linnaeus 1758 L58
                  |         T. marilandicus Linnaeus 1758 L58
                  |         T. orientalis Linnaeus 1758 L58
                  |         T. urogalloides C01
                  |--+--T. mlokosiewiczi JT12
                  |  `--T. tetrix Linnaeus 1758 JT12, CC10 (see below for synonymy)
                  |       |--T. t. tetrix CC10
                  |       `--‘Lyrurus’ t. britannicus Witherby & Lönnberg 1913 CC10
                  `--+--T. parvirostris JT12
                     `--T. urogallus Linnaeus 1758 JT12, M02 (see below for synonymy)

Bonasa bonasia (Linnaeus 1758) M02 [=Tetrao bonasia M02, Tetrastes bonasia T89; incl. Tetrastes praebonasia Jánossy 1974 M02, B. praebonasia M02, Lagopus voinstvenskii Ganea 1972 M02]

Tetrao tetrix Linnaeus 1758 JT12, CC10 [=Lyrurus tetrix CC10; incl. Lagopus balcanicus Boev 1995 M02, Lyrurus tetrix longipes Mourer-Chauviré 1975 M02, Ly. partium Kretzoi 1962 M02, Tetrao partium M02]

Tetrao urogallus Linnaeus 1758 JT12, M02 [incl. T. conjugens Jánossy 1974 M02, T. macropus Jánossy 1976 M02, T. praeurogallus Jánossy 1969 M02, T. rhodopensis Boev 1998 M02]

Tympanuchus phasianellus (Linnaeus 1758) BKB15, CC10 [=Tetrao phasianellus CC10, Pedioecetes phasianellus CC10]

Tympanuchus phasianellus columbianus (Ord in Guthrie 1815) [=Phasianus columbianus, Pedioecetes phasianellus columbianus, Podioecetus (l. c.) columbianus] CC10

*Type species of generic name indicated

REFERENCES

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[CC10] Checklist Committee (OSNZ). 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica 4^th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.

[C01] Csiki, E. 1901. Utivázlat [Reiseskizze]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. xii–xli. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[MMJ03] Mayr, G., A. Manegold & U. S. Johansson. 2003. Monophyletic groups within ‘higher land birds’—comparison of morphological and molecular data. Journal of Zoological Systematics and Evolutionary Research 41: 233–248.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[T89] Takeshita, N. 1989. Nihon no Yachoo. Kogakukan: Tokyo.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

Last updated: 15 June 2019.

Francolinus

Grey francolin Francolinus pondicerianus interpositus, copyright Jugal Tiwari.

Belongs within: Phasianidae.

Francolinus, the francolins, is a diverse genus of quail- and partridge-like birds found in Africa and southern Asia. Members of the genus have streaked, barred or vermiculated plumage dorsally, and scutellate scalation on the rear of the tarsus (Dinesen et al. 1994).

Recent phylogenetic studies indicate that the genus Francolinus as previously recognised is broadly polyphyletic. In particular, the junglefowl of the genus Gallus, including the domestic chicken G. gallus, may be nested within Francolinus. Junglefowl are distinguished from other members of the Phasianidae by the presence in males of well-developed wattles and a comb on the head, and a strongly arched tail. They are currently found in tropical Asia, though fossils also indicate the presence of chickens in Europe during the warm interglacials of the Pleistocene (Mlíkovský 2002). The domestic chicken, of course, has been carried by humans around the world.

<==Francolinus Stephens 1819 [Francolinini] B94
    |--F. francolinus BKB15
    |    |--F. f. francolinus L81
    |    `--F. f. asiae L81
    `--+--+--F. coqui BKB15
       |  `--F. lathami JT12
       |--+--F. sephaena BKB15
       |  `--+--F. gularis JT12
       |     `--F. pondicerianus JT12
       |          |--F. p. pondicerianus L81
       |          `--F. p. interpositus L81
       `--+--+--F. levaillantii BKB15
          |  `--+--F. finschi BKB15
          |     `--+--F. shelleyi BKB15
          |        `--+--F. africanus BKB15
          |           `--F. levaillantoides BKB15
          |                |--F. l. levaillantoides B93
          |                `--F. l. jugularis (see below for synonymy) B93
          `--+--Bambusicola JT12
             |    |--B. fytchii JT12
             |    `--B. thoracica WBSJ82
             |         |--B. t. thoracica WBSJ82
             |         `--B. t. sonorivox WBSJ82
             `--Gallus Brisson 1760 BKB15, CC10 [incl. Pliogallus Gaillard 1939 M02; Gallini]
                  |  i. s.: G. beremendensis Jánossy 1976 M02
                  |         G. europaeus Harrison 1978 M02
                  |         G. moldovicus Burčak-Abramovič, Ganea & Šušpanov 1993 M02
                  |--*G. gallus (Linnaeus 1758) CC10, KB14, CC10 (see below for synonymy)
                  |    |--G. g. gallus SMD01
                  |    |--G. g. bankiva Temminck 1813 L81, CC10
                  |    |--G. g. domesticus (Linnaeus 1758) SMD01
                  |    |--G. g. ferrugineus L81
                  |    |--G. g. jabouillei RN72
                  |    |--G. g. murghii L81
                  |    `--G. g. spadiceus RN72
                  `--+--G. varius KB14
                     `--+--G. lafayetii BKB15
                        `--G. sonneratii BKB15

Francolinus incertae sedis:
  F. ahantensis JT12
  F. albogularis DL94
  F. camerunensis JT12
  F. chinensis S89
  F. clappertoni Children 1826 [=Perdix clappertoni; incl. F. rueppellii Gray 1844] S05
    |--F. c. clappertoni S05
    |--F. c. koenigseggi Madarász 1914 S05
    |--F. c. nigrosquamatus Neumann 1902 S05
    `--F. c. sharpii Ogilvie-Grant 1892 S05
  F. harwoodi JT12
  F. icterorhynchus L81
  F. jacksoni JT12
  F. nobilis JT12
  F. ochropectus FP64
  F. pictus DL94
  F. pintadeanus JT12
  F. psilolaemus JT12
  F. rufopictus JT12
  F. schlegelii DL94
  F. streptophorus JT12
  F. swierstrai FP64

Inorganic: Gallus gallus minilorientalus Okamura 1987 O87

Nomina nuda: Gallus steinheimensis Fraas in Engel 1908 M02

Francolinus levaillantoides jugularis [incl. Scleroptila jugularis cunenensis Roberts 1932, S. jugularis gilli Roberts ex Brooke 1993 (n. n.)] B93

*Gallus gallus (Linnaeus 1758) CC10, KB14, CC10 [=Phasianus gallus CC10; incl. *Pliogallus crassipes Gaillard 1939 M02, G. domesticus fossilis Giebel 1847 (n. n.) M02, P. kormosi Gaillard 1939 M02, G. gallus levantinus Pichon & Tchernov 1987 M02]

*Type species of generic name indicated

REFERENCES

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[B93] Brooke, R. K. 1993. Annotated catalogue of the Aves type specimens in the South African Museum. Annals of the South African Museum 102 (10): 327–349.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[CC10] Checklist Committee (OSNZ). 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica 4^th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.

[DL94] Dinesen, L., T. Lehmberg, J. O. Svendsen, L. A. Hansen & J. Fjeldså. 1994. A new genus and species of perdicine bird (Phasianidae, Perdicini) from Tanzania, a relict form with Indo-Malayan affinities. Ibis 136: 2–11.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[KB14] Kimball, R. T., & E. L. Braun. 2014. Does more sequence data improve estimates of galliform phylogeny? Analyses of a rapid radiation using a complete data matrix. PeerJ 2: e361.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[RN72] Rutgers, A., & K. A. Norris (eds) 1972. Encyclopaedia of Aviculture vol. 1. London, Blandford Press.

[S89] Salvadori, T. 1889. Viaggio di Leonardo Fea nella Birmania e nelle regioni vicine. XIX.—Uccelli raccolti nei Monti Carin a nord-est di Tounghoo, nel Pegù presso Rangoon e Tounghoo e nel Tenasserim presso Malewoon. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 369–438.

[SMD01] Skoracki, M., W. Magowski & J. Dabert. 2001. Picobia polonica sp. n. (Acari: Prostigmata: Syringophilidae), a new species of quill mite from the domestic hen, Gallus gallus domesticus (Aves: Phasianidae). Folia Parasitologica 48: 154–158.

[S05] Steinheimer, F. D. 2005. Eduard Rüppel’s avian types at the Natural History Museum, Tring (Aves). Senckenbergiana Biologica 85 (2): 233–264.

[WBSJ82] Wild Bird Society of Japan. 1982. A Field Guide to the Birds of Japan. Kodansha International Ltd.: Tokyo.

Last updated: 16 June 2019.

Tinamidae

Small-billed tinamou Crypturellus parvirostris, copyright Dario Sanches.

Belongs within: Aves.

The Tinamidae, tinamous, are superficially partridge-like birds found in South America. They are the only living flighted members of the palaeognath clade. Tinamous are small to medium-sized, plump terrestrial birds with short, strong legs and a rudimentary tail (Internet Bird Collection).

<==Tinamidae [Crypturi, Crypturidae, Dromaeomorphae, Pantinamidae, Tinami, Tinamiformes]
    |  i. s.: Nothocercus JT12
    |           |--N. bonapartei JT12
    |           |--N. julius JT12
    |           `--N. nigrocapillus JT12
    |         Taoniscus nanus JT12
    |         Crypturus soui S18
    |           |--C. s. soui S18
    |           `--C. s. modestus S18
    |--+--+--Eudromia Geoffroy St.-Hilaire 1832 JT12, B94 [Eudromiidae]
    |  |  |    |--E. elegans HK08
    |  |  |    `--E. formosa JT12
    |  |  `--Tinamotis Vigors 1837 JT12, B94 [Tinamotidae, Tinamotididae]
    |  |       |--T. ingoufi JT12
    |  |       `--T. pentlandii JT12
    |  `--+--Nothura JT12
    |     |    |--N. boraquira M03
    |     |    |--N. chacoensis JT12
    |     |    |--N. darwinii C04
    |     |    |--N. maculosa ZJM03
    |     |    `--N. minor JT12
    |     `--Nothoprocta JT12
    |          |  i. s.: N. ornata JT12
    |          |         N. pentlandii JT12
    |          |         N. taczanowskii JT12
    |          |--+--N. cinerascens BKB15
    |          |  `--Rhynchotus Spix 1825 BKB15, B94 [Rhynchotidae]
    |          |       |--R. maculicollis JT12
    |          |       `--R. rufescens M03
    |          `--+--N. curvirostris O93
    |             |--N. perdicaria BKB15
    |             |    |--N. p. perdicaria L81
    |             |    `--N. p. sanborni L81
    |             `--‘Tinamus’ tao BKB15
    `--+--Tinamus Hermann 1783 JT12, B94 [incl. Cryptura Illiger 1811 B94]
       |    |--T. castaneiceps S18
       |    |--T. guttatus HK08
       |    |--T. major (Gmelin 1789) [=Tetrao major] C04
       |    |--T. osgoodi JT12
       |    |--T. robustus [=T. major robustus] T62
       |    `--T. solitarius M03
       `--Crypturellus JT12
            |  i. s.: C. atrocapillus JT12
            |         C. bartletti JT12
            |         C. berlepschi JT12
            |         C. boucardi ZJM03
            |         C. brevirostris JT12
            |         C. casiquiare JT12
            |         C. cinereus JT12
            |         C. cinnamomeus M03
            |         C. duidae JT12
            |         C. erythropus JT12
            |         C. kerriae JT12
            |         C. noctivagus C04
            |         C. ptaritepui JT12
            |         C. transfasciatus JT12
            |--C. soui BKB15
            `--+--+--C. strigulosus BKB15
               |  `--C. undulatus BKB15
               `--+--+--C. obsoletus BKB15
                  |  `--C. tataupa JT12
                  `--+--C. parvirostris BKB15
                     `--C. variegatus BKB15

*Type species of generic name indicated

REFERENCES

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[C04] Clarke, J. A. 2004. Morphology, phylogenetic taxonomy, and systematics of Ichthyornis and Apatornis (Avialae: Ornithurae). Bulletin of the American Museum of Natural History 286: 1–179.

[HK08] Hackett, S. J., R. T. Kimball, S. Reddy, R. C. K. Bowie, E. L. Braun, M. J. Braun, J. L. Chojnowski, W. A. Cox, K.-L. Han, J. Harshman, C. J. Huddleston, B. D. Marks, K. J. Miglia, W. S. Moore, F. H. Sheldon, D. W. Steadman, C. C. Witt & T. Yuri. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320: 1763–1768.

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[M03] Mayr, G. 2003. The phylogenetic affinities of the shoebill (Balaeniceps rex). Journal für Ornithologie 144: 157–175.

[O93] O'Hara, R. J. 1993. Systematic generalization, historical fate, and the species problem. Systematic Biology 42 (3): 231–246.

[S18] Stone, W. 1918. Birds of the Panama Canal Zone, with special reference to a collection made by Mr. Lindsey L. Jewel. Proceedings of the Academy of Natural Sciences of Philadelphia 70: 239–280.

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: Revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7–460.

[ZJM03] Zeffer, A., L. C. Johansson & Å. Marmebro. 2003. Functional correlation between habitat use and leg morphology in birds (Aves). Biological Journal of the Linnean Society 79: 461–484.

Monotropoideae

Umbellate wintergreen Chimaphila umbellata, copyright Christian Fischer.

Belongs within: Ericaceae.

The Monotropoideae are a Holarctic clade of herbaceous plants. Members of the tribes Pterosporeae and Monotropeae lack chlorophyll, acquiring nutrients through parasitism on soil fungi (Kron et al. 2002).

Characters (from Kron et al. 2002): Echlorophyllous mycotrophic herbs usually blackening on drying, stems annual, with spiral, entire, sessile, non-differentiated leaves convolute in bud. Pith homogeneous. Leaf epidermal cells not lignified, fibres not associated with midrib. Indumentum none, rarely with multicellular, obscurely glandular, hairs. Inflorescence terminal, racemose or flower single, terminal; bracts present, bracteoles none, basal or apical; calyx articulated with pedicel or not. Flowers (4-)5(-6)-merous, actinomorphic. Calyx lobes small, more or less persistent; corolla sympetalous or polypetalous, campanulate or tubular, or perianth uniseriate, more or less tubular, parts free, basally saccate, glabrous or with hairs adaxially. Stamens 10, the filaments not dilated basally, straight, glabrous or with short hairs; anther dehiscing by terminal pores or more or less elongated slits, inverting just before anthesis, appendages uncommon, small, smooth, paired, at anther-filament junction, or anthers hippocrepiform, dehiscing by slits, appendages none, surface usually smooth, endothecium at most poorly developed, pollen monadinous. Ovary (4-)5(-8)-locular, with axile to parietal placentation and many ovules per locule, superior; style usually impressed, little longer than ovary; stigma barely expanded, rarely lobed. Fruit a loculicidal capsule, berry or irregularly dehiscent; seeds with isodiametric cells moderately to massively thickened on inner surfaces, or tailed, with moderately elongated and unthickened cells; embryo minute, undifferentiated, with <30 cells; n = 8, 13, 26, 32 (x = 8, 13?).

<==Monotropoideae [Pyroloideae]
    |  i. s.: Andresia H03
    |--Pterosporeae KJ02
    |    |--Pterospora andromeda KJ02
    |    `--Sarcodes sanguinea KJ02
    |--Pyroleae [Pyrolaceae] KJ02
    |    |--Moneses uniflora KJ02
    |    |--Orthilia secunda KJ02
    |    |--Pyrola KJ02
    |    |    |--P. picta KJ02
    |    |    `--P. rotundifolia KJ02
    |    `--Chimaphila KJ02
    |         |--C. maculata KJ02
    |         |--C. menziesii KJ02
    |         `--C. umbellata KJ02
    `--Monotropeae [Monotropaceae] KJ02
         |--Allotropa virgata KJ02
         |--Hemitomes congestum KJ02
         |--Monotropsis odorata KJ02
         |--Pityopus californicus KJ02
         |--Pleuricospora fimbriolata KJ02
         |--Cheilotheca KJ02
         |--Hypopitys KJ02
         |--Monotropastrum KJ02
         `--Monotropa KJ02
              |--M. hypopithys KJ02
              `--M. uniflora KJ02

*Type species of generic name indicated

REFERENCES

[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311–449.

[KJ02] Kron, K. A., W. S. Judd, P. F. Stevens, D. M. Crayn, A. A. Anderberg, P. A. Gadek, C. J. Quinn & J. L. Luteyn. 2002. Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68: 335–423.

Arbutoideae

Sonoma manzanita Arctostaphylos densiflora 'Howard McMinn', copyright Pete Veilleux.

Belongs within: Ericaceae.

The Arbutoideae are a Holarctic clade of shrubs and trees, including the strawberry trees and manzanitas, with fleshy, brightly coloured fruits.

Characters (from Kron et al. 2002): Trees or shrubs, usually evergreen. Leaves usually alternate, non-ericoid, entire or serrate, convolute in bud. Leaf epidermal cells not lignified. Indumentum of stalked glandular hairs and sometimes elongate, non-glandular hairs. Inflorescence terminal, racemose or paniculate; bracts present, bracteoles paired, more or less basal; calyx not articulated with pedicel. Flowers (4-)5-merous, actinomorphic. Calyx lobes small, persistent; corolla sympetalous, urceolate, with long unicellular hairs on the inside, the lobes small. Stamens (8-)10, included, the filaments more or less strongly dilated basally, more or less straight, unicellular-pubescent or smooth; anthers tetrasporangiate, inverting only just before anthesis, slightly roughened, with a pair of dorso-apical spurs, lacking a fibrous endothecium, lacking disintegration tissue, dehiscing by terminal pores or slit-like pores. Pollen in tetrads, without viscin threads. Ovary 4-10-locular, with axile placentation and only 1 to few ovules per locule, superior, sometimes with fleshy papillae; style continuous with the ovary; stigma truncate. Fruit more or less fleshy with bony or fibrous endocarp, a drupe or drupe-like berry; seeds with testa cells more or less elongated; embryo with 2 cotyledons.

<==Arbutoideae [Arbuteae]
    |--Comarostaphylis arbutoides KJ02
    |--Ornithostaphylos oppositifolia KJ02
    |--Arbutus KJ02
    |    |--A. andrachne DH98
    |    |--A. canariensis KJ02
    |    |--A. menziesii KJ02
    |    |--A. unedo KJ02
    |    `--A. xalapensis SZ03
    `--Arctostaphylos KJ02
         |--A. alpina KJ02
         |--A. densiflora KJ02
         |--A. mariposa D51
         |--A. patula D51
         |--A. pungens KJ02
         |--A. tomentosa KJ02
         `--A. uva-ursi KJ02

*Type species of generic name indicated

REFERENCES

[DH98] Danin, A., & I. C. Hedge. 1998. Contributions to the flora of Jordan 2. A new species of Satureja (Labiatae) and some new records. Willdenowia 28: 135–142.

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

[KJ02] Kron, K. A., W. S. Judd, P. F. Stevens, D. M. Crayn, A. A. Anderberg, P. A. Gadek, C. J. Quinn & J. L. Luteyn. 2002. Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68: 335–423.

[SZ03] Sobek, E. A., & J. C. Zak. 2003. The Soil FungiLog procedure: method and analytical approaches toward understanding fungal functional diversity. Mycologia 95 (4): 590–602.

Plecotus

Brown long-eared bat Plecotus auritus, copyright Dragiša Savić.

Belongs within: Vespertilioninae.

Plecotus, the long-eared bats, is a Holarctic genus of bats with a narrow rostrum and ears much longer than wide (Miller 1907).

Characters (from Miller 1907): Dental formula i2/3, c1/1, pm2/3, m3/3, total 36. Upper incisors well devoloped, each with a distinct secondary cusp, the inner tooth much smaller than the outer, and separated from canine by a space about equal to its diameter; lower incisors subequal, trifid, the crowns increasing in size from first to third, and forming a continuous, strongly convex row between canines. Upper canines rather small and weak, but not peouliar in form; lower canine with cingulum produced into a distinct though small anterior basal cusp. Cheek teeth normal, though rather small; anterior upper premolar (pm²) in contact with canine, but separated from large premolar (pm⁴) by a narrow space; posterior premolar with distinct though small inner cusp; upper molars short on inner side, the spaces between the crowns unusually large; m¹ and m² with no trace of hypocone; m³ with crown area about half that of m², its metacone and third commissure small but distinct; lower molars with all the cusps present and well developed, the protoconid noticeably higher than the hypoconid. Skull with large, elongate and rounded braincase, but with rostrum slender and reduced, its upper surface convex in lateral view; lachrymal ridges well developed (subgenus Plecotus) or lachrymal region smoothly rounded (Corynorhinus); audital bullae large, rounded, their greatest diameter equal to about three times the distance between them. Ears very large, much longer than head, joined across forehead; nostrils opening upward, their orifice continued backward by a slit-like prolongation.

<==Plecotus Geoffroy 1813 [incl. Macrotus Leach 1816 (n. n.) non Gray 1843] M07
    |  i. s.: P. austriacus IT07
    |         P. mexicanus (Allen 1916) MB86
    |         P. rafinesquei Lesson 1827 [=Corynorhinus rafinesquei] K92
    |         P. taivanus IT07
    |         P. teneriffae IT07
    |--*P. (Plecotus) auritus (Linnaeus 1758) M07, K92 [=Vespertilio auritus M07]
    `--P. (Corynorhinus Allen 1865) G69, M07 (see below for synonymy)
         |--P. (*C.) macrotis M07
         `--P. (C.) townsendii Cooper 1837 G69, K92
              |--P. t. townsendii MB86
              |--P. t. australis (Handley 1955) MB86
              |--P. t. ingens BP87
              `--P. t. virginianus BP87

Nomen nudum: Macrotus europaeus Leach 1816 M07

Plecotus (Corynorhinus Allen 1865) G69, M07 [=Corynorhynchus Peters 1865 M07; incl. Synotus Allen 1864 non Keyserling & Blasius 1839 M07]

*Type species of generic name indicated

REFERENCES

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i–vi, 1–154.

[MB86] Matson, J. O., & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[M07] Miller, G. S., Jr. 1907. The families and genera of bats. Smithsonian Institution, United States National Museum, Bulletin 57: i–xvii, 1–282, pl. I–XIV.

Phyllodoceae

Pink mountain-heath Phyllodoce empetriformis, copyright Walter Siegmund.

Belongs within: Ericaceae.

The Phyllodoceae are a clade of boreal shrubs with flowers on articulated pedicels.

Characters (from Kron et al. 2002): Shrubs. Leaves evergreen, alternate and spiral, decussate, or whorled, ericoid or non-ericoid, entire to serrulate, convolute or revolute in bud. Leaf epidermal cells not lignified. Indumentum of glandular hairs and elongate non-glandular hairs, and sometimes papillae. Inflorescence terminal, rarely axillary, spicate, racemose, or corymbose, sometimes reduced; bracts paired and more or less basal, bracteoles usually present; calyx articulated with pedicel. Flowers 4–6-merous, actinomorphic. Calyx lobes variable in size, withering persistent or persistent; corolla sympetalous or choripetalous, rotate to urceolate, campanulate, the lobes small to large. Stamens 5–10, included or exserted, the filaments straight to curved, unicellular-pubescent or smooth; anthers smooth or roughened, lacking appendages, dehiscing by slits or terminal pores. Pollen usually with viscin threads. Ovary 2–6-locular, with axile placentation; style impressed or not; stigma truncate, sometimes slightly expanded. Fruit a septicidal capsule, sometimes partly loculicidal as well; seeds with testa cells more or less isodiametric to elongated; embryo with two cotyledons.

<==Phyllodoceae [Cladothamneae, Epigaeae, Leiophylleae, Phyllodocoideae]
    |--Elliottia KJ02
    |    |--E. bracteata KJ02
    |    `--E. racemosa KJ02
    `--+--Kalmia [incl. Leiophyllum, Loiseleuria] KJ02
       |    |--K. angustifolia KJ02
       |    |--K. polifolia KJ02
       |    `--+--K. latifolia KJ02
       |       `--+--K. buxifolia (Bergius) Gift, Kron & Stevens in Kron, Judd et al. 2002 KJ02
       |          `--K. procumbens (Linnaeus) Gift, Kron & Stevens in Kron, Judd et al. 2002 (see below for synonymy) KJ02
       `--+--+--Epigaea repens KJ02
          |  `--Rhodothamnus chamaecistus KJ02
          `--+--Kalmiopsis leachiana KJ02
             `--Phyllodoce KJ02
                  |--P. breweri W81
                  |--P. caerulea KJ02
                  |--P. empetriformis KJ02
                  |--P. nipponica KJ02
                  `--P. taxifolia [incl. Menziesia coerulea] C55

Kalmia procumbens (Linnaeus) Gift, Kron & Stevens in Kron, Judd et al. 2002 [=Azalea procumbens, Loiseleuria procumbens] KJ02

*Type species of generic name indicated

REFERENCES

[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. 2. Librairie de Victor Masson: Paris.

[KJ02] Kron, K. A., W. S. Judd, P. F. Stevens, D. M. Crayn, A. A. Anderberg, P. A. Gadek, C. J. Quinn & J. L. Luteyn. 2002. Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68: 335–423.

[W81] West, E. W. 1981. Adaptive patterns in behavior of the Sierran pika, Ochotona princeps. Dissertation Abstracts International B 41: 4374.

Iodeae

Flowering twig of Iodes sp., copyright Leonardo L. Co.

Belongs within: Garryales.

The Iodeae have been recognised as a tribe within the pantropical flowering plant family Icacinaceae, though phylogenetic analysis indicates that it is paraphyletic with regard to the Phytocreneae and Sarcostigma (Kårehed 2001).

Synapomorphies (from Kårehed 2001, as unnamed clade): Leaves with prominent major veins present; flowers functionally unisexual; fruit a laterally compressed drupe; pollen with spines originating from the foot-layer; columellae more or less long and fine, often heterogeneous.

<==Iodeae
    |--Mappianthus K01
    `--+--+--+--Rhyticaryum K01
       |  |  `--+--Sarcostigma [Sarcostigmateae] K01
       |  |     `--Stachyanthus K01
       |  `--+--Hosiea K01
       |     `--+--Natsiatopsis K01
       |        `--Natsiatum K01
       `--+--+--Polyporandra K01
          |  `--Iodes K01
          |       |--I. africana K01
          |       |--I. cirrhosa K01
          |       `--I. ovalis K01
          `--Phytocreneae K01
               |--+--Miquelia K01
               |  `--Phytocrene K01
               `--+--Polycephalium K01
                  `--+--Chlamydocarya K01
                     `--Pyrenacantha K01
                          |--P. grandifolia K01
                          `--P. malvifolia K01

*Type species of generic name indicated

REFERENCES

[K01] Kårehed, J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. American Journal of Botany 88 (12): 2259–2274.

Mormoopidae

Parnell's moustached bat Pteronotus parnellii, copyright Brock Fenton.

Belongs within: Noctilionoidea.

The Mormoopidae, moustached bats and ghost-faced bats, are family of bats found in tropical and subtropical parts of the New World, characterised by complex outgrowths of the lower lip that take the place of the nose leaf found in other bat groups.

Characters (from Miller 1907): Teeth normal; humerus without secondary articulation with scapula, the tubercles short, rising scarcely above level of head, epitrochlea small, hardly more than base to large spinous process; muzzle without nose leaf; lower lip with plate-like outgrowths.

<==Mormoopidae (see below for synonymy)
    |--Mormoops Leach 1821 [incl. Aello Leach 1821] M07
    |    |--*M. blainvillii M07
    |    |--‘*Aello’ cuvieri M07
    |    |--M. magna Silva Taboada 1974 AC98
    |    `--M. megalophylla Peters 1864 MB86
    `--Pteronotus Gray 1838 K92, C57 (see below for synonymy)
         |--*P. davyi Gray 1838 [=Chilonycteris davyi, Dermonotus davyi] C57
         |    |--P. d. davyi MB86
         |    `--P. d. fulvus (Thomas 1892) MB86 [incl. P. suapurensis calvus Goodwin 1958 G69]
         |--P. gymnonotus (Wagner 1843) K06, C57 (see below for synonymy)
         |--P. macleayii (Gray 1839) K92 [=*Chilonycteris macleayii C57; incl. Lobostoma cinnamomeum G66]
         |--P. parnellii MB86 [=*Phyllodia parnellii M07]
         |    |--P. p. parnellii MB86
         |    `--P. p. mexicanus (Miller 1902) MB86 [=P. rubiginosa mexicana G69]
         |--P. personata (Wagner 1843) G69, C57 [=Chilonycteris personata C57, Chylonycteris personatus C57]
         |    |--P. p. personatus G69
         |    `--P. p. psilotis (Dobson 1878) G69, C57 [=Chilonycteris psilotis C57]
         |--P. pristinus Silva Taboada 1974 AC98
         |--P. quadridens IT07
         `--P. rubiginosa (Wagner 1843) G69, C57 [=Chilonycteris rubiginosus C57, C. rubiginosa G69]
              |--P. r. rubiginosa G69
              `--P. r. fusca (Allen 1911) G69, C57 [=Chilonycteris rubiginosa fusca C57]

Mormoopidae [Chilonycteriinae, Chilonycterinae, Lobostomidae, Lobostominae, Mormopes, Mormophidae, Mormopida, Mormopidae, Mormopinae, Mormopini, Mormopsina, Phyllodiana] ST01

Pteronotus Gray 1838 K92, C57 [incl. Chilonycteris Gray 1839 G69, C57, Dermonotus Gill 1901 C57, Lobostoma Gundlach 1840 C57, Phyllodia Gray 1843 C57]

Pteronotus gymnonotus (Wagner 1843) K06, C57 [=Chilonycteris gymnonotus C57; incl. Dermonotus suapurensis Allen 1904 C57, Pteronotus suapurensis K06]

*Type species of generic name indicated

REFERENCES

[AC98] Alcover, J. A., X. Campillo, M. Macias & A. Sans. 1998. Mammal species of the world: additional data on insular mammals. American Museum Novitates 3248: 1–29.

[C57] Cabrera, A. 1957. Catalogo de los mamiferos de America del Sur. I (Metatheria—Unguiculata—Carnivora). Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto Nacional de Investigacion de Las Ciencias Naturales, Ciencias Zoológicas 4 (1): 1–307.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[G66] Gray, J. E. 1866. Notice of the examination of the type specimen of Aëllo cuvieri of Leach. Proceedings of the Zoological Society of London 1866: 147–148.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i–vi, 1–154.

[K06] Kwiecinski, G. G. 2006. Phyllostomus discolor. Mammalian Species 801: 1–11.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[M07] Miller, G. S., Jr. 1907. The families and genera of bats. Smithsonian Institution, United States National Museum, Bulletin 57: i–xvii, 1–282, pl. I–XIV.

Musteloidea

Channel Islands spotted skunk Spilogale gracilis amphiala, photographed by Brian Kentosh.

Belongs within: Carnivora.
Contains: Conepatus, Procyonidae, Mustelidae.

The Musteloidea is a clade of mostly small-bodied carnivorans, including the weasels, raccoons and related taxa. Previously divided between the families Procyonidae and Mustelidae, more recent phylogenetic analyses have agreed in placing the skunks (Mephitidae) and red panda Ailurus fulgens outside the clade formed by the former two taxa. The skunks are omnivorous animals native to North and South America, commonly with contrasting black and white colour patterns and with defensive anal scent glands that produce a noxious spray.

Synapomorphies (from Flynn et al. 1988): Auditory bulla with caudal ectotympanic ventrally and posteriorly expanded, with relative shortening of carotid canal (reversed in some sublineages); alisphenoid canal absent; third upper and lower molars both absent.

<==Musteloidea [Melinae] MJ11
    |  i. s.: Megalictis Matthew 1907 D07
    |           `--M. ferox D07
    |         Bavarictis RDT09
    |         Aelurocyon FNT88
    |         Zodiolestes FNT88
    |--+--Mydaus FS15
    |  |    |--M. javanensis FS15
    |  |    `--M. marchei FS15
    |  `--Mephitidae [Mephitinae] MJ11
    |       |  i. s.: Miomephitis pilgrimi MJ11
    |       |--+--Brachyprotoma obtusata FS15
    |       |  `--Conepatus FS15
    |       `--+--Mephitis FS15
    |          |    |--M. macroura Lichtenstein 1832 MB86
    |          |    |    |--M. m. macroura [incl. M. macroura vittata] G69
    |          |    |    `--M. m. milleri Mearns 1897 MB86
    |          |    `--M. mephitis DD06
    |          |         |--M. m. mephitis B75
    |          |         |--M. m. avia B75
    |          |         |--M. m. hudsonica Richardson 1829 [=M. americana var. hudsonica] B75
    |          |         `--M. m. mesomelas B75
    |          `--Spilogale FS15
    |               |--S. pygmaea FS15
    |               `--+--S. gracilis FS15
    |                  |    |--S. g. gracilis DD06
    |                  |    |--S. g. amphiala DD06
    |                  |    `--S. g. leucoparia Merriam 1890 MB86
    |                  `--+--S. angustifrons Howell 1902 FS15, MB86 [=S. gracilis angustifrons MB86]
    |                     `--S. putorius FS15
    |                          |--S. p. putorius B75
    |                          |--S. p. ambarvalis J68
    |                          |--S. p. interrupta (Rafinesque 1820) [=Mephitis interrupta] B75
    |                          `--S. p. tropicalis G69
    `--+--Mustelictis olivieri MJ11
       `--+--Ailuridae MJ11
          |    |--Amphictis ambigua SA06, MJ11
          |    `--+--Simocyon batalleri SA06
          |       `--Ailurinae SA06
          |            |--Magerictis imperialensis SA06
          |            `--+--Pristinailurus Wallace & Wang 2004 WW04
          |               |    `--*P. bristoli Wallace & Wang 2004 WW04
          |               `--+--Ailurus fulgens WW04
          |                  |    |--A. f. fulgens R84
          |                  |    `--A. f. styani R84
          |                  `--Parailurus WW04
          |                       |--P. anglicus WW04
          |                       `--P. hungaricus WW04
          `--+--+--Procyonidae MJ11
             |  |--Broiliana WW04
             |  |--Pseudobassaris riggsi MJ11
             |  `--Stromeriella RDT09
             `--+--+--+--Mustelidae MJ11
                |  |  `--Plesiogale RDT09
                |  `--Promartes RDT09
                `--Plesictis Pomel 1846 RDT09, D07
                     |--P. genettoides D07
                     |--P. himilidens D07
                     |--P. mayeri D07
                     `--P. vireti D07

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[DD06] Doty, J. B., & R. C. Dowler. 2006. Denning ecology in sympatric populations of skunks (Spilogale gracilis and Mephitis mephitis) in west-central Texas. Journal of Mammalogy 87 (1): 131–138.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[FNT88] Flynn, J. J., N. A. Neff & R. H. Tedford. 1988. Phylogeny of the Carnivora. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 73–116. Clarendon Press: Oxford.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[J68] Johnson, M. L. 1968. Application of blood protein electrophoretic studies to problems in mammalian taxonomy. Systematic Zoology 17 (1): 23–30.

[MB86] Matson, J. O., & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[MJ11] Meredith, R. W., J. E. Janečka, J. Gatesy, O. A. Ryder, C. A. Fisher, E. C. Teeling, A. Goodbla, E. Eizirik, T. L. L. Simão, T. Stadler, D. L. Rabosky, R. L. Honeycutt, J. J. Flynn, C. M. Ingram, C. Steiner, T. L. Williams, T. J. Robinson, A. Burk-Herrick, M. Westerman, N. A. Ayoub, M. S. Springer & W. J. Murphy. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334: 521–524.

[R84] Russell, J. K. 1984. Other procyonids. In: Macdonald, D. (ed.) All the World’s Animals: Carnivores pp. 98–99. Torstar Books Inc.: New York.

[RDT09] Rybczynski, N., M. R. Dawson & R. H. Tedford. 2009. A semi-aquatic Arctic mammalian carnivore from the Miocene epoch and origin of Pinnipedia. Nature 458: 1021–1024.

[SA06] Salesa, M. J., M. Antón, S. Peigné & J. Morales. 2006. Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas. Proceedings of the National Academy of Sciences of the USA 103: 379–382.

[WW04] Wallace, S. C., & X. Wang. 2004. Two new carnivores from an unusual late Tertiary forest biota in eastern North America. Nature 431: 556–559.

Last updated: 7 August 2018.

Ceratomorpha

Skeleton of Metamynodon planifrons, from Wikimedia Commons.

Belongs within: Perissodactyla.
Contains: Tapiroidea, Hyracodontidae, Rhinocerotidae.

The Ceratomorpha, tapirs and rhinoceroses, are a group of heavily built odd-toed hoofed mammals. Early representatives were small and possibly include the European Eocene genus Pachynolophus, alternatively placed with the Palaeotheriidae. Karagalax mamikhelensis, known from the early Eocene of Pakistan, is more firmly established as a ceratomorph, resembling later forms in the reduction of the post-canine diastema.

Synapomorphies (from Hooker & Dashzeveg 2004): M₃ hypolophid complete; P₃ paraconid strong and approaching height of paraconid; upper molars with metaconal fold consistently joined to metaconule forming complete metaloph; lower preultimate molar with hypolophid complete, comprising equal buccal and lingual segments joined into long unnotched loph, with hypoconulid median; post P₁ diastema absent.

<==Ceratomorpha [Rhinocerotoidea]
    |--Pachynolophus [Pachynolophidae] HD04
    |    |--P. hookeri Godinot in Godinot et al. 1987 HD04
    |    |--P. lavocati F02
    |    `--P. livinerensis F02
    `--+--Karagalax mamikhelensis HD04
       `--+--Tapiroidea RH14
          `--+--+--Deperetellidae RH14
             |  |    |--Deperetella cristatum RH14
             |  |    |--Pachylophus Tong & Lei 1984 SM93
             |  |    |--Haagella Heissig 1978 SM93
             |  |    `--Teleolophus Matthew & Granger 1925 SM93
             |  |         `--T. medius RH14
             |  `--Lophialetidae RH14
             |       |--Lophialetes expeditus RH14
             |       |--Parabreviodon Reshetov 1975 SM93
             |       |--Kalakotia Ranga Rao 1972 SM93
             |       |--Eoletes Biryukov 1974 SM93
             |       |--Breviodon Radinsky 1965 SM93
             |       |--Simplaletes Qi 1980 SM93
             |       `--Schlosseria Matthew & Granger 1926 SM93
             |            `--S. magister RH14
             `--Rhinocerotoidea W96
                  |  i. s.: Amynodontopsis bodei Stock 1933 W96
                  |--Hyracodontidae W96
                  |--Rhinocerotidae FS15
                  `--Amynodontidae [Amynodontinae] G88
                       |--Gigantamynodon CND05
                       |--Zaisanamynodon Beliaeva 1971 H96
                       |--Megalamynodon regalis H96, P96
                       |--Paramynodon H96
                       |--Procadurcodon Gromova 1960 H96
                       |    `--*P. lrientalis Gromova 1960 H96
                       |--Amynodon Marsh 1877 SM93
                       |    |--A. advenus (Marsh 1875) [incl. A. intermedius] W96
                       |    `--A. reedi Stock 1939 W96
                       |--Cadurcodon Kretzoi 1942 R06, D07
                       |    |--C. ardynense D07
                       |    `--C. saisanensis D07
                       `--Metamynodon Scott & Osborn 1887 [Metamynodontini] D07
                            |--M. chadronensis D07
                            |--M. mckinneyi D07
                            `--M. planifrons D07

Ceratomorpha incertae sedis:
  Alicornops complanatum CS04
  Didermoceros sumatrensis A71
  Amynodentopsis JJ84
  Parelasmotherium DW04
    |--P. linxiaense DW04
    `--P. simplum DW04
  Sinotherium DW04
  Shansirhinus Kretzoi 1942 DW04
    |--*S. brancoi (Schlosser 1903) [=Rhinoceros brancoi] DW04
    `--S. ringstromi Kretzoi 1942 DW04
  Menoceras Troxell 1921 D07
    |--M. arikarensis D07
    `--M. barbouri D07
  Elasmotherium Fischer 1808 FS15, D07
    |--E. caucasicum D07
    |--E. inexpectatum D07
    `--E. sibiricum FS15

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[CS04] Chaimanee, Y., V. Suteethorn, P. Jintasakul, C. Vidthayanon, B. Murandat & J.-J. Jaeger. 2004. A new orang-utan relative from the Late Miocene of Thailand. Nature 427: 439–441.

[CND05] Clarke, J. A., M. A. Norell & D. Dashzeveg. 2005. New avian remains from the Eocene of Mongolia and the phylogenetic position of the Eogruidae (Aves, Gruoidea). American Museum Novitates 3494: 1–17.

[DW04] Deng T., Wang X., Ni X. & Liu L. 2004. Sequence of the Cenozoic mammalian faunas of the Linxia Basin in Gansu, China. Acta Geologica Sinica (English Edition) 78 (1): 8–14.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[F02] Froehlich, D. J. 2002. Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla). Zoological Journal of the Linnean Society 134: 141–256.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[H96] Hanson, C. B. 1996. Stratigraphy and vertebrate faunas of the Bridgerian-Duchesnean Clarno Formation, north-central Oregon. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 206–239. Cambridge University Press.

[HD04] Hooker, J. J., & D. Dashzeveg. 2004. The origin of chalicotheres (Perissodactyla, Mammalia). Palaeontology 47 (6): 1363–1386.

[JJ84] Janis, C., & P. J. Jarman. 1984. The hoofed mammals. In: Macdonald, D. (ed.) All the World’s Animals: Hoofed Mammals pp. 28–39. Torstar Books: New York.

[P96] Prothero, D. R. 1996. Magnetic stratigraphy and biostratigraphy of the Middle Eocene Uinta Formation, Uinta Basin, Utah. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 3–24. Cambridge University Press.

[R06] Rose, K. D. 2006. The Beginning of the Age of Mammals. John Hopkins University Press: Baltimore.

[RH14] Rose, K. D., L. T. Holbrook, R. S. Rana, K. Kumar, K. E. Jones, H. E. Ahrens, P. Missiaen, A. Sahni & T. Smith. 2014. Early Eocene fossils suggest that the mammalian order Perissodactyla originated in India. Nature Communications 5: 5570.

[SM93] Stucky, R. K., & M. C. McKenna. 1993. Mammalia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 739–771. Chapman & Hall: London.

[W96] Walsh, S. L. 1996. S. Middle Eocene mammal faunas of San Diego County, California. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 75–119. Cambridge University Press.

Last updated: 9 February 2022.