Lepidagathis

Lepidagathis cristata, copyright Jayesh Patil.


Belongs within: Acanthoideae.

Lepidagathis is a pantropical genus of shrubs and subshrubs with flowers borne in dense spikes or thyrses.

Characters (from Kameyama 2008): Small decumbent to erect subshrubs, or erect to sprawling shrubs. Inflorescences simple (thyrse or spike) or compound indeterminate; most commonly secundiflorous spikes with two rows of sterile bracts and two rows of fertile bracts. Calyx zygomorphic, formed by five unequal sepals, lateral ones narrower and very often shorter than others, posterior or upper sepal wider, two anterior sepals generally fused to each other to greater extent than with lateral sepals. Corolla bilabiate, upper lip entire or shallowly bilobed, lower lip trilobed. Two or four fertile stamens, anterior anthers bithoecous, posterior anthers (if present) mono- or bithecous or reduced to staminodia. Pollen 3-colporate, reticulate, heterobrochate, lumina verrucate, pilate or baculate. Ovary with two ovules at each locule; stigma subcapitate, shallowly bilobed; nectariferous disk cupulate, covering one quarter to almost half of ovary; base of style with ring of long, straight hairs. Capsule outline oblong, elliptic to ovate, apex acute to acuminate, subquadrangular, with relatively thin walls and thick septum, usually with four seeds (sometimes two). Seeds lenticular, with hygroscopic trichomes that expand when moist.

Lepidagathis Willdenow 1800 K08 (see below for synonymy)
    |--*L. cristata Willdenow 1800 K08
    |--L. alopecuroides K08
    |--L. alvarezia K08
    |--L. aristata E09
    |--L. bandraensis PP07
    |--L. callistachys Kameyama 2008 K08
    |--L. chiapensis (Acosta) Kameyama 2008 [=Lophostachys chiapensis Acosta 1985] K08
    |--L. cuneiformis Kameyama 2008 K08
    |--L. cyanea (Leonard) Kameyama 2008 [=Lophostachys cyanea Leonard 1959] K08
    |--‘Lophostachys’ falcata SV00
    |--L. fasciculata VD03
    |--L. floribunda [=Lophostachys floribunda Pohl 1831] K08
    |--L. grandidieri K08
    |--L. guatemalensis (Donn. Sm.) Kameyama 2008 [=Lophostachys guatemalensis Donn. Sm. 1901] K08
    |--L. hyalina SV00
    |--L. laxifolia (Nees) Kameyama 2008 (see below for synonymy) K08
    |--L. meridionalis Kameyama 2008 K08
    |--L. montana (Mart.) Kameyama 2008 (see below for synonymy) K08
    |--L. nemoralis (Mart. ex Nees) Kameyama 2008 (see below for synonymy) K08
    |--L. paraensis Kameyama 2008 K08
    |--L. pohliana K08
    |--L. pseudoaristata Ensermu 2009 E09
    |--L. riedeliana K08
    |--L. scariosa SR07
    |--L. sessilifolia [=Lophostachys sessilifolia Pohl 1831] K08
    |--L. soconuscana (Daniel) Kameyama 2008 [=Lophostachys soconuscana Daniel 1993] K08
    |--L. trinervis PP07
    |--L. uxpanapensis (Acosta) Kameyama 2008 [=Lophostachys uxpanapensis Acosta 1985] K08
    |--L. villosa Hedrén 1993 K08
    `--L. wasshausenii Kameyama 2008 K08

Lepidagathis Willdenow 1800 K08 [incl. Apolepsis SV00, Lindauea SV00, Liberatia SV00, Lophostachys Pohl 1831 K08, Teliostachya Nees 1847 K08, Volkensiophyton SV00]

Lepidagathis laxifolia (Nees) Kameyama 2008 [=Lophostachys laxifolia Nees in Wall. 1832; incl. Lo. patula Mart. ex Nees in Mart. 1847] K08

Lepidagathis montana (Mart.) Kameyama 2008 [=Lophostachys montana Mart. ex Nees in Mart. 1847; incl. Lo. villosa Pohl 1831 non Lepidagathis villosa Hedrén 1993] K08

Lepidagathis nemoralis (Mart. ex Nees) Kameyama 2008 [=Lophostachys nemoralis Mart. ex Nees in Mart. 1847; incl. Lo. semiovata var. minor Nees in Mart. 1847, Lo. semiovata Nees in Mart. 1847] K08

*Type species of generic name indicated

REFERENCES

[E09] Ensermu K. 2009. Three new species of Acanthaceae from Ethiopia. Kew Bulletin 64 (1): 57–65.

[K08] Kameyama, C. 2008. New species, nomenclatural changes and lectotypifications in Neotropical Lepidagathis Willd. (Acanthaceae). Kew Bulletin 63 (4): 565–581.

[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.

[SR07] Sankar, R. V., K. Ravikumar, N. M. Ganesh Babu & D. K. Ved. 2007. Botany of Anapady MPCA, Palghat district, Kerala with special emphasis on species of conservation concern. Bulletin of the Botanical Survey of India 49: 165–172.

[SV03] Scotland, R. W., & K. Vollesen. 2000. Classification of Acanthaceae. Kew Bulletin 55 (3): 513–589.

[VD03] Venu, P., & P. Daniel. 2003. Is Hemigraphis venosa C.B. Clarke (Acanthaceae) extinct? Bulletin of the Botanical Survey of India 45: 91–96.

Mimulus section Paradanthus

Seep monkeyflower Mimulus guttatus, copyright Walter Siegmund.


Belongs within: Phrymaceae.

The section Paradanthus of the genus Mimulus has been recognised as including monkeyflowers with a deciduous corolla with an open mouth, calyx lobes much shorter than the tube, and with the overall plant being glabrous, puberulent or with more or less straight hairs (Hickman 1993). Phylogenetic analysis indicates that this group is non-monophyletic with North American species of Paradanthus paraphyletic to the sections Erythranthe and Simiolus (Beardsley et al. 2004).

<==Mimulus sect. Paradanthus
    |  i. s.: M. acutidens BS04
    |         M. diffusus BS04
    |         M. evanescens BS04
    |         M. grayi BS04
    |--M. inconspicuus BS04
    `--+--+--+--+--M. montiodes BS04 [incl. M. barbatus H93, M. discolor H93]
       |  |  |  `--M. suksdorfii BS04
       |  |  `--+--+--M. gracilipes BS04
       |  |     |  `--M. palmeri BS04
       |  |     `--+--M. purpureus BS04
       |  |        `--+--M. androsaceus BS04
       |  |           `--M. shevockii BS04
       |  `--+--M. primuloides BS04
       |     |    |--M. p. ssp. primuloides [incl. M. primuloides var. pilosellus] H93
       |     |    `--M. p. ssp. linearifolius H93
       |     `--+--+--M. rubellus BS04
       |        |  `--+--M. brewerii BS04
       |        |     `--+--M. bicolor BS04
       |        |        `--M. filicaulis [incl. M. bioletti] BS04
       |        `--M. sect. Erythranthe BS04
       |             |--+--M. eastwoodiae BS04
       |             |  `--M. nelsoni BS04
       |             |--M. lewisii BS04
       |             |--M. parishii BS04
       |             |--M. rupestris BS04
       |             `--+--M. cardinalis BS04
       |                `--M. verbenaceus BS04
       `--+--+--M. alsinoides BS04
          |  `--+--M. dentatus BS04
          |     `--M. sessilifolius BS04
          `--+--M. gemmiparus BS04
             `--+--+--+--M. bodinieri BS04
                |  |  `--+--M. nepalensis BS04
                |  |     `--M. tenellus BS04
                |  `--+--M. latidens BS04
                |     `--+--+--+--M. moschatus BS04 [incl. M. moschatus var. moniliformis H93]
                |        |  |  `--M. pulsiferae BS04
                |        |  `--+--M. dudleyii BS04
                |        |     `--+--M. floribundus BS04 [incl. M. arenarius H93, M. floribundus ssp. subulatus H93]
                |        |        `--M. norrisi BS04
                |        `--+--+--M. ampliatus BS04
                |           |  `--+--M. hymenophyllus BS04
                |           |     `--M. patulus BS04
                |           `--+--M. breviflorus BS04
                |              `--+--M. jungermannioides BS04
                |                 `--M. washingtonensis BS04
                `--+--M. exiguus BS04
                   `--M. sect. Simiolus BS04
                        |  i. s.: M. laciniatus BS04
                        |         M. microphyllus BS04
                        |         M. nasutus BS04
                        |         M. whipplei BS04
                        |--M. wiensii BS04
                        `--+--M. dentilobus BS04
                           `--+--M. glabratus BS04
                              |    |--M. g. ssp. glabratus BS04
                              |    |--M. g. ssp. fremontii BS04
                              |    `--M. g. ssp. utahensis BS04
                              |--+--M. luteus [incl. M. rivularis] BS04
                              |  `--+--M. cupreus BS04
                              |     `--M. depressus BS04
                              `--+--M. glaucescens BS04
                                 |--M. tilingii BS04
                                 |--M. yeocorensis BS04
                                 `--+--M. guttatus BS04 (see below for synonymy)
                                    |--M. micranthus BS04 [=M. guttatus ssp. micranthus H93]
                                    `--M. nudatus BS04

Mimulus guttatus BS04 [incl. M. guttatus ssp. arenicola H93, M. guttatus ssp. arvensis H93, M. guttatus ssp. litoralis H93]

*Type species of generic name indicated

REFERENCES

[BS04] Beardsley, P. M., S. E. Schoenig, J. B. Whittall & R. G. Olmstead. 2004. Patterns of evolution in western North American Mimulus (Phrymaceae). American Journal of Botany 91 (3): 474–489.

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

Phrymaceae

Southern bush monkeyflower Mimulus aurantiacus ssp. australis, copyright Seán A. O'Hara.


Belongs within: Lamiales.
Contains: Mimulus section Paradanthus, Mimulus section Eunanus.

The Phrymaceae are a smaller group of herbs and subshrubs with tubular, bilaterally symmetrical flowers, most of which were historically classified within the related family Scrophulariaceae.

Characters (from the Jepson Herbarium): Annual to shrub. Stem with cross-section generally round. Leaves opposite, simple, generally entire or toothed; stipules absent. Inflorescence a spike, raceme, or panicle, bracted, or flowers 1–2 in axils. Flower bisexual; calyx radial or more or less bilateral, tube long, generally ribbed, lobes 5; corolla generally bilateral, generally 2-lipped, upper 2- (rarely 1-)lobed, lower 3-lobed; stamens 4 in 2 pairs, epipetalous, included or exserted; pistil 1, ovary superior, chambers 1–2, placentas parietal or axile, style 1, stigma lobes 2, flat, folding together when touched. Fruit generally a capsule, generally ellipsoid, loculicidal (rarely indehiscent, 1-seeded).

<==Phrymaceae
    |--Phryma leptostachya C55
    |--Microcarpaea minima LK14
    `--Mimulus Linnaeus 1753 BS04, A61 [=M. subg. Mimulus BS04, M. subg. Synplacus BS04; Mimuleae, Mimulinae]
         |  i. s.: M. prostratus BR65
         |         M. repens Br. 1810 [incl. M. colensoi Kirk 1871] A61
         |--M. sect. Mimulus BS04
         |    |--M. uvedaliae BS04
         |    |    |--M. u. var. uvedaliae LK14
         |    |    `--M. u. var. lutea LK14
         |    `--+--M. gracilis BS04
         |       `--M. ringens BS04
         `--+--+--Leucocarpus perfoliatus BS04
            |  `--M. sect. Paradanthus BS04
            `--+--+--Berendtiella rugosa BS04
               |  `--Hemichaena fruticosa BS04
               `--M. subg. Schizoplacus BS04
                    |--Mimulus (sect. Mimuloides) pilosus [=Mimetanthe pilosa] BS04
                    `--+--+--M. parryi BS04
                       |  `--M. rupicola BS04
                       `--+--M. sect. Eunanus BS04
                          `--+--M. (sect. Mimulastrum) pictus BS04
                             `--M. sect. Oenoe BS04
                                  |  i. s.: M. brandegei BS04
                                  |         M. latifolius BS04
                                  |         M. traskiae BS04
                                  |--+--M. pygmaeus BS04
                                  |  `--+--M. tricolor BS04
                                  |     `--+--M. angustatus BS04
                                  |        `--M. pulchellus BS04
                                  `--+--+--M. torreyi BS04
                                     |  `--+--M. kelloggii BS04
                                     |     `--+--M. congdonii BS04
                                     |        `--M. douglasii BS04
                                     `--M. sect. Diplacus BS04
                                          |--M. aridus BS04
                                          |--M. aurantiacus BS04 (see below for synonymy)
                                          |    |--M. a. ssp. aurantiacus BS04
                                          |    `--M. a. ssp. australis BS04
                                          |--M. bifidus BS04
                                          |    |--M. b. ssp. bifidus BS04
                                          |    `--M. b. ssp. fasciculatus BS04
                                          |--M. clevelandii BS04
                                          |--M. flemingii BS04
                                          |--M. longiflorus BS04
                                          |    |--M. l. ssp. longiflorus BS04
                                          |    `--M. l. ssp. calycinus BS04
                                          `--M. puniceus BS04

Mimulus aurantiacus BS04 [incl. M. aurantiacus var. lompocensis H93, M. longiflorus var. rutilus H93]

*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).

[BS04] Beardsley, P. M., S. E. Schoenig, J. B. Whittall & R. G. Olmstead. 2004. Patterns of evolution in western North American Mimulus (Phrymaceae). American Journal of Botany 91 (3): 474–489.

[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. 2. Librairie de Victor Masson: Paris.

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

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

Palaeognathae

Reconstruction of Lithornis, copyright Sheatherius.


Belongs within: Ornithurae.
Contains: Struthio, Apteryx, Aepyornithidae, Casuariiformes, Dinornithiformes, Tinamidae.

The Palaeognathae are a clade of birds found primarily on the southern continents, including the ratites and tinamous. Members of this clade are united by the lack of fusion between the maxillar process of the nasal bone and the maxillary bone, as well as the flat dorsal surface and presence of a pair of furrows on the ventral surface of the mandibular symphysis (Mayr 2009).

Palaeognathae (see below for synonymy)
    |--+--+--Struthio WD17
    |  |  `--Palaeotis Lambrecht 1928 H86, M02 [Palaeotididae M05]
    |  |       `--*P. weigelti Lambrecht 1928 (see below for synonymy) M02
    |  `--+--+--+--Apteryx PB15
    |     |  |  `--Aepyornithidae ML14
    |     |  `--+--Casuariiformes WD17
    |     |     `--+--Dinornithiformes WD17
    |     |        `--Tinamidae WD17
    |     `--Rheidae [Opisthodactylidae, Rheiformes] BKB15
    |          |  i. s.: Opisthodactylus patagonicus Ameghino 1891 AH03
    |          |         Heterorhea dabbeni Rovereto 1914 U93
    |          |--Diogenornis fragilis Alvarenga 1983 ML14, M09
    |          `--+--Pterocnemia pennata L81
    |             `--Rhea Brisson 1760 B94
    |                  |--R. americana JT12
    |                  `--R. pennata JT12
    `--Lithornithidae [Eoaves, Lithornithidae, Lithornithiformes] WD17
         |--Pseudocrypturus Houde 1988 M05
         |    `--P. cercanaxius Houde 1988 M09
         |--Pediorallus Harrison & Walker 1977 M05
         |    `--*P. barbarae Harrison & Walker 1977 M02
         |--Promusophaga Harrison & Walker 1977 M05
         |    `--*P. meini Harrison & Walker 1977 M02
         |--Fissuravis Mayr 2007 M07
         |    `--*F. weigelti Mayr 2007 M07
         `--Lithornis Owen 1840 M05
              |  i. s.: *L. vulturinus Owen 1840 [incl. Promusophaga magnifica Harrison & Walker 1977] M02
              |         L. celetius Houde 1988 M09
              |         L. hookeri (Harrison 1984) [=Pediorallus hookeri] M02
              |         L. nasi (Harrison 1984) M09 [=Pediorallus nasi M02]
              |--L. promiscuus Houde 1988 WD17, M09
              `--+--L. plebius Houde 1988 WD17, M07
                 `--Paracathartes Harrison 1979 WD17, M07
                      `--P. howardae (Harrison 1979) M09

Palaeognathae incertae sedis:
  Remiornis Lemoine 1881 M02 [Remiornithidae M05, Remiornithiformes]
    `--*R. heberti Lemoine 1881 M02
  Eleutherornithidae [Eleutherornithini] U93
    |--Eleutherornis Schaub 1940 M02
    |    `--*E. helveticus Schaub 1940 M02
    `--Proceriavis Harrison & Walker 1979 U93, M02
         `--*P. maritini Harrison & Walker 1979 M02
  Eremopezus Andrews 1904 ML14, RS01 [Eremopezidae, Eremopezithes]
    `--*E. eocaenus Andrews 1904 [incl. Stromeria fajumensis Lambrecht 1929, S. fayumensis] RS01
  Stromeria Lambrecht 1929 U93

Inorganic: Rhea americana minilorientalis Okamura 1987 O87

Palaeognathae [Apteryges, Apterygimorphae, Brevipennes, Palaeognathiformes, Panpalaeognathae, Panratitae, Ratitae, Ratiti, Struthiones, Struthioni, Struthioniformes, Struthionimorphae, Struthionoidea, Timamomorphae]

*Palaeotis weigelti Lambrecht 1928 [incl. Paleogrus geiseltalensis Lambrecht 1935, Ornitocnemus geiseltalensis] M02

*Type species of generic name indicated

REFERENCES

[AH03] Alvarenga, H. M. F., & E. Höfling. 2003. Systematic revision of the Phorusrhacidae (Aves: Ralliformes). Papéis Avulsos de Zoologia 43 (4): 55–91.

[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.

[H86] Houde, P. 1986. Ostrich ancestors found in the Northern Hemisphere suggest new hypothesis of ratite origins. Nature 324: 563–565.

[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.

[M05] Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515–542.

[M07] Mayr, G. 2007. The birds from the Paleocene fissure filling of Walbeck (Germany). Journal of Vertebrate Paleontology 27 (2): 394-408.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[ML14] Mitchell, K. J., B. Llamas, J. Soubrier, N. J. Rawlence, T. H. Worthy, J. Wood, M. S. Y. Lee & A. Cooper. 2014. Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344 (6186): 898–900.

[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.

[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.

[RS01] Rasmussen, D. T., E. L. Simons, F. Hertel & A. Judd. 2001. Hindlimb of a giant terrestrial bird from the Upper Eocene, Fayum, Egypt. Palaeontology 44 (2): 325–337.

[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.

[WD17] Worthy, T. H., F. J. Degrange, W. D. Handley & M. S. Y. Lee. 2017. The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres). Royal Society Open Science 4: 170975.

Mimulus section Eunanus

Skunky monkeyflower Mimulus mephiticus, copyright Lars Rosengreen.


Belongs within: Phrymaceae.

The section Eunanus of the genus Mimulus is a group of monkeyflowers found in western North America, characterised by a divided placenta, pubescent styles, and small, nearly regular, corollas (Beardsley et al. 2004).

<==Mimulus sect. Eunanus
    |  i. s.: M. angustifolius BS04
    |         M. ovatus [=M. bigelovii var. ovatus] BS04
    |         M. spissus BS04
    |         M. subsecundus BS04
    |--M. nanus BS04
    |--+--M. clivicola BS04
    |  |--M. cusickii BS04
    |  `--M. jepsonii BS04
    `--+--+--M. rattanii BS04 [incl. M. rattanii ssp. decurtatus H93]
       |  |    |--M. r. ssp. rattanii BS04
       |  |    `--M. r. ssp. decurrens BS04
       |  `--+--M. brevipes BS04
       |     `--+--M. fremontii BS04
       |        `--M. johnstonii BS04
       `--+--M. mohavensis BS04
          `--+--+--M. layneae BS04 [incl. M. brachiatus H93]
             |  `--M. viscidus BS04
             `--+--M. mephiticus BS04 [=M. nanus var. mephiticus BS04; incl. M. coccineus H93, M. densus H93]
                `--+--+--M. bigelovii BS04
                   |  |    |--M. b. var. bigelovii BS04
                   |  |    `--M. b. var. cuspidatus BS04 [incl. M. bigelovii var. panamintensis H93]
                   |  `--M. bolanderi BS04 [incl. M. platylaemus H93]
                   `--+--M. leptaleus BS04
                      `--+--M. constrictus BS04
                         `--M. whitneyi BS04

*Type species of generic name indicated

REFERENCES

[BS04] Beardsley, P. M., S. E. Schoenig, J. B. Whittall & R. G. Olmstead. 2004. Patterns of evolution in western North American Mimulus (Phrymaceae). American Journal of Botany 91 (3): 474–489.

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

Dinornithiformes

Reconstruction of heavy-footed moa Pachyornis elephantopus, copyright Paul Martinson.


Belongs within: Palaeognathae.

The Dinornithiformes, moas, are a group of moderately to extremely large flightless birds known from New Zealand (an historical record of a species from Australia was based on a specimen of mislabelled provenance) that became extinct following human colonisation of the region.

Characters (from Worthy & Holdaway 2002): Skull with small orbits, large olfactory cavities, broad across occipital region. Basitemporal platform with prominent basipterygoid processes. Basiparasphenoidal rostrum long, with triangular expansions extending laterally beneath antorbital plate. Maxilla usually with large dorsal inflation enclosing cavity (maxillary antrum). Beak short, may be pointed or rounded, curved down at tip. 21–23 cervical, six dorsal, 18 sacral, and eleven caudal vertebrae, no pygostyle. Pelvis broad at and posterior to acetabulum. Wing entirely absent, pectoral girdle represented only by vestigial scapulocoracoid bone without glenoid facet. Sternum with well-developed lateral processes. Femur with concave articular surface for antitrochanter on pelvis, tibiotarsus with osseous bridge over tendinal groove, tarsometatarsus with two hypotarsal ridges.

<==Dinornithiformes [Anomalopterygidae, Anomalopteryginae, Dinornithes, Dinornithoidea, Emeidae]
    |--Megalapteryx Haast 1886 BH05, CC10 [incl. Palaeocasuarius Rothschild 1907 CC10]
    |    |--M. didinus (Owen 1883) (see below for synonymy) CC10
    |    `--M. benhami Archey 1941 BKB15, CC10
    `--+--Dinornis Owen 1843 BH05, CC10 (see below for synonymy)
       |    |--*D. novaezealandiae Owen 1843 CC10 (see below for synonymy)
       |    `--D. robustus Owen 1846 (see below for synonymy) CC10
       `--+--Anomalopteryx Reichenbach 1853 ML14, CC10 [=Anomalornis Hutton 1897; incl. Graya Bonaparte 1856]
          |    `--*A. didiformis (Owen 1843) (see below for synonymy) CC10
          `--+--Pachyornis Lydekker 1891 ML14, CC10 [incl. Mauiornis Oliver 1949 WH02, Pounamua Oliver 1949 WH02]
             |    |--P. geranoides (Owen 1848) ML14, CC10 (see below for synonymy)
             |    `--+--*P. elephantopus (Owen 1856) CC10, BKB15, CC10 (see below for synonymy)
             |       `--P. australis Oliver 1949 BKB15, CC10
             `--Emeinae CC10
                  |--Emeus Reichenbach 1853 BH05, CC10 (see below for synonymy)
                  |    `--*E. crassus (Owen 1846) (see below for synonymy) CC10
                  `--Euryapteryx Haast 1874 (see below for synonymy) CC10
                       |--*E. gravis (Owen 1870) CC10 (see below for synonymy)
                       `--E. curtus (Owen 1846) (see below for synonymy) CC10

*Anomalopteryx didiformis (Owen 1843) [=Dinornis didiformis, *Anomalornis didiformis; incl. Anomalopteryx antiqua Hutton 1893, Anomalopteryx antiquus Hutton 1892, Dinornis dromaeoides Owen 1844, Anomalopteryx dromaeoides, *Graya dromaeoides, D. dromioides Owen 1846, D. (Palapteryx) dromioides, Anomalopteryx fortis Hutton 1893, Anomalornis gracilis Hutton 1897, Dinornis oweni Haast 1886, Anomalornis owenii, Pachyornis owenii, Dinornis parvus Owen 1883, Anomalopteryx parva, Anomalopteryx parvus] CC10

Dinornis Owen 1843 BH05, CC10 [=Megalornis Owen 1843 non Gray 1841 CC10; incl. Moa Reichenbach 1853 CC10, Movia Reichenbach 1853 CC10, Owenia Gray 1855 CC10, Palapteryx Owen 1846 CC10, Tylopteryx Hutton 1891 CC10; Dinornithidae]

*Dinornis novaezealandiae Owen 1843 CC10 [=D. novaezelandiae CC10, *Megalornis novaezealandiae CC10; incl. D. excelsus Hutton 1891 CC10, D. firmus Hutton 1891 CC10, D. gazella Oliver 1949 CC10, D. giganteus Owen 1843 CC10, *Moa gigantea CC10, D. gigas Owen 1846 CC10, D. gracilis Owen 1854 CC10, *Tylopteryx gracilis CC10, D. hercules Oliver 1949 CC10, D. ingens Owen 1844 CC10, *Movia ingens CC10, *Palapteryx ingens CC10, D. struthioides Lydekker 1891 CC10, D. struthoides Owen 1844 CC10, *Owenia struthoides CC10, Tylopteryx struthoides WH02]

Dinornis robustus Owen 1846 [=D. ingens var. robustus, Palapteryx robustus; incl. D. altus Owen 1879, D. maximus Haast 1869, Palapteryx plenus Hutton 1891, D. potens Hutton 1891, D. strenuus Hutton 1893, D. torosus Hutton 1891, D. validus Hutton 1891] CC10

Emeus Reichenbach 1852 BH05, CC10 [incl. Meionornis Haast 1874 CC10, Mesopteryx Hutton 1891 CC10, Syornis Reichenbach 1853 CC10]

*Emeus crassus (Owen 1846) [=Dinornis crassus, Euryapteryx crassa, Eu. crassus, Syornis crassus; incl. Dinornis casuarinus Owen 1846, Anomalopteryx casuarina, Cela casuarinus, Emeus casuarinus, *Meionornis casuarinus, Mesopteryx casuarina, *Syornis casuarinus, Dinornis huttoni Owen 1879, Emeus huttonii, Megalapteryx huttoni, *Mesopteryx huttoni, Dinornis rheides Owen 1870, Cela rheides] CC10

Euryapteryx Haast 1874 [incl. Cela Reichenbach 1853 non Moehring 1758, Celeus Bonaparte 1856 non Boie 1831, Zelornis Oliver 1949] CC10

Euryapteryx curtus (Owen 1846) [=Dinornis curtus, Anomalopteryx curta, A. curtus, *Cela curtus, *Celeus curtus; incl. Euryapteryx exilis Hutton 1897, *Zelornis exilis, E. tane Oliver 1949] CC10

*Euryapteryx gravis (Owen 1870) CC10 [=Dinornis gravis CC10; incl. Emeus boothi Rothschild 1907 CC10, Euryapteryx boothi WH02, Eu. compacta Hutton 1893 CC10, Emeus gravipes Lydekker 1891 CC10, Eu. gravipes CC10, Emeus haasti Rothschild 1907 CC10, Zelornis haasti CC10, Euryapteryx kuranui Oliver 1930 CC10, Emeus parkeri Rothschild 1907 CC10, Euryapteryx pygmaeus Hutton 1891 CC10, Pachyornis pygmaeus CC10]

Megalapteryx didinus (Owen 1883) [=Dinornis didinus, Anomalopteryx didina, Meionornis didinus, Mesopteryx didina, Mes. didinus; incl. Palaeocasuarius elegans Rothschild 1907, *P. haasti Rothschild 1907, Megalapteryx hamiltoni Rothschild 1907, *M. hectori Haast 1886, M. tenuipes Lydekker 1891, Palaeocasuarius velox Rothschild 1907] CC10

*Pachyornis elephantopus (Owen 1856) CC10, BKB15, CC10 [=Dinornis elephantopus CC10, Euryapteryx elephantopus CC10; incl. D. elephantopus var. major Hutton 1875 CC10, Pachyornis immanis Lydekker 1891 CC10, Euryapteryx immanis CC10, Pachyornis inhabilis Hutton 1893 CC10, Dinornis elephantopus var. major Hutton 1875 CC10, Pachyornis (Pounamua) murihiku Oliver 1949 WH02, Euryapteryx ponderosus Hutton 1891 CC10, E. ponderosa CC10, Dinornis queenslandiae De Vis 1884 CC10, Dromiceius queenslandiae CC10, Pachyornis queenslandiae CC10, Pachyornis rothschildi Lydekker 1892 CC10, Pachyornis valgus Hutton 1893 CC10]

Pachyornis geranoides (Owen 1848) ML14, CC10 [=Palapteryx geranoides CC10, Anomalopteryx geranoides CC10, Cela geranoides CC10, Dinornis geranoides CC10, Euryapteryx geranoides CC10, Dinornis expunctus Archey 1927 CC10; incl. Pachyornis mappini Archey 1941 BKB15, CC10, Pac. (Mauiornis) mappini CC10, Pac. (*M.) septentrionalis Oliver 1949 CC10]

*Type species of generic name indicated

REFERENCES

[BH05] Baker, A. J., L. J. Huynen, O. Haddrath, C. D. Millar & D. M. Lambert. 2005. Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: the giant moas of New Zealand. Proceedings of the National Academy of Sciences of the USA 102 (23): 8257–8262.

[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 4th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.

[ML14] Mitchell, K. J., B. Llamas, J. Soubrier, N. J. Rawlence, T. H. Worthy, J. Wood, M. S. Y. Lee & A. Cooper. 2014. Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344 (6186): 898–900.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

Neognathae

Hoatzin Opisthocomus hoazin, copyright Francesco Veronesi.


Belongs within: Ornithurae.
Contains: Pelagornithidae, Galloanseres, Mirandornithes, Gruiformes, Charadriiformes, Caprimulgiformes, Columbaves, Telluraves, Phaethontiformes, Aequornithia.

The Neognathae are one of the primary clades of living birds, excluding only the ratites and tinamous. Members of the clade are characterised by an intrapterygoidal joint allowing greater mobility of the palate, as well as by a conjoint auditory tube and caudally closed ilioischiadic foramina in the pelvis (Mayr 2009). Within the Neognathae, most recent studies agree on a basal division between the galloanserines (gamefowl and waterfowl) and remaining species in the clade Neoaves. Basal relationships within the Neoaves remain highly contentious, with many large-scale analyses differing significantly in their results; some authors have even suggested that the base of Neoaves may be an insoluble hard polytomy. Particularly contentious has been the position of the hoatzin Opisthocomus hoazin, a medium-sized folivorous bird found in the Amazon basin. The Neotropical sunbittern Eurypyga helias and the New Caledonian kagu Rhynochetos jubatus together form another isolated clade that may be related to the tropicbirds of the Phaethontidae.

Neognathae [Amblypterae, Ciconii, Insessores, Odontoanserae, Pullastrae] HK08
    |--+--Pelagornithidae M11
    |  `--Galloanseres PB15
    `--Neoaves (see below for synonymy) HK08
         |  i. s.: Protopelicanus Reichenbach 1852 M05, M02
         |           `--*P. cuvieri Reichenbach 1852 M02
         |         Onychopteryx [Onychopterygidae] M09
         |           `--O. simpsoni Cracraft 1971 M09
         |--+--Juncitarsus Olson & Feduccia 1980 M05, M02 [Juncitarsinae]
         |  |    |--*J. gracillimus Olson & Feduccia 1980 M02
         |  |    `--J. merkeli Peters 1987 M02
         |  |--Mirandornithes KF-V21
         |  `--Palaelodidae [Palaeolontidae] M09
         |       |--Adelalopus hoogbutseliensis Mayr & Smith 2002 M09
         |       |--Palaelodus Milne-Edwards 1863 BV-R98 [=Paloelodus (l. c.) M02; incl. Probalearica Lambrecht 1933 M02]
         |       |    |--*P. ambiguus Milne-Edwards 1869 BV-R98 (see below for synonymy)
         |       |    |--P. crassipes Milne-Edwards 1863 BV-R98, M02
         |       |    |--P. gracilipes Milne-Edwards 1863 BV-R98, M02 (see below for synonymy)
         |       |    |--P. pledgei Baird & Vickers-Rich 1998 BV-R98
         |       |    `--P. wilsoni Baird & Vickers-Rich 1998 BV-R98
         |       `--Megapaloelodus Miller 1944 M02
         |            |--*M. connectens Miller 1944 M02
         |            |--M. goliath (Milne-Edwards 1868) BV-R98, M02 (see below for synonymy)
         |            `--M. opsigonus Brodkorb 1961 U93
         `--+--+--+--Gruiformes KF-V21
            |  |  `--Charadriiformes KF-V15
            |  `--+--+--Caprimulgiformes KF-V15
            |     |  `--Opisthocomidae [Opisthocomi, Opisthocomiformes] KF-V15
            |     |       |--Hoazinoides magdalenae Miller 1953 U93
            |     |       |--Hoazinavis lacustris JM14
            |     |       `--Opisthocomus Illiger 1811 PB15, B94
            |     |            `--O. hoazin (Müller 1776) ME04
            |     `--Columbaves KF-V15
            `--+--Telluraves KF-V15
               `--+--Phaethontimorphae JM14
                  |    |--Phaethontiformes KF-V15
                  |    `--Eurypygiformes [Eurypygae] JM14
                  |         |--Rhynochetos Verreaux & des Murs 1860 (see below for synonymy) L98
                  |         |    |--R. jubatus L98
                  |         |    `--R. orarius Balouet & Olson 1989 U93
                  |         `--Eurypyga Illiger 1811 [Eurypygidae, Eurypygides] L98
                  |              `--E. helias JM14
                  |                   |--E. h. helias RN72
                  |                   |--E. h. major RN72
                  |                   `--E. h. meridionalis RN72
                  `--Aequornithia KF-V21

Inorganic: Opisthocomidus opisthocomidus minilorientalus Okamura 1987 O87

Megapaloelodus goliath (Milne-Edwards 1868) BV-R98, M02 [=Palaelodus goliath M02, Paloelodus (l. c.) goliath M02]

Neoaves [Aequorlitornithes, Anomalogonates, Charadriimorphae, Ciconiae, Ciconiiformes, Ciconiimorphae, Ciconioidea, Coccyges, Columbea, Coronaves, Cuculimorphae, Cursorimorphae, Dendrornithes, Eurypygoidea, Gaviae, Gavii, Gaviomorphae, Grallae, Impennes, Incessores, Inopinaves, Longipennes, Metaves, Oxypterae, Paludicolae, Passerea, Passeres, Pelecani, Picae, Plethornithes, Psittacimorphae, Pygopodes, Pygopodotubinares, Ralliformes, Simplicirostres, Terrestrornithes, Turniciformes] HK08

*Palaelodus ambiguus Milne-Edwards 1869 BV-R98 [incl. Grus miocenicus Grigorescu & Kessler 1977 M02, G. problematica Milne-Edwards 1871 M02, *Probalearica problematica M02]

Palaelodus gracilipes Milne-Edwards 1863 BV-R98, M02 [incl. incl. Paloelodus (l. c.) minutus Milne-Edwards 1868 BV-R98, M02]

Rhynochetos Verreaux & des Murs 1860 [Rhynocheti, Rhynochetidae, Rhynochetides, Rhynochetoidea] L98

*Type species of generic name indicated

REFERENCES

[BV-R98] Baird, R. F., & P. Vickers-Rich. 1998. Palaelodus (Aves: Palaelodidae) from the Middle to Late Cainozoic of Australia. Alcheringa 22 (2): 135–151.

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

[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.

[JM14] Jarvis, E. D., S. Mirarab, A. J. Aberer, B. Li, P. Houde, C. Li, S. Y. W. Ho, B. C. Faircloth, B. Nabholz, J. T. Howard, A. Suh, C. C. Weber, R. R. de Fonseca, J. Li, F. Zhang, H. Li, L. Zhou, N. Narula, L. Liu, G. Ganapathy, B. Boussau, M. S. Bayzid, V. Zavidovych, S. Subramanian, T. Gabaldón, S. Capella-Gutiérrez, J. Huerta-Cepas, B. Rekepalli, K. Munch, M. Schierup, B. Lindow, W. C. Warren, D. Ray, R. E. Green, M. W. Bruford, X. Zhan, A. Dixon, S. Li, N. Li, Y. Huang, E. P. Derryberry, M. F. Bertelsen, F. H. Sheldon, R. T. Brumfield, C. V. Mello, P. V. Lovell, M. Wirthlin, M. P. Cruz Schneider, F. Prosdocimi, J. A. Samaniego, A. M. Vargas Velazquez, A. Alfaro-Núñez, P. F. Campos, B. Petersen, T. Sitcheritz-Ponten, A. Pas, T. Bailey, P. Scofield, M. Bunce, D. M. Lambert, Q. Zhou, P. Perelman, A. C. Driskell, B. Shapiro, Z. Xiong, Y. Zeng, S. Liu, Z. Li, B. Liu, K. Wu, J. Xiao, X. Yinqi, Q. Zheng, Y. Zhang, H. Yang, J. Wang, L. Smeds, F. E. Rheindt, M. Braun, J. Fjeldså, L. Orlando, F. K. Barker, K. A. Jønsson, W. Johnson, K.-P. Koepfli, S. O'Brien, D. Haussler, O. A. Ryder, C. Rahbek, E. Willerslev, G. R. Graves, T. C. Glenn, J. McCormack, D. Burt, H. Ellegren, P. Alström, S. V. Edwards, A. Stamatakis, D. P. Mindell, J. Cracraft, E. L. Braun, T. Warnow, W. Jun, M. T. P. Gilbert & G. Zhang. 2014. Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346 (6215): 1320–1331.

[KF-V21] Kuhl, H., C. Frankl-Vilches, A. Bakker, G. Mayr, G. Nikolaus, S. T. Boerno, S. Klages, B. Timmermann & M. Gahr. 2021. An unbiased molecular approach using 3′-UTRs resolves the avian family-level tree of life. Molecular Biology and Evolution 38 (1): 108–127.

[L98] Livezey, B. C. 1998. A phylogenetic analysis of the Gruiformes (Aves) based on morphological characters, with an emphasis on the rails (Rallidae). Philosophical Transactions of the Royal Society of London Series B—Biological Sciences 353: 2077–2151.

[M05] Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515–542.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[M11] Mayr, G. 2011. Cenozoic mystery birds—on the phylogenetic affinities of the bony-toothed birds (Pelagornithidae). Zoologica Scripta 40: 448–467.

[ME04] Mayr, G., & P. G. P. Ericson. 2004. Evidence for a sister group relationship between the Madagascan mesites (Mesitornithidae) and the cuckoos (Cuculidae). Senckenbergiana Biologica 84 (1–2): 1–17.

[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.

[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.

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

[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.

Last updated: 29 January 2022.

Chalcophaps

Female (left) and male common emerald doves Chalcophaps indica, copyright Sham Edmond.


Belongs within: Treroninae.

Chalcophaps, the emerald doves, is a genus of stocky doves with brownish neck and underparts and bright green wings, found between southern and eastern Asia and northern and eastern Australia.

<==Chalcophaps Gould 1843 [incl. Monornis Hodgson in Gray 1844; Chalcophabinae] CC10
    |--C. indica (Linnaeus 1758) CC10 (see below for synonymy)
    |    |--C. i. indica CC10
    |    |--C. i. longirostris Gould 1848 CC10
    |    |--C. i. natalis Lister 1888 CC10
    |    |--C. i. rogersi Mathews 1912 [=C. chrysochlora rogersi; incl. C. chrysochlora kempi Mathews 1912] CC10
    |    |--‘Chalcospilos’ i. timorensis T62
    |    `--C. i. yamashinai Hachisuka 1939 I92
    `--C. stephani J06
         |--C. s. stephani J06
         `--C. s. wallacei J06

Chalcophaps indica (Linnaeus 1758) CC10 [=Columba indica CC10, Chalcospilos indica T62; incl. Columba chrysochlora Wagler 1827 CC10, *Chalcophaps indica chrysochlora CC10, *Monornis perpulchra CC10]

*Type species of generic name indicated

REFERENCES

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

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[J06] Johnstone, R. E. 2006. The birds of Gag Island, Western Papuan islands, Indonesia. Records of the Western Australian Museum 23 (2): 115–132.

[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.

Gallicolumba

Luzon bleeding heart Gallicolumba luzonica, photographed by Kao-Tai.


Belongs within: Treroninae.

Gallicolumba, the bleeding hearts, is a genus of ground-dwelling doves found in insular south-east Asia and New Guinea, a number of species of which have a prominent red patch on the breast that inspires the vernacular name.

<==Gallicolumba Heck 1849 (see below for synonymy) CC10
    |--*G. luzonica CC10, JT12 [=Columba luzonica CC10, *Plegoenas luzonica CC10]
    |--G. canifrons JT12
    |--G. crinigera JT12 [=Peristera criniger CC10, *Pampusanna criniger CC10]
    |--G. cruenta CC10
    |--G. erythroptera JT12
    |    |--G. e. erythroptera HSS13
    |    `--G. e. pectoralis HSS13
    |--G. ferruginea HSS13
    |--G. hoedtii JT12
    |--G. keayi JT12
    |--G. kubaryi JT12
    |--G. menagei JT12
    |--G. norfolciensis (Latham 1802) [=Columba norfolciensis, Carpophaga norfolciensis] CC10
    |--G. platenae JT12
    |--G. rubescens JT12
    |--G. rufigula JT12
    |--G. salamonis HSS13
    |--G. sanctaecrucis JT12
    |--G. stairi JT12
    |--G. tristigmata JT12
    `--G. xanthonura JT12

Gallicolumba Heck 1849 [=Plegoenas Reichenbach 1851; incl. Pampusanna Pucheran in Dumont d’Urville 1853; Gallicolumbinae] CC10

*Type species of generic name indicated

REFERENCES

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

[HSS13] Hirschfeld, E., A. Swash & R. Still. 2013. The World's Rarest Birds. Princeton University Press: Princeton (New Jersey).

[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.

Strisores

Little nightjar Caprimulgus parvulus, copyright Jairmoreirafotografia.


Belongs within: Neognathae.
Contains: Caprimulgus, Chordeiles, Nyctibiidae, Podargidae, Daedalornithes.

The Strisores is a clade of birds including the members of the commonly recognised orders 'Caprimulgiformes' and Apodiformes with the former now recognised as paraphyletic to the latter. Members of the clade are united by possession of a greatly elongated leg (crus longum) of the ulnar carpal bone (Mayr 2009). All members of the traditional Caprimulgiformes are crepuscular or nocturnal birds with wide beaks, a large gape, and soft cryptically-coloured plumage, that feed on the wing (most are insectivores; the oilbird Steatornis caripensis is frugivorous); Apodiformes have lost some of these characters, but still feed on the wing. Morphological support for the total group of Caprimulgiformes (including Apodiformes) is, however, uncertain.

Within the Strisores, the Caprimulgidae include the nightjars, a cosmopolitan group of nocturnal insectivorous birds. Nightjars are cryptically coloured in mottled brown. The family has historically been divided between subfamilies Caprimulginae and Chordeilinae based on the presence of strong rictal bristles around the mouth of the former, but this division has not been supported by recent phylogenetic analyses. Instead, the eared nightjars of the genus Eurostopodus represent the most basal members of the Caprimulgidae; these large Australasian nightjars lack rictal bristles and possess more or less developed ear tufts.

The Steatornithidae are represented in the modern fauna by a single species, the oilbird Steatornis caripensis of northern South America. This large bird, reddish-brown with white spots on the nape and wings, nest colonially in caves, using echolocation to find their way in the dark. The Fluvioviridavidae are wide-billed, fossil birds known from the Eocene of Europe and North America. The Archaeotrogonidae of the Oligocene of Europe were originally described as relatives of the trogons but later regarded as more similar to caprimulgids.

<==Strisores [Caprimulgi, Caprimulgimorphae, Cypselomorphae, Podarginae] PB15
    |--Caprimulgidae [Aereae, Caprimulgiformes, Caprimulginae, Glabrirostres, Setirostres, Terricolae] PB15
    |    |  i. s.: Lyncornis temminckii S55
    |    |         Nannochordeiles pusillus S55
    |    |           |--N. p. pusillus S55
    |    |           `--N. p. septentrionalis S55
    |    |         Semeiophorus Gould 1838 B94
    |    |           `--S. vexillarius MMJ03
    |    |         Scotornis Swainson 1837 [Scotornithinae] B94
    |    |           `--S. climacurus S55
    |    |                |--S. c. climacurus S55
    |    |                `--S. c. clarus S55
    |    |         Stenopsis S66a
    |    |           |--S. bifasciata S66b (see below for synonymy)
    |    |           |--S. cayanensis S66b [=Caprimulgus cayanensis S66a; incl. C. cayanus S66a, C. leopetes S66a]
    |    |           |    |--S. c. cayanensis S18
    |    |           |    `--S. c. albicauda S18
    |    |           |--S. langsdorfi Pelzeln in Sclater 1866 S66b
    |    |           |--S. platura Pelzeln in Sclater 1866 [=Caprimulgus platurus] S66b
    |    |           `--S. ruficervix Sclater 1866 S66a
    |    |         Macropsalis Sclater 1866 S66a
    |    |           |--M. forcipata JT12 (see below for synonymy)
    |    |           |--M. lyra [=Hydropsalis (Macropsalis) lyra] S66a
    |    |           |--M. segmentata Cassin 1849 [=Hydropsalis (Macropsalis) segmentata] S66a
    |    |           `--M. ypanemae von Pelzeln 1865 [=Hydropsalis (Macropsalis) ypanemae] S66a
    |    |         Antrostomus S66a
    |    |           |--A. cubanensis Lawrence 1860 S66b
    |    |           |--A. macromystax [=Caprimulgus macromystax] S66a
    |    |           |--A. maculicaudus [=Stenopsis maculicaudus] S66b
    |    |           |--A. nuttalli [=Caprimulgus nuttalli] S66a
    |    |           |--A. ocellatus (see below for synonymy) S66a
    |    |           |--A. ornatus Sclater 1866 S66b
    |    |           |--A. rutilus S66b
    |    |           `--A. sericeocaudatus S66b
    |    |         Ventivorus Mourer-Chauviré 1988 M09, M02
    |    |           `--*V. ragei Mourer-Chauviré 1988 M02
    |    |--‘Eurostopodus’ macrotis BKB15
    |    `--+--Eurostopodus Gould 1838 BKB15, B94 [Eurostopodidae]
    |       |    |  i. s.: E. albogularis R87
    |       |    |         E. archboldi JT12
    |       |    |         E. diabolicus FP64
    |       |    |         E. guttatus (Vigors & Horsfield 1827) [=Caprimulgus guttatus] WS48
    |       |    |         E. temminckii H03
    |       |    |--E. papuensis JT12
    |       |    `--+--E. argus JT12
    |       |       `--E. mystacalis JT12
    |       `--+--‘Caprimulgus’ enarratus BKB15
    |          `--+--+--+--Caprimulgus JT12
    |             |  |  `--Chordeiles BKB15
    |             |  `--+--Siphonorhis BKB15
    |             |     |    |--S. americana (Linnaeus 1758) JT12, L58 [=Caprimulgus americanus S66a]
    |             |     |    `--S. brewsteri JT12 [=S. americana brewsteri FP64]
    |             |     `--+--Nyctiphrynus JT12
    |             |        |    |--N. rosenbergi JT12
    |             |        |    `--+--N. mcleodii JT12
    |             |        |       `--+--N. ocellatus JT12
    |             |        |          `--N. yucatanicus JT12
    |             |        `--+--Phalaenoptilus nuttallii JT12
    |             |           `--+--+--‘Caprimulgus’ saturatus BKB15
    |             |              |  `--‘Caprimulgus’ vociferus JT12 [=Antrostomus vociferus S66a]
    |             |              `--+--‘Caprimulgus’ ridgwayi BKB15
    |             |                 `--+--‘Caprimulgus’ salvini BKB15
    |             |                    `--+--‘Caprimulgus’ carolinensis JT12 [=Antrostomus carolinensis S66a]
    |             |                       `--‘Caprimulgus’ rufus JT12 [=Antrostomus rufus S66a; incl. Ca. cortopao S66a]
    |             `--+--‘Caprimulgus’ sericocaudatus BKB15
    |                `--+--Lurocalis BKB15
    |                   |    |--L. rufiventris BKB15
    |                   |    `--+--L. nattereri BKB15 [=Caprimulgus nattereri S66a, Chordeiles nattereri S66a]
    |                   |       `--L. semitorquatus BKB15 (see below for synonymy)
    |                   `--+--Nyctiprogne JT12
    |                      |    |--N. leucopyga JT12
    |                      |    `--N. vielliardi JT12
    |                      `--+--+--‘Caprimulgus’ nigrescens JT12 (see below for synonymy)
    |                         |  `--+--‘Caprimulgus’ anthonyi JT12
    |                         |     `--Nyctidromus Gould 1838 JT12, B94 [Nyctidrominae]
    |                         |          `--N. albicollis (see below for synonymy) S66a
    |                         |               |--N. a. albicollis FS55
    |                         |               `--N. a. intercedens FS55
    |                         `--+--Uropsalis JT12
    |                            |    |--U. lyra JT12
    |                            |    `--U. segmentata JT12
    |                            `--+--+--‘Caprimulgus’ longirostris BKB15 (see below for synonymy)
    |                               |  `--+--‘Caprimulgus’ parvulus Gould 1837 BKB15, S66a (see below for synonymy)
    |                               |     `--‘Caprimulgus’ whitelyi BKB15
    |                               `--+--Eleothreptus BKB15
    |                                  |    |--E. anomalus (Gould 1837) JT12, S66a (see below for synonymy)
    |                                  |    `--E. candicans JT12 (see below for synonymy)
    |                                  `--+--‘Caprimulgus’ maculicaudus BKB15
    |                                     `--+--‘Caprimulgus’ cayennensis JT12
    |                                        `--Hydropsalis JT12
    |                                             |  i. s.: H. (Diplopsalis) S66a
    |                                             |           |--H. (D.) schomburgki S66a
    |                                             |           `--H. (D.) trifurcata [incl. H. climacocerca] S66a
    |                                             |--H. climacocerca BKB15
    |                                             `--+--H. brasiliana BKB15
    |                                                `--H. (Hydropsalis) S66a
    |                                                     |--H. (H.) furcifera [=Caprimulgus furcifer] S66a
    |                                                     |--H. (H.) pallescens von Pelzeln 1865 S66a
    |                                                     `--H. (H.) torquata BKB15, S66a (see below for synonymy)
    `--+--+--Nyctibiidae BKB15
       |  `--Steatornithidae [Steatornithes, Steatornithiformes, Steatornithinae] BKB15
       |       |--Prefica Olson 1987 M03, M02 [Preficinae]
       |       |    `--P. nivea Olson 1987 M03
       |       `--Steatornis Humboldt 1814 PB15, B94
       |            `--S. caripensis [incl. Caprimulgus steatornis] S66a
       `--+--+--Podargidae PB15
          |  `--Fluvioviridavidae M05b
          |       |--Fluvioviridavis Mayr & Daniels 2001 PB15, M05b
          |       |    `--F. platyrhamphus Mayr & Daniels 2001 M05b
          |       `--Eurofluvioviridavis Mayr 2005 M05b
          |            `--*E. robustipes Mayr 2005 M05b
          `--Daedalornithes PB15

Strisores incertae sedis:
  Archaeotrogonidae M04
    |--Hassiavis Mayr 1998 M04
    |    `--*H. laticauda Mayr 1998 M02
    `--Archaeotrogon Milne-Edwards 1892 M02
         |--*A. venustus Milne-Edwards 1892 M02, M09
         |--A. cayluxensis (Lydekker 1891) [=Palaeortyx cayluxensis, Ludiortyx cayluxensis] M02
         |--A. hoffstetteri Mourer-Chauviré 1980 M09
         |--A. nocturnus Mlíkovský 2002 (see below for synonymy) M02
         `--A. zitteli Gaillard 1908 M09
  Microena Harrison & Walker 1977 M05a
    `--*M. goodwini Harrison & Walker 1977 M02
  Protocypselomorphus manfredkelleri Mayr 2005 M09

Antrostomus ocellatus [=Caprimulgus ocellatus; incl. C. brasilianus, C. brasiliensis, C. lunulatus Natt. in Sclater 1866 (n. n.)] S66a

Archaeotrogon nocturnus Mlíkovský 2002 [=A. cayluxensis Gaillard 1908 non Palaeortyx cayluxensis Lydekker 1891] M02

‘Caprimulgus’ longirostris BKB15 [=Antrostomus longirostris S66a, Stenopsis longirostris S66a]

‘Caprimulgus’ nigrescens JT12 [=Antrostomus nigrescens S66a, Stenopsis nigrescens S66a]

‘Caprimulgus’ parvulus Gould 1837 BKB15, S66a [=Antrostomus parvulus S66a; incl. C. mixtus S66a]

Eleothreptus anomalus (Gould 1837) JT12, S66a [=Amblypterus anomalus S66a, Heleothreptus anomalus S66a]

Eleothreptus candicans JT12 [=Caprimulgus candicans BKB15, Stenopsis candicans S66b; incl. C. leucurus S66b]

Hydropsalis (Hydropsalis) torquata BKB15, S66a [=Caprimulgus torquatus S66a; incl. H. azarae Wagl. 1883 S66a, Psalurus macropterus S66a, Caprimulgus psalurus S66a, Hydropsalis psalurus S66a]

Lurocalis semitorquatus BKB15 [=Caprimulgus semitorquatus S66a, Chordeiles semitorquatus S66a; incl. Podager gouldi S66a]

Macropsalis forcipata JT12 [=Caprimulgus forcipatus S66a, Hydropsalis (Macropsalis) forcipata S66a; incl. H. limbatus S66a, Caprimulgus megalurus Licht. in Sclater 1866 (n. n.) S66a]

Nyctidromus albicollis [=Caprimulgus albicollis; incl. N. affinis, N. americanus Cassin 1851, N. derbyanus, N. grallarius, Caprimulgus guianensis, N. guianensis] S66a

Stenopsis bifasciata S66b [=Caprimulgus bifasciatus S66a; incl. C. conterminus S66a, C. decussatus S66a]

*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.

[FS55] Felten, H., & J. Steinbacher. 1955. Zur Vogelfauna von El Salvador. Senckenbergiana Biologica 36 (1–2): 9–19.

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

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

[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.

[M03] Mayr, G. 2003. On the phylogenetic relationships of trogons (Aves, Trogonidae). Journal of Avian Biology 34 (1): 81–88.

[M04] Mayr, G. 2004. New specimens of Hassiavis laticauda (Aves: Cypselomorphae) and Quasisyndactylus longibrachis (Aves: Alcediniformes) from the Middle Eocene of Messel, Germany. Courier Forschungsinstitut Senckenberg 252: 23–28.

[M05a] Mayr, G. 2005a. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515–542.

[M05b] Mayr, G. 2005b. A Fluvioviridavis-like bird from the Middle Eocene of Messel, Germany. Canadian Journal of Earth Sciences 42: 2021–2037.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[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.

[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.

[R87] Ramsay, E. P. 1887. Descriptions of Australian birds' eggs. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1141–1152, pl. 19.

[S66a] Sclater, P. L. 1866a. Notes upon the American Caprimulgidae. Proceedings of the Zoological Society of London 1866: 123–145.

[S66b] Sclater, P. L. 1866b. Additional notes on the Caprimulgidae. Proceedings of the Zoological Society of London 1866: 581–590.

[S55] Steinbacher, J. 1955. Über die Schwanzmauser der Eulen (Strigidae) und Nachtschwalben (Caprimulgidae). Senckenbergiana Biologica 36: 235–240.

[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.

[WS48] Whittell, H. M., & D. L. Serventy. 1948. A systematic list of the birds of Western Australia. Public Library, Museum and Art Gallery of Western Australia, Special Publication 1: 1–126.