Abies

Nikko fir Abies homolepis, copyright MPF.

Belongs within: Pinaceae.

Abies, the firs, is a genus of conifers with erect seed cones native to the Northern Hemisphere. Leaf scars are smooth, round and flush with the surface of the stem (Hickman 1993).

Characters (from Hickman 1993): Young bark smooth, with resin blisters, mature bark thick, deeply furrowed; young branches appearing whorled; twig glabrous or hairy; leaf scars smooth, round, flush with surface; bud generally more or less spheric, generally <1 cm, more or less resinous. Leaves 2–9 cm, sessile, twisted at base to become two-ranked, often curved upward on upper twigs, generally more or less flat; upper surface with two longitudinal, whitish bands, midrib sometimes depressed; lower surface with or without whitish bands, midrib sometimes ridge-like. Seed cone erect, <23 cm, maturing first season; stalk generally absent; bracts, scales deciduous; bract included in scale or exserted, free from scale; axis persistent on stem. Seed with obvious resin deposits on surface; wing <2.5 cm. 2n=24 (where known).

<==Abies
    |--A. alba Miller 1768 SM14
    |--A. amabilis W85
    |--A. balsamea HP03
    |--A. bituminosa P92
    |--A. bracteata [incl. A. venusta] H93
    |--A. cephalonica RCT96
    |--A. concolor [incl. A. concolora var. lowiana] H93
    |--A. excelsa C55
    |--A. fabri MM12
    |--A. firma MM12
    |--A. grandis C84
    |--A. holophylla MM12
    |--A. homolepis MM12
    |--A. lasiocarpa H93
    |--A. magnifica F56
    |    |--A. m. var. magnifica H93
    |    `--A. m. var. shastensis H93
    |--A. mariesii T85
    |--A. nebrodensis R-CT01
    |--A. nephrolepis S00
    |--A. numidica EH06
    |--A. pectinata [incl. Pinus picea] C55
    |--A. pindrow N02
    |--A. procera M87
    |--A. religiosa C-MM-R07
    |--A. sachalinensis T03
    `--A. veitchii MM12
         |--A. v. var. veitchii A70
         `--A. v. var. shikokiana A70

*Type species of generic name indicated

REFERENCES

[A70] Aoki, J. 1970. Descriptions of oribatid mites collected by smoking of trees with insecticides. I. Mt. Ishizuchi and Mt. Odaigahara. Bulletin of the National Science Museum, Tokyo 13 (4): 585–602.

[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.

[C-MM-R07] Castaño-Meneses, G., & B. E. Mejía-Recamier. 2007. Community structure of mites in Tillandsia violacea (Bromeliaceae) from a temperate forest of central Mexico. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 657–664. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[C84] Cokendolpher, J. C. 1984. Revision of the harvestman genus Leptobunus and dismantlement of the Leptobunidae (Arachnida: Opiliones: Palpatores). Journal of the New York Entomological Society 92 (4): 371–402.

[EH06] Eckert, A. J., & B. D. Hall. 2006. Phylogeny, historical biogeography, and patterns of diversification for Pinus (Pinaceae): phylogenetic tests of fossil-based hypotheses. Molecular Phylogenetics and Evolution 40 (1): 166–182.

[F56] Foote, R. H. 1956. Gall midges associated with cones of western forest trees (Diptera: Itonididae). Journal of the Washington Academy of Sciences 46 (2): 48–57.

[HP03] Hauck, M., A. Paul, S. Gross & M. Raubuch. 2003. Manganese toxicity in epiphytic lichens: chlorophyll degradation and interaction with iron and phosphorus. Environmental and Experimental Botany 49: 181–191.

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

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793–7798.

[M87] McAlpine, J. F. 1987. Pallopteridae. In: McAlpine, J. F. (ed.) Manual of Nearctic Diptera vol. 2 pp. 839–843. Research Branch, Agriculture Canada.

[N02] Negi, H. R. 2002. Abundance and diversity of moss communities of Chopta-Tunganath in the Garhwal Himalaya. Journal of the Bombay Natural History Society 99 (3): 418–433.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[RCT96] Ragusa Di Chiara, S., & H. Tsolakis. 1996. A. survey of phytoseiid mites (Phytoseiidae) associated with various plants in Sicily (Italy). In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 253–256. Ohio Biological Survey: Columbus (Ohio).

[R-CT01] Ragusa-di Chiara, S., & H. Tsolakis. 2001. Phytoseiid faunas of natural and agricultural ecosystems in Sicily. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 522–529. CSIRO Publishing: Melbourne.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2^nd ed. CABI Publishing: Wallingford (UK).

[SM14] Smith, G. B., & J. M. McRae. 2014. New species of subterranean silverfish (Zygentoma: Nicoletiidae: Atelurinae) from Western Australia's semi-arid Pilbara region. Records of the Western Australian Museum 29: 105–127.

[T85] Tsurusaki, N. 1985. Geographic variation of chromosomes and external morphology in the montanum-subgroup of the Leiobunum curvipalpe-group (Arachnida, Opiliones, Phalangiidae) with special reference to its presumable process of raciation. Zoological Science 2: 767–783.

[T03] Tsurusaki, N. 2003. Phenology and biology of harvestmen in and near Sapporo, Hokkaido, Japan, with some taxonomical notes on Nelima suzukii n. sp. and allies (Arachnida: Opiliones). Acta Arachnologica 52: 5–24.

[W85] Walter, D. E. 1985. The effects of litter type and elevation on colonization of mixed coniferous litterbags by oribatid mites. Pedobiologia 28: 383–387.

Last updated: 18 July 2021.

Dicranophyllales

Reconstruction of Dicranophyllum hallei, from here.

Belongs within: Coniferophyta.

The Dicranophyllales are a group of arborescent plants known from the Carboniferous and Permian, possibly related to modern coniferophytes. They are defined by the having stomata confined to sunken furrows on the abaxial side of the leaf (Rothwell & Mapes 2001).

See also: The Dicranophyllales: an early branch of the conifers?

<==Dicranophyllales
    |--Slivkovia RM01
    |    |--S. lorata RM01
    |    `--S. petschorensis RM01
    |--Entosovia RM01
    |    |--E. kungurica RM01
    |    `--E. rarisulcata RM01
    |--Mostotchkia RM01
    |    |--M. gomankovii RM01
    |    `--M. longifolia RM01
    `--Dicranophyllum RM01
         |--D. effusum RM01
         |--D. gallicum RM01
         |--D. glabrum A38
         |--D. hallei RM01
         |--D. longifolium RM01
         `--D. striatum RM01

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1–65.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637–667.

Miroviaceae

Preserved leaf sections of Mirovia gothanii, from Gomez (2002).

Belongs within: Pinopsida.

The Miroviaceae is an extinct group of conifers known from the mid-Jurassic to the Late Cretaceous, with linear leaves similar in appearance to the modern umbrella pine Sciadopitys verticillata. They differ from modern conifers in having the closely placed stomata restricted to a sunken groove with a protruding margin on the abaxial side of the leaf (Gomez 2002).

<==Miroviaceae
    |--Tritaenia G02
    |--Holkopitys Bose & Manum 1990 G02
    |    `--H. hoegii G02
    |--Oswaldheeria Bose & Manum 1990 G02
    |    |--O. arctica G02
    |    |--O. hallei G02
    |    |--O. macrophylla G02
    |    `--O. scotica G02
    |--Sciadopityoides G02
    |    |--S. crameri G02
    |    |--S. greenboana G02
    |    |--S. ikorfatensis G02
    |    |--S. microphylla G02
    |    |--S. nathorstii G02
    |    |--S. ukrainensis G02
    |    |--S. uralensis G02
    |    `--S. variabilis G02
    `--Mirovia G02
         |--*M. szaferi Reymanówna 1985 G02
         |--M. capbohemanensis G02
         |--M. florinii G02
         |--M. gothanii Gomez 2002 G02
         |--M. groenlandica G02
         |--M. ineffecta G02
         |--M. lagerheimii G02
         |--M. persulcata G02
         `--M. sibirica G02

*Type species of generic name indicated

REFERENCES

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761–773.

Cheirolepidiaceae

Frenelopsis hoheneggeri, from the Peabody Museum of Natural History.

Belongs within: Pinopsida.

The Cheirolepidiaceae are a group of fossil conifers known from the Mesozoic. They are characterised by a distinct pollen type (assigned to the form genus Classopollis) with a complex, tectate-columellate wall structure (Axsmith et al. 2004). Shoots were often opposite-decussate or spiral, and leaves (which may be dimorphic) were typically reduced or scale-like, and imbricate (Pole 2000).

<==Cheirolepidiaceae [Frenelopsidaceae]
    |--Alvinia bohemica AKW04
    |--Tomaxellia biforme P00
    |--Otwayia Pole 2000 P00
    |    |--*O. tetragona (Cantrill & Douglas) Pole 2000 [=Geinitzea tetragona Cantrill & Douglas 1988] P00
    |    `--O. cudgeloides Pole 2000 P00
    |--Classostrobus Alvin, Spicer & Watson 1978 AKW04 [incl. Pseudofrenelopsis AKW04, Suturovagina GM-C02]
    |    |--*C. comptonensis Alvin, Spicer & Watson 1994 AKW04 (see below for synonymy)
    |    |--C. arkansensis Axsmith, Krings & Waselkov 2004 AKW04
    |    |--C. cathayanus [incl. Pseudofrenelopsis intermedia] AKW04
    |    |--C. turolensis Gomez, Martín-Closas et al. 2002 GM-C02
    |    `--C. ugnaensis Gomez, Martín-Closas et al. 2002 GM-C02
    |--Classopollis Pflug 1953 AKW04
    |    |--*C. classoides Pflug 1953 GM-C02
    |    |--C. brasiliensis Herngreen 1975 I02
    |    |--C. mirabilis Reyre 1970 GM-C02
    |    |--C. noeli Reyre 1970 GM-C02
    |    `--C. obidosensis Groot & Groot 1962 GM-C02
    |--Cupressinocladus Seward 1919 AKW04 [incl. Watsoniocladus GM-C02]
    |    |--C. asferensis Barale & Ouaja 2002 BO02
    |    |--C. itieri (Saporta) Barale 1981 BO02
    |    |--C. leptocladoides (Berry) Pons 1990 BO02
    |    |--C. micromerum (Heer) Pais 1974 BO02
    |    |--C. pompeckji (Salfeld) Pons 1990 BO02
    |    |--C. pseudoexpansum Barnard & Miller 1976 BO02
    |    `--C. soltanensis Barale, Ouaja & Philippe 2000 B)2
    `--Frenelopsis Schenck 1869 AKW04, GM-C02
         |--*F. hoheneggeri (Ettingshausen) Schenck 1869 [=Thuites hoheneggeri Ettinghausen 1852] GM-C02
         |--F. alata Feistmantel 1881 AKW04
         |--F. choshiensis GM-C02
         |--F. elegans GM-C02
         |--F. harrisii Doludenko & Reymanówna 1978 GT02
         |--F. kaneviensis Barale & Doludenko 1985 GM-C02
         |--F. occidentalis GM-C02
         |--F. oligostomata AKW04 [incl. Classoidites glandis GM-C02]
         |--F. pombetsuensis Saiki 1997 GM-C02
         |--F. ramosissima GM-C02
         |--F. rubiesensis Barale 1973 GM-C02
         |--F. sifloana Watson 1983 GM-C02
         |--F. teixeirae GM-C02
         |--F. turolensis Gomez, Martín-Closas et al. 2002 GM-C02
         |--F. ugnaensis Gomez, Martín-Closas et al. 2002 GM-C02
         `--F. veneta Gomez, Thévenard et al. 2002 GT02

*Classostrobus comptonensis Alvin, Spicer & Watson 1994 AKW04 [incl. Classopollis martinottii GM-C02, Pseudofrenelopsis parceramosa (Fontaine) Watson 1977 AKW04]

*Type species of generic name indicated

REFERENCES

[AKW04] Axsmith, B. J., M. Krings & K. Waselkov. 2004. Conifer pollen cones from the Cretaceous of Arkansas: implications for diversity and reproduction in the Cheirolepidiaceae. Journal of Paleontology 78: 402–409.

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707–737.

[GM-C02] Gomez, B., C. Martín-Closas, G. Barale, N. Solé de Porta, F. Thévenard & G. Guignard. 2002. Frenelopsis (Coniferales: Cheirolepidiaceae) and related male organ genera from the Lower Cretaceous of Spain. Palaeontology 45 (5): 997–1036.

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671–685.

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775–788.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153–197.

Coniferophyta

Maidenhair tree Ginkgo biloba, copyright Liné1.

Belongs within: Spermatophyta.
Contains: Dicranophyllales, Pinopsida.

The Coniferophyta has been recognised as a clade of gymnosperms including conifers, the ginkgo Ginkgo biloba, and the Palaeozoic Cordaitales. Members of this clade have simple, dichotomously veined leaves, simple strobili in the axils of the leaves, and platyspermic seeds (Doyle 1998).

<==Coniferophyta
    |  i. s.: Carpentieria frondosa RM01
    |         Dichophyllum RM01
    |         Ginkgophyllum RM01
    |         Polyspermophyllum sergii RM01
    |         Sphenobaiera RM01
    |         Vojnovskyales RM01
    |           |--Sergeia neuburgii RM01
    |           `--Vojnovskya [Vojnovskyaceae] RM01
    |                `--V. paradoxa RM01
    |--Ginkgoopsida [Ginkgophyta] RY03
    |    |  i. s.: Ginkgoites RY03
    |    |         Phoenicopsis RY03
    |    |         Trichopitys heteromorpha RM01
    |    |         Karkenia D98
    |    |         Baiera D98
    |    `--Ginkgoales FC07
    |         |--Nehvizdya penalveri G02
    |         |--Ginkgo [Ginconinae, Ginkgoideae] CD07
    |         |    |--G. baiera GT12
    |         |    |--G. biloba Linnaeus 1771 CD07
    |         |    |--G. huttoni PL12
    |         |    `--G. liaoningensis PL12
    |         |--Eretmophyllum PL12
    |         |--Pseudotorellia PL12
    |         `--Nehvizdyella bipartita PL12
    `--Pan-Coniferae CD07
         |--+--Dicranophyllales RMH-C05
         |  `--Pinopsida FC07
         `--Cordaitales [Cordaitanthales] FC07
              |  i. s.: Cordaites F71
              |           |--C. australis F71
              |           |--C. borassifolius [=Flabellaria borassifolia] F04
              |           `--C. robbii [incl. C. borassifolius] A38
              |         Cordaitanthus devonicus A38
              |--Shanxioxylon FC07
              `--+--Mesoxylon priapii FC07, RMH-C05
                 `--Cordaixylon FC07
                      |--C. dumusum RMH-C05
                      `--C. iowensis MM12

*Type species of generic name indicated

REFERENCES

[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1-65.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[D98] Doyle, J. A. 1998. Phylogeny of vascular plants. Annual Review of Ecology and Systematics 29: 567-599.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549-552.

[F04] Fritsch, A. 1904. Palaeozoische Arachniden. Selestverlag: Prague.

[GT12] Giribet, G., A. L. Tourinho, C. Shih & D. Ren. 2012. An exquisitely preserved harvestman (Arthropoda, Arachnida, Opiliones) from the Middle Jurassic of China. Organisms, Diversity & Evolution 12: 51-66.

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761-773.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793-7798.

[PL12] Peñalver, E., C. C. Labandeira, E. Barrón, X. Delclòs, P. Nel, A. Nel, P. Tafforeau & C. Soriano. 2012. Thrips pollination of Mesozoic gymnosperms. Proceedings of the National Academy of Sciences of the USA 109 (22): 8623-8628.

[RY03] Ren, D., & J. Yin. 2003. New ‘osmylid-like’ fossil Neuroptera from the Middle Jurassic of Inner Mongolia, China. Journal of the New York Entomological Society 111 (1): 1-11.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637-667.

[RMH-C05] Rothwell, G. W., G. Mapes & G. R. Hernandez-Castillo. 2005. Hanskerpia gen. nov. and phylogenetic relationships among the most ancient conifers (Voltziales). Taxon 54 (3): 733-750.

Pinaceae

Norway spruce Picea abies, photographed by MPF.

Belongs within: Coniferophyta.
Contains: Abies, Pinus subg. Pinus, Pinus subg. Strobus.

The Pinaceae include the pines (Pinus), spruces (Picea), cedars (Cedrus) and related taxa. This is a primarily Northern Hemisphere family, with some species being the dominant tree species within their range. Pinus species differ from other members of the family in having cones that are biennial or triennial, while other genera grow new cones each year. Members of the genus Picea, the spruces, have whorled branches with needles attached singly in a spiral pattern. Larix, larches, are deciduous, dropping their needles over winter.

Characters (from J. W. Thieret): Trees (occasionally shrubs), evergreen (annually deciduous in Larix), resinous and aromatic, monoecious. Bark smooth to scaly or furrowed. Lateral branches well developed and similar to leading (long) shoots or reduced to well-defined short (spur) shoots (Pinus, Larix); twigs terete, sometimes clothed by persistent primary leaves or leaf bases; longest internodes less than 1cm; buds conspicuous. Roots fibrous to woody, unspecialized. Leaves (needles) simple, shed singly (except whole fascicles shed in Pinus), alternate and spirally arranged but sometimes proximally twisted so as to appear 1- or 2-ranked, or fascicled, linear to needlelike, sessile to short-petiolate; foliage leaves either borne singly (spirally) on long shoots or in tufts (fascicles) on short shoots; juvenile leaves (when present) borne on long shoots, scalelike; resin canals present. Pollen cones maturing and shed annually, solitary or clustered, axillary, ovoid to ellipsoid or cylindric; sporophylls overlapping, bearing 2 abaxial microsporangia (pollen sacs); pollen spheric, 2-winged, less commonly with wings reduced to frill (in Tsuga sect. Tsuga), or not winged (in Larix and Pseudotsuga). Seed cones maturing and shed in 1-3 seasons or long-persistent, sometimes serotinous (not opening upon maturity but much later: Pinus), compound, axillary, solitary or grouped; scales overlapping, free from subtending included or exserted bracts for most of length, spirally arranged, strongly flattened, at maturity relatively thin to strongly thickened and woody (in Pinus), with 2 inverted, adaxial ovules. Seeds 2 per scale, elongate terminal wing partially decurrent on seed body (wing short or absent in some species of Pinus); aril lacking; cotyledons 2-12[-18].

Pinaceae [Abietaceae, Abietinae, Pinales]
    |  i. s.: Pinites succinifer Göppert 1836 E01 [=Pinus succinifera P02]
    |         Nothotsuga CD07
    |         Compsostrobus neotericus MM12
    |         Schizolepidopsis HS17
    |--Cedrus EH06
    |    |--C. atlantica C55
    |    |--C. brevifolia D30
    |    |--C. deodara EH06
    |    `--C. libani C55
    `--+--+--+--Abies EH06
       |  |  `--Keteleeria EH06
       |  |       |--K. davidiana EH06
       |  |       `--K. evelyniana MM12
       |  `--+--Pseudolarix amabilis EH06
       |     `--Tsuga EH06
       |          |--T. canadensis SS96
       |          |--T. chinensis CP02
       |          |--T. dumosa SN88
       |          |--T. heterophylla C90
       |          |--T. mertensiana EH06
       |          `--T. sieboldii [incl. Abies tsuga] LO98
       `--+--+--Pseudotsuga EH06
          |  |    |--P. douglasii B88
          |  |    |--P. macrocarpa B14
          |  |    |--P. menziesii EH06
          |  |    `--P. taxifolia SW03
          |  `--Larix EH06
          |       |--L. decidua EH06
          |       |--L. gmelinii L02
          |       |    |--L. g. var. gmelinii F92
          |       |    `--L. g. var. japanica F92
          |       |--L. groenlandii TH03
          |       |--L. himalaica SN88
          |       |--L. kaemferi F92
          |       |--L. koreana F92
          |       |--L. laricina TH03
          |       |--L. leptolepis F92
          |       |--L. occidentalis MM12
          |       |--L. olbensis F92
          |       |--L. sibirica F92
          |       `--L. subczewii F92
          `--+--+--Cathaya argyrophylla EH06
             |  `--Picea EH06
             |       |--P. abies EH06
             |       |--P. alcoquiana [incl. Abies menziessi] LO98
             |       |--P. breweriana H93
             |       |--P. canadensis R38
             |       |--P. contorta F56
             |       |    |--P. c. var. contorta F56
             |       |    `--P. c. var. latifolia F56
             |       |--P. engelmannii H93
             |       |--P. excelsa M47
             |       |--P. glauca BFD01
             |       |--P. glehnii M03
             |       |--P. jezoensis JKW03
             |       |--P. mariana BFD01
             |       |--P. morrisonicola MM12
             |       |--P. orientalis U01
             |       |--P. rubens SR03
             |       |--P. sitchensis MM12
             |       `--P. smithiana MM12
             `--+--Pityostrobus bernissartensis MM12
                `--Pinus EH06
                     |  i. s.: P. abies S00
                     |         P. ayacahuite CG05
                     |         P. cooperi CG05
                     |         P. fenzeliana L02
                     |         P. kochiana CK05
                     |         P. larix S00
                     |         P. luchuensis SM03
                     |         P. maritima C55
                     |         P. montana SR03
                     |         P. occidentalis J87
                     |         P. palaeostrobus S89
                     |         P. pallasiana CK05
                     |         P. pringlei CG05
                     |         P. quadrifolia H93
                     |         P. rudis CG05
                     |         P. torreyana [incl. P. torreyana ssp. insularis] H93
                     |         P. uncinata C55
                     |--P. subg. Pinus EH06
                     `--P. subg. Strobus EH06

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[BFD01] Bowman, J., G. J. Forbes & T. G. Dilworth. 2001. The spatial component of variation in small-mammal abundance measured at three scales. Canadian Journal of Zoology 79: 137–144.

[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.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1–E44.

[CK05] Chemeris, A. N., & N. M. Kovblyuk. 2005. A contribution to the knowledge of the harvestman fauna of the Crimea (Arachnida: Opiliones). Arthropoda Selecta 14 (4): 305–328.

[CP02] Chung, S.-W., & C.-I. Peng. 2002. Senecio kuanshanensis (Asteraceae), a new species from southern Taiwan. Botanical Bulletin of Academia Sinica 43: 155–159.

[CG05] Cognato, A. I., N. E. Gillette, R. C. Bolaños & F. A. H. Sperling. 2005. Mitochondrial phylogeny of pine cone beetles (Scolytinae, Conophthorus) and their affiliation with geographic area and host. Molecular Phylogenetics and Evolution 36: 494–508.

[C90] Crawford, R. L. 1990. Discovery of Caviphantes saxetorum in North America; status of Scironis tarsalis (Araneida, Linyphiidae). Journal of Arachnology 18: 235–236.

[D30] Druce, G. C. 1930. Account of a botanical tour in Cyprus. Proceedings of the Linnean Society of London 141: 50–52.

[EH06] Eckert, A. J., & B. D. Hall. 2006. Phylogeny, historical biogeography, and patterns of diversification for Pinus (Pinaceae): phylogenetic tests of fossil-based hypotheses. Molecular Phylogenetics and Evolution 40 (1): 166–182.

[E01] Engel, M. S. 2001. A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History 259: 1–192.

[F92] Fan Z. 1992. Key to the Common Flies of China 2^nd ed. Science Press: Beijing.

[F56] Foote, R. H. 1956. Gall midges associated with cones of western forest trees (Diptera: Itonididae). Journal of the Washington Academy of Sciences 46 (2): 48–57.

[HS17] Herrera, F., G. Shi, N. Ichinnorov, M. Takahashi, E. V. Bugdaeva, P. S. Herendeen & P. R. Crane. 2017. The presumed ginkgophyte Umaltolepis has seed-bearing structures resembling those of Peltaspermales and Umkomasiales. Proceedings of the National Academy of Sciences of the USA: E2385–E2391.

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

[JKW03] Jacobs, K., T. Kirisits & M. J. Wingfield. 2003. Taxonomic re-evaluation of three related species of Graphium, based on morphology, ecology and phylogeny. Mycologia 95 (4): 714–727.

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

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263–276.

[L02] Li F. 2002. Psocoptera of China vol. 2. Science Press: Beijing.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793–7798.

[M47] Mayr, E. 1947. Ecological factors in speciation. Evolution 1: 263–288.

[M03] Mockford, E. L. 2003. New species and records of Psocoptera from the Kuril Islands. Mitt. Mus. Nat.kd. Berl., Dtsch. Entomol. Z. 50 (2): 191–230.

[P02] Ponomarenko, A. G. 2002. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 164–176. Kluwer Academic Publishers: Dordrecht.

[R38] Ross, H. H. 1938. The Nearctic species of Pikonema, a genus of spruce sawflies (Hymenoptera, Tenthredinidae). Proceedings of the Entomological Society of Washington 40 (1): 17–20.

[SM03] Saldarriaga, J. F., M. L. McEwan, N. M. Fast, F. J. R. Taylor & P. J. Keeling. 2003. Multiple protein phylogenies show that Oxyrrhis marina and Perkinsus marinus are early branches of the dinoflagellate lineage. International Journal of Systematic and Evolutionary Microbiology 53: 355–365.

[S00] Saussure, C. de. 1800. De l'influence du sol sur quelques parties constituantes des végétaux. Bulletin des Sciences, par la Societé Philomathique de Paris 2 (40): 124–125.

[SS96] Simberloff, D., & P. Stilling. 1996. How risky is biological control? Ecology 77 (7): 1965–1974.

[SR03] Skovgaard, K., S. Rosendahl, K. O’Donnell & H. I. Nirenberg. 2003. Fusarium commune is a new species identified by morphological and molecular phylogenetic data. Mycologia 95 (4): 630–636.

[SW03] Song, D., & Q. Wang. 2003. Systematics of the longicorn beetle genus Coptomma Newman (Coleoptera: Cerambycidae: Cerambycinae). Invertebrate Systematics 17: 429–447.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 73–76.

[SN88] Suzuki, M., & S. Noshiro. 1988. Wood structure of Himalayan plants. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 341–379.

[TH03] Tedford, R. H., & C. R. Harington. 2003. An Arctic mammal fauna from the Early Pliocene of North America. Nature 425: 388–390.

[U01] Urhan, R. 2001. Two new species of the genus Zercon Koch from Turkey (Acari: Gamasida: Zerconidae). Genus 12 (4): 589–597.

Last updated: 2 December 2018.

Cupressaceae

Coastal redwoods Sequoia sempervirens, copyright Acroterion.

Belongs within: Cupressophyta.
Contains: Taxodioideae, Callitroideae, Cupressoideae.

The conifer family Cupressaceae includes cypresses (Cupressus), junipers (Juniperus) and related taxa. The family has previously been divided between the Cupressaceae sensu stricto (with leaves opposite or whorled, comprising Callitroideae and Cupressoideae) and Taxodiaceae (with leaves mostly alternate), but the latter is now recognised as paraphyletic to the former and most recent authors combine the two. Many species are significant timber trees. The giant redwood Sequoiadendron giganteum is the world's largest (though not tallest) tree.

Members of the Cupressaceae include the east Asian genera Cunninghamia and Taiwania, both of which bear seed cones with thin, leathery scales; Cunninghamia has linear, tapering leaves whereas Taiwania has scale-like leaves. Both these genera have more extensive distributions in the fossil record, with close relatives of both known from Europe and North America.

Characters (from Watson & Eckenwalder): Trees or shrubs, evergreen (rarely deciduous), generally resinous and aromatic, monoecious (sometimes dioecious). Bark fibrous and furrowed (occasionally smooth or exfoliating in plates). Lateral branches well developed, similar to leading shoots, twigs terete, angled, or flattened dorsiventrally (with structurally distinct lower and upper surfaces), densely clothed by scalelike leaves or by decurrent leaf bases; longest internodes to 1 cm; buds usually undifferentiated and inconspicuous. Roots fibrous to woody (rarely bearing aboveground "knees"). Leaves simple, usually persisting 3-5 years and shed with lateral shoots (cladoptosic) (rarely shed annually), alternate and spirally arranged but sometimes twisted so as to appear 2-ranked, or opposite in 4 ranks, or whorled, deltate-scalelike to linear, decurrent, sessile or petioled; adult leaves appressed or spreading, often differing between lateral and leading shoots (twigs heterophyllous), sometimes strongly dimorphic on each twig with lateral scale-leaf pairs conspicuously keeled; juvenile leaves linear, flattened, spreading; often with solitary abaxial resin gland; resin canal present. Pollen cones maturing and shed annually, solitary, terminal (rarely in clusters of 2-5, axillary, or in terminal panicles), simple, spheric to oblong; sporophylls overlapping, bearing 2-10 abaxial microsporangia (pollen sacs); pollen spheric, not winged. Seed cones maturing in 1-2 seasons, shed with short shoots or persisting indefinitely on long-lived axes (rarely shattering at maturity), compound, solitary, terminal (rarely in clusters of 2-5 or axillary); scales overlapping or abutting, fused to subtending bracts with only bract apex sometimes free; each scale-bract complex peltate, oblong or cuneate, at maturity woody or fleshy, with 1-20 erect (rarely inverted with age), adaxial ovules. Seeds 1-20 per scale, not winged or with 2-3 symmetric or asymmetric wings; aril lacking; cotyledons 2-9.

Cupressaceae [Taxodiaceae, Taxodinae, Thujopsidae]
    |--Sewardiodendron laxum MM12
    |--Austrohamia MM12
    |    |--*A. minuta MM12
    |    `--A. acanthobractea MM12
    |--+--Cunninghamiostrobus yubariensis MM12
    |  |--Cunninghamites MM12
    |  |    |--C. elegans GT02
    |  |    `--C. lignitum MM12
    |  `--Cunninghamia [Cunninghamioideae] MM12
    |       |--C. hornbyensis MM12
    |       |--C. konishii MM12
    |       `--C. lanceolata MM12
    `--+--Taiwania [Taiwanioideae] MM12
       |    |--T. cretacea MM12
       |    |--T. cryptomerioides K02
       |    |--T. flousiana GT02
       |    `--T. schaeferi MM12
       `--+--Athrotaxidoideae MM12
          |    |--Athrotaxites berryi MM12
          |    `--Athrotaxis MM12
          |         |  i. s.: A. ungeri MM12
          |         |--A. cupressoides MM12
          |         `--+--A. laxifolia MM12
          |            `--A. selaginoides MM12
          `--+--+--Taxodioideae MM12
             |  `--+--Callitroideae MM12
             |     `--Cupressoideae MM12
             `--Sequoioideae MM12
                  |--Metasequoia MM12
                  |    |--M. foxii MM12
                  |    |--M. glyptostroboides MM12
                  |    |--M. milleri MM12
                  |    `--M. occidentalis MM12
                  `--+--Austrosequoia wintonensis MM12
                     |--Quasisequoia MM12
                     |    |--Q. couttsiae (Heer) Kunzmann 1999 (see below for synonymy) HL08
                     |    `--Q. florinii MM12
                     `--+--Sequoiadendron giganteum MM12
                        `--Sequoia MM12
                             |--S. ambigua GT02
                             |--S. carbonaria JD05
                             |--S. concinna GT02
                             |--S. gigantea H04
                             |--S. langsdorfi S89
                             |--S. reichenbachii MM12
                             `--S. sempervirens GT02

Cupressaceae incertae sedis:
  Geinitzia reichenbachii GT02
  Taxodioxylon gypsaceum JD05
  Cupressoconus rhenanus JD05
  Otakauia Pole 1995 P00
    |--*O. lanceolata P00
    `--O. macchatiensis Pole 2000 P00
  Cupressinoxylon (nom. cons.) JD05 [=Retinodendron (nom. rej.) S03]
    |--C. glyptostrobium JD05
    |--C. polonicum JD05
    |--C. ponderosum JD05
    `--C. sequoianum JD05
  Hughmillerites juddii MM12
  Elatides williamsonii MM12

Quasisequoia couttsiae (Heer) Kunzmann 1999 [=Sequoia couttsiae Heer 1862, Athrotaxis couttsiae (Heer) Gardner 1884] HL08

*Type species of generic name indicated

REFERENCES

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671–685.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[HL08] Henniger, M., & R. Leder. 2008. Eozäne Makroflora der Querfurter Mulde. Mauritiana 20 (2): 229–251.

[JD05] Junge, F. W., M. Dolezych, H. Walther, T. Böttger, A. Kühl, L. Kunzmann, P. Morgenstern, T. Steinberg & R. Stange. 2005. Ein Fenster in Landschaft und Vegetation vor 37 Millionen Jahren: Lithologische, sedimentgeochemische und paläobotanische Befunde aus einem Paläoflusssystem des Weißelsterbeckens. Mauritiana 19 (2): 185–273.

[K02] Kvaček, Z. 2002. Novelties on Doliostrobus (Doliostrobaceae), an extinct conifer genus of the European Palaeogene. Časopis Národního Muzea, Řada Přírodovědná 171 (1–4): 47–62.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793–7798.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153–197.

[S03] Skog, J. E. 2003. Report of the Committee for Fossil Plants: 4. Taxon 52: 341.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 73–76.

Pinopsida

Chilean rimu Lepidothamnus foncki, copyright Jeff Bisbee.

Belongs within: Coniferophyta.
Contains: Miroviaceae, Doliostrobaceae, Voltziales, Pinaceae, Cheirolepidiaceae, Cupressophyta.

The name Pinopsida is used here for a clade uniting the living conifers with the Palaeozoic Voltziales (probably a paraphyletic grouping). Unique morphological synapomorphies have not yet been identified for this clade (Rothwell & Mapes 2001), but the crown group Coniferae is supported relative to other living plants by their needlelike leaves, tiered proembryos and loss or extreme reduction of one copy of the inverted repeat in the chloroplast genome (Cantino et al. 2007). Recent phylogenies have mostly placed the Pinaceae as the sister clade to the remaining living conifers.

<==Pinopsida [Coniferales, Coniferopsida, Conopsida]
    |--Voltziales FC07
    `--Coniferae [Pinoideae] CD07
         |  i. s.: Lepidothamnus D03
         |           |--L. foncki D03
         |           `--L. laxifolius W91
         |--Pinaceae FC07
         `--+--Cheirolepidiaceae FC07
            `--Cupressophyta CD07

Pinopsida incertae sedis:
  Tylodendron H-CRM01
  Palaeotaxites praecursor H-CRM01
  Paranocladus H-CRM01
  Ugartecladus genoensis H-CRM01
  Szecladia multinervia H-CRM01
  Dadoxylon H-CRM01
    |--D. farleyensis Walkom 1928 F71
    `--D. owangondianum H-CRM01
  Araucarioxylon arizonicum H-CRM01, GE05
  Walchiopremnon valdajolense H-CRM01
  Culmitzschia H-CRM01
  Buriadia heterophylla H-CRM01, RM01
  Curionia H-CRM01
  Feysia H-CRM01
  Gomphostrobus bifidus H-CRM01, RM01
  Walkomia H-CRM01
  Podocarpites ZBH03
  Cephalotaxopsis ZBH03
  Samaropsis pincombei Walkom 1928 Z02, F71
  Bilsdalea dura Z02
  Heidiphyllum Z02
  Podozamites Braun 1843 [Podozamitaceae] BO02
    |--P. gracilis [=Araucarites gracilis] F71
    `--P. guttiformis BO02
  Pityophyllum RY03
  Rhipidiocladus RY03
  Elatocladus RY03
    |--E. australis F71
    `--E. planus F71
  Schizolepis RY03
  Cyparissidium gracile GT02
  Hirmerellaceae Z02
    |--Pagiophyllum GT02
    |--Brachyphyllum GT02
    |    |--B. graciliforme BO02
    |    |--B. hegewaldia BO02
    |    `--B. speciosa BO02
    `--Hirmeriella muensteri (Schenk) Jung 1968 S05
  Sequoites GT02
  Miroviaceae G02
  Sciadopityoxylon PM02
  Xenoxylon PM02
  Podocarpoxylon Gothan 1905 JD05
    |--P. angustiporosum JD05
    |--P. helmstedtianum Gottwald 1966 K02
    `--P. knowltonii JD05
  Protopodocarpoxylon S03
  Doliostrobaceae K02
  Dammara K02
    |--D. borealis K02
    |--D. intermedia Ettingshausen 1886 F71
    |--D. macrosperma K02
    `--D. podozamioides Ettingshausen 1886 F71
  Chamaecyparites hardtii K02
  Embothrites cuneatus Ettinghausen 1868 K02
  Rufloria [Rufloriaceae] RM01
  Pityanthus jefferyi RM01
  Eutactoides Pole 2000 P00
    `--*E. wintonensis Pole 2000 P00
  Circoporoxylon Kraeusel 1949 JD05

*Type species of generic name indicated

REFERENCES

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707-737.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549-552.

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761-773.

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671-685.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H-CRM01] Hernandez-Castillo, G. R., G. W. Rothwell & G. Mapes. 2001. Thucydiaceae fam. nov., with a review and reevaluation of Paleozoic walchian conifers. International Journal of Plant Science 162 (5): 1155-1185.

[JD05] Junge, F. W., M. Dolezych, H. Walther, T. Böttger, A. Kühl, L. Kunzmann, P. Morgenstern, T. Steinberg & R. Stange. 2005. Ein Fenster in Landschaft und Vegetation vor 37 Millionen Jahren: Lithologische, sedimentgeochemische und paläobotanische Befunde aus einem Paläoflusssystem des Weißelsterbeckens. Mauritiana 19 (2): 185-273.

[K02] Kvaček, Z. 2002. Novelties on Doliostrobus (Doliostrobaceae), an extinct conifer genus of the European Palaeogene. Časopis Národního Muzea, Řada Přírodovědná 171 (1-4): 47-62.

[PM02] Philippe, M., H. Méon, G. Lambert, B. Erdei, F. Thevenard & B. Gomez. 2002. A palm-tree and Sciadopitys swamp-forest from the Neogene of Bresse (eastern France). Comptes Rendus Palevol 1: 221-225.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153-197.

[RY03] Ren, D., & J. Yin. 2003. New ‘osmylid-like’ fossil Neuroptera from the Middle Jurassic of Inner Mongolia, China. Journal of the New York Entomological Society 111 (1): 1-11.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637-667.

[S05] Säilä, L. K. 2005. A new species of the sphenodontian reptile Clevosaurus from the Lower Jurassic of south Wales. Palaeontology 48 (4): 817-831.

[S03] Skog, J. E. 2003. Report of the Committee for Fossil Plants: 4. Taxon 52: 341.

[Z02] Zherikhin, V. V. 2002. Insect trace fossils. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 303-324. Kluwer Academic Publishers: Dordrecht.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807-814.

Araucariaceae

Norfolk Island pine Araucaria excelsa, photographed by Danielle Langlois.

Belongs within: Cupressophyta.

The Araucariaceae is a family of thirty-five living species of conifer found in South America and the south-western Pacific region. Members of the genus Agathis have hard and smooth bark and petiolate leaves; species of Agathis are found between the Philippines, eastern Australia and Polynesia with a single species A. australis, the kauri, found in northern New Zealand. Species of Araucaria and Wollemia nobilis have rough, exfoliating bark and sessile leaves that are spirally arranged in Araucaria but opposite in Wollemia nobilis. Some are popular ornamental trees; the Norfolk Island pine Araucaria excelsa is particularly popular for its remarkably geometrical, conical shape.

Characters (from Allan 1961): Plants mostly dioecious. Male strobili with numerous sporophylls. Female cones woody, terminal on short branchlets. Carpidia numerous, spirally arranged; two or four cotyledons present.

Araucariaceae [Araucarinae]
    |  i. s.: Araucarioides P00
    |         Araucarites Presl 1838 BO02
    |           |--A. cutchensis F71
    |           |--A. falsanii BO02
    |           |--A. indica Lele 1962 C93
    |           |--A. janaianus BO02
    |           `--A. parsorensis Lele 1955 C93
    |         Woodworthia arizonica Jeffrey 1910 CA04
    |--+--Wollemia nobilis MM12
    |  `--Agathis Salisbury 1807 MM12, A61
    |       |--A. alba JD05
    |       |--A. australis Salisb. 1807 (see below for synonymy) A61
    |       |--A. dammara MM12
    |       |--A. palmerstoni H42
    |       `--A. robust W95
    `--Araucaria MM12
         |--A. angustifolia CL-Z03
         |--A. araucana (Molina) Koch 1873 CD07
         |--A. balansae GT02
         |--A. balcombensis P00
         |--A. bidwillii M94
         |--A. biramulata M94
         |--A. brasiliensis MM96
         |--A. carinatus P00
         |--A. columnaris M94
         |--A. cunninghamii GT02
         |--A. excelsa GT02
         |--A. fletcheri Selling 1950 F71
         |--A. heterophylla NM11
         |--A. hunsteinii K77
         |--A. imbricata V01
         |--A. macrophylla GT02
         `--A. mirabilis MM12

Agathis australis Salisb. 1807 [=Dammara australis Don in Lamb. 1824; incl. Podocarpus zamiaefolius Rich. 1832] A61

*Type species of generic name indicated

REFERENCES

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

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707–737.

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[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793–7798.

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Last updated: 26 June 2020.

Podocarpaceae

Kahikatea Dacrycarpus dacrydioides, from here.

Belongs within: Cupressophyta.

The Podocarpaceae are a family of more than 150 living species of mostly Southern Hemisphere conifers. Some are dominant forest species within their range and are used for timber. Many species produce fleshy 'pseudo-fruits' (really an enlarged seed scale) with seeds that are animal-dispersed as with many true fruits. In Podocarpus and Dacrydium, the ovule is exposed atop the seed scale; in Prumnopitys, the seed scale encloses the ovule. Members of the genus Phyllocladus, the celery pines, are distinctive in having the leaves largely replaced by broad, flattened, leaf-like photosynthesising stems. The genus Nageia, found in tropical and subtropical parts of south-east Asia and Australasia, has broad flat leaves more similar to those of Agathis (kauri) species than other Podocarpaceae.

Characters (from Eckenwalder 2009): Evergreen trees or shrubs with scale-, claw- or flattened, needlelike or broader leaves, usually with single undivided midvein. Leaves attached individually in spiral arrangement or in pairs radiating around twig or distichously in more or less flat rows on either side of predominantly horizontal branchlets. Branching typically with extra weaker branches along trunk between main tiers of three to five major branches. Pollen cones small to moderate, single at tips of branchlets or single to clustered in axils of foliage leaves or bracts on otherwise ordinary branchlets or on specialised reproductive shoots, often with one or a few rows of distinctive bracts at base. Pollen scales usually numerous to very numerous, attached spirally or apparently so by a (usually) short stalk, with two (or four) round to slightly elongated pollen sacs attached to stalk and to lower, outer edge of upturned more or less triangular blade. Pollen grains usually with two (or three) variously large to small air bladders. Seed cones single or clustered and often upright on short or long specialised reproductive branchlets in axils of leaves or at tips of otherwise ordinary shoots, almost always with some kind of fleshy structure(s). Seed scales typically few, often just a single fertile scale, with only a little fusion between the sometimes fleshy lower bract portion (podocarpium if fleshy) and the more frequently fleshy upper ovuliferous scale portion (epimatium). Seeds almost always single, folded within epimatium, small to large, plump and wingless, usually animal-dispersed themselves or as part of intact seed cone, with fleshy seed coat, aril, epimatium or podocarpium or combination thereof. Cotyledons normally two, each with two veins. Chromosome base numbers x = 9-19.

Podocarpaceae
    |--Phyllocladus L.C. & Rich. 1826 MM12, A61 [Phyllocladinae]
    |    |--P. alpinus Hook. f. 1853 [=P. trichomanoides var. alpina Parl. in DC. 1868] A61
    |    |--P. aspleniifolius J00 [=Podocarpus aspleniifolius BR00]
    |    |--P. asplenioides Ettingshausen 1886 F71
    |    |--P. glaucus Carr. 1855 [=P. trichomanoides var. glauca Parl. in DC. 1868] A61
    |    |--P. hypophyllus P88
    |    `--P. trichomanoides Don in Lamb. 1832 [incl. P. rhomboidalis Rich. 1832 non L.C. & Rich. 1826] A61
    `--+--Acmopyle MM12
       |    |--A. pancheri MM12
       |    `--A. sahniana MM12
       `--+--Dacrycarpus MM12
          |    |--D. compactus C78
          |    |--D. cumingii H03
          |    |--D. dacrydioides WH02
          |    |--D. imbricatus MM12
          |    |--D. kinabaluensis H03
          |    `--D. steupii H03
          `--+--Nageia nagi MM12
             `--Podocarpus L’Heritier 1807 MM12, A61
                  |--P. acutifolius Kirk 1884 A61
                  |--P. aristulatus J87
                  |--P. atjehensis H03
                  |--P. chinensis MM12
                  |--P. dacrydioides Rich. 1832 (see below for synonymy) A61
                  |--P. drouynianus GK00
                  |--P. falcatus K09
                  |--P. ferrugineus Benn. ex Don in Lamb. 1832 A61
                  |--P. hallii Kirk 1889 (see below for synonymy) A61
                  |--P. imbricatus P88
                  |--P. latifolius MM12
                  |--P. lawrencei MM04
                  |--P. × loderi Ckn. 1932 [P. hallii × P. totara] A61
                  |--P. macrophyllus (Thunberg) Sweet 1818 CD07 [=P. macrophylla LO98]
                  |--P. micropedunculatus H03
                  |--P. nagi MH98
                  |--P. nivalis Hook. 1843 [incl. P. montanus Col. 1895] A61
                  |    |--P. n. var. nivalis A61
                  |    `--P. n. var. erectus Ckn. 1916 A61
                  |--P. novae-caledoniae MM12
                  |--P. nubigena D03
                  |--P. oleifolius R96
                  |--P. praecupressina Ettingshausen 1886 F71
                  |--P. rotundus S03
                  |--P. rumphii H03
                  |--P. spathoides H03
                  |--P. spicatus Br. ex Mirbel 1825 (see below for synonymy) A61
                  |--P. totara Benn. ex Don in Lamb. 1832 A61
                  `--P. urbanii M93

Podocarpaceae incertae sedis:
  Eromangia Pole 2000 P00
    `--*E. douglasii Pole 2000 P00
  Thargomindia Pole 2000 P00
    `--*T. wordsmithiana Pole 2000 P00
  Prumnopitys MM04
    |--P. ferruginea MM04
    `--P. taxifolia WH02
  Dacrydium Sol. ex Forst. f. 1786 A61
    |--D. beccarii P88
    |--D. bidwillii Hook. f. ex Kirk 1878 (see below for synonymy) A61
    |--D. biforme (Hook.) Pilger 1903 (see below for synonymy) A61
    |--D. colensoi Hook. 1843 [incl. D. westlandicum Kirk ex Hook. f. 1877] A61
    |--D. comosum P88
    |--D. cupressinoides Ettingshausen 1886 F71
    |--D. cupressinum Lamb. 1807 [=Thalamia cupressina Spreng. 1826] A61
    |--D. dacrydioides BKS98
    |--D. elatum P88
    |--D. intermedium Kirk 1878 [incl. D. intermedium var. gracilis Kirk 1885] A61
    |--D. kirkii Muell ex Parl. in DC 1868 A61
    |--D. laxifolium Hook. f. 1845 A61
    |--D. magnum H03
    |--D. medium H03
    |--D. pectinatum P88
    `--D. pendulum A27

Dacrydium bidwillii Hook. f. ex Kirk 1878 [incl. D. bidwillii var. erecta Kirk 1889, D. bidwillii var. reclinata Kirk 1889] A61

Dacrydium biforme (Hook.) Pilger 1903 [=Podocarpus biformis Hook. 1843; incl. D. colensoi Hook. f. 1853 non Hook. 1843] A61

Podocarpus dacrydioides Rich. 1832 [incl. Dacrydium excelsum Don in Lamb. 1832, Nageia excela Kuntze 1891, Podocarpus excelsus (Don) Druce 1917, Dacrydium ferrugineum Houtte ex Gord. 1858 A61, P. thujoides Br. in Benn. 1838, D. thuioides Banks & Sol. ex Carr. 1855] A61

Podocarpus hallii Kirk 1889 [=P. totara var. hallii (Kirk) Pilger 1903; incl. P. cunninghamii Colenso 1884] A61

Podocarpus spicatus Br. ex Mirbel 1825 [=Prumnopitys spicata Kent in Veitch 1900; incl. Dacrydium mai Cunn. 1838, Podocarpus matai Lamb. ex Hook. f. 1867, D. mayi Houtte ex Gord. 1858, D. taxifolium Banks & Sol. ex Don in Lamb. 1824 non Po. taxifolia Kunth 1817] A61

*Type species of generic name indicated

REFERENCES

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