EMPIRE BIOTA

                                                   a Comprehensive Taxonomy

Table 12. Plantae 
      
        I have introduced series (srs.) at this high level due to the large number of ranks, which is sometimes used for subgeneric levels, but was used by August Pulle for divisions of dicots;  all the groups are stat. nov. except Spermatophyta; synapomorphies are in parenthses as are the totals; *indicates extinct groups.

      sbk./phyl. Volvocophyta (Oltsmann 1904, as order) stat. nov. (2 classes, 500 gen., 8000 sp.)
      sbk. Metaphyta stat. nov. (unilateral rootlets, glycolate degradation by glycolate oxidase)
            infk./phyl. Chlorokybophyta stat. nov. (1 species)
            infk. Neophyta stat. nov., nom. nov. (rosette complex)
                  mck./phyl. Gamophyta stat. nov. (1  order or 2 orders, 3 families, 60 gen., 4600 sp.)
                  mck. Streptophyta (Jeffrey 1967) stat. nov. (Phragmaphyta nom. nov.) (cell plate)
                        nnk./phyl. Coleochetophyta stat. nov. (2 gen., 15 sp.)
                        nnk. Apicophyta stat. nov., nom. nov. (apical growth, C6-C3 phenylpropanoid-derived flavonoid 
                                                                                            compound,  similarities in mitotic mechanisms)
                              sbnnk./phyl. Charophyta (Rabenhorst 1863) stat. nov. (stoneworts and musk grasses)
                              sbnnk. Embryophyta (Endlicher 1836) stat. nov. (Cormophyta Endlicher 1836) (embryo, stem,
                                                 multicellular sporophytes,  archegonia, lignins, meiospores with exine,  sporangium,
                                                 sporopollenin )
                                    infnnk. Hepaticophyta
                                          phyl. Hepaticophyta (liverworts) (6000 sp.)
                                    infnnk. Stomophyta nom. nov.(stomates, guard cells, collumate sporanguim, D-methionine)
                                          spsrs. Cerotophyta nom. nov. 
                                               phyl. Cerotophyta  (hornworts)(100 sp.)
                                          spsrs. Coscinophyta nom. nov. (leptoid (nutrient-conducting cell)-sieve-element
                                                             homology)(Gk. "coscinos" ( sieve)),  axial gametophyte,
                                                             terminal gametangiums, persistent and      internally differentiated sporophytes
                                                srs. Muscophyta nom. nov.
                                                      phyl. Muscophyta (mosses)(10, 000 sp.)
                                                srs. Polysporangiophyta (multiple sporangia (sporangial branching), columella,
                                                                     sunken   archegonium, independent alternation of generations)
                                                      sbsrs. Aglaophyta*
                                                            phyl. Aglaophyta (1 genus)
                                                      sbsrs. Stelophyta Pia 1931 (Tracheophyta Eames 1936) (stele (vascular cylinder),
                                                           vascular  system, water-conducting cells with annular-helical thickenings (tracheids),
                                                             lignin)
                                                            hprggph. Horneophyta* (protracheophytes)
                                                                  phyl Horneophyta (3 genera)
                                                            hprggph. Eustelophyta ( thick lignified, decay-resistant  wall layer in the tracheids,
                                                                             pitlets  between thickenins or with pits, sterome)
                                                                  spggph. Rhyniophyta* (4 genera) (protracheophytes)
                                                                        phyl. Rhyniophyta Banks 1958 stat. nov.
                                                                  spggph. Neostelophyta nom. nov., stat. nov. (endarchy (centrifugal xylem
                                                                                 maturation), terminal branching, lateral and endogenous root branching
                                                                                 pattern, terminal   sporangia)
                                                                        ggph. Lycophyta Eichler 1883
                                                                              phyl. Lycophyta (club mosses and quillworts(Merlin's grass))(15 gen.,
                                                                                             1000 sp.)
                                                                        ggph. Telomophyta (telomation, longitudinal sporangial dehiscence, centrarch
                                                                                         or        mesarch xylem)
                                                                              mgph. Trimerophyta*
                                                                                    phyl. Trimerophyta
                                                                              mgph. Megaphylla (Euphyllophyta Kenrick and Crane 1997) 
                                                                                        ( large leaf with web of veins midrib, and multiple traces,
                                                                                         pseudopodial or  monopodial and helical branching, recurvation
                                                                                         of branch apices, trachieds with  scalariform bordered pits,
                                                                                         paired sporangiums grouped into terminal trusses, radially aligned
                                                                                         xylem in larger axes, multiflagellate spermatozoids)
                                                                                    hprph. Calamophyta Haeckel 1866 
                                                                                        phyl. Calamophyta (ferns, whisk ferns, and horsetails)(12,000 sp.)
                                                                                    hprph. Lignophyta Doyle and Donaghue 1986 (secondary wood,
                                                                                                 phlem (bast), wood rays)
                                                                                           spph. Aneurophyta* ( progymnosperms, 1 gen.)
                                                                                           spph. Metalignophyta (heterospory)
                                                                                                phyl. Archeophyta* ( progymnosperms, 1 gen.)
                                                                                                phyl. Spermatophyta Eichler 1883 (Phanerogamae auct;
                                                                                                        Siphonogama Engler and Diels 1936) (80 orders (6
                                                                                                         others extinct),  430 families, 14,000 genera, and a
                                                                                                         good quarter  million species) (seeds, siphonogamy)

Spermatophyta (only extant forms)

     Cycadophyta
     Platysperma (platyspermy,  biradial symmetry of  the seed, saccate pollen, sealing off of the micropyle after pollination)
          Pinophyta (conifers and ginkyos) (the 2nd g is an orthographic error originating with Kaempfer in 1712)
          Diplophyta (double fertilization, heteroxylous wood, columnar tracheid, perianth)
               Chlamydospermae (joint fir, welwichia, ephedra)
               Angiospermae (Florideae) (flowering plants)
 
Anthophyta ( Angiospermae or  Florideae)
 
        spcl Monocolpatae
              cl. Magnoliopsida (Nympheales (water lily) and Magnoliales (magnolia))
              cl. Liliopsida (Monocotyledonae)
                   sbcl.  Alismatidae (water plantain)
                   sbcl Lilidae (lily)
         spcl Tricolpatae (Eudicotyledonae)
              cl. Ranuculopsida (buttercup) 
              cl. Rosopsida
                   sbcl Rosidae (rose)
                   sbcl Asteridae (aster)


Table 15. Animalia

 

Here I  used section (sct.) and series (srs.), which have been introduced due to the many ranks occuring in this kingdom. Branch and grade are also used but grade is not an acceptable term in phylogenetics and all are branches anyways; synaomorphies are in parentheses; figures in parentheses are generally approximations and are for classes, orders, families, genera, and species, in that order; extinct groups are designated with an asterisk.

sbk. /phyl. Choanoflagellata Kent 1880 (Craspedophyceae Chadefaud 1960, Choanomastigota, Choanozoa Cavalier-Smith
                                                             1989) (collar flagellates) 
sbk. Metazoa Haeckel 1877
   infk. /phyl. Archeocyatha* Bornemann 1884
   infk. Neozoa nom. nov.
      mck. phyl. Spongiae (syns. Spongiaria de Blainville 1816, Parazoa Sollas 1884, Porifera Grant
                                                    1835) (3, 28, 100, 350, 9000)
      mck.  Epitheliozoa nom. nov. ( epithelium and belt desmosomes)
         nnk./phyl. Placozoa Grell 1971 (only 1 species)
         nnk. Euzoa (Enterozoa Lankester 1878, Eumetazoa) ( symmetry, organs, organ systems, gap junctions,
                                     basement  membranes, and hemidesmosomes)
            sbnnk./phyl. Cnidaria (Celenterata) (4, 27, 236, 927, 9000)
            sbnnk. Argazoa nom. nov.   ( multiciliate epithelium, acetylcholine as neurotransmitter, unidirectional synaptic
                                                             impulse transmission, nerve fibers with insulating sheath )
               hprsrs./phyl. Ctenophora Eschscholtz 1829
               hprsrs. Artiozoa (Bilateria Schimkewitsch 1891, Triploblastica Lankester 1900) ( bilateral symmetry, 3 germ layers,
                                     well   organized tisssue formation, CNS (central nervous system), protonephridia, mobility, cephaly)
                  spsrs. Protostomia Grobben 1908 (monociliate epithelia, ventral longitudinal nerves paired or secondarily fused,
                                                                     protonephridia)  
                      srs. Ecdysyzoa Aguinaldo et al 1997 (molting cuticle)
                             hprph/phyl. Cephalorhyncha (cls. Kinorhyncha Reinhard 1887 (Echinodera Zelinka 1928), Priapula
                                         (Priapuloidea     Sedgewick 1898), and Lorcifera) 
                             hprph. Sensillozoa nom. nov. (sensillium) (or Nematozoa + Cepalorhyncha forming Introverta)
                                spph./phyl. Nematohelminthes Gegenbauer 1859 (Nematozoa) (Nematoda +Nematomorpha) (1000 gen.,
                                                 12, 000 sp.)
                                spph. Panarthropoda Neilsen 1985 
                                   phyl. Onychophora Grube 1851 (velvet worms)
                                   phyl. Arthropoda von Siebold and Stannius 1845  
                     srs.  Spiralia Neilsen 1995 s.l. (spiral cleavage with 4-d mesoderm)
                            sbsrs. Platyzoa
                                 phyl. Trochata (cl. Rotifera (Rotatoria Ehrenberg 1838), cl. Acanthocephala Rudolphi 1808)
                                                (or with  Ecdyzyzoa forming Aschelminthes Neilsen 1995)
                                 phyl. Platymorpha nom. nov. (cls. Gastrotricha Metschnikoff 1864, Gnathostoma,
                                          Platyhelminthes     (flatworms)) (2000 gen., 20,000 sp.)
                            sbsrs. Trochozoa (trochophore larva)
                                   mgph. /phyl. Kamptozoa Cori 1929 (syns. Calyssozoa, Entoprocta Nitsche 1870)(+Cycliphora?) 
                                   mgph. Eutrochozoa  
                                        hprph./ phyl. Echiuroidea Sedgewick 1898 s.s.  (cls. Echiura Savigny 1817 as 
                                                Echiurida (spoon  worms), Pogonophora Johansson 1937, and Chetifera)
                                        hprph. Molluscoidia nom. nov.
                                           spph/phyl. Nemertina Oersted 1844 (Rhynchocela Schulze 1850)(ribbon worms) 
                                           spph. Molluscomorpha nom. nov. 
                                               phyl. Sipuncula Raffinesque 1814 
                                               phyl. Mollusca Linneus 1758 (Malacozoa) (7, 34, 530, 3000, 50 000) 
                  spsrs. Deuterostomia Grobben 1908 (Enterocelia Hatschek 1811, Epineuralia Cuénot 1940, Notoneuralia
                                         Ulrich 1951, Oligomeria, Irregularia) (blastopore develops posteriorly, enterocely, and radial,
                                             indeterminate,  and regular cleavage)
                                    srs. Chetognatha Leuckart 1854 (arrow worms) 
                                    srs. Enterocela Lankester 1876 (Trimera) (trimery (prosome, mesosome, metasome), dorsal
                                              CNS,  upstream collecting larval ciliary bands)
                                      sbsrs.  Lophophorata Hyman 1959 (Tentaculata Hatschek 1891; Polyzoa Thompson 1830)
                                            phyl. Bryozoa Ehrenberg 1831 (Ectoprocta Nitsche 1870, Zoophyta Cuvier 1812) (moss
                                                     animals) (2,     10, 108, 300, 10 000)                                         
                                            phyl. Phoronozoa  Zrzavy et al 1998  (cls. Brachiopoda Cuvier 1802,
                                                Phoronida Hatschek 1888 (Pygocaulia Thompson 1927)  
                                      sbsrs. Epithelioneuria  (axial complex, epithelioneural system)
                                                ggph./phyl. Echinodermata Laske 1778 (6, 36, 145, 250, 7000)
                                                ggph. Stomochordata  (stomochorda)
                                                 mgph./phyl. Pterobranchia Lankester 1877
                                                    mgph. Cyrtotreta Neilsen 1995 (gill slits, dorsal neural tube, collagenous skeleton)
                                                     hprph./  phyl. Enteropneusta Gegenbaur 1870 (acorn worms)
                                                          hprph. Chordata Haeckel 1874 (Chordonia auct.) (chorda, dorsal neural tube,  true tail,
                                                                    dorsal longitudinal muscles,  endostyle organ)
                                                            spph. /phyl. Urochordata (Tunicata Lamarck 1816)
                                                            spph. Notochordata (chorda extending along the whole dorsal side, myomery,
                                                                     mesoderm segmentation, open capillary junctions, grey  and white differntiaton
                                                                     in the neural tube)
                                                                   phyl. Cephalochordata Owen 1846 (Acrania Haeckel 1866) (lancelets)
                                                                   phyl. Vertebrata Cuvier 1817 (Craniata Lankester 1877)
                                                                           (?, 100 ( 100 more extinct),  800, 8,000,  44 000)

spcoh Holometabola Martynov 1925 (Oligoneoptera auct., Endopterygota Sharp 1893)
 
          coh Coleopterida  Handlirsch 1908
                ord Coleoptera Linneus 1758 s.s. (beetles)
          coh Telomerida Boudreaux 1979
                 sbcoh Hymenopterida Handlirsch 1908 , emend. from Hymenopteroida
                      ord. Hymenoptera Linneus 1758 (wasps, bees, ants)
                 sbcoh Meronida Boudreaux 1979
                      infcoh Neuropterida Handlirsch 1908, emend. from Neuropteroidea
                           spord Raphidida Handlirsch 1908
                                ord Raphidiodea Burmeister 1839 ( snake flies)
                           spord Sialida Handlirsch 1908, emend. from Sialoidea
                                ord Neuroptera Linneus 1758 s.s. (=Plannipennes Latreille 1817) (lace-wings)
                                ord Megaloptera Latreille 1802 s.s. (alder flies and allies)
                      infcoh Panorpida Handlirsch 1908
                           hprord Mecopterida (=Siphonoptera + Antliophora, Hennig 1969) stat. nov.
                                spord Mecopterodea Boudreaux 1979)
                                     ord Mecoptera Packard 1886 (scorpion flies)
                                     ord Siphonaptera Latreille 1825  (fleas)
                                spord Hanstellodea Clairville 1798, s.s., emend. from Haustellata
                                     ord Diptera Linneus 1758 (2-winged flies)
                                     ord Strepsiptera Kirby 1813
                           hpord Trichopterida Boudreaux 1979 (Amphiesmenophora Hennig 1969) stat. nov.
                                    ord Trichoptera Kirby 1826 (caddis flies)
                                    ord Lepidoptera Linneus 1758 (moths and butterflies)

Based on Hennig 1969 Boudreaux 1979, Kristensen 1991 and Whiting et al 1997. Boudreaux places Coleopterida on top and includes Strepsiptera but this inclusion  is not well supported  by morphology nor molecules (Whiting et al , 1997). Antliophora, Mecopteromorpha, Dipteromorpha, and Amphiesmenophora are all strongly supported by both types of data  (Whiting et al , 1997).

        The amazing success and diversity of winged insects is attributable to water loss prevention (gas exchange system, spiracular closing mechanism, cuticular exoskeleeton and impermeable waxy cuticle, laying of waterproof eggs, and Malpighian tubules),  ability to fly, ability to bite and chew food, rapid generation, small size, malleable structures, and, in Holometabola, a specialized larval (pupal) stage, and, in beetles, chitinized armour to protect against predators and parasites.
        In Holometabola, are found the essential pollinators, the Big 4, all named by Linneus, which were long thought to have co-evolved with flowers, but new evidence--a comprehensive study of insect evolution (Sepkoski and Labandeira, 1993) and the discovery of bees' nests from the Triassic in the Petrified Forest of Arizona (Haciotis, 1995), which go back 220 mln. yrs. indicating an origin 140 mln. yrs. earlier than previously believed-- strongly suggests that bees were already pollinating gymnosperms and that flowers evolved showy petals and scents to compete with them for the attention of insects so that the tradtional view is probably erroneous.

Boudreaux, H. B. 1979. Arthropod Phylogeny, with special reference to insects.  Wiley.
Hasiotis, S.T. and Dubiel R.F. 1995. Termite (Insecta: Isoptera) nest ichnofossils from the upper Triassic Chinle formation,
     Petrified   Forest National Park, Arizona. Ichnos 4: 126-130.
Kristensen. N.P.  1991. Phylogeny of extant hexapods. In: The Insects of Australia, I.D. Nauman et al, eds., pp. 125-40.
     University   Press, Melbourne.
Sepkoski, J.J., Jr., and Labandeira, C.C.  1993. Insect diversity in the fossil record. Science 261: 310-15.
Whiting, M.F., Carpenter, J.C., Wheeler, Q.D., Wheeler, W.C. 1997. The Strepsiptera problem: Phylogeny of the
     holometabolous   insect orders inferred from 18S and 28SrDNA sequences and morphology. Syst. Biol. 46: 1-68.

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