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РЕФЕРАТ.
Проведено исследование эволюционной истории гиперразнообразной клады, подсемейства муравьёв Myrmicinae (Hymenoptera: Formicidae), основанное на анализе матрицы данных 251 вида и 11 ядерных генов. Вместо 25 триб оставлено только 6 новых по объёму триб.
Under both maximum
likelihood and Bayesian methods of inference, we recover a robust phylogeny that
reveals six major clades of Myrmicinae, here treated as newly defined tribes and occurring
as a pectinate series: Myrmicini, Pogonomyrmecini trib.n., Stenammini, Solenopsidini,
Attini and Crematogastrini. Because we condense the former 25 myrmicine tribes
into a new six-tribe scheme, membership in some tribes is now notably different, especially
regarding Attini. We demonstrate that the monotypic genus Ankylomyrma is neither
in theMyrmicinae nor even amember of themore inclusive formicoid clade – rather
it is a poneroid ant, sister to the genus Tatuidris (Agroecomyrmecinae). Several
species-rich myrmicine genera are shown to be nonmonophyletic, including Pogonomyrmex,
Aphaenogaster, Messor, Monomorium, Pheidole, Temnothorax and Tetramorium.
We propose a number of generic synonymies to partially alleviate these problems
(senior synonym listed first): Pheidole = Anisopheidole syn.n.=Machomyrma syn.n.;
Temnothorax = Chalepoxenus syn.n.=Myrmoxenus syn.n.=Protomognathus syn.n.;
Tetramorium = Rhoptromyrmex syn.n.=Anergates syn.n. = Teleutomyrmex syn.n. The
genus Veromessor stat.r. is resurrected for the New World species previously placed
in Messor; Syllophopsis stat.r. is resurrected from synonymy under Monomorium to
contain the species in the hildebrandti group; Trichomyrmex stat.r. is resurrected from
synonymy under Monomorium to contain the species in the scabriceps- and destructor-
groups; and the monotypic genus Epelysidris stat.r. is reinstated for Monomorium
brocha. Bayesian divergence dating indicates that the crown group Myrmicinae originated
about 98.6Ma (95% highest probability density 87.9–109.6Ma) but the six major
clades are considerably younger, with age estimates ranging from 52.3 to 71.1Ma.
Although these and other suprageneric taxa arose mostly in the middle Eocene or earlier,
a number of prominent, species-rich genera, such as Pheidole, Cephalotes, Strumigenys,
Crematogaster and Tetramorium, have estimated crown group origins in the late Eocene
or Oligocene. Most myrmicine species diversity resides in the two sister clades, Attini
and Crematogastrini, which are estimated to have originated and diversified extensively
in the Neotropics and Paleotropics, respectively. The newly circumscribed Myrmicini is
Holarctic in distribution, and ancestral range estimation suggests a Nearctic origin. The
Pogonomyrmecini and Solenopsidini are reconstructed as being Neotropical in origin,
but they have subsequently colonized the Nearctic region (Pogonomyrmecini) and many
parts of the Old World as well as the Nearctic region (Solenopsidini), respectively. The
Stenammini have flourished primarily in the northern hemisphere, and are most likely
of Nearctic origin, but selected lineages have dispersed to the northern Neotropics and
the Paleotropics. Thus the evolutionary history of the Myrmicinae has played out on a
global stage over the last 100 Ma, with no single region being the principal generator of
species diversity.
ТАКСОНОМИЯ.
Veromessor stat.r. восстановлен из синонимии с
Messor, generating the following revived combinations in
Veromessor: andrei, chamberlini, chicoensis, julianus, lariversi,
lobognathus, pergandei, smithi and stoddardi.
Syllophopsis stat.r. восстановлен из синонимии с
Monomorium, to contain the Monomorium species in the
hildebrandti-group (sensu Heterick, 2006; this includes the
fossulatum-group of Bolton, 1987). Ireneidris is removed
from synonymy under Monomorium, and newly synonymized
(syn.n.) under Syllophopsis. This creates the following new
combinations in Syllophopsis: adiastolon, aureorugosa, australica,
dentata, ferodens, fisheri, gongromos, hildebrandti,
infusca, kondratieffi, malamixta, sechellensis, sersalata, subcoeca,
thrascolepta and vitiensis; and the following revived
combinations in Syllophopsis: arnoldi (= jonesi, unnecessary
replacement name), cryptobia, elgonensis and modesta.
Род Epelysidris stat.r. восстановлен из синонимии с Monomorium to contain the single known species
E. brocha (comb.r.). Its putative sister species ‘Monomorium’
latinode is very different morphologically and a new genus
will be erected for it in a forthcoming publication (B. Bolton,
personal communication).
Авторы поддерживают синонимизацию Carebarella under Solenopsis,
generating the following new combinations in Solenopsis:
alvarengai, condei. The type species of Carebarella (bicolor)
was already combined with Solenopsis by Pacheco & Mackay
(2013). Although they declined to propose new generic synonymy,
their new combination effectively did this, although it
left the other two names orphaned.
Anisopheidole синонимизирован (syn.n.) under Pheidole, generating
the following revived combination in Pheidole: antipodum.
Machomyrma синонимизирован (syn.n.) under Pheidole, generating
the following new combination in Pheidole: dispar.
Chalepoxenus синонимизирован (syn.n.) under Temnothorax,
generating the following new combinations in Temnothorax:
brunneus, hyrcanus, kutteri, muellerianus, spinosus, tarbinskii,
tauricus, tramieri and zabelini. Two of these new combinations
become secondary junior homonyms, T. spinosus (Arnoldi) and
T. tauricus (Radchenko), which are here replaced (nom.n.) with
T. sentosus and T. inquilinus, respectively.
Myrmoxenus синонимизирован (syn.n.) under Temnothorax,
generating the following new combinations in Temnothorax:
adlerzi, africana, algeriana, bernardi, birgitae, corsica, gordiagini,
kraussei, ravouxi, stumperi, tamarae, and zaleskyi. Two
of these new combinations, T. kraussei (Emery) and T. tamarae
(Arnoldi), create secondary junior homonyms among two
species already in Temnothorax, T. kraussei (Emery) and T.
tamarae (Radchenko), which are here replaced (nom. n.) with
T. mediterraneus and T. georgicus, respectively.
Protomognathus синонимизирован (syn. n.) under Temnothorax,
generating the following new combination in Temnothorax:
americanus.
Rhoptromyrmex синонимизирован (syn.n.) под Tetramorium,
generating the following new combinations (unless otherwise
noted) in Tetramorium: caritum, critchleyi, globulinodis, mayri,
melleum (comb.r.), opacum, rawlinsoni, schmitzi, transversinodis
and wroughtonii. One of these, T. mayri Forel, creates a
secondary junior homonym of a species already in Tetramorium,
T. mayri (Mann), which is here replaced (nom.n.) with
T. solomonensis. The new combination T. transversinodis
(Mayr) is a secondary senior homonym of an invalid name,
T. transversinodis (Enzmann), which therefore does not require
a replacement name.
Anergates синонимизирован (syn.n.) under Tetramorium, generating
the following new combination in Tetramorium: atratulum.
Teleutomyrmex синонимизирован (syn.n.) under Tetramorium,
generating the following new combinations in Tetramorium:
kutteri, schneideri. Tetramorium schneideri (Kutter) becomes a
secondary junior homonym, here replaced with T. inquilinum
(nom.n.).
Strongylognathus Mayr также синонимизирован с Tetramorium
Mayr, but it has priority over Tetramorium. Transfer of the hundreds
of valid Tetramorium species names to Strongylognathus,
with consequent changes in spelling to many of them, would
create considerable confusion. In this instance a ruling by the
International Commission on Zoological Nomenclature would
be required to validate a reversal of precedence (ICZN Article
23.9.3). In the interim we maintain current usage and refrain
from synonymizing the two genera.
Trichomyrmex stat.r. восстановлен из синонимии под
Monomorium, to contain the Monomorium species in the scabriceps-
and destructor-groups (sensu Bolton, 1987; see also
Radchenko, 1997). Holcomyrmex and Parholcomyrmex are
removed from synonymy under Monomorium, and newly synonymized
(syn.n.) under Trichomyrmex. These actions generate
the following new combinations in Trichomyrmex: aberrans,
abyssinicus, chobauti, chobauti ajjer, criniceps, destructor,
emeryi, emeryi laevior, epinotale, glaber, lameerei, mayri,
muticus, oscaris, perplexus, robustior, santschii (= santschianum,
unnecessary replacement name) and scabriceps; and
the following revived combinations in Trichomyrmex: rogeri
and wroughtoni (= wroughtonianum, unnecessary replacement
name).
ТРИБЫ.
Триба Myrmicini Lepeletier de Saint-Fargeau 1835
Родовой состав. Manica, Myrmica, †Plesiomyrmex and
†Protomyrmica.
Примечание. Eutetramorium, Huberia, Hylomyrma, Pogonomyrmex
и Secostruma исключены из Myrmicini.
Триба Pogonomyrmecini trib.n.
Родовой состав. Hylomyrma and Pogonomyrmex.
Триба Stenammini Ashmead 1905
= Aphaenogastrini Enzmann 1947 syn.n.
Родовой состав. Aphaenogaster, Goniomma, Oxyopomyrmex,
Messor, †Paraphaenogaster, Stenamma и Veromessor.
Примечание. Veromessor is resurrected from synonymy (above).
The reclassification of ‘Aphaenogaster’ awaits more detailed
phylogenetic analysis of lineage diversity within Stenammini
Триба Solenopsidini Forel 1893
= Myrmicariini Forel 1893 syn.n.
= Monomoriini Emery 1914
= Megalomyrmecini Dlussky & Fedoseeva 1988
= Stegomyrmecini Wheeler 1922 syn.n.
= Adelomyrmecini Fernández 2004 syn.n.
Родовой состав. Adelomyrmex, Anillomyrma, Austromorium,
Baracidris, Bariamyrma, *Bondroitia, Cryptomyrmex,
Dolopomyrmex, Epelysidris, Kempfidris, Megalomyrmex,
Monomorium, Myrmicaria, Oxyepoecus, Rogeria, Solenopsis,
Stegomyrmex, Syllophopsis, Tropidomyrmex и Tyrannomyrmex.
Примечание. Carebarella is considered a synonym of Solenopsis
(above). Monomorium remains a heterogeneous, nonmonophyletic
group. Из состава Solenopsidini исключены рода Adlerzia, Allomerus, Carebara, Diplomorium,
Mayriella и Tranopelta.
Триба Attini F. Smith 1858
= Pheidolini Emery 1877 (= Anergatidini Emery 1922) syn.n.
= Dacetini Forel 1892 (= Epopostrumina Brown 1952
= Orectognathina Brown 1952=Strumigenina Brown 1952)
syn. n.
= Ochetomyrmecini Emery 1914 syn.n.
= Cephalotini M. R. Smith 1949 (= Cryptoceridae F. Smith
1853) syn.n.
= Basicerotini Brown 1949 syn.n.
= Phalacromyrmecini Dlussky & Fedoseeva 1988 syn.n.
= Blepharidattini Wheeler & Wheeler 1991 syn.n.
= Lenomyrmecini Bolton 2003 syn.n.
Родовой состав. Acanthognathus, Acromyrmex,
Allomerus, Apterostigma, Atta, †Attaichnus, Basiceros,
Blepharidatta, Cephalotes, *Chimaeridris, Colobostruma,
*Cyatta, Cyphomyrmex, Daceton, Diaphoromyrma,
Epopostruma, Eurhopalothrix, *Ishakidris, Kalathomyrmex,
Lachnomyrmex, Lenomyrmex, Mesostruma, Microdaceton,
*Mycetagroicus, Mycetarotes, *Mycetophylax, Mycetosoritis,
Mycocepurus, Myrmicocrypta, Ochetomyrmex, Octostruma,
Orectognathus, *Paramycetophylax, Phalacromyrmex, Pheidole,
Pilotrochus, Procryptocerus, Protalaridris, *Pseudoatta,
Rhopalothrix, *Sericomyrmex, Strumigenys, Talaridris, Trachymyrmex,
Tranopelta и Wasmannia.
Триба Crematogastrini Forel 1893
=Formicoxenini Forel 1893 (= Cardiocondylini Emery
1914 = Leptothoracini Emery 1914 = Stereomyrmecini
Emery 1914 = Podomyrmini Emery 1924 = Solenomyrmini
Donisthorpe 1943) syn.n.
= Cataulacini Emery 1895 syn.n.
= Tetramoriini Emery 1895 (= Anergatini Emery 1914
= Teleutomyrmecini Kutter 1950) syn.n.
= Melissotarsini Emery 1901 syn.n.
= Myrmecinini Ashmead 1905 syn.n.
= Metaponini Forel 1911 syn.n.
= Lophomyrmecini Emery 1914 syn.n.
= Meranoplini Emery 1914 syn.n.
= Ocymyrmecini Emery 1914 syn.n.
= Pheidologetonini Emery 1914 syn.n.
= Proattini Forel 1917 syn.n.
= Hypopomyrmecina Brown 1952 syn.n.
= Calyptomyrmecini Dlussky & Fedoseeva 1988 syn.n.
= Liomyrmecini Bolton 2003 syn.n.
= Paratopulini Bolton 2003 syn.n.
Родовой состав. Acanthomyrmex, Adlerzia, Ancyridris,
Atopomyrmex, Calyptomyrmex, Cardiocondyla, Carebara,
Cataulacus, Crematogaster, Cyphoidris, Dacatria, Dacetinops,
Dicroaspis, Dilobocondyla, Diplomorium, †Enneamerus,
†Eocenomyrma, Eutetramorium, Formicoxenus, *Formosimyrma,
Gauromyrmex, *Goaligongidris, Harpagoxenus,
Huberia, †Hypopomyrmex, *Indomyrma, Kartidris,
*Lasiomyrma, Leptothorax, Liomyrmex, †Lonchomyrmex,
Lophomyrmex, Lordomyrma, Malagidris, Mayriella, Melissotarsus,
Meranoplus, Metapone, Myrmecina, Myrmisaraka,
Nesomyrmex, Ocymyrmex, †Oxyidris, †Parameranoplus,
Paratopula, Perissomyrmex, *Peronomyrmex, Podomyrma, Poecilomyrma,
Pristomyrmex, Proatta, Propodilobus, Recurvidris,
Rhopalomastix, Romblonella, *Rostromyrmex, Rotastruma,
Royidris, Secostruma, *Stereomyrmex, †Stigmomyrmex,
†Stiphromyrmex, Strongylognathus, Temnothorax, Terataner,
Tetheamyrma, Tetramorium, Trichomyrmex, Vitsika, Vollenhovia,
Vombisidris and Xenomyrmex.
Ископаемые таксоны без уточнения трибы
†Afromyrma, †Attopsis, †Bilobomyrma, †Boltonidris,
†Brachytarsites, †Cephalomyrmex, †Clavipetiola,
†Electromyrmex, †Eocenidris, †Eomyrmex, †Fallomyrma,
†Fushunomyrmex, †Ilemomyrmex, †Miosolenopsis,
†Myrmecites, †Orbigastrula, †Quadrulicapito,
†Quineangulicapito, †Sinomyrmex, †Solenopsites,
†Sphaerogasterites, †Wumyrmex and †Zhangidris.
Исключённые из состава Myrmicinae
Триба Ankylomyrmini Bolton и её единственный род Ankylomyrma
Bolton, перенесён в подсемейство Agroecomyrmecinae, т.к.
Ankylomyrma является сестринским к роду Tatuidris.
Myrmica, Formica, Lasius,
и Camponotus
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