Robert W. Taylor.
Taxonomy and Parataxonomy of Some Fossil Ants (Hymenoptera-Formicidae).
Psyche 71:134-141, 1964.

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TAXONOMY AND PARATAXONOMY OF
SOME FOSSIL ANTS
P HYMENOPTERA-FORMICIDAE)
BY ROBERT W. TAYLOR
Biological Laboratories, Harvard University In current revisionary studies of the ant tribe Ponerini it has become necessary to re-examine the status of various fossils previously placed in the genus Ponera. This taxon dates to 1804 and conse- quently has an unusually complex conceptual and nomenclatural his- tory. The included fossils require special treatment to unravel their part in the resulting snarl.
Thirty-six fossil ants have been placed as Ponera or Ponera-like by earlier authors but little confidence in the generic assignment of most of them is possible. Some are certainly ponerine, and occasional placement in tribe Ponerini is reasonable. Most species, however, cannot be satisfactorily placed, even to subfamily. The fact is that, to some authors, Ponera has served as a "catch-all" for small, possibly ponerine ant fossils, or wing impressions with venation similar to that of Ponera.
It is proposed here to review these species and to attempt their allocation into various categories : ( I ) Formicidae incertae generis; (2) Ponerinae incertae generis; (3) Ponera; (4) (?) Ponera; or
(5) the form-genus Poneropsis Heer, 1867 - as redefined below. The result of sorting the fossil "Ponera" in this way has, I believe, some utility relative to evolutionary studies. Species are either placed definitely or reasonably certainly in a known taxon. rendered 'incertae" at the level at which they begin to be uncertain in diag- nostic features; or allocated to the phylogenetically meaningless limbo of the parataxon Poneropsis. My category "( ?) Ponera'' in general contains species equally well placed in Ponera or Hypoponeraz, al- though smaller members of other genera of tribe Ponerini may be included.
'Based on research supported by the U.S. National Science Foundation, Grant No. GB 163%
'Santschi's subgenus Ponera (Hypoponera) (1938, Bull. Soc. Ent. France, 43: 8-80) has recently been elevated to full generic status (Taylor, mss.). It contains the majority of the living species currently assigned to Ponera, and many of its species are superficially Ponera-like, Manuscript received by the Editor May 29, 1964.



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Taylor - Fossil ants
The form-genus Poneropsis Heer.
In his study of the fossil Hymenoptera of Oeningen and Radoboj, Heer (1867) proposed the use of a formicid form-genus Poneropsis, which was defined as follows: ". . . Die fossilen Ameisen welche drei Cubitalzellen in den Oberflugeln und einen einknotigen Hinterleibs- stiel, aber keine Einschniirung beim zweiten Hinterleibssegment haben. Sie stimmen im Flugelgeader und dem einknotigen Stiel mit Ponera uberein, daher ich sie fruher dieser Gattung zugerechnet habe; in der Bildung des Hinterleibes weichen sie aber bedeutend von den Poneren ab, namentlich die Arten mit rundem, dickem Hinterleib." Heer's figures show that his "drei Cubitalzellen" are those now referred to as the first and second cubital cells, with the discoidal cell.
Sixteen species were allocated to Poneropsis at its inception, in- cluding some previously placed in Ponera by Heer ( I 849). No better placement of any of them is possible on the basis of the published data. There appears to be much species-level synonomy among these forms and judging from their size most do not seem close to Ponera. Since the venational type specified for Poneropsis is convergently developed in many lines of ant evolution, this '(genus'' could con- ceivably contain wing impressions of members of almost every ant subfamily3. Moreover the convergent types cannot be separated on the basis of wing venation alone. Accordingly it is pointless to assign such wings indiscriminately to recent taxa to whch they might, at present, be referable. It is far better to assign them definitely to a parataxonomic form-genus which need not be considered in phylo- genetic, paleo-zoogeographic, or other studies, rather than to place them randomly in a true taxonomic genus, with presumed affinities to other taxa, extinct or living.
It may be argued that this procedure offers little in comparison with a simple "Formicidae incertae generis" allocation. This is partly true, but since Heer's parataxon is available, use of it may as well be maintained, at least until a complete revision of fossil ants is possible. At that time the problem of the use of ant-wing form-genera will 'For example, all the following recent genera possess wing venation of the "Poneropsis" type: Gnamptogenys, Eciton, Pseudomyrmex, Messor, An- euretus, Dolichoderus, Hypoclinea (See figures of Brown and Nutting, 1950, and Wilson et.al., 1956). Extinct ants with this venation pattern include: Trachymesopus succinea (Mayr), Aphaenogaster mayri Carp., Pheidole ter- tiaria Carp., Dolichoderus antiquus Carp., Iridomyrmex florissantius Carp., Liometopum microcephalus Carp., and members of the genera Protazteca and Elaeomyrmex (see Wheeler, 1914 and Carpenter, 1930).



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136 Psyche [September
need careful consideration. We must consider the fact that Poneropsis, as defined here, contains wings all of which are at ap- proximately the same evolutionary grade of venational reduction (Brown and Nutting, ~gso), and that certain genera of ants can be excluded from it, as they never possess such venation. Under such terms we are actually designating fossils more precisely by placing them in Poneropsis rather than considering them simply as "Formi- cidae incertae generis". Moreover, and this is an important con- sideration, use of this parataxon allows convenient placement of such fossils in a single group easily referred to by those seeking examples of such venational types for other studies. I propose the following redefinition of Poneropsis. The nomencla- ture used for wing veins is that of Brown and Nutting (1950). Form-genus, Poneropsis Heer, I 867
Hyrnenopterous forewings, apparently belonging to family Formi- cidae, and either alone or attached to fossils otherwise unclassifiable, and of a type not known to be associated with remains yielding more satisfactory placement.
Two closed, fully separated, cubital cells (the 1st and 2nd) present. First discoidal cell always closed ; second discoidal open or closed. Ra- dial cell open or closed. The adventitious longitudinal vein Rsx, and the first radial cross vein (ir), or a stub of it, ab~ent.~ Second radial cross vein (2s) usually arising near the anterior base of the radio-medial cross vein (r-m), and always reaching the stigma at a point distal to the first quarter of its posterior b~rder.~ The sec- ond free abscissa of the median vein may be contracted, so that the posterior end of Rs + M2 lies adjacent to the anterior end of the (first) medio-cubital cross vein (m-cu) ; or fusion of elements in this area may cause the base of the former vein to lie distal to that of the latter. First abscissa of median vein (M~I) lying proximal, distal, or adjacent to the anterior base of the cubital anal cr'oss vein (cu-a) where it meets CUA.~
Specimens with a two-segmented petiole and Poneropsis-type wing "'Wings referable to primitive ponerines and myrmeciines such as Platy- thyrea, M~rmecia, and some Amblyoponini are, therefore, excluded, (Brown and Nutting, 1950; Brown, 1960).
'This clause allows distinction of Eoponera Carpenter (1929) - see Brown and Nutting, fig. 6.
"As Brown and Nutting point out, it is possible that origin of Mfl well proximal of cu-a is a key character identifying doryline ants. If this should prove to be so, the above diagnosis could be easily modified to preclude wings of fossil Dorylinae.




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19 641 Taylor - Fossil ants 137
venation must be placed in the Myrmicinae or one of the other ap- plicable subfamilies. If the node is one-segmented and other charac- ters of the gaster (presence of sting, etc.) are visible, then placement to subfamily should be possible.
The many qualifications made to the simple basic diagnosis, "two closed cubital cells, and a single closed discoidal," allow inclusion in Poneropsis of virtually all known ants with these primary characters. I do not wish to imply that study of wing vein patterns, such as was pioneered by Brown and Nutting, should not be applied to ant fossils. These authors have shown, however, that 'extreme parallelism may take place in the details of venational reduction in the various ant subfamilies, with the result that amazingly similar wings may be produced in divergent lines. The various ranges specified in my diagnosis simply cover all stages in venational reduction known to show such parallelism in wings with two cubital cells and at least one closed discoidal cell.
With the possible exception of the feature discussed in footnote 6 of the diagnosis, no' alternative condition in these venational charac- ters, or combination of conditions, is currently known to diagnose un- equivocally any ant taxon.
Ponera and Poneropsis species described by Heer (1849, 1867). In 1849, Heer described nine extinct species in Ponera from the Miocene of Radoboj, Oeningen and Parschlung, Croatia. In his 1867 paper four of these were referred to the newly defined form- genus Poneropsis, and thirteen further specific or infraspecific forms were also described, all in Poneropsis.
I have been unable to justify any of the generic assignments in Ponera, and find that most of Heer's species, both of Ponera and Poneropsis, can be assigned to Poneropsis as defined above, thus con- veniently disposing of them. Others, including some placed by Heer in Poneropsis, do not appear referable there on the basis of his figures, since the wing venation is too incompletely shown in the fossils or the wings appear to have had only a single cubital cell. The history and present status of Heer's (1849) Ponera species is summarized in the following Table. The two species considered here to be "Formicidae incertae generisJ' were based on remains too incomplete to allow better allocation.
Mayr (1867) and Popov (1932) have both referred to some of these species, assigning them with or without query to Ponera. Re- petition of Mayr's names serves no purpose; most of them were ori- ginally placed in Poneropsis (by Heer) and so Mayr's combinations do not constitute nomenclatural occupation in Ponera, since none of



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Psyche
[September
Species placed
in Ponera by
Heer I 849
affinIS
crassinervis
cr oatica
el0 ngatula
fuliginosa (with subspecies
oeningensis and radoboj)
globosa
longaeva
nitida
ventrosa
Species placed
in Ponero psis
by Heer 1867
afin is
elongatula
f uligin osa
-
nitida
Current
assignment
Formicidae incertae
generis
Poneropsis
Poneropsis
Formicidae incertae
generis
Poneropsis
Poneropsis
Poneropsis
Poneropsis
Poneropsis
them are now considered to belong in the genus. Popov's citations are important, however, as he used some of the names originally assigned to Ponera by Heer, thus firmly establishing them in modern systematic nomenclature. Those involved are croutica, crarsinervis (incorrectly spelled as crassicornis) , ventrosa, longaeva and globosa. All of the additional thirteen species described in Poneropsis in I 867 appear to be satisfactorily placed, except elongata, anthracina, imhoff, and siygia in which the wings are too incompletely preserved to allow allocation - they should be considered "Formicidae incertaf generis".
A further species, Ponera veneraria, was described by Heer in his Urwelt der S h e k ( 1865). This species was later transferred to Poneropsis in the 1879, second edition of the same work. On the basis of Heer's 1865 figure I concur with Handlirsch ( 1908) that this species is best placed as Formicidae incertae generis. The name was misspelled "vernaria" by Handlirsch. Fossil Ponera described by authors other than Heer. The following list, as far as I am aware, includes all ant fossils allocated to Ponera by authors other than Heer. This includes those which have since been placed elsewhere by previous authors, whose reassignments are discussed below with my own opinions on the proper placement of all the species listed here. The appropriate sef- erences may be obtained in the bibliography.



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19641 Taylor - Fossil ants 139
I. Ponera atavia Mayr, 1868: 72, figs. 66-69, female, male. Oli- gocene - Baltic Amber. Wheeler, 1914: 38, fig. 9. worker. 2. Ponera brodiei, Giebel, 1856: 173. This forewing fragment, originally described as an ant, Formicium brodiei, by Westwood (1854) has been subsequently placed in the Jurassic siricoid family Anaxyelidae (Maa, I 949).
3. Ponera gracilicornis Mayr, 1868: 72, worker, Baltic Amber. 4. Ponera hendersoni Cockerell, 1906, female. Miocene-Florissant. 5. Ponera hypolitha Cockerell, 1915: 483, plate 64, figs. 3-4, wing impression. Oligocene - Gurnet Bay, Isle of Wight. 6. Ponera ( ?) leptocephala Emery, I 89 I : 8, plate I, figs. 3, 4, female. Miocene - Sicilian Amber.
7. Ponera minuta Donisthorpe, 1920: 85, plate 5, fig. 4, male ( ?). Oligocene, Gurnet Bay, Isle of Wight. 8. Ponera rhenana Meunier, I 91 7, wing impression. Oligocene
- Bavaria.
9. Ponera scitula Clark,
1934, listed from Tertiary, Allendale,
Australia by Oke ( 1957).
10. Ponera succinea Mayr, 1868: 72, female. Oligocene- Baltic
Amber.
I I. Ponerah( ?) umbra Popov I 933 : I 7, fig. I, female. Miocene -
Kuban Caucasas.
Of these species only one, P. atavia Mayr, is considered here to be satisfactorily referred to Ponera. P. succinea Mayr was trans- ferred to Euponera (Trachymesopus) - now Trachymesopus - by Wheeler ( I 9 I 4), on grounds which are entirely acceptable. P. gra- cilicornis Mayr is too large to be considered a Ponera (Wheeler, 1914)~ but Mayr's assignment of the species to the Ponerinae is prob- ably dependable - the species is considered here as "Ponerinae in- certae generis". ( ?) P. leptocepfzala Emery is best assigned with reservation to Ponera. This form is evidently close to Ponera or Hy- popont7ra, but has very long legs and antennae, and the eyes appear to be placed exceptionally far back on the head. It may belong to a distinct genus as yet undiagnosed, but it would be premature to so assign it on the basis of Emery's description and figures. P. ( ?) umbra Popov also seems best assigned to (?) Ponera. It appears close to Ponera although it could equally well be a Hypoponera or a member of some other small genus of the tribe Ponerini. 1 propose the following NEW COMBINATIONS in Poneropsis: Poneropsis hypolitha ( Cockerell), and Poneropsis rhenana (Meu- nier), these are both wing impressions and cannot be assigned more satisfactorily at present. P. minuta is considered "Formicidae incertae



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Psyche
[September
generis" ; no reason whatsoever was presented by Donisthorpe to
justify its placement in Ponm, and no satisfactory diagnostic charac- ters are given in his figure or description. P. hendersoni Cockerel1 has been shown by Carpenter (1930) to be referable to the extinct genus Protazteca. The recent Australian species, Hypoponera scitula (Clark) (NEW COMBINATION from Ponera), was listed as a tertiary fossil from Allendale, Victoria, under the name Ponera scitula, by Oke ( 1957). I have not seen the specimens involved, but since they were determined by Clark, the assignment is presumably trustworthy.
Fossil names and their nomenclatural status. According to the principle of homonomy certain of the specific names given above are no longer available for use in Ponera. The eleven names assigned by Heer in 1849 (see list, p. 138), and the ten species, excluding brodei, assigned by subsequent authors and listed above on page 139 are in this category, as is the specific name veneraria Heer ( I 865).
BROWN, W. L., JR.
1960. Contributions toward a reclassification of the Formicidae. Ill tribe Amblyoponini. Bull. Mus. Comp. 2001. Harv., 122 (4) : 145-230.
BROWN, W. L., JR., and W. L. NUTTING
1950. Wing venation and the phylogeny of the Formicidae. Trans. Amer. Ent. SOC., 75: 113-132.
CARPENTER, F. M.
1929. A fossil ant from the lower Eocene (Wilcox) of Tennessee. J. Wash. Acad. Sci., 19: 300-301.
1930.
The fossil ants of North America. Bull. Mus. Comp. 2001. Harv., 70 (1) : 1-66, 11 pis.
COCKERELL, T. D. A.
1906.
A new fossil ant. Ent. News, Philad., 17: 27-28. 1915. British fossil insects. Proc. U.S. Nat. Mus., 49: 469-499, 6 pis. DONISTHORPE, H. ST. J. K.
1920.
British Oligocene ants. Ann. Mag. Nat. Hist., 6: 81-94, 1 pi. EMERY, C.
1891. Le formiche dell' ambra Siciliana nel museo mineralogico dell' universitadi Bologna. Mem. R. Ace. Bologna, 5 (1) : 141-165, 3 pis.
GIEBEL, C. G.
1856.
Insecten und Spinnen der Vorwelt,, Brodhaus, Leipzig., pp. 1-511. HANDLIRSCH, 0.
1908. Die Fossilen Insekten. W. Engelmann, Leipzig., pp. 1-1430.



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19641 Taylor - Fossil ants 141
HEER, 0.
1849. Die Insektenfauna der Tertiargebilde von Oeningen und von Radoboj in Croatien.
11. Neue Denkschr. Allgem. Schweiz. Geol. Ges. Naturw., 11: 1-264, 17 pis.,
(pp. 145-153).
1867.
Fossile Hymenoptera aus Oeningen und Radoboj. Ibid., 22 (4) : 1-42, 17 pis.
MAA, T.
1949.
A synopsis of Asiatic Siricoidea with notes on certain exotic and fossil forms. Notes dlEnt. Chinoise, Shanghai, 13 (2) : 11-189. MAYR, G. L.
- 1867. Vorlaufige Studien uber die Radoboj-formiciden. Jahrb. Geol. RchsAnst. Wien., 17: 47-62, 1 pi.
1868. Die Ameisen des baltischen Bernsteins. Beitr. Naturk. Preuss., 1: 1-102, 5 pis.
MEUNIER, F.
1917. Sur quelques insectes des lignites de 1'Aquitanien de Rott sept Montagnes (Preusse rhbnane). Verh. Akad. Wet. Amst., (2) 20 (I) : 1-17.
OKE, C.
1957. Fossil Insecta from Cainozoic Resin at Allendale, Victoria. Proc. R. Soc. Victoria. n. s., 69: 29-31.
POPOV, V.
1932. Two new fossil ants from Caucasus (Hymenoptera-Formicidae) . Trav. Inst. Paliozool. Acad. Sci. U.S.S.R., 2: 17-21. TAYLOR, R. W.
mss.
A monographic revision of the ant genus Ponera (Hymenoptera- Formicidae).
To appear in Bull. Mus. Comp. 2001. Harv. WESTWOOD, J. 0.
1854. Contributions to fossil entomology. Quart. J. Geol. Soc. Lond.. 10: 378-396.
WHEELER, W. M.
1914. The ants of the Baltic Amber. Schrift. Physick. Oken. GeseII. Konigsberg, 55: 1-142, 66 figs.
WILSON, E. O., and T. EISNER, 6. C. WHEELER, J. WHEELER 1956. Aneuretus simoni Emery, a major link in ant evolution. Bull. Mus. Comp. 2001. Harv.,
115 : 81-99, 3 pis.




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