Contributions toward a reclassification of the Formicidae. V. Ponerinae, tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini.
Author
Brown, W. L.
text
Search: Agriculture; Cornell University Agricultural Experiment Station
1975
15
1
115
http://antbase.org/ants/publications/6751/6751.pdf
journal article
6751
45422C7B-83F2-4F5A-9EE4-74C51F2C2BFE
Sphinctomyrmex
>
Sphinctomyrmex
Mayr, 1866: 895, [[ queen ]]. Type:
Sphinctomyrmex stali
, 1866, monobasic. ----- Borgmeier, 1957: 104, [[ worker ]] ', discussion of
S. stali
.
[[ ... ]]
Cerapachys
: Forel, 1893: 461, not F. Smith.
> Eusphinctus Emery, 1893, Ann. Soc. Entomol. France, 61 (1892): cclxxv, [[ worker ]]. Type:
Eusphinctus furcatus
, 1893, by original designation, monobasic. ----- 1895 a: pl. 14, fig. 4, [[ male ]]: genitalia.
>
Sphinctomyrmex
subgenus
Eusphinctus
: Emery, 1894 b: 457. ----- 1911: 7, [[ worker ]] [[ queen ]], species list.
>
Eusphinctus
: Wheeler, 1918: 219, 224, review of Australian spp. with key.
>
Eusphinctus
subgenus
Eusphinctus
: Wheeler, 1918: 219, diagnosis; 224 - 231, review of Australian spp. with key.
>
Eusphinctus
subgenus
Zasphinctus
Wheeler, 1918: 219, 233. Type:
Eusphinctus (Zasphinctus) turneri
Forel, monobasic.
>
Euphinctus
subgenus
Nothosphinctus
Wheeler, 1918: 219, 225, 233 - 239, diagnosis and review of Australian spp. with key. Type:
Sphinctomyrmex froggatti Forel
, by designation of Donisthorpe, 1943, Ann. Mag. Natur. Hist., (11) 10: 675.
>
Aethiopopone
Santschi, 1930: 49, a. Type:
Sphinctomyrmex rufiventris Santschi
, monobasic.
Worker: With characters of
Cerapachys
, but gastric segments IV, V, and VI separated by distinct constrictions and (except for
S. furcatus
and
S. taylori
) nearly equal in length. Eyes reduced to very small size or altogether lacking (except in s.
turneri
, which has fairly large, convex eyes). Antennae with 11 or 12 segments; palpi usually segmented 3,3. Sculpture of round piligerous foveolae, sparse or crowded, with smooth or finely roughened interspaces. Hairs short, simple, sparse to fairly dense; pubescence sometimes present on petiole, postpetiole, or succeeding gastric segments. Color black, brown, red, or yellow.
Queen: A morphocline ranges from perfect winged (or dealate) females with large eyes, ocelli, etc., to blind subdichthadiiform queens that are workerlike except for larger size and broader head and petiole, and sometimes vestigial eyes and 1 ocellus. Intermediates exist in, e. g.,
S. steinheili
and
S. asper
; these may have compound eyes of modest size and 3 ocelli, but workerlike trunk. It seems likely that the species with subdichthadiiform queens have monogynous colonies, while at least some of the dealate or intermediate forms are found several (to more than 20) to a single colony. It is not known whether the multiple queens are all functional reproductives.
Male: Differs from
Cerapachys
male by the distinctly constricted subequal segments of the gaster, and even here a difficulty exists because some American
Cerapachys
have narrowed bases to the main gastric segments. The male of
S. furcatus
, like its worker, has the gastric segments unequal, with the first much the largest. Antennal segments 13, rarely (
S. furcatus
) 12.
Aside from s.
furcatus
, males of
Sphinctomyrmex
divide into 2 classes. The first class, associated in a few cases with workers and / or queens (
S. steinheili
,
S. turneri
, S
. asper
) has slender males with triangular mandibles, and usually distinct notauli. The second class, consisting of males taken at light, probably belongs with the species having subdichthadiiform queens (froggatti group); these have long, tapered, falcate mandibles and lack distinct notauli; they tend to be larger and relatively robust, and have denser, softer, more regularly arranged pilosity, particularly on head and mandibles (figs. 97, 98); see also fuller discussion at [67].
The genus
Sphinctomyrmex
was first described by Mayr (1866: 895) from a single dealate queen with 12 - segmented antennae (
S. stali
) from southeastern Brasil. Although at least one winged queen of
Sphinctomyrmex
had already been collected in West Africa in the last century [70], the genus was not to be reported in print from that continent again until Santschi's record of 1915. Forel described forms from Australia (as
Cerapachys
) in 1893 and 1895, and in 1895 Emery introduced the new generic name
Eusphinctus
for a worker from Burma with 11 - segmented antennae that he named
E. furcatus
. New
Eusphinctus
species were added from New Guinea by Emery (1897) and from India and Australia by Forel, who both saw that workers of some species had 11, others 12, antennal segments, and therefore regarded
Eusphinctus
only as a subgenus of
Sphinctomyrmex
. Andre (1905) noted that the Indo-Australian species known at the time differed from the lone neotropical species,
S. stali
, in that they had ergatoid queens, whereas the
S. stali
queen had well-developed thoracic sclerites and had obviously once borne wings. On this basis, he recommended that
Sphinctomyrmex
and
Eusphinctus
be recognized as separate genera. Emery, in his 1911 ponerine fascicle of the Genera Insectorum, kept
Eusphinctus
at subgeneric level. In 1918, however, Wheeler not only embraced Andre's generic separation, but went on to subdivide
Eusphinctus
into 3 subgenera:
1.
Eusphinctus
s. str. Workers and females with 11 - jointed antennae, with entire or emarginate pygidium, the workers blind, the females with eyes and ocelli. Habits hypogaeic.
2.
Nothosphinctus
subgen. nov.
Workers and females with 12 - jointed antennae; the former blind and with entire pygidium, the latter with emarginate pygidium and either blind or with very minute eyes and the anterior ocellus. Habits hypogaeic.
3.
Zasphinctus
subgen. nov.
Workers large, dark colored, with 12 - jointed antennae and well developed eyes, but without ocelli. Females unknown. Habits probably epigaeic.
Meanwhile, Santschi (1915) had described a species of Sphinctomyrmex from males taken at light in West Africa. Wheeler (1918) dismissed Santschi's generic assignment as " open to doubt. " But when Santschi read Wheeler's 1918 revision, he displayed his carefree notion of generic-level taxonomy by making
S. rufiventris
the type of a new genus,
Aethiopopone
, and justified his action as follows (Santschi En 1915, quand je decrivis cette espece en la rapportant au genre
Sphinctomyrmex
, l'habitat de celui-ci etait considere comme etant le Bresil, l'Australie et l'Indie, et le S etait encore inconnu. Or, le [[ male ]] de
S. rufiventris
avec ses caracteres de Prodorylinas, surtout ses segments abdominaux etrangles, son habitat intermediaire, me fit risquer une identification pareille a celle que firent Forel et Emery en rapportant a ce genre les
Eusphinctus
du vieux monde. Ce n'est du reste qu'en 1918 que Wheeler fit la distinction generique, ne considerant plus que comme
Sphinctomyrmex
la [[ queen ]]
S. stali Mayr
du Bresil. En elevant au rang de genre le sous-genre
Eusphinctus
Em. pour les especes indo-australiennes, il le divise en trois sous-genres:
Eusphinctus
,
Notosphinctus
et
Zasphinctus
. Cela etant donne, je me vois oblige de creer un nouveau genre pour l'espece africaine. Les caracteres de celle-ci tiennent das G.
Sphinctomyrmex
,
Cerapachys
et
Simopone
.
This house of cards should have been fluttered away in 1923 by Clark when he described
Eusphinctus occidentalis
from Western Australia. This species turned out to have a dealate queen with 11 - segmented antennae associated with blind workers much like those of the eastern Australian
E. steinheili
and
E. duchaussoyi
.
Further discoveries were the worker caste of
S. stali (Borgmeier, 1957)
[69] and the winged
Sphinctomyrmex
queen collected long ago by Afzelius in West Africa [70], already mentioned above, and also workers of
Sphinctomyrmex
dug out of the Ivorian savanna by Jean Levieux [70]. Now another species has been found by Gotwald in rotting wood in Gabon (unpublished).
It seems from a survey of all the old and new material that we are dealing with a single genus in which the functional queens vary, according to species, from " normal " winged (when virgin) to ergatoid or even dichthadiiform types. This arrangement has already been tacitly accepted by Wilson (1957), and it would be well to deal with the diagnostic characters (Wheeler, 1918: 219) one at a time.
Specialization of queen. The most primitive kind of queen (Africa, Brasil, Western Australia) has wings that are lost, presumably in the usual fashion, before the colony is founded, or at least before the queen joins an established colony. Queens of species such as
S. steinheili
,
S. asper
, and
S. duchaussoyi
are wingless and ergatoid in thoracic structure, but have 3 distinct ocelli and a pair of compound eyes. In
imbecilis
, the queen has undergone further specialization; the head is broadened and its sides rounded, the ocelli are reduced to one, and the compound eyes are reduced to minute vestiges; this ant is clearly on its way towards becoming a dichthadiiform, and it is important to note that the development of wings and pterothorax is not the only criterion on which the queens may be sorted.
Habits " hypogaeic " vs. " probably epigaeic. " Wheeler's use of this ethological character, even as stated, is obviously speculative. The meager evidence available fails to support his classification. I found
S. steinheili
travelling aboveground in full daylight during a raid on a nest of
Stigmacros
in Victoria, Australia, although the raiders used cracks in the soil for some parts of their trail (Wilson, 1958: 136). On the other hand, I collected a worker of
S. turneri
foraging under leaves in dark rain forest in northern Queensland. At the very least, the distinction between these two species and their respective subgenera on hypogaeic vs. epigaeic foraging habits must be considered as blurred. In fact, Wheeler's classification is based on the presence or absence of worker eyes, a trait that may no more reflect foraging habits than it does in the true army ants. By the same token, it is possible, even probable, that some of the species with blind workers really are hypogaeic in foraging habits.
Antennae 12 - merous vs. 11 - merous in worker and queen. This character was weak to begin with, because the workers of forms with winged / dealate queens can have either 12 or 11 segments,
S. furcatus
, though not very close to the Australian "
Eusphinctus
s. str., " has 11 segments in worker and queen. Now we have
S. asper
, which is very close to the Australian Eusphinctus s. str. but has 12 antennal segments instead of 11. As in
Cerapachys
, antennal segment number apparently has undergone independent reductions that make it a poor group character.
Pygidium notched vs. entire. Wheeler (1918: 219) noted that: the worker of
E. cribratus Emery
of New Guinea has an entire pygidium and belongs with the Australian species in Eusphinctus s. str., but the workers of the Indian species
E. furcatus Emery
and
taylori Forel
have a notched pygidium. Should future investigation show that the pygidial characters are correlated with other peculiarities or with different types of female, it may be advisable to restrict the subgenus
Eusphinctus
to the two Indian species and to suggest a new subgeneric name for the Australian and Papuan forms with 11 - jointed antennae. The emarginate pygidium itself is not a very impressive group character, since it is partly linked to size. "
Zasphinctus
"
turneri
has it in the worker, and "
Nothosphinctus
" queens have it, while workers do not. Also, emargination occurs to different degrees among the species that have it, and even the angle of view affects its distinctiveness.
The Indian species
furcatus
[68] has now been found to possess an ergatoid female with eyes modestly larger than the minute ones of its worker, although its overall body size hardly differs from that of the worker in the same colony. The differences in the proportions of the gastric segments between
furcatus
(fig. 100) and other
Sphinctomyrmex
seem not to have attracted much notice, though they weaken the separation between
Sphinctomyrmex
and
Cerapachys
. Now that we have the presumed males of
furcatus
, we note another difference in that this specimen has only 12 antennal segments, whereas all other known cerapachyine males have 13. The
furcatus
male also has the gastric segments proportioned about the same as in the worker, and its terminalia are fairly distinctive (although terminalia for related species are poorly known).
Forgetting about the emarginate pygidium, then, we might still make a fair case for separating
furcatus
and
taylori
into a genus
Eusphinctus
apart from both Sphinctomyrmex and
Cerapachys
, but it seems unwise to do this in a group so poorly known as
Sphinctomyrmex
, in which males have yet to be matched up to workers or queens in most species, and in which many species doubtless remain undescribed from any caste. It seems that problems in this group will best be solved by rearing live colonies to get the males in association with the female castes.
bionomics: Discussed previously under the tribe.
distribution:
Sphinctomyrmex
is best represented in numbers of species, and is decidedly more common, in Australia than in other parts of the world, but the genus also occurs in New Guinea, New Caledonia, and southern Asia.
At least 2, and probably more, species exist in west and central Africa, where the 2 were originally described from males, and 2 distinctly different, undescribed species are known in the worker caste, plus 1 undescribed form known from a winged queen. Elsewhere,
Sphinctomyrmex
is represented only by a single rare species from southeastern Brasil.