Cataulacus intrudens

Cataulacus intrudens nests in twigs and branches of trees and shrubs. Arnold (1917:391) records it from hollow twigs of an acacia, Prins (1965:104) found it nesting in a branch of a red bush-willow, Combretum apiculatum.

Identification
A member of the intrudens group.

Distribution based on Regional Taxon Lists
Afrotropical Region: Angola, Ethiopia, Kenya, Malawi, Mozambique, Namibia, Somalia, South Africa, United Republic of Tanzania, Zimbabwe. Malagasy Region: Madagascar.

Abundance
The most common Cataulacus species in southern and eastern Africa.

Nomenclature

 *  intrudens. Meranoplus intrudens Smith, F. 1876d: 609, pl. 11, fig. 7 (w.q.m.) SOUTH AFRICA. Combination in Cataulacus: Mayr, 1886c: 364. Senior synonym of batonga, baumi, densipunctatus, foveosquamosus, gazanus, hararicus, intermedius, johannae, pseudotrema, rugosus, subrugosus, umbilicatus and material of the unavailable names krugeri, tangana referred here: Bolton, 1974a: 43; of bulawayensis: Bolton, 1995b: 138.
 * hararicus. Cataulacus hararicus Forel, 1894b: 79 (w.) ETHIOPIA. Junior synonym of intrudens: Bolton, 1974a: 43.
 * rugosus. Cataulacus intrudens var. rugosus Forel, 1894b: 78 (w.) MOZAMBIQUE. Mayr, 1895: 129 (q.). Raised to species: Mayr, 1895: 129; Prins, 1963: 104. Junior synonym of intrudens: Bolton, 1974a: 43.
 * johannae. Cataulacus johannae Forel, 1895c: 250 (w.q.) MADAGASCAR. Junior synonym of intrudens: Bolton, 1974a: 43.
 * baumi. Cataulacus baumi Forel, 1901d: 304 (w.q.m.) ANGOLA. [Also described as new by Forel, 1903e: 560.] Junior synonym of intrudens: Bolton, 1974a: 43.
 * batonga. Cataulacus baumi r. batonga Forel, 1913a: 114 (w.) ZIMBABWE. Junior synonym of intrudens: Bolton, 1974a: 43.
 * subrugosus. Cataulacus rugosus var. subrugosus Santschi, 1914e: 26 (w.) SOUTH AFRICA. Santschi, 1937d: 236 (q.). Junior synonym of intrudens: Bolton, 1974a: 43.
 * bulawayensis. Cataulacus baumi var. bulawayensis Arnold, 1917: 391 (w.q.) ZIMBABWE. [First available use of Cataulacus baumi r. batonga var. bulawayensis Forel, 1914d: 218; unavailable name (Bolton, 1974a: 43).] Junior synonym of intrudens: Bolton, 1995b: 137.
 * intermedius. Cataulacus intrudens st. intermedius Santschi, 1917b: 287 (w.) ZIMBABWE. Junior synonym of intrudens: Bolton, 1974a: 43.
 * densipunctatus. Cataulacus johannae r. densipunctatus Stitz, 1923: 163 (w.) NAMIBIA. Junior synonym of intrudens: Bolton, 1974a: 43.
 * pseudotrema. Cataulacus baumi st. pseudotrema Santschi, 1926b: 244 (w.) TANZANIA. Junior synonym of intrudens: Bolton, 1974a: 43.
 * gazanus. Cataulacus baumi var. gazanus Santschi, 1928f: 208 (w.) MOZAMBIQUE. Junior synonym of intrudens: Bolton, 1974a: 43.
 * foveosquamosus. Cataulacus foveosquamosus Santschi, 1937a: 58, figs. 8, 9 (q.) SOUTH AFRICA. Junior synonym of intrudens: Bolton, 1974a: 43.
 * umbilicatus. Cataulacus umbilicatus Santschi, 1937a: 59, figs. 10-12 (q.) MOZAMBIQUE. Junior synonym of intrudens: Bolton, 1974a: 43.

Bolton (1974) - The tremendous variation of this species accounts for the many synonymized names. The majority of these forms were described from variations in head shape or sculpturation, some from even more minor details such as differences in propodeal spine length or in pronotal denticulation. Large workers with female-shaped heads and foveolate sculpturation were responsible for baumi and its subspecies and varieties whilst variation in size and sculpturation account for rugosus and johannae with coarse sculpture on the one hand, and subrugosus and intermedius with finer sculpture on the other. Many of the forms were stated in their original descriptions to be close to Cataulacus intrudens or stated as links between two or more forms. For example, both intermedius and subrugosus were given as intermediates between rugosus and intrudens. As this species is the most common of the genus in southern and eastern Africa it is probably the one which Arnold (1917) had in mind when he wrote that, 'many of the so-called species and races are very closely allied, so much so that I believe a study based on more abundant material will later on serve to reduce the present number of species to a much smaller figure'. In fact, when part 4 of his monograph of South African ants was published, Arnold (1920:403) went so far as to state that there were probably only the nuclei of two species in the area, one of which included baumi, batonga, bulawayensis and intermedius. This observation is now known to be accurate. Santschi (1937a) described the female of the species twice, from two separate localities, basing his descriptions to a large extent upon the very characters which had proved so misleading in the past.

The same mistake was made by Prins who, as late as 1965 described a variety krugeri, differentiating it from rugosus by its, 'slightly longer and more acute epinotal spines and by the sculpture of the abdomen which is less developed'. These are the trifling and intrinsically variable characters so much used by earlier authors and which are responsible for the great proliferation of valueless names in the more variable species of this genus.

Worker
Bolton (1974) - TL 4.3 – 5.1, HL 1.04 – 1.48, HW 1.14 – 1.56, CI 103 - 110, EL 0.42 – 0.54, OI 34 - 40, IOD 0.86 – 1.16, SL 0.52 – 0.68, SI 43 - 47, PW 0.94 – 1.30, AL 1.16 – 1.62, MTL 0.54 – 0.82 (15 measured).

Occipital crest absent, vertex and occiput meeting through an angle, not normally confluent in a curved surface. Occipital corners dentate, with a second tooth or denticle internally upon the margin; often also with a few minute denticles on the angle separating vertex and occiput. Sides of head behind eyes usually denticulate, in a majority of cases noticeably so but sometimes the denticles minute, sometimes reducing in size from occipital corner to eye. Rarely denticles are completely absent. When this occurs the workers generally have a female head-shape. Shape of head very variable. On the one hand is a form which may be regarded as extreme worker-shape, and on the other as extreme female-shape, the latter approaching or the same as that found in most queens of this species. Numerous intermediate forms are known and on occasion both extreme forms may be found in the same nest, along with intermediates. The 'worker-shaped' head usually occurs in smaller individuals, more rarely in large, and is characterized by strong denticulation of the sides behind the eyes, strong anterior convergence of the frontal carinae and convexity of the sides of the head behind the eyes. The 'female-shaped' head always occurs in large individuals and shows reduced or absent denticulation of the sides behind the eyes (variable between workers from the same nest series), a reduced tendency for convergence of the frontal carinae anteriorly and a marked lack of convexity in the sides behind the eyes, so that in some individuals these are almost parallel. Alitrunk marginate laterally, denticulate, the denticles variable in size amongst workers, especially upon the pronotal margins. Propodeum with a pair of spines of variable configuration, but which are usually quite broad and somewhat flattened dorsoventrally. Dorsal alitrunk without sutures. Petiole in profile with a steeply sloping anterior face and a usually less steep posterior face, the latter sometimes shallowly concave. These two faces meeting in an acute angle dorsally and occasionally forming a weak ridge at their junction, the petiole without a developed dorsal surface. Postpetiole with a narrow, rounded dorsal surface separating the anterior and posterior faces. Subpetiolar process simple, usually with the antero- and posteroventral angles blunt, more rarely acute but never with the latter extended as a heel or spur. Subpostpetiolar process small and simple or virtually absent. First gastral tergite not marginate laterally.

Sculpturation extremely variable. The most common form of sculpturation consists of a dense rugoreticulum upon the head and pronotal dorsum with reticulate-punctate interspaces and usually with the reticular meshes shagreened. The rugulae usually tend to assume a more longitudinal direction upon the mesonotum and are often reduced or absent in the middle of this sclerite, being replaced wholly or in part by a fine, dense reticulate-puncturation. Propodeal dorsum usually quite strongly longitudinally rugose with few or no cross-meshes; these rugae noticeably more coarse than those upon the pro- or mesonotum. Propodeal declivity with transverse rugae between the spines. In some specimens the cross-meshes of the cephalic rugoreticulum are very reduced, so that the sculpturation is predominantly longitudinal. Variation from this common sculpturation usually occurs by reduction or intensification of one or more of the components. The most coarsely sculptured forms have the dorsal alitrunk and to a lesser extent the head covered with very coarse, strong, predominantly longitudinal rugae. The most weakly sculptured have the head shagreened with scattered foveolae, and the major portion of the dorsal alitrunk similarly sculptured. Traces of rugation are usually maintained on the propodeal dorsum. In large workers the cephalic sculpturation is often somewhat modified. Many of the rugulae are expanded and flattened, obliterating the smaller interspaces. This results in the appearance of coarse foveolae set in a shagreened surface. The pedicel segments usually show longitudinal rugae but some may be irregularly rugose. First gastral tergite varying from finely reticulate-punctate throughout to coarsely longitudinally rugose with reticulate-punctate interspaces.

Erect hairs present on all dorsal surfaces of head, body and appendages but very short and inconspicuous upon the clypeus and cephalic dorsum, and also on the pronotum, where they are usually reduced to very short, stout stubs or even stud-like vestiges.

Queen
Bolton (1974) - TL 6.2 – 7.4, HL 1.20 – 1.60, HW 1.24 – 1.66, CI 96 - 103, EL 0.46 – 0.60, OI 35 - 39, IOD 1.00 - 1.30, SL 0.60 - 0.76, SI 46 - 48, PW 1.20 - 1.50, AL 1.70 – 2.10, MTL 0.66 – 0.86 (5 measured).

Head shape as described above for large female-like workers, very rarely shaped similarly to the typical head form of smaller workers. Denticulation of the sides of the head behind the eyes reduced or absent, as is the denticulation of the alitrunk margins. Propodeal spines reduced, proportionately broader and shorter than in the worker. Sculptural variation as described above.

Male
Bolton (1974) - TL 5.3 -5.7, HL 1.12- 1.20, HW 1.22 – 1.28, CI 106 - 109, EL 0.44, OI 34 - 36, IOD 0.94 – 0.96, SL 0.54 - 0.56, SI 43 - 44, PW 1.06 – 1.08, AL 1.74 – 1.82, MTL 0.68 – 0.72 (3 measured).

Head shape similar to small worker, the sides of the head behind the eyes dentate. Pronotum marginate laterally but not denticulate, similarly with the propodeal margins. Propodeal spines short, blunt and very stout. Parapsidal furrows distinct, the notauli acutely V-shaped rather than Y -shaped, with the posterior portion distinct to the margin of the scutellum. Dorsum of head finely and very densely reticulate-rugose, the longitudinal constituents predominating and more emphasized than the transverse; the interspaces reticulate-punctate. Pronotum similarly but more loosely sculptured, the mesoscutum with only a few weak rugulae, predominantly reticulate-punctate. Scutellum and propodeal dorsum with a rather coarse rugoreticulum. Pedicel longitudinally rugose, gaster finely and densely reticulate-punctate. All dorsal surfaces of head, body and appendages with numerous hairs.

Type Material
Bolton (1974) - LECTOTYPE worker, SOUTH AFRICA: Natal, Durban, Weenen District, in acacia thorns (J. M. Hutchinson) (BMNH), here designated [examined].

During the search for the types of this species a single specimen (a male) was found at UM, Oxford which was marked 'type'. However, the date of collection upon the labels was 1879, proving that it was not of the type series as the species was described in 1876.

Amongst the numerous unnamed specimens of the F. Smith collection in the BMNH were five specimens of intrudens (3 workers, 2 males) each bearing only a single small data label, inscribed '76, 48. Natal'. This reference was checked against volume 4 of the British Museum Zoological Accessions (1864 1881), and on page 214, under '1876 no. 48' was the information: "Meranoplus intrudens. Natal. Presented by F. Smith. This species was found by Mr J. Monkhouse Hutchinson inhabiting thorns of a species of Acacia in the Weenen District, Natal." This is exactly the information given by Smith in the original description of the species and it is concluded that these five specimens represent the type-series.

These have presently been designated lectotype (worker) and paralectotypes (2 workers, 2 males) and will be deposited in BMNH and MCZ, Boston. There is no trace of the female described by Smith, and this is presumed lost or destroyed.

References based on Global Ant Biodiversity Informatics

 * Arnold G. 1917. A monograph of the Formicidae of South Africa. Part III. Myrmicinae. Annals of the South African Museum. 14: 271-402.
 * Bolton B. 1974. A revision of the Palaeotropical arboreal ant genus Cataulacus F. Smith (Hymenoptera: Formicidae). Bulletin of the British Museum (Natural History). Entomology 30: 1-105.
 * Bolton B. 1982. Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax, Melissotarsus, Messor and Cataulacus (Formicidae). Bulletin of the British Museum (Natural History). Entomology 45: 307-370.
 * Fisher B. L. 1997. Biogeography and ecology of the ant fauna of Madagascar (Hymenoptera: Formicidae). Journal of Natural History 31: 269-302.
 * Fisher B. L. 2003. Formicidae, ants. Pp. 811-819 in: Goodman, S. M.; Benstead, J. P. (eds.) 2003. The natural history of Madagascar. Chicago: University of Chicago Press, xxi + 1709 pp.
 * Forel A. 1903. Einige neue Ameisen aus Sud-Angola. Pp 559-564, in: Baum, H. Kunene-Sambesi-Expedition, 1903. Berlin: Verlag des Kolonial-Wirtschaftlichen Komitees, 593pp.
 * Garcia F.H., Wiesel E. and Fischer G. 2013.The Ants of Kenya (Hymenoptera: Formicidae)Faunal Overview, First Species Checklist, Bibliography, Accounts for All Genera, and Discussion on Taxonomy and Zoogeography. Journal of East African Natural History, 101(2): 127-222
 * IZIKO South Africa Museum Collection
 * Menozzi C. 1930. Formiche della Somalia italiana meridionale. Memorie della Società Entomologica Italiana. 9: 76-130.
 * Menozzi C., M. Consani. 1952. Missione biologica Sagan-Omo diretta dal Prof. E. Zavattari. Hymenoptera Formicidae. Rivista di Biologia Coloniale 11: 57-71.
 * Prins A. J. 1963. A list of the ants collected in the Kruger National Park with notes on their distribution. Koedoe 6: 91-108.
 * Prins A. J. 1964. Revised list of the ants collected in the Kruger National Park. Koedoe 7: 77-93.
 * Prins A. J. 1967. The ants of our National Parks. Koedoe - African Protected Area Conservation and Science 10(1): 63-81.
 * Prins A. J., and J. J. Cillie. 1968. The ants collected in the Hluhluwe and Umfolozi game reserves. The Lammergeyer 8: 40-47.
 * Santschi F. 1914. Meddelanden från Göteborgs Musei Zoologiska Afdelning. 3. Fourmis du Natal et du Zoulouland récoltées par le Dr. I. Trägårdh. Göteborgs Kungliga Vetenskaps och Vitterhets Samhälles Handlingar. 15: 1-44.
 * Santschi F. 1937. Résultats de la Mission scientifique suisse en Angola (2me voyage) 1932-1933. Fourmis angolaises. Revue Suisse de Zoologie. 44: 211-250.
 * Santschi, F.. "Résultats de la Mission scientifique suisse en Angola, 1928-1929. Formicides de l'Angola." Revue Suisse de Zoologie 37 (1930): 53-81.
 * Viehmeyer H. 1923. Wissenschaftliche Ergebnisse der mit Unterstützung der Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommenen zoologischen Expedition nach dem anglo-ägyptischen Sudan (Kordofan) 1914. VII. Hymenoptera A. Formicidae. Denkschriften der Akademie der Wissenschaften in Wien. Mathematisch-Naturwissenschaftliche Klasse 98: 83-94.
 * Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. IX. A synonymic list of the ants of the Malagasy region. Bulletin of the American Museum of Natural History 45: 1005-1055
 * Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VIII. A synonymic list of the ants of the Ethiopian region. Bulletin of the American Museum of Natural History 45: 711-1004