The Ants of Egypt SUBFAMILY MYRMICINAE - Genus Pheidole Westwood in the Mediterranean area and the Middle East - taxonomic history

Pheidole main page - Genus Pheidole Westwood in the Mediterranean area and the Middle East - References

SUBFAMILY MYRMICINAE - Genus Pheidole Westwood in the Mediterranean area and the Middle East - taxonomic history

The work of Gustav L. Mayr, starting with his seminal publication on the Ants of Austria (1855), laid much of the foundations of modern ant taxonomy. For the first time, clear and detailed descriptions were written and the dichotomous key was adopted. For example, whereas Nylander (1849: 42) had described Pheidole pallidula (as Myrmica pallidula) in about 12 lines of Latin, Mayr (1853: 145) unknowingly re-described the species as Oecophthora subdentata in a way that would satisfy most modern taxonomists. Mayr (1855: 455) himself recognised O. subdentata as a junior synonym of pallidula and subsequently (Mayr, 1861: 70) accepted Pheidole Westwood (1839) as the senior genus . For most of his long career, in which he was a school master of some distinction and, it appears, never a paid taxonomist, Mayr primarily described new species, with occasional later publications (the last being in 1907) including subspecies.

It was the influence of Auguste Forel, from almost the start of his taxonomic writings, that led to the practice of using trinomials with descriptions of subspecies and beyond to variations (Forel, 1874). This was taken to extremes in the otherwise tremendous works of Felix Santschi (see Wehner, 1990: 304). It is some of the resulting obfuscation and confusion that this paper seeks to rectify. The opposite path follows the dogma of Wilson & Brown (1953) who stated "the subspecies concept is the most critical and disorderly area of modern systematic theory" arguing the subspecies trinomial is "both illusory and superfluous". It seems clear, however, that Wilson & Brown, who wrote of subspecies "as a formal means of registering geographical variation within the species", did not go back to Forel's argument. The latter appears to have been disturbed at the vast number of species being named, as by Mayr, who Forel praises and almost damns in the same breath. Thus, Forel adopted sous-espèces or confusingly races where he felt transitions between species were unclear. Quite what Forel meant is itself unclear.

What happens today, at least within the ranks of ant taxonomists, is that synonymization of old subspecies, etc., often appears to be adopted as an absolute and, yet, minute differences are accepted as meriting the naming of new species. The Wilson & Brown school, however, perhaps neglects the need of ecologists, behavioural scientists, and others trying to sort collections of ostensibly similar specimens. A recent and informative review of the wider significance of subspecies, together with a summary of the biological species concept versus the phylogenetic species concept, is that by Garnett & Chistidis (2007).

It is ironic that the taxonomic works of Brown and even perhaps Wilson's recent publication on Pheidole (Wilson, 2003), do not always provide truly precise and clear descriptions. Other common failings are a lack of geographical information and inadequate labelling of specimens shown in drawings and photographs.

From a wide study of around 500 species of ant from Africa, I have observed how widely distributed species, for instance P. megacephala, commonly are very consistent in shape and form and vary most in the intensity or shade of the base colour. Beyond that the use of the term "plastic species" or terms such as "very variable" are as misleading as the Forel cult of bestowing infraspecific names.

Historical records of North African Pheidole

Fabricius (1793) described Formica megacephala from the "Isle de France", Mus. Dom. Bosc. Latreille (1802c) gave a slightly longer description and illustrations of a major worker and a queen. There is no mention of a minor worker in either Fabricius or Latreille. Losana (1834) described Myrmica megacephala with reference to "la megacephala di Latreille" but noted that although the specimens, from his own garden in Piedmont, looked similar to those in the Latreille drawings the description was not sufficient to be sure he (Losana) had the same species. From Losana's details it is clear he was describing P. pallidula, for example the ants were lion-coloured.

Sykes (1835: 103) described and illustrated Atta providens; renamed Pheidole providens by Westwood (1839: 219).

Heer (1852: 15) described in detail and illustrated a new species from Madeira that he named Oecophthora pusilla. A translation was published later (Heer, 1856). The synonymy of P. pusilla under P. megacephala is mysterious. Bolton (1995: 328) cites Wheeler (1922: 812) but there is no evidence that Wheeler did any taxonomic research to justify his listings. For instance, Wheeler cites Emery (1919) but there P. pusilla appears as a subspecies of P. megacephala not as a synonym. A similar situation appeared in Santschi (1920, see below).

F. Smith (1855: 130) described a worker of Myrmica laevigata he had found at Battersea in London. From the description and illustration it is clearly a minor of P. megacephala. Three years later (F Smith, 1858a), however, he reported how he had re-examined his earlier Myrmica laevigata and compared it with specimens of "pallidula" sent to him by Dr. Mayr. Although brief, Smith's descriptions seem indubitably P. pallidula, notably the worker-major with darker mandibles and emarginate posterior to the head; the queen with only small tubercles on the propodeum and the dark male. The form listed in Bolton (1995: 323) as P. pusilla junior synonym janus (F Smith, 1858b: 175) from Sri Lanka clearly is not P. megacephala. The note (F Smith, 1859) only a short time later on another minor named as Myrmica suspiciosa from Aru, Indonesia appears confusing but the confusion seems to be due to the fact that Smith only had minors of laevigata and suspiciosa . The description (F Smith, 1861b) of specimens from Bachian, Celebes, collected by A. R. Wallace, is weird and obviously not of P. megacephala. The description of Myrmica jucunda F Smith (1861a) from Israel could be a minor worker of the locally dark "P. teneriffana" [note, we now feel the species present in Israel is Pheidole indica].

Mayr (1853) described a new species, Oecophthora subdentata, with major and minor workers, from specimens collected at several locations in Austria and compared the morphology to Oecophthora pusilla Heer. Two years later (Mayr, 1855), he listed further findings from Dalmatia west to Spain, Sicily and Sardinia . In his review of ants in Europe (Mayr, 1861), he recognised the use of Pheidole and noted two species were known, living under stones and in houses . Curiously, he listed only P. pusilla, as in Andalucia, Africa and Madeira, and P. megacephala in south Europe below the isotherm 21°C. This is clarified later in the text, where he gave a key to separate the two species and had P. pallidula as a synonym of P. megacephala, referring to F Smith (1858a). This key, however, appears to confuse exotic specimens of megacephala with the indigenous pallidula and the enigmatic pusilla. Mayr (1862) described P. sinaitica from the Sinai Peninsula, Egypt, comparing it to P. pusilla. Mayr (1865) changed the listing again, as he had "P. megacephala F. (pusilla Heer)" (p 93). This seems to have stemmed from Roger (1859), but that was corrected by Roger (1862), see below.

Roger (1859) gave Pheidole Westwood (Oecophthora) pusilla Heer as synonymous with laevigata Smith, and, "Pheidole pallidula Nyl (megacephala Los.)". Roger (1862) clarified the status of Myrmica megacephala by Losana (1834), noting this was a junior synonym of P. pallidula and not of P. megacephala.

Saulcy (1874) named specimens from Jericho, Palestine [Israel], collected by M. Plochard, as Pheidole jordanica, giving a cursory description and brief notes on it being the host of a Paussid.

In his guide to the ants of Europe, André (1874) gave a key (p 198) to separate P. pallidula, from southern Europe, and P. megacephala, solely from Spain. Further (p 204) he listed pallidula, Nyl., with synonyms megacephala Los. and subdentata Mayr. Under megacephala Fabr. he synonymized pusilla Heer. The main distinguishing character he gave for the megacephala minor, however, was the propodeum with strong raised teeth. This is a variable character in megacephala but they are rarely strong. He had the propodeal teeth on the major twice as long as the basal width. That also does not ring true. It is possible that he actually described specimens of P. teneriffana or P. pusilla as a distinct species.

In his contribution to the results of Fedtschenko's expedition to Turkestan, Mayr (1877) again used P. pusilla Heer and it seems from the Russian text that he regarded P. pallidula as a synonym.

Emery & Forel (1879) compliled a Catalogue of European ants and had P. pallidula as a junior synonym of P. megacephala.

André (1881) listed findings of minor and major workers of P. pallidula from Jaffa, Israel, without comment. Under the heading P. sinaitica he recorded minor and major workers from Alexandria, Egypt. He noted, however, that the major worker had the head striated to the back and a wide postpetiole, like pallidula, stating that it might be a separate new species but with only a single specimen he could not be sure.

Mayr (1886) reported his study of the ant collection in the British Museum in London and stated P. janus was identical with P. pusilla Heer. His subsequent key and text (Mayr, 1887) had P. pusilla as the senior synonym for P. laevigata Mayr, 1862, and P. megacephala of the Emery & Forel (1879) Catalogue.

Earlier we described how Forel (1874) had expressed misgivings over the proliferation of new species and decided to adopt a practice of trinomial and even quadrinomial names. Thus, Forel (1890) reporting his own travel, from 26.iii to 23.iv.1889 in Tunisia and eastern Algeria, had Pheidole megacephala i. sp. Fab., dark variety, from Sfax, and Pheidole pallidula Nyl., type form everywhere on the plain up to about 1000m, plus an entirely pale yellow variety, the major with yellow red mandibles [ possibly what we now regard as P. recticeps], from Djebel Ozmor, near Tébessa to the summit of a mountain near Souk-Ahras. Forel (1892) reported P. megacephala Fabr. rasse P. pallidula Nyl. from Dubitzna, and other places in Bulgaria.

Emery (1889) examined the Saulcy type of P. jordanica and expanded the description, confirming that it was quite distinct from a type specimen of P. sinaitica he had been sent by the Vienna Museum. He made other comparisons with "P. megacephala (pusilla)".

There is an odd record by Emery (1891b), writing about ants from Tunisia, in which he noted: 30. Pheidole capensis Mayr - "For a long time I have had a worker and soldier coming from Egypt. The French explorers had taken this form at Gafsa, El-Oued and Douz, in Tunisia. This is probably a form of the plain and the oases. Pheidole capensis is excessively close to P. megacephala, with the difference that it is a little larger and the postpetiole is widened to a point on each side" (my translation). This record was corrected by Emery in his 1915 review (p. 227) where he placed the specimens under Ph. pallidula var. recticeps.

Forel (1893) gave the first description of Pheidole teneriffana from the Canary Islands, Laguna Ténériffe, collector M. Medina, a major, plus a separate minor worker from Las Palmas, collector M. Cabrera y Diaz. Forel's description was based wholly on comparisons with P. megacephala. He did, however, point out some similarity with P. striativentris Mayr, from India, but said the head shape was closer to that of P. rhombinoda Mayr, 1879: 675 (also from India). Bingham (1903) described the head of P. rhombinoda as rectangular, slightly longer than wide.

Mayr (1893) recording ants from Tanzania, East Africa, listed P. megacephala as a distinct species as had other authors, for example Emery (1893).

Forel (1894) then changed track and named P. pallidula as common in Oran Province, Algeria. Writing on southern Palaearctic ants, including specimens from Madeira, Forel (1895, pp 4 & 5) stated P.E. Schmitz believed Heer's "Hausameise Madeira" was P. megacephala Fabr. adding that the ants were no longer found in houses and only in wild places. In the same paper, Forel listed P. megacephala and P. pallidula from Catalonia, plus P. pallidula from eastern Mediterranean countries and Biskra (Sahara). Forel also described Pheidole santschii, n. sp. (now synonymized with P. sinaitica), from La Mecque (Kairouan, Tunisia).

Mayr (1904) described Pheidole sinaitica Mayr, subsp. laticeps nov. subspec. from under a stone at Cairo, Egypt. The major had a head wider than the type sinaitica but not as wide as P. megacephala. He also recorded P. megacephala from Abba Island, White Nile, in Sudan, commenting that the species, found under a rotting tree trunk, was from the warmer tropics.

Ruzsky (1905: 694), in Russian, gave an illustrated description of what he called Pheidole pallidula Nyl., together with a new variety, arenarum, and a new subspecies koshewnikovi.

Mayr (1907) recorded P. megacephala findings from Tanzania, and referred to following the synonymy of C. G. Dalla Torre (1893). From that, however, it is impossible to know the extent of the taxonomy, if any, done by C. G. Dalla Torre and he wrote few other ant papers. Quite why Mayr, who had studied ants in such detail should suddenly accept an opinion contrary to his own, e.g. Mayr (1886, 1887), is curious. Dalla Torre's list included Formica (Myrmica) trinodis Losana (1834) as a synonym of P. megacephala, when Losana's description clearly shows it to be something other than a Pheidole, possibly a Tetramorium, and undermines the credibility of the listing. The year before, a paper by K. W. Dalla Torre (1892) stated, without any reasoning, that the name P. pusilla, "die Hausameise Madeiras", had to follow the name P. megacephala Fabricius (1793) but should not be confused with the P. megacephala of F Smith (1860) from Bachian, Indonesia. Dalla Torre gave the latter the name Pheidole megacephalotes, itself now a junior synonym of Pheidologeton diversus Jerdon (see Bolton, 1995: 325). It seems that the two sets of initials C.G. and K.W. are of the same person.

(Forel, 1907) described P. pallidula Nyl., var. tristis from workers, TL 2.6-2.8 mm, which were dark brown with the mandibles, antennae and tarsi yellow-brown, collected by Dr Santschi, at Kairouan, Tunisia.

Santschi (1908) reported collections from the Canary Islands, by Cabrera & Diaz, including P. megacephala; from Cairo, Egypt, by M Borcard, including P. teneriffana; and, from Tunisia, by himself, including P. teneriffana and P. pallidula Nyl. r. tristis Forel. He described the teneriffana queen and noted that, as it seemed to be found only near ports, it was a cosmopolitan, tramp, species. On P. pallidula tristis he described the major for the first time.

Forel (1909) recorded P. pallidula from Andalusia, Escorial, near Madrid, Spain, at 1000m. Forel (1911a) described P. schmitzi n.sp. (now synonymized with P. jordanica) from Jerusalem, Israel, collected by Pater Direktor Schmitz. From Smyrna (western Turkey), several places in Greece and Corfu, Forel (1911b) listed P. pallidula, he also gave P. teneriffana from Smyrna. Forel (1913) lastly listed P. pallidula from Corfu and P. teneriffana, major from Cairo, collectors U. & J. Sahlberg.

Karavaiev (1910) gave a brief note and illustrations of P. pallidula but it is not clear whether he was dealing with the type form of eastern varieties. Karavaiev (1911) recorded P. sinaitica Mayr var. laticeps Mayr, major and minor workers from Khartoum and Port Sudan, without details.

Krausse (1912) named P. pallidula ssp. pallidula var emeryi as a dark form of the type species from Sardinia.

Emery (1915a) listed ants from the Italian Islands with P. pallidula as the only genus member. Emery (1915b) recorded P. sinatica from Nefassit, Eritrea. With a subsequent correction (Emery, 1919), he then tried to clarify the status of P. megacephala and its relatives (Emery, 1915c). Sadly, despite giving clear notes on the sometimes vivid differences between them, he failed to shake off the Forel shackles and retained the confusion of subspecies and varieties for P. pallidula. He confirmed the status of P. sinaitica, with its synonym santschii, and, apparently without examining any specimens, "latifrons", i.e. the laticeps of Mayr (1903). He also confirmed the status of P. jordanica, with its synonym schmitzi. On the megacephala group, most of them from sub-Saharan Africa, Emery's analysis again retains the proliferation of trinomials. It is noteworthy, however, that he gave P. pusilla as a subspecies of P. megacephala and not as a synonym, writing of the distribution of the two as quite distinct. There is no evidence, however, that he saw the original Heer specimens. An interesting final paragraph places P. teneriffana Forel, P. minima Mayr (1901, from West Africa), P. squalida Santschi (1910, from the Congo Basin) and P. prelli Forel (1911, from East Africa) as among a group of numerous species belonging principally to India, the indica group. This group has the head entirely or almost entirely striate, with the frontal carinae as long as the scape. Emery (1916) published a treatise on the ants of Italy. He regarded only one species of Pheidole as indigenous to Europe, plus two or three imported species. Only P. pallidula type form and variety emeryi were found in mainland Italy, although the subspecies arenarum and its variety orientalis, which he gave as synonymous with the P. pallidula of Ruzsky, were named; with orientalis as in southern Istria, the Balkan peninsula and Asia Minor, etc.

Emery (1915j: 234) commented that Ph. pallidula with its subspecies arenarum, Ph. jordanica and Ph. sinaitica formed an interesting series of ants, which transit from the slender Ph. sinaitica with its long appendages, to the Ph. pallidula type. The source of the group seemed to run from tropical Africa through the Nile Basin to the Mediterranean region. Ph. teneriffana, of a different group, probably had the same origin with the name being inappropriate as far as the origin was concerned.

Stitz (1917), reporting findings from Teneriffe, Morocco, Algeria and Egypt, listed P. pallidula, type form (Morocco), P. pallidula v. inermis n.v. (Algeria), P. pallidula v. cicatricosa n.v. (Algeria), P. megacephala, type form (Teneriffe), and, P. megacephala v. obtusa n.v. (Egypt). He gave no descriptions or other notes but did give drawings of the three new varieties which, with now available photographs of the specimens, show them to be P. sinaitica, P. tristis and P. jordanica respectively (see below).

Bondroit (1918) catalogued the ants of France and recorded only P. pallidula, as common in the southern part of the country, noting the dark variety emeryi (?) Krausse was found at Banyuls on the Mediterranean coast.

Santschi (1919a) recorded species from Samoa. These included P. teneriffana and drew the comment that it had been found quite widely in ports or the surrounding areas, from the Canary Islands east along North Africa and down the East African coast but, without giving any details, it was known also from far up the Nile in Egypt and Khartoum. Santschi (1919b) listed P. pallidula Nyl. v. cicatricosa Stitz from Spain, queen and workers.

Santschi (1920a) listed P. teneriffana from Mombasa, Kenya, collector G Arnold. In a report on maladies of cocoa on Sao Thomé Island (Santschi, 1920b), he separately listed P. megacephala F. and P. megacephala F. v. pusilla Heer, citing both as cosmopolitan.

On ants from Asia Minor, Santschi (1921) listed P. pallidula Nyl. st. arenarum Ruzs. v. orientalis Em.

Santschi (1928) reported collections from Pacific Islands, principally Fiji, and noted strong similarities between P. oceanica Mayr and its varieties and P. teneriffana. He recorded P. megacephala as found on several islands, apparently as a recent introduction.

Karavaiev (1926) gave an analysis of the P. pallidula varieties arenarum, tristis and orientalis.

Finzi (1930) reported P. pallidula arenarum var. recticeps For. major and minor from Azrou, Morocco. He described this form in detail from Sinai, Egypt (Finzi, 1936). He later reported it also from Libya, numerous localities (Finzi, 1940). Also in his Sinai paper, he gave notes on P. jordanica, adding that the Stitz obtusa could be ascribed to jordanica. He also gave illustrated descriptions of the queens of P. teneriffana and P. sinaitica, together with a key to separate the four species.

Wheeler (1926) recorded P. pallidula from the Balearic Islands. He then reported ants from the Canary Islands (Wheeler, 1927) including P. megacephala and P. teneriffana. Perhaps misleadingly, he cited the descriptions by Heer of "Pheidole megacephala subsp. pusilla" from Madeira. Heer, of course, described Pheidole pusilla as a new species.

Alfieri (1931) gave many findings of major and minor workers of P. teneriffana from Egypt, x-xi.1925, 151 tubes of specimens determined by W.C. Crawley. Aguayo (1932) named the subspecies Pheidole teneriffana taina from 5 major workers taken at Holguin, Cuba, August 1930.

Collections in the western Mediterranean, Morocco and Tunisia by H & H Lindberg, in 1924-26, were reported by Santschi (1931). P. pallidula was the only genus member found. Santschi (1932) recorded P. jordanica as collected in Tibesti, northern Chad-southern Libya, by the Dalloni expedition in 1931. On a cruise in mid-August 1933, Santschi (1934) visited places in the eastern Mediterranean. Among the ants he collected were P. pallidula Nyl. st. arenarum Ruzs. v. orientalis Em, from Istanbul and Rhodes, and P. teneriffana from Alexandria, Egypt, in public gardens. He commented that the latter is rarely found far from ports. Santschi (1935) reported ants from the Lake Omo Region of northern Kenya, with P. sinaitica from the Turkana Desert, 4 workers, and P. megacephala at Nanoropus, on the shore of Lake Rudolph. Reporting ants from Morocco, Santschi (1936) included P. pallidula type, from several locations; v. inermis from Marais d'Esmir, major and minor workers and queen, with the eyes smaller than the type, pale, the major with the head more rounded; v. tristis from six locations; v. obscura Sants. = (v. emeryi Krausse, 1912, nom. preoc.), with the body, specially the head and gaster more or less dark, the genae and appendages still pale, differing from tristis by not having the postpetiole wider than the type form and cicatricosa by its colour and larger propodeal spines; v. cicatricosa similar to tristis but paler; "the group of forms reported to Pheidole pallidula need a revision". Santschi (1939) recorded P. pallidula Nyl. v. inermis Stitz from Asni and the Forest of Mamora, Morocco.

Bernard (1945) recorded species from the Mamora Forest in Morocco. He noted that only the type form P. pallidula was found in the black humus soil of the often shady forest. He commented the type form was widespread in the Mediterranean, often in houses. The variety tristis seemed to be limited to clay soils and, such as in the Barbary valleys. On the contrary the subspecies arenarum Ruzsky was limited to the dry plateaux and the Sahara. Bernard (1948) wrote how five species were known from the "Berbérie", that is the Mediterranean coast from Morocco eastwards to Egypt. In the southwestern Libya area of Fezzan, essentially very dry saharan with oases the sole species was Pheidole pallidula (Nyl.) sbsp. arenarum Ruzsky var. recticeps Forel. Bernard expressed surprise at not finding P. jordanica or P. sinaitica but thought this might be due to the salinity of the soil. Bernard (1950) wrote on ants from the Aïr Massif in north central Niger, ca 19ºN 8º50'E, collected in 1947 by Chopard & Villiers. Minor workers were found of P. megacephala, although Bernard admitted that minors were not easy to determine. P. sinaitica was found at Mount Tarraouaji, 900 m, 9-12-ix.1947. Pheidole pallidula (Nyl.) sbsp. arenarum Ruzsky var. recticeps Forel was represented by a single major similar to those from Fezzan. Bernard (1956) reporting the presence of P. pallidula in the Balearic Islands, added that the species is common throughout the Mediterranean, except in very shaded forests, and that it probably originated in western Asia. On the Pelagie Islands off northern Tunisia, solely P. pallidula was present at one location, Lampedusa (Bernard, 1959). On "l'île de Port-Cros", Bernard (1959b) found P. pallidula. On Corsica (Bernard, 1960) P. pallidula was conspicuous by its near absence. In his grand opus on the ants of western (Italy westwards) and northern Europe, Bernard (1967) gave only the indigenous P. pallidula and the exotic P. megacephala. He provided illustrations of all four castes of P. pallidula and noted that a dark almost black form could be found near Lyons. In his introduction, he was severely critical of the confused taxonomy stemming from Forel and even exaggerated by Santschi.

Baroni Urbani (1964a) reporting findings of P. pallidula from Izmir, Turkey, vii.1963, major and minor workers, referred to the discussion in Emery (1916) on the distribution of the subspecies and varieties but concluded he (Baroni Urbani) did not have enough evidence to contribute an answer. He next reported numerous findings from Sicily and the Appenine Mountains but without comments (Baroni Urbani, 1964b, 1964c). From the island of Malta (Baroni Urbani, 1968) he reported P. teneriffana and described, with illustrations, morphological variations between specimens from the Canary Islands, Tunisia, Eritrea and Malta. He also gave a full description of the queen and noted the species was found relatively rarely. In the same report, he attempted an analysis of P. pallidula, using head length as a determinant for specimens from around the Mediterranean. He, however, found no significant differences. In his catalogue of ants from Italy (Baroni Urbani, 1971), he listed a great number findings of P. pallidula; a single finding, at Lampedusa (an island closer to Tunisia than to Sicily) of P. sinaitica; and, a single finding of P. teneriffana from Malta.

Collingwood (1961a) raised P. pallidula arenarum to species status as P. arenarum Ruzsky, commenting on the elongated head and deep emargination of the occiput. He gave it as a species of Central Asia, with collection records from Afghanistan. Petrov & Collingwood (1962), on the ants of Yugoslavia, listed P. megacephala, P. orientalis Emery and P. pallidula Nyl.

Cagniant (1964) studied some ants in Morocco, and gave findings of P. pallidula and P. sinaitica. On the former, he added this is very common throughout the Barbary, from 0-300 m, but the subspecies of some authors do not have much significance, because the major and minor workers are very variable from the same nest. He gave similar results in other papers but those do not add to this.

Collingwood & Yarrow (1969) listed only P. pallidula from the Iberian Peninsula.

Yarrow (1967) reported P. megacephala from the Azores, noting that it seems to be extremely rare and adding "In Madeira this ant was at one time a real domestic menace", presumably misquoting from Heer, or more likely from a second-hand version of Heer's paper.

Pisarski (1967: 385) listed many findings of Pheidole pallidula arenarum Ruzsky, 1905, from northern but not southern Afghanistan and gave the overall range as the mountains of Central Asia, the Kopet Dag, Hindu Kush and Tian-Chan.

On a wider front, Wilson & Taylor (1967) listed P. megacephala among the ants of Polynesia.

Wetterer (2007) has examined and reviewed data on the occurrence of P. megacephala in Pacific Islands. Wetterer states that Wilson & Taylor "concluded that the P. teneriffana specimens listed by Santschi (1919) were actually P. megacephala". They actually wrote "Listed from Pago Pago as P. teneriffana Forel by Santschi (1919)" and gave no evidence to support their statement. The report by Santschi clearly was about what he thought to be P. teneriffana and not P. megacephala. He was well acquainted with both (Santschi, 1908) and in a later paper, not listed by the modern authors, he reported collections from Pacific Islands, principally Fiji, noting strong similarities between P. oceanica Mayr and its varieties and P. teneriffana (Santschi, 1928).

Tarbinsky (1976, in Russian) provided notes and illustrations of what he gave as P. pallidula type form and P. pallidula koshewnikovi from Kirghizia (Republic of Kyrgyzstan). The illustrations of the type form, however, confirm with the description of P. pallidula arenarum.

Aktaç (1976) listed numerous findings from Turkey but without comment.

Agosti & Collingwood (1987) listed P. pallidula as found in all the Balkan states.

Schembri & Collingwood (1981) listed P. pallidula and P. teneriffana as recorded from the Maltese Islands. From Saudi Arabia, Collingwood (1985) listed P. jordanica, P. megacephala, P. sinaitica and P. teneriffana.

Bolton (1995) cites Atanassov & Dlussky (1992) as re-synonymizing P. arenarum and P. pallidula. Yassen Dobrov of Bulgaria has helped by translating from the Bulgarian. He (pers. comm.) wrote on the "Bulgarian Fauna. 22", there is a short paragraph on the synonymy there. Here is what it says: "In the range of Pheidole pallidula one observes weak climatic changes. All geographic forms which have been described since the beginning of the century (pallidula s. str. - Western Europe, North Africa; orientalis - the Balkans, Caucasus; arenarum - the north part of Middle Asia; koshewnikovi - the eastern part of the range) gradually merge in each other and it is thus inappropriate to describe subspecies".

Collingwood (1993) reporting on species from five Greek islands, commented that P. megacephala was found only once but P. teneriffana forms strong colonies in urban areas and appears to be spreading throughout the Middle East and the Mediterranean coast. The native P. pallidula was common everywhere.

The prior knowledge of Pheidole species from the circum-Mediterranean area, together with the Spanish Atlantic Islands, generally is taken to be as summarised in Bolton (1995). The entry in his Genus-Group Taxa section, however, lists no modern analysis. His species listing shows three well-defined species, that is without a plethora of trinomial names. These are P. jordanica, P. sinaitica and P. teneriffana. The listings under P. megacephala and P. pallidula show many trinomials and even quadrinomials but no modern evidence.

The P. megacephala information in, for example, Eguchi (2001) and Wetterer (2007) is taken from Bolton. Bolton himself credited the synonymy of P. pusilla under P. megacephala to Wheeler (1922: 812). In his introduction, Wheeler (p 711) makes it clear that an undefined proportion of his synonymization was based "on one's own interpretation" and that he used the term "subspecies" as all-embracing for the "race" or "stirps" terminology of Forel, Santschi and Arnold. Much of the cataloguing was done by Dr. J. Bequaert. Wheeler's inclusion of such as the Formica (Myrmica) trinodis of Losana show that examination of the source papers, at least, was incomplete.

Eguchi (2001) had P. megacephala among the species from Borneo. Eguchi (2008) listed P. megacephala from North Vietnam, with synonymization of Myrmica agilis F Smith, 1857.

Pheidole main page - Genus Pheidole Westwood in the Mediterranean area and the Middle East - References

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