Pheidole fervens main page; Pheidole providens main page;

Revisionary notes on "Pheidole teneriffana", Pheidole fervens and Pheidole providens (j. syn. indica)

Note - we have discovered that Pheidole indica, as referred to by many authors cited below, almost certainly is a junior synonym of Pheidole providens Sykes (as linked above).

"Pheidole teneriffana Forel, 1893"

The long held view is that P. teneriffana is the sole Pheidole from west of the Indian subcontinent to the islands of the eastern Atlantic with major workers that have longitudinal striations over the whole length of the head.

The listing in Bolton (1995: 331) is - Pheidole teneriffana Forel, 1893: 465, major and minor workers; Santschi, 1908: 521, queen. Holotype major worker. CANARY ISLANDS, Teneriffe, collector M. Medina. Minor worker thought to be of the species, Las Palmas, collector M. Cabrera y Diaz. Pheidole teneriffana taina Aguayo, 1932: 219, 5 major workers. CUBA.

Aguayo (1932) separated the taina major as having the sides of the head more convex and narrow posteriorly, the occipital impression deeper and wider, the postpetiole larger and the striation of the head feebler.

Alfieri (1931) gave many findings of major and minor workers from Egypt, x-xi.1925, 151 tubes of specimens determined by W.C. Crawley.

André (1881) listed a single major worker from Alexandria, Egypt, collector M. Abeille de Perrin. This André listed as P. sinaitica Mayr but remarked how the specimen had the postpetiole widened as in P. pallidula but the head was striated right up to the posterior. As it had relatively large eyes and "non transversaux" (not longer than wide) segments of the antennae, he attached it to P. sinaitica whilst feeling it could constitute a new species but he could not decide on a single specimen. He separated three species of Pheidole, sinaitica, pallidula and megacephala, as then known in his great catalogue (1881-6). The last, however, was separated from pallidula solely on the basis of having large propodeal teeth. From his note it seems sound to conclude André (1881) reported a specimen of P. teneriffana.

Forel's (1893) description of P. teneriffana 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 of the minor was closer to that of P. rhombinoda Mayr (also from India).

Emery (1915c) had an interesting final paragraph that places P. teneriffana Forel, Pheidole minima Mayr (1901, from West Africa), Pheidole squalida Santschi (1910, from the Congo Basin) and P. prelli Forel (1911b, from East Africa) as among a group of numerous species belonging principally to India, the indica group. Emery stated this group has the head entirely or almost entirely striate, with the frontal carinae as long as the scape. The description of P. squalida by Santschi (1910) has the striations fading out before the posterior third of the head and P. minima is similar.

Santschi (1919a) recorded ant species from Samoa which included P. teneriffana. He commented 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.

Snelling (1992) reported the discovery of P. teneriffana from California and Wilson (2003) included it as among the tramp species found in the New World,.

Espadaler & Bernal (2003) commented that, although originally described from Teneriffe, the species is certainly not very abundant in the Canaries.

Wetterer et al. (2007) reported it as newly recorded from Ascension Island and St. Helena in the South Atlantic. interestingly, they cite Mellish (1875) on a "larger, quite black" species as found in addition to P. megacephala.

Examining our collections of fresh specimens from locations in Egypt, Israel, Canary Islands and Iran, indicated they could be separated into two sets. This led to a wider search of the literature and the internet resources. First we found, the images of P. fervens shown on the Japanese Ant Image Database website, PCD0571-80 (http://ant.edb.miyakyo-u.ac.jp/E/Taxo/F40601.html) are an exact match for the P. teneriffana specimens from Hawamdyia, Egypt, not least including the configuration of the hypostomal processes of the major. The information then revealed an assemblage of Pheidole species from southern and eastern Asia that have the major worker head bearing striations that stretch back for the whole length, or almost so, and long frontal carinae. A further apparently consistent characteristic of this assemblage is a strong transverse carina or welt on the mesonotum. There is a second assemblage with sculpturation on the gaster, including P. striativentris Mayr, 1879: 678, major; and, P. fossulata Forel, 1902: 181, major, 196, minor [raised to species Bingham, 1903: 257, and Menozzi, 1939: 298 (missed by Bolton, 1995: 321)], but our specimens all have smooth shiny gasters. The member of the assemblage described first was Pheidole fervens F Smith, 1858b: 176, but as with most of F Smith's works, the description was poor and lacking in detail. The type location was Singapore and the two main keys to Pheidole from India, Forel (1902) and Bingham (1903), do not include P. fervens. The check list of the ants (Hymenoptera: Formicidae) of Asia by Chapman & Capco (1951) had only the F Smith record.

Mayr (1879: 679) described Pheidole indica - major, TL 4.5-5.5 mm; castaneo-fuscus (dark chestnut brown), alitrunk part ferruginous, antennae and legs fusco-testaceous (dusky yellow); head strongly longitudinally striate. We found a major labelled P. indica shown on http://ant.edb.miyakyo-u.ac.jp/J/Taxo/F40603.html, where it is separated from P. fervens, by the dark-coloured body, larger eyes and upwardly directed propodeal spines in the major and minor workers. Unfortunately, no size scale is given and the colour appears falsely orange. The P. teneriffana specimens we have from Qaliobyia, Abuzabal, Egypt, however, are a match for those specimens.

Following the key to Pheidole of India given by Forel (1902) from the separation of species with a mesonotal welt and the whole of the head sculptured led to:
P. striativentris (outside this group). Then:
Major with a very distinct antennal scrobe, with very fine sculpture; the scrobe recurves posteriorly and runs back down towards the eye; the frontal striation apparently not continuing right to the posterior; reddish but gaster femora and scapes brown, tarsi and funiculi yellowish; TL 3.5-4.0 mm, scapes short, P. magrettii Emery, 1887.
Without a strong scrobe.
Tibiae and scapes with pubescence and no elongate hairs. P. jucunda Forel, 1885.
Tibiae and scapes with erect hairs.
Small species TL max 3.5 mm, P. javana Mayr, 1867.
Larger species.
Head strongly cordate with highly convex sides, clypeus protruding forward, medially impressed and bidentate, P. peguensis Emery, 1895.
TL 4.8 mm, clypeus medially impressed but not protruding and not bidentate; propodeal spines very long, robust and pointed, P. roberti n.sp..
TL 3.4-3.6 mm; rugae behind the eyes less regular, occiput rugo-reticulate, gaster with base punctulate, P. feae Emery, 1895
Tibiae and scapes with only oblique hairs, TL 4.2-4.5 mm, pitch brown, alitrunk part reddish, legs and antennae yellow-brown, P. indica Mayr.

In the description of Pheidole magrettii Emery, 1887: 462, major and minor worker. JAVA, Emery made comparison with P. parva Mayr, 1865: 98, but the major of magrettii quite clearly is different without the extended striations on the head and without a mesonotal welt. On that, the photographs shown on http://ant.edb.miyakyo-u.ac.jp/J/P/PCD3017/37.html are not P. parva.

With the omission of P. peguensis, Bingham's (1903) key to Pheidole from India, separated on the basis of the presence of lateral tubercles on the pronotum. If absent, the key led to the same four species with P. fervens as Forel. These fell into two pairs,
P. feae (text p 260) and P. roberti (text p 259) with the medial portion of the clypeus smooth and shining;
P. jucunda (text p 256) and, P. javana (text p 262) with the medial portion of the clypeus opaque and longitudinally striate.
In his text, Bingham gave P. javana as with head nearly square and TL 3.5-4.0 mm, scape of antenna falling short of the occiput by more than half its length, castaneous red, appendages more yellowish, gaster yellow-brown. P. jucunda was similar but larger, major TL 4-4.5 mm, with longer scapes, falling short of the occiput by only one-fifth of its own length, brownish cinnamon-red, the gaster dark brown, with the other characters matching P. teneriffana. P. roberti major, TL 4.5-5 mm, was bright light red, antenna, legs and gaster brownish, head not deeply striate with the striae breaking into reticulations posteriorly. The P. feae page is missing in the HNS copy but the key separated it as having only weak scrobes, with longitudinal internal striations, from P. roberti with strong scrobes but with fine internal sculpture. When tubercles are present on the pronotum, Bingham's key led to P. plagiaria F Smith, 1860: 112, from INDONESIA, which has longitudinally striate scrobes; P. binghami Forel, 1902: 184, from INDIA, with the postpedicel with acute lateral cones; and, P. indica, with the postpedicel laterally rounded.

Wilson & Taylor (1967) gave an illustrated report of P. fervens and P. oceanica from the Pacific Islands. Without giving precise descriptions or measurements, they separated the species despite noting P. oceanica shows great variability in the size, scape length, form and intensity of sculpture. In their key the defining characters were given as: P. oceanica major with the area between the antennal insertion and the eye longitudinally rugose, propodeal spine of minor distinctly longer than the greatest width of the propodeal spiracle. P. fervens major with the area between the antennal insertion and the eye rugoreticulate, propodeal spine of minor only about as long as the greatest width of the propodeal spiracle. They gave a considerable number synonymies under P. fervens but provided no supporting evidence and nothing to show they examined any type specimens. The same applies to the synonymies under P. oceanica.

On p 46, Wilson & Taylor stated (reiterated by Wetterer, 2007), that the Santschi (1919) record of P. teneriffana from Pago Pago, American Samoa, was P. megacephala. This we find untenable as Santschi, perhaps the greatest ant taxonomist of his era, was familiar with both species. It also overlooks the Santschi (1928) description of Pheidole oceanica subsp. nigriscapa, var. tahitiana (see below). The puzzle really is to why Wilson & Taylor did not look into P. teneriffana as a possible tramp. The original description of P. oceanica by Mayr (1866) has the major "lateribus pone oculos reticulo-striatum", which translates as laterally by the eyes reticulo-striate - exactly the opposite of the Wilson & Taylor statement. The latter claimed P. oceanica is larger than P. fervens but Santschi's upoluana was TL 4.0 mm (Wilson & Taylor placed this with oceanica) and nigriscapa was TL 4.3 mm (Wilson & Taylor placed it with fervens). Mayr's type oceanica was TL 4.4 mm; the type size for fervens is imprecise but F Smith's "2 lines" = 4.2 mm, and most varietal descriptions give TL ca 3.8 mm. Smith described the major as having an oblong head, narrowest in front and deeply marginate behind, the lateral angles rounded and with a central channel extending to the base of the clypeus.

Ogata (1982) on Pheidole from Japan keyed out three species with the major worker having the occipital area sculptured, antennal scrobes long, distinct, beyond the level of the eyes; in the minor worker the apical segment of the antenna was longer, usually 0.28 mm or more in length. These were P. noda F Smith, 1874: 407, P. indica and P. fervens. P. noda was separated by the large postpetiole, which is distinctly higher than the petiole. P. indica was separated by the major with robust propodeal spines, directed upward and outward, a large eye, about 0.24 mm in diameter; the minor with the apical segment of the antenna 0.26-0.30 mm and antennal segment X 0.15-0.18 mm long. P. fervens was separated by the major with slender pointed propodeal spines, directed upward and curved backwards, the eye smaller, about 0.18 mm in diameter; the minor with the apical segment of the antenna 0.31-0.32 mm and antennal segment X 0.19-0.20 mm long. Ogata's drawings of the male aedagus (Figs. 6-11) showed a close similarity between P. indica and P. noda with P. fervens quite distinct from both.

Zhou & Zheng (1999) gave a key, apparently following that of Ogata (1982) to separate Pheidole from Guangxi, China. Those with rugose heads were P. noda, P. feae, P. fervens, P. indica and P. longiscapa sp. nov. Apart from the key, they gave information solely on the last and that is quite distinctive in many ways. Not least is the dense yellow pilosity on the major and the long scapes, SI 164-167, of the minor.

Eguchi (2001) on Bornean species of Pheidole uses the hypostomal armature as a useful character for separating major workers. Note the reference list reveals that he did not read many of the earlier publications, as he lists them as "indirectly cited from Bolton (1995)". Under his description of P. fervens, Eguchi shows he had examined only a major and 2 minor syntypes of the type form, plus seven syntypes of the variety desucta.

Eguchi (2004) went a stage further, however, elucidating and separating several of the Wilson & Taylor synonymies, particularly from P. fervens. His descriptions and illustrations leave two species, P. fervens and P. indica as relevant here. From Eguchi's key, these share the following features: major - postpetiole much shorter than the petiole, propodeal dorsum with standing hairs, lateral face of occipital lobe strongly rugo-reticulate; minor - dorsum of promesonotal dome without lateral tubercles, the dome also almost smooth and shiny. He separated P. fervens by the major with the eye no longer than the 10th segment of the antenna, the propodeal spines relatively narrow based and slightly curved apically; the minor with the eye no longer than the 10th segment of the antenna. P. indica by the major with the eye much longer than the 10th segment of the antenna, the propodeal spines relatively broad based and not curved apically; the minor with the eye as long as or longer than the 10th segment of the antenna.

From all the foregoing, we feel our two sets of "P. teneriffana" specimens can be matched to P. fervens and P. providens. We note that Eguchi (2004) gave the "median hypostomal processes ill developed or almost absent". Unfortunately, in none of his works does Eguchi appear to have looked beyond the remit of Asian and a few Pacific Island specimens.

Notes. The CASENT images have a specimen from Japan (CASENT0008638) that matches Bingham's description of P. javana, in colour and the short scapes. Forel (1902) had noted the scapes only reached the 3/4 point from their insertion to the occiput. The head shape matches the F Smith short description of P. fervens, viz. narrowest in front, deeply emarginate behind, the lateral angles rounded and a central channel extending to the base of the clypeus. The California specimen of a major (CASENT000579), however has long scapes, as with Bingham's jucunda, and the Eguchi drawing. If the foregoing is correct, and it is hard to believe otherwise, the Wilson & Taylor (1967) determinations and illustrations have to be transposed with their "P. oceanica" becoming P. fervens and their "P. fervens", perhaps, becoming P. jucunda. To reiterate, Bingham (1903) had P. jucunda and P. javana with the medial portion of the clypeus opaque and longitudinally striate (shown in the Wilson & Taylor drawings). P. javana had the head nearly square and TL 3.5-4.0 mm, scape of antenna falling short of the occiput by more than half its length, castaneous red, appendages more yellowish, gaster yellow-brown. P. jucunda was similar but larger, major TL 4-4.5 mm, with longer scapes, falling short of the occiput by only one-fifth of its own length, brownish cinnamon-red, the gaster dark brown, with the other characters matching P. teneriffana.

Pheidole teneriffana Forel

Type location Canary Islands (Forel, 1893d: 465, soldier & worker; Santschi: 1908: 521, queen) from Teneriffe, collector M. Medina; subspecies taina (Aguayo, 1932: 219, soldier) from Cuba ; soldier and worker only described (Bolton, 1995).

Forel's (1893d) description is at . Santschi's (1908) illustrated description of the queen (the illustration is labelled male but that was wrong) is at . The specimens were sent to Santschi from Cairo, Egypt, collected by Borcard, and he reports his own finding at Sousse, Tunisia, in the public garden close to the port, suggesting to him that it is a cosmopolitan species. Nests occur in the ground and under rocks. In a personal visit to Alexandria in 1933, Santschi (1934d: 276) collected workers in the public garden, noting that this cosmopolitan species rarely seems to be found far from the ports. Baroni Urbani (1968b: 438) described specimens from Malta and provided a number of comparative illustrations. Among those is a specimen from Eritrea, Massua Belli, i.e. from those reported by Emery (1901e), collected by V Belli. Also other from Tunisia. Baroni Urbani also gave a description and illustration of the queen, essentially matching that we show below.

Santschi (1918e: 63; repeated in 1919a) gave a brief note on the remarkable distribution of teneriffana is at . Suggesting the point of origin may have been the regions neighbouring the "Haut Nile", he listed its successive discovery in the canary islands, at Cairo, at Souste (Tunisia), at Smyrna, at Khartoum, in Eritrea, at Mombasa and lastly in the Samoan Islands of the Pacific. His suggested place of origin, the only non-seaport area ("having descended the river to expand") seems unlikely from current knowledge.

Aguayo's (1932) description of taina is at . He commented on the wide distribution and listed Tunis, Smyrna, Khartoum, Mombasa, Samoa and China as among the known findings. Finzi's (1936) description and illustration of the queen is at . It is tempting to suggest that Myrmica jucunda F Smith (1861a: 34) is a minor worker of teneriffana, his description is at .

Pheidole teneriffana descriptions (translated from Forel, 1893d)
MAJOR - TL 3.8 mm. Head as wide as it is long; with strong parallel striations for the entire length. The head more or less parallel sided and not narrowed towards the occiput. Central area of the clypeus smooth and shiny. Frontal carinae extending back to the extremitiy of the scapes; the latter are lodged in a distinct gutter, the scrobe, which has weak reticulate sculpture. Pronotum subtuberculate; mesonotum with a strong transverse impression [in addition to the metanotal groove]. The propodeum with pyramidal teeth or spines. Petiole node quite thick, obtuse, entire and rounded at the summit, much thicker than megacephala . Postpetiole twice as wide as it is long. Variable brown, the anterior gaster brownish-yellow the rest dark brown. Laguna, Teneriffe, collector M. Medina.
MINOR - TL 2.8 mm. Relative to megacephala the head is much bigger, the postpetiole is wider than long and the mesonotum has a distinct transverse impression. The head is smooth and shiny, with several anterior striations. Gaster dark brown, with the anterior third yellowish. Pilosity slightly more erect and longer than megacephala.Las Palmas, Canary Islands, collectors M. Cabrera & Diaz - probably but not certainly the same species as the major.

Wilson (2003) noted Pheidole teneriffana as one of only two Old World Pheidole known from the Americas, several widely scattered locations. He thought it may be of Mediterranean origin. Santschi (1920d: 378) reported it from Mombasa, Kenya, collected by G Arnold, and noted it as (my translation) - "an African species which has become cosmopolitan".

Wilson, whose illustration is shown right, had the following inadequate description (based on specimens from Cuba) -
Major - TL ?, HL 1.34, HW 1.34, CI 100, SL 0.82, SI 61, PW 0.64, PW/HW 48; head with overall yellowish-brown with head, mandibles and gaster slightly darker
Minor - occiput slightly narrowed, no nuchal collar; colour as major.
These appear top match those we regard as P. fervens.

NOTE - most minor specimens shown on our Minor workers page and http://www.antweb.org/specimen.do?name=casent0005778&shot=p1&project= have oval heads, with a very distinct but narrow nuchal collar; they also have larger eyes than shown by Wilson.