Впервые ревизована таксономия муравьёв рода Hypoponera Santschi (Hymenoptera: Formicidae: Ponerinae) Афротропики и Западной Палеарктики. Для этих регионов указано 56 видов, включая описания 34 новых для науки видов. 33 названия (главным образом, инфравидовые формы) сведены в синонимы. Приводится диагнозы всех основных и промежуточных (эргатоидных) каст (24 признака для рабочих), определительные таблицы и 129 цветных фотографий.
      РЕФЕРАТ. The taxonomy of the ponerine ants of the genus Hypoponera is revised for the Afrotropical and West Palaearctic regions. A combined key to both faunae is presented, and the West Palaearctic species are also keyed separately. Fifty-six species are recognised in total, of which 51 are Afrotropical endemics and two are restricted to the West Palaearctic; three tramp species occur in both regions. Thirty-four Afrotropical species are described as new while 33 names, including a number of infraspecific taxa and extralimital forms, are relegated to synonymy. Two previously described Afrotropical Hypoponera names are regarded as species inquirendae (lea, petiolata). Two extralimital species are mentioned with respect to the studied fauna: gibbinota, described from a casual introduction in a Palaearctic hothouse, and the Oriental assmuthi, which is provisionally raised to species to dissociate it from the Palaearctic abeillei, to which it was previously linked as an infraspecific taxon.

      Key words: Ponerinae, Hypoponera, Afrotropical, West Palaearctic, taxonomy.

      Новый вид Hypoponera importuna sp. n.

Fig. 55. Рабочий муравей Hypoponera importuna Bolton & Fisher sp. n. (Африка: Габон, Кения, ЦАР).
(по: ©Bolton& Fisher, 2011)

      ВВЕДЕНИЕ.

      Among the Afrotropical ants Hypoponera represents the epitome of morphological monotony and taxonomic confusion, and it is one of the genera that has actively initiated avoidance behaviour among taxonomists. To cite a significant instance of this, there is no mention at all of Ponera (as it was then) in the taxonomic section of Wheeler’s (1922) huge volume on ants collected in the territory that was then called the Belgian Congo, despite the fact that numerous specimens must have been collected. This omission must reflect a deliberate decision by Wheeler to ignore the genus. We can be fairly sure of this because in a survey of leaf litter ants conducted by Belshaw & Bolton (1994) in Ghana the number of Hypoponera specimens retrieved was high. The total number of individuals referable to the tribe Ponerini (sensu Bolton, 2003) in that survey was 3554, which was about 8% of the total number of ants sampled. Of these, 2410 (about 68% of Ponerini) were Hypoponera, which in terms of the total number of ants found in the survey represents a very significant 5.5%. Further, a single species of Hypoponera, H. dulcis, on its own represented 4.2% of all the ants collected in the survey, a remarkable 51% of all the Ponerini. In other words, Hypoponera specimens are very common in West Africa, as indeed they are in other tropical regions (Ward, 2000). In addition to these figures from Ghana, many samples of the most common species of the region, in particular dulcis and coeca, have been collected by hand or Winkler bag from all across the forest zone of West and Central Africa, so Wheeler’s omission of Hypoponera was probably deliberate. The main reason for his decision almost certainly lay in the remarkable uniformity of the component species of Hypoponera and the difficulty in separating them. These similarities led to their superficial and shabby taxonomic treatment throughout most of the taxonomic history of the group in this region. Original descriptions of the Afrotropical forms are uniformly poor, so poor in fact that hardly any single species can be recognised from its original description alone. The original descriptions of type-material contain much pointless repetition of insignificant features and irrelevant comparisons of features already published by earlier authors, regardless of the fact that many of those characters lacked taxonomic value. Any attempt to analyse characters for usefulness across a number of species, or to investigate other characters for novelty, is conspicuous by its absence. The poverty of endeavour in the genus has truly been monumental. Coupled with this, and complicating the taxonomic task even more, is the fact that we have never before encountered a genus in which the type-material of the various taxa has been in such degraded condition. Much is missing, and much of what remains is damaged, very poorly mounted, or both. The type-series of several taxa were found to be mixed, containing either two different Hypoponera species, or in one instance a Hypoponera plus a member of a separate genus. Original descriptions based on such series are incomprehensible, obviate any later attempts at identification and only serve to cloud the taxonomic waters to the point of total opacity. A final factor that contributed to the overall confusion was the largely unrecognised ability of some common Afrotropical species to produce intercastes between queens and workers, and the ability of one commonly collected species to produce ergatoid males. On some occasions the large-eyed but worker-like intercastes, and the very worker-like ergatoid males, were described as workers of separate species, or as infraspecific forms. The combination of all these circumstances accounts for most of the taxonomic confusion that descended upon the Afrotropical fauna of Hypoponera.

      The West Palaearctic fauna is depauperate compared with the 54 Afrotropical species. Individuals are far less frequently encountered, which is not surprising as Hypoponera is primarily a tropical genus. Only five species are known from the sub-region, three of which (eduardi, punctatissima, ragusai), all tramps, are shared with the Afrotropical and also with other regions. Of the two exceptions, abeillei belongs to a primarily Afrotropical group but has not yet been detected in that region; the other, nivariana, appears to be a genuine endemic species of the Canary Islands. Most West Palaearctic species are found in the circum-Mediterranean zone and numbers rapidly fall off with increased latitude, except for synanthropic discoveries of tramp species.

      As a genus-rank taxon Hypoponera dates back only to Taylor (1967), before which it had been treated as a subgenus of Ponera since it was first described by Santschi (1938). Since 1967 very little taxonomic work has been devoted to Hypoponera. What has been done has mostly been based on single countries, where each study has accounted for no more than two or three species. Notable exceptions include the work of Wilson & Taylor (1967), who revised and keyed the Polynesian species, and contributions to the species of Japan by Onoyama (1989), Morisita, et al. (1989) and Imai, et al. (2003), that have effectively revised the country’s fauna. Other territories covered include Armenia (Arakelian, 1994), the Balkans (Agosti & Collingwood (1987b), Bulgaria (Atanassov & Dlussky, 1992), China (Wu & Wang, 1995), Cuba (Alayo, 1974), European Russia (Arnold’i & Dlussky, 1978), Korea (Kim, et al. 1998), the Mediterranean Basin (Bernard, 1967), Portugal (Collingwood & Prince, 1998), Saudi Arabia (Collingwood & Agosti, 1996), Spain (Collingwood, 1978) and Turkmenistan (Dlussky, et al. 1990). Prior to Taylor (1967), when Hypoponera and Ponera were regarded as a single genus, useful information on some Hypoponera species and local faunae can still be gleaned from Andre (1882), Bingham (1903), Creighton (1950), Emery (1896, 1900, 1909, 1916), Forel (1900), Gallardo (1918), Mann (1921) and Wilson (1958); other, older references are summarised in Bolton (2003). Despite the fact that Ponera and Hypoponera appear morphologically very similar, and were treated as a single genus prior to 1967, recent molecular analyses (Schmidt, 2009 and Philip S. Ward (pers. com.)) indicate that the two are not sister-groups, nor are they particularly closely related within the Ponerini.

      Apart from the Polynesian study of Wilson & Taylor (1967) and the repetitive coverage of a couple of Holarctic species whose references are above, this is the first attempt at a revision of the Hypoponera fauna of an entire zoogeographical region that contains more than a small handful of species. Previous works on the Hypoponera of sub-Saharan Africa, beside the scattered literature of original descriptions of new taxa, include only the studies by Arnold (1915) for South Africa and Bernard (1953) for West Africa. Arnold’s work is now considerably out of date and Bernard’s contains many misinterpretations. Bernard (1953: 197) at least attempted to draw up a key to species, ostensibly of West Africa but also including some but by no means all extralimital species. Unfortunately this key is largely unworkable because of the confused taxonomy in use at that time.

      Since its inception Hypoponera has been included in the tribe Ponerini of subfamily Ponerinae. A diagnosis of the tribe, together with taxonomic and historical references, are provided in Bolton (2003). More recent molecularbased phylogenetic studies by Ouellette, et al. (2006), Brady, et al. (2006) and Schmidt (2009) all retrieve the genus as a monophyletic group within Ponerini, although admittedly all are based on very few species.

      In the Afrotropical region species of the genus Hypoponera are characteristically members of the fauna of leaf litter, topsoil (directly or under stones), rotten wood on or in the ground and among the roots of plants, with nests built in all of these places as well as under flakes of tree bark at soil level. They may also be found, but apparently uncommonly, within the structure of termitaries (Dejean, et al. 1997), usually after the termitarium has been abandoned by its builders. To the best of our knowledge no Afrotropical species are arboreal, for example, none was recorded in Watt, et al. (2002), but the possibility that nests might still be found in soil pockets on trees, or in rotten wood high up a tree, must be entertained. Members of Hypoponera are generally considered to be predators of small arthropods, but apart from a record by Lйvieux (1983) of a species “near coeca” preying on collembolans, definite records are not known to exist. In short, general biological information about the species is conspicuous by its absence.

      Recently, Serna & MacKay (2010) have published a morphological paper that proposes a number of new terms. One of them, cinctus, is adopted here. In terms of this paper the cinctus is the constriction between the presclerites and postsclerites of the second gastral segment (= abdominal segment IV), that has previously been called the “girdling constriction” of that segment (as defined in Bolton, 1994: 195).


      ПРОМЕРЫ И ИНДЕКСЫ.

      Measurements were taken using an optical micrometer, to the nearest 0.01 mm, on a Wild M5 microscope. All measurements are expressed in millimetres.

ПРОМЕРЫ.

      Head Length (HL). The length of the head capsule excluding the mandibles; measured in full-face view in a straight line from the mid-point of the anterior clypeal margin to the mid-point of the posterior margin. Where the posterior margin is concave the measurement is taken from the midpoint of a transverse line that spans the apices of the projecting posterior corners.

Head Width (HW). The maximum width of the head, measured in full-face view.

Head Size (HS). HL + HW, divided by 2.

Scape Length (SL). The maximum straight-line length of the scape, excluding the basal constriction or neck that occurs just distal of the condylar bulb.

Pronotal Width (PrW). The maximum width of the pronotum in dorsal view.

Weber’s Length of Mesosoma (WL). The diagonal length of the mesosoma in profile, from the angle at which the pronotum meets the cervix to the posterior basal angle of the metapleuron.

Hind Femur Length (HFL). The maximum straight-line length of the metafemur.

Petiole Height (PeH). The vertical height of the petiole measured in profile from the lowest point of the subpetiolar process to a line that intersects the highest point of the dorsal outline.

Petiole Node Length (PeNL). In profile, the maximum length of the petiole node, measured in a straight horizontal line from immediately above the dorsal base of the anterior petiolar tubercle to the posterior margin.

Petiole Node Width (PeNW). The maximum width of the petiole node in dorsal view.

Petiole Size (PeS). PeH + PeNL + PeNW, divided by 3.


ИНДЕКСЫ.

      Cephalic Index (CI). HW divided by HL, × 100.

Mandibular Index (MI). Length of closed mandible in full-face view from apex to midpoint of clypeal margin, divided by HL, × 100.

Scape Index (SI). SL divided by HW, × 100.

Petiole Node Index (PeNI). PeNW divided by PrW, × 100.

Lateral Petiole Index (LPeI). PeNL divided by PeH, × 100.

Dorsal Petiole Index (DPeI). PeNW divided by PeNL, × 100.


ФОТОГРАФИИ.

      Digital colour images were created using a JVC KY-75 digital camera and Syncroscopy Auto-Montage (v. 5.0) software.

     
Hypoponera Santschi

      Hypoponera Santschi, 1938: 79 [as subgenus of Ponera]. Type-species: Ponera abeillei Andrй, 1881: 61 and xlviii, by original designation. [Raised to genus: Taylor, 1967: 9.]

     
Diagnosis of worker.

      Members of subfamily Ponerinae, tribe Ponerini (sensu Bolton, 2003) that principally inhabit rotten wood, leaf litter and topsoil.

      1 Mandible triangular and stout, short to moderate in length (MI 27–35), without a basal groove and without a basal pit; with a basal angle between masticatory and basal margins. When mandibles are fully closed there is no space between the masticatory margins, nor between basal margins of mandibles and the clypeus.

      2 Masticatory margin of mandible with 7 to about 18 teeth and denticles in total.

      3 Palp formula 1,1 or 1,2, the maxillary palp usually minute.

      4 Clypeus simple, without extended lobes or teeth on either the median or the lateral portions and usually unarmed anteromedially. Median portion of clypeus inserted as a small narrow triangle between the extreme anterior ends of the frontal lobes.

      5 Frontal lobes small, almost confluent medially, separated only by a median longitudinal impression; the frontal lobes not raised or specialised in any way and their anterior margins well behind the anterior clypeal margin. Frontal carinae and antennal scrobes absent.

      6 Eyes absent or present; when present always small (generally of 1 to about 20 ommatidia), lateral and located well in front of the midlength of the head.

      7 Antenna with 12 segments, the apical 4–6 antennomeres gradually incrassate; only extremely rarely with a sharply differentiated club.

      8 Dorsum of mesosoma with or without a metanotal groove.

      9 Mesopleuron without a distinct transverse sulcus that conspicuously divides it into anepisternum and katepisternum.

      10 Epimeral sclerite usually absent.

      11 Metapleural gland orifice small and simple, opening posteriorly.

      12 Metasternal process small and simple.

      13 Propodeal spiracle small, circular to slightly elliptical, located far down on the side, usually close to the bulla of the metapleural gland.

      14 Propodeum unarmed; propodeal lobes vestigial to absent.

      15 Mesotibiae, metatibiae, mesobasitarsi and metabasitarsi all without spines and without enlarged prominent setae to enhance traction on their dorsal (outer) surfaces.

      16 Mesotibia and metatibia each with only a single spur; metatibial spur always pectinate.

      17 Pretarsal claws small and simple, without preapical teeth.

      18 Petiole tergite nodiform to squamiform, always unarmed.

      19 Subpetiolar process usually simple, a rounded to angulate ventral lobe; never with paired prominent teeth at the posteroventral corner of the lobe; subpetiolar process usually without an anterior fenestra or thin-spot but a fenestra present in some abeillei group members.

      20 Articulation of petiole to helcium simple.

      21 Helcium arises low down on anterior face of first gastral tergite (Abd. III) and is always simple; anterior surface of first gastral tergite forms a tall vertical surface above the helcium.

      22 Prora usually present (absent in only one Afrotropical species): an arched tranverse crest that extends across the first gastral sternite below the helcium; usually the prora extends up the anterior face of the first sternite on each side, so that the entire prora is broadly U-shaped in anterior view.

      23 Cinctus of second gastral tergite (Abd. IV) present, usually conspicuous.

      24 Stridulitrum absent or present on pretergite of abdominal segment IV.

      In addition, fine sculpture is predominantly present, uncommonly entirely absent. The sculpture is expressed as various forms of punctation, that varies in density and intensity on different parts of the body and sometimes differs between species. Coarse, dense sculpture, such as rugae, costulae, or strong striation, appears never to be developed.

     
Diagnosis of worker-queen intercaste. (= ergatoid gyne)

      Characters as worker but always with much larger eyes than conspecific worker (intercastes with 7–30 ommatidia in Afrotropical species in which workers have 0–7 ommatidia), but without ocelli; often with a shorter petiole node in profile and a somewhat enlarged gaster; sometimes with a gyne-like transverse sulcus on mesopleuron. One intercaste of punctatissima was dissected: a spermatheca was present and the ovaries were enlarged, though much smaller than in the gyne (see also Yamauchi, et al. (1996)).

      Intercastes have been confirmed in the following species that occur in the Afrotropical region. H. abeillei group: austra, importuna, lepida, producta. H. punctatissima group: eduardi, ragusai, punctatissima. Possible intercastes are also suspected in ignavia and occidentalis (see discussions of those species). It seems reasonable to assume that intercastes occur in many more species but are not currently represented in collections. It is equally obvious that some species, such as the extremely common dulcis, do not produce them. In some previous publications, intercastes have been termed major workers or ergatoid females (e.g. Forel (1894), Le Masne (1956), Brown (1958)). Observations on the reproductive biology of intercastes can be found in Le Masne (1956), Yamauchi, et al. (1996) and Yamauchi, et al. (2001).

     
Diagnosis of queen (gyne).

      Characters as listed for workers except for worker characters 6, 8, 9; with the following differences. 1 Eyes always present and large, usually obviously with > 50 ommatidia. Eyes are located in front of midlength of head and all species examined have small setae that project between the ommatidia.

      2 Ocelli present.

      3 Mesopleuron with a well developed transverse suture that divides it into anepisternum and katepisternum.

      4 Mesosoma with a full complement of flight sclerites (alate when virgin).

      5 Jugal lobe absent from hindwing.

      6 Venation almost complete (only cross-vein 1r-rs absent); with 8 or 9 closed cells including the pterostigma (8 cells in those species with Cu2 incomplete or absent); Rs.f5 meets R1.f3 on the anterior margin (i.e. marginal cell always closed); cross-veins 2r-rs, 2rs-m, 1m-cu and cu-a all present; cu-a arises from M+Cu (i.e. proximal of point where M+Cu divides into M and Cu; 2rs-m distal of 2r-rs; a free abscissa of M (M.f2) present between Rs+M and 1m-cu; an angle or bend sometimes present in Rs.f2&3; a fenestra present in cu-a and fenestrae sometimes visible in Rs.f2&3 and 2rs-m, but not in minute species.

      7 Petiole node in profile is usually more slender, and often more tapered dorsally, than in the conspecific worker.

      Generally slightly larger than conspecific worker; gaster sometimes distinctly larger.

     
Diagnosis of alate male.

      1 Mandible lobiform to unidentate (apical tooth only present), not meeting at full closure. Basal cavity of mandible extends to its front face and is visible in full-face view. 2 Eyes large and conspicuous, with minute setae projecting from between the ommatidia. Three distinct ocelli present.

      3 Antenna with 13 segments, filiform.

      4 Scape short, shorter than second funicular segment.

      5 Second funicular segment longer than the first and also longer than the third.

      6 Palp formula 1,1; 1,2; 1,3; 1,4; very rarely maxillary palp of 2 segments.

      7 Mesonotum in profile not overhanging pronotum.

      8 Mesoscutellum convex in profile.

      9 Notauli absent; parapsidal grooves present but sometimes very faint.

      10 Epimeral sclerite absent.

      11 Mesotibia and metatibia each with a single spur; metatibial spur always pectinate.

      12 Pretarsal claws simple.

      13 Venation as alate gyne.

      14 Jugal lobe absent from hindwing.

      15 Petiole unspecialised ventrally; helcium very low on anterior face of first gastral segment.

      16 Prora present, small.

      17 Cinctus of second gastral tergite (Abd. IV) present.

      18 Tergite of abdominal segment VIII (pygidium) without a median downcurved spine.

      19 Pygostyles (= cerci) present.

     
Diagnosis of ergatoid male.

      1 Body form extremely worker-like but male genitalia present.

      2 Mandibles reduced (similar to alate males) or worker-like.

      3 Antenna with 12 or 13 segments, worker-like or specialised but without the basal segment arrangement of alate males (male characters 4 and 5, above). Scape distinctly shorter than in conspecific worker.

      4 Eyes present or absent.

     
Comments on ergatoid male characters.

      The production of ergatoid males appears to be restricted to species of the punctatissima group (see below). The characters above are based on eduardi, punctatissima and ragusai, but ergatoid males are also produced by extralimital members of the group, such as opacior (Forel), opaciceps (Mayr) and nubatama Terayama & Hashimoto.

      At first glance ergatoid males can easily be mistaken for workers, especially as the characteristic male genitalia are retractile and may be almost entirely concealed within the body. The ergatoid male of eduardi is monomorphic, with reduced mandibles, 13-segmented antennae and small eyes present. The ergatoid male of punctatissima is dimorphic. Both morphs have worker-like mandibles and head shape and 12-segmented antennae, but the larger morph is brown and has small eyes, while the smaller morph is yellow and lacks eyes. Ergatoid males of ragusai have not yet been found in the Afrotropical region, but specimens from the U.S.A. have a worker-like head and mandibles, as punctatissima, but possess 13-segmented antennae.

     
Separation of Hypoponera from convergent genera.

      1 Ponera
No native species of Ponera has ever been found in sub-Saharan Africa and it is probably now safe to say that the genus is not represented by endemic species in this region. However, some Ponera species have tramping ability and one small, as yet unidentified, species has been intercepted in East Africa (BMNH), which is the only confirmed record from the entire Afrotropical region. There are also three species found on islands in the Southwest Indian Ocean, all of which are assumed to be introduced. The West Palaearctic contains two indigenous species of Ponera, coarctata (Latreille) and testacea Emery (Csцsz, 2003; Csцsz & Seifert, 2003). At first glance Ponera workers may be difficult to separate from Hypoponera, but Ponera workers (and queens) have a 2-segmented maxillary palp and a specialised subpetiolar process on which an anterior fenestra is present and the posteroventral angles of which project into a pair of sharp teeth (Taylor, 1967). Sometimes the petiolar sternite itself is bifurcated posteriorly. In Hypoponera the maxillary palp is 1-segmented at most, there is usually no fenestra (but see worker comment 19, above) and the posteroventral angle of the subpetiolar process, no matter how acute, never terminates in a pair of teeth. The males of Ponera always have the tergite of abdominal segment VIII produced into an elongate, downcurved median spine, whereas in male Hypoponera abdominal tergite VIII is a simple triangular sclerite.

      2 Cryptopone
Only a single Afrotropical species seems correctly placed in Cryptopone, C. hartwigi Arnold (1948) from South Africa. Brown (1963: 6) regarded hartwigi as a member of Ponera, saying that true Cryptopone had not been taken in Africa south of the Sahara. This appears to have led Taylor (1967: 12) to transfer hartwigi to Hypoponera as it did not conform to his diagnosis of Ponera, but it is unlikely that either author had seen an actual specimen at that time. Genuine hartwigi workers have a basal mandibular pit, a characteristic of Cryptopone that is absent from all Hypoponera and Ponera. A second African species, originally described as Cryptopone angustata, properly belongs in Hypoponera and is discussed below. There is a single West Palaearctic species of Cryptopone, ochracea (Mayr), that has a distinct basal mandibular pit.

      3 Pachycondyla
In the Afrotropical region there is a discrete group of at least three small, yellow Pachycondyla species that bear a superficial close resemblance to Hypoponera. This group currently contains Pachycondyla gulera Ozdikmen, 2010: 992 (a replacement name for Ponera ambigua Weber, 1942: 46, which was a junior secondary homonym of Pachycondyla ambigua Andre, 1890: 316), P. weberi (Bernard, 1953: 194), and a third, apparently undescribed species from Cameroun and Gabon (CASC). P. gulera and the undescribed species will run to Hypoponera in standard keys to genera because they have lost the anterior mesotibial and metatibial spurs. The absence of these features gives them, convergently, the same spur formula as Hypoponera; the anterior spurs on both tibiae are retained in weberi. Taken together the three species are quickly distinguished from Hypoponera by their elongate triangular mandibles (MI ca 45–50) and palp formula of 3,3.

      In West and Central Africa there are also two apparently undescribed species of eyeless but darkly coloured Pachycondyla (BMNH, CASC) in which the subpetiolar process terminates posteroventrally in a Ponera-like pair of teeth. These species retain mesotibial and metatibial anterior spurs and have an epimeral sclerite, and by in situ count appear to have a palp formula higher than 2,2. At first glance they are reminiscent of the very common and widespread P. brunoi Forel (see Brown, 1963), but are considerably different in detail.

     
СПИСОК ВИДОВ.

      abeillei group

    Section 1
abeillei (Andre, 1881)
angustata (Santschi, 1914)
austra Bolton & Fisher sp. n.
blanda Bolton & Fisher sp. n.
bulawayensis (Forel, 1913) stat. n.
camerunensis (Santschi, 1914)
coeca (Santschi, 1914)
    = myrmicariae (Wasmann, 1918) syn. n.
defessa Bolton & Fisher sp. n.
exigua Bolton & Fisher sp. n.
fatiga Bolton & Fisher sp. n.
hawkesi Bolton & Fisher sp. n.
inaudax (Santschi, 1919)
juxta Bolton & Fisher sp. n.
lassa Bolton & Fisher sp. n.
lepida Bolton & Fisher sp. n.
meridia Bolton & Fisher sp. n.
natalensis (Santschi, 1914)
obtunsa Bolton & Fisher sp. n.
orba (Emery, 1915)
perparva Bolton & Fisher sp. n.
pulchra Bolton & Fisher sp. n.
regis Bolton & Fisher sp. n.
rigida Bolton & Fisher sp. n.
traegaordhi (Santschi, 1914)

    Section 2
aprora Bolton & Fisher sp. n.
comis Bolton & Fisher sp. n.
dema Bolton & Fisher sp. n.
dis Bolton & Fisher sp. n.
faex Bolton & Fisher sp. n.
hebes Bolton & Fisher sp. n.
importuna Bolton & Fisher sp. n.
jeanneli (Santschi, 1935)
    = jeanneli st. abyssinica (Santschi, 1938) syn. n.
    = coarctata subsp. imatongica (Weber, 1942) syn. n.
jocosa Bolton & Fisher sp. n.
mixta Bolton & Fisher sp. n.
molesta Bolton & Fisher sp. n.
occidentalis (Bernard, 1953)
    = intermedia (Bernard, 1953) syn. n.
odiosa Bolton & Fisher sp. n.
producta Bolton & Fisher sp. n.
quaestio Bolton & Fisher sp. n.
segnis Bolton & Fisher sp. n.
surda Bolton & Fisher sp. n.
tecta Bolton & Fisher sp. n.
tristis Bolton & Fisher sp. n.
ursa (Santschi, 1924)
venusta Bolton & Fisher sp. n.

      dulcis group
dulcis (Forel, 1907)
    = dulcis var. uncta (Santschi, 1914) syn. n.
    = rothkirchi (Wasmann, 1918) syn. n.
    = lotti (Weber, 1947) syn. n.
    = muscicola (Weber, 1947) syn. n.
    = lamottei (Bernard, 1953) syn. n.
    = mandibularis (Bernard, 1953) syn. n.
    = villiersi (Bernard, 1953) syn. n.

      punctatissima group
eduardi (Forel, 1894)
    = antipodum (Forel, 1895)
    = dideroti (Forel, 1913) syn. n.
    = opaciceps r. chilensis (Forel, 1914) syn.n.
ragusai (Emery, 1894)
    = gleadowi (Forel, 1895) syn. n.
    = gleadowii r. decipiens (Forel, 1899) syn. n.
    = gleadowi subsp. aethiopica (Forel, 1907) (junior homonym) syn. n.
    = ragusai v. santschii (Emery, 1909) syn. n.
    = japonica r. formosa (Forel, 1913) syn. n.
    = lesnei (Bondroit, 1916)
    = parva (Bondroit, 1918) (junior homonym)
    = massiliensis (Bondroit, 1920) (first replacement name)
    = gyptis (Santschi, 1921) (second replacement name)
    = oblongiceps (Smith, M.R., 1939) syn. n.
nivariana (Santschi, 1908)
punctatissima (Roger, 1859)
    = androgyna (Roger, 1859)
    = tarda (Charsley, 1877)
    = punctatissima r. jugata (Forel, 1892)
    = ergatandria (Forel, 1893)
    = kalakauae (Forel, 1899)
    = punctatissima subsp. schauinslandi (Emery, 1899) syn. rev.
    = ergatandria subsp. bondroiti (Forel, 1911) syn. n.
    = dulcis var. aemula (Santschi, 1911) syn. n.
    = ergatandria st. cognata (Santschi, 1912) syn. n.
    = ragusai var. sordida (Santschi, 1914) syn. n.
    = incisa (Santschi, 1914) syn. n.
    = sulcatinasis r. durbanensis (Forel, 1914) syn. n.
    = ergatandria r. petri (Forel, 1916) syn. n.
    = brevis (Santschi, 1921) syn. n.
    = punctatissima v. exacta (Santschi, 1923)
    = mina (Wheeler, 1927)
    = argonautorum (Arnol’di, 1932) syn. n.
    = mumfordi (Wheeler, 1933)
    = mesoepinotalis (Weber, 1942) syn. n.
    = breviceps (Bernard, 1953) syn. n.
    = ursoidea (Bernard, 1953) syn. n.
    = sulcitana (Stefani, 1970) syn. n.
sinuosa (Bernard, 1953)


      boerorum group
boerorum (Forel, 1901)
ignavia Bolton & Fisher sp. n.
spei (Forel, 1910)
    = spei r. devota var. sancta (Forel, 1914) unavailable name
    = spei var. fidelis (Santschi, 1926) syn. n.
sulcatinasis (Santschi, 1914)
    = spei st. devota (Santschi, 1914) syn. n.
transvaalensis (Arnold, 1947)

      Species inquirendae
lea (Santschi, 1937)
petiolata (Bernard, 1953)

      Extralimital species
gibbinota (Forel, 1912)